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Volume 42 1988 Number 1

ISSN 0024-0966

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

16 March 1988

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

JERRY A. POWELL, President JEAN-FRANCOIS LANDRY, Vice
DoucLas C. FERGUSON, Immediate Past President

President ATUHIRO SIBATANI, Vice
JACQUELINE Y. MILLER, Vice President President
RICHARD A. ARNOLD, Secretary JAMES P. TUTTLE, Treasurer

Members at large:

MIRNA M. CASAGRANDE M. DEANE BOWERS JULIAN P. DONAHUE
EDWARD C. KNUDSON RICHARD L. BROWN JOHN E. RAWLINS
FREDERICK W. STEHR PAUL A. OPLER Jo BREWER

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Cover illustration: Computer-drawn figure of Dakota skipper, Hesperia dacotae (Skin. ),
nectaring on narrow-leaved purple coneflower, Echinacea angustifolia DC. Drawing
originally executed freehand in the Macintosh application Fullpaint and embellished with
FatBits, exported to SuperPaint and further embellished with LaserBits, then produced
on a LaserWriter. Cover version is 50% width of LaserWriter version. Submitted by
Ronald A. Royer, Division of Science, Minot State University, Minot, North Dakota 58701.

JouRNAL OF
Tue LeErPiIpoprTreERIstTs’ SOCIETY

Volume 42 1988 Number 1

Journal of the Lepidopterists’ Society
42(1), 1988, 1-13

SPEYERIA ATLANTIS IN COLORADO: REARING STUDIES
CONCERNING THE RELATION BETWEEN SILVERED
AND UNSILVERED FORMS

JAMES A. SCOTT
60 Estes Street, Lakewood, Colorado 80226

ABSTRACT. Speyeria atlantis in the SE Rocky Mts. occurs in two forms, silvered
and unsilvered, that could be mere forms or separate species. Nine wild females laid eggs
and produced adults in the laboratory. Offspring resembled mothers in most cases, except
for two mothers about half silvered and one mother about one-third silvered that produced
nearly unsilvered offspring. The two forms have the same courtship, without obvious
courtship barriers between them, and male pheromones smell the same. Silvered and
unsilvered adults have differently colored larvae. The two forms can differ in habitat,
and adults actively select different habitats. The two are probably forms of the same
species.

Additional key words: Nymphalidae, habitat selection, polymorphism, courtship.

The relation between silvered and unsilvered forms of Speyeria at-
lantis (Edw.) has puzzled many people (Scott 1986b). Thus Grey et al.
(1963) discussed the two forms in the Black Hills of South Dakota,
where the silvered form with chocolate ventral hindwing (a. atlantis)
predominates in wet meadow areas, and the unsilvered form with
reddish-brown ventral hindwing (a. hesperis Edw. = a. lurana dosP.
& G.) prevails in drier areas. From a locality with 44% silvered adults,
W. Evans (in Grey et al. 1963:146) reared 8 silvered offspring with
chocolate ventral hindwing from silvered mothers with chocolate ven-
tral hindwing, and 26 unsilvered plus at least 1 silvered offspring with
a reddish-brown ventral hindwing from unsilvered mothers with a
reddish-brown ventral hindwing. The exact number of mothers con-
tributing was not known, but was probably one or two for each form.
Evans noted that the double dorsal stripes were light brown on atlantis
larvae, grayish white on hesperis larvae, and that hesperis pupae have
more light-brown shading on the wing case than do atlantis. Grey et
al. (1968) suggested that the two could be treated as separate species,

2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

though they retained them in one species because they seem to inter-
grade in other western U.S. regions.

A similar situation occurs in SW Manitoba where a dark variety of
silvered a. atlantis (a. hollandi F. & R. Cherm.) with chocolate ventral
hindwing flies in mountains and forest, whereas a very pale silvered a.
dennisi dosP. & G. usually with light-brown ventral hindwing flies on
tallgrass prairie. They occur near each other. At Duck Mountain, ad-
jacent populations show no intergradation (J. Troubridge pers. comm.).
In this area, they behave as separate species, although westward they
intergrade at Meadow Lake Park, Saskatchewan (Hooper 1973).

In Colorado E of the continental divide, the unsilvered form (a.
hesperis) prevails in the mountain foothills, and as one goes higher in
the mountains the silvered form (a. atlantis, = a. electa Edw.) gradually
increases in frequency until it predominates in the Canadian Zone.
Silvered forms in Colorado’s Front Range usually have a chocolate-
brown ventral hindwing, whereas unsilvered forms usually have a red-
dish-brown ventral hindwing, although this association sometimes breaks
down; thus some silvered adults have a red-brown ventral hindwing,
and some unsilvered ventral hindwing adults have a darker reddish-
brown ventral hindwing. Females have a slightly darker ventral hind-
wing than males; a bilateral gynandromorph from Critchell, for in-
stance, has a very red-brown ventral hindwing on the male side, a
darker red-brown ventral hindwing on the female side.

The silvered or unsilvered color is due to light reflection from in-
dividual scales. Silver scales appear transparent through a microscope,
but their surfaces reflect a white sheen (evidently due to structural
interference of light) which causes the silver appearance. Unsilvered
scales are cream in color because they appear to be filled with cream
pigment, and their surfaces do not reflect light; their scale structure
could be the same as silvered scales if the internal pigment blocks
transmission of light through the scale to prevent light interference. So
the difference between silvered and unsiivered scales could result solely
from absence or presence of internal cream pigment. A given wing
spot can be entirely cream (unsilvered), or it can be cream with a few
silver scales, or the entire spot can be covered with silver scales. Po-
tentially silvered spots occur in four series on the ventral hindwing:
basal, postbasal, postmedian, and submarginal. In the basal series, the
dot in the discal cell is more likely to have silver scales than the other
spots. The postbasal series of spots is less likely to be silvered than the
other series, and the basal and marginal series are most likely to be
silvered in the mostly unsilvered forms.

To determine the relation between the forms in Colorado, I reared
the eggs oi selected females, especially those females of a form rare in

VOLUME 42, NUMBER 1 3

their population because these would have the greatest likelihood of
mating with a male of the opposite form.

REARING METHODS

To obtain eggs, females were collected from Colorado Front Range
localities, brought to the laboratory, and placed in jars with Viola
nephrophylla Green leaves and fed honey-water once per day. Most
females lived about a week and laid several dozen eggs. Eggs hatch
readily, but first-stage larvae diapause in nature, so to prevent diapause
they were placed under constant light in tiny vials with a slice of green
violet leaf. After a few days or weeks some larvae ended diapause and
started to feed; these fed steadily until pupation on V. nephrophylla
leaves. Three months were required to raise offspring of one female.
Voucher specimens including larvae, pupal shells, and reared silvered
and unsilvered adults are in the National Museum of Natural History,
Washington, D.C.

RESULTS
Silvering of Mothers and Offspring

A total of 104 adult offspring were reared from 9 mothers from 6
Colorado sites. Each site is described below.

Tinytown (2120 m), Jefferson Co., isa Transition Zone foothills valley
bottom with ponderosa pine, douglasfir, willow, alder, honeysuckle,
etc., along the creek; the hostplants Viola canadensis L. and V. adunca
Smith (Scott 1986a) are common on the shaded gulch bottom and the
base of the N-facing slope. Here 92% of adults had a reddish-brown
ventral hindwing with mostly unsilvered spots, 6% were partly silvered
(N = 6 half silvered, N = 1 mostly silvered), and 2% were fully silvered
with a chocolate-brown ventral hindwing (N = 117). If the fully silvered
mother mated at random, the father was probably unsilvered; yet all
offspring were silvered (Table 1).

Corwina Park (2120 m), Jefferson Co., is a Transition Zone foothills
wooded gulch draining N; the hostplants V. adunca and probably V.
canadensis are in gulch-bottom shade and E-facing shaded slopes. Here
91% of adults were unsilvered with a red-brown ventral hindwing, 9%
silvered with a chocolate-brown ventral hindwing (N = 21). If the
completely silvered mother mated at random, the father was probably
unsilvered; yet all offspring were fully silvered (Table 1).

O’Fallon Park (2100 m), Jefferson Co., is near Corwina Park, and is
also a Transition Zone foothills wooded gulch draining N with the
hostplants V. adunca and V. canadensis in gulch-bottom shade and
E-facing shaded slopes. Here 83% were unsilvered with a red-brown
ventral hindwing, 18% silvered with a chocolate-brown ventral hind-

TABLE 1. Extent of silvering on ventral hindwing spots, and color of basal two-thirds
of ventral hindwing, of mothers and offspring. Numbers are proportions: for example,
“1” under “base” means all scales on wing base spots are silvered, “1/5” under “post-
median” means 20% of scales of postmedian spots are silvered, “0” under “‘submarginal”’
means no scales of submarginal spots are silvered, etc., “gyn’’ is bilateral gynandromorph,
“fis female, and “m”’ is male.

Material Sex Ventral hindwing Base Postbasal Postmed. Submarg.

Tinytown, Jefferson Co., mother caught 20 July 1984

Mother liek chocolate 1 1 1 1
Offspring 27 m chocolate I 1 il il
Offspring 19 f chocolate 1 1 1 1
Corwina Park, Jefferson Co., mother caught 13 July 1985
Mother et dark choc-brown 1 1 1 i.
Offspring 2m dark red-brown i 1 1 i
Offspring Pt choc-brown 1 1 I 1
Offspring lf dark choc-brown 1 Il 1 1
O'Fallon Park, Jefferson Co., mother caught 12 August 1985
Mother Ib it red-brown 2/38 Wis 12) 2
Offspring lm very red-brown 0 0 1/4 1/3
Offspring Gh very red-brown 0 0 0 1/5
Offspring Ike very red-brown 2/3 0 0 1/2
Critchell, Jefferson Co., mother caught 3 August 1985
Mother lf red-brown We 1/10 V2, 2
Offspring 8 f very red-brown 0 0 0 0
Offspring 6 f very red-brown 0 0 0 0
Offspring l gyn very red-brown 0 0 0 0
Mt. Judge female B, Clear Creek Co., mother caught 8 August 1985
Mother i at red-brown 28 1/8 is 1/8
Offspring lm very red-brown 0 0 0 1/6
Offspring 6m very red-brown 0 0 0 1/10
Offspring lm very red-brown 0 0 0 Vs
Offspring lm very red-brown 1/10 0 0 1/5
Offspring IL sem very red-brown 1/5 0 0 1/10
Offspring 2a very red-brown 0 0 0 1/10
Offspring Si very red-brown 0 0 0 0
Cherry Gulch, Jefferson Co., mother caught 17 July 1984
Mother ef red-brown 2/3 1/5 0 1/3
Offspring lef very red-brown 1/4 0 0 1/3
Mt. Judge female D, Clear Creek Co., mother caught 8 August 1985
Mother Lak dark red-brown yd 0 0 1/2
Offspring 2m very red-brown 0 0 0 0
Mt. Judge female F, Clear Creek Co., mother caught 8 August 1985
Mother at red-brown LS 0 0 1/10
Offspring 1m very red-brown 0 0 0 1/10
Offspring 1m very red-brown 1/10 0 0 1/5
Mt. Judge female A, Clear Creek Co., mother caught 8 August 1985
Mother hag red-brown 0) 0) 0 IAs
Offspring 9m very red-brown 0 0 0 0
Offspring 3f dark red-brown ) 0 0 1/10
Offspring lg dark red-brown 0 0 0 WG
Offspring ie) red-brown 0 0 0 0
Offspring I red-brown 0 0 0 1/6
Offspring Zit very red-brown 0 0 0 0

VOLUME 42, NUMBER 1 5

wing, and 4% intermediate (N = 19). If the nearly half-silvered mother
mated at random, the father was probably unsilvered; all offspring were
nearly unsilvered (Table 1).

Critchell (2370 m), Jefferson Co., is a shaded E—W streamside in the
upper Transition Zone foothills, with ponderosa pine, douglasfir, various
shrubs, grassy glades, and V. canadensis and V. adunca. Here 88%
were unsilvered with a reddish brown ventral hindwing, 7% fully sil-
vered, and 5% intermediate (N = 2 half silvered, N = 1 mostly silvered)
(N = 53). If the nearly half-silvered mother mated at random, the father
probably was unsilvered; all offspring were completely unsilvered (Ta-
ble 1).

Cherry Gulch (2100 m), Jefferson Co., is a Transition Zone foothills
gulch at the base of a N-facing slope covered with douglasfir, Holo-
discus, Physocarpus, other shrubs, and Viola canadensis. Here 97%
were unsilvered with a reddish brown ventral hindwing, 3% silvered
with a brown ventral hindwing (N = 69). If the mostly unsilvered mother
mated at random, the father was probably unsilvered; the single off-
spring was less silvered than the mother (Table 1).

Mt. Judge (2 km NE, 2770 m), Clear Creek Co., is a Canadian Zone
valley bottom, with forest (spruce, pine, douglasfir, some aspen) beside
grassy meadows, a tiny creek on the valley bottom, and V. canadensis
and V. nephrophylla. Silvered adults with a chocolate ventral hindwing
were most common, with a few silvered adults with a reddish brown
ventral hindwing; but unsilvered adults with a red-brown ventral
hindwing were also found, a few unsilvered adults with a brown ventral
hindwing, and a few variably silvered intermediates. The upperside
black lines vary from narrow to wide independent of ventral hindwing
variation. Shape of silver spots varies between individuals, as does amount
of black at the base of each silver spot, but this variation is also inde-
pendent of degree of silvering. Four females from this site labeled A,
B, D, and F, produced offspring (Table 1). If the Mt. Judge mothers
mated at random, they probably mated with silvered males because
74% of males here were silvered (Table 2). However, because of habitat
selection at this site (described in next section), and because all four
mothers were found in mixed woods away from the creek where only
38% of males were silvered (Table 2), the mothers probably mated with
unsilvered fathers. Mother B was about one-third silvered; her offspring
were almost completely unsilvered. Mothers A, D, and F, and their
offspring, were almost completely unsilvered.

Habitat Selection and Movements

The Mt. Judge site displayed habitat selection by the forms (Table
2). In several meadows along the tiny creek 90% of adults were silvered,

6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 2. Frequency of color forms at Mt. Judge site, based on nine visits 1984 to
1987.

Mixed woods away from creek Meadows along creek

Silvering No. male No. female No. male No. female
Mostly unsilvered 24 13 let 2
Half silvered 1 0 1 0
Completely silvered 15 8 2 53

10% unsilvered. In contrast, at the habitat edge near the head of the
valley, away from the creek in mixed woods—tiny meadows edging the
large meadow and in the adjacent meadow-edge, one-third (38%) of
adults were silvered, and two-thirds (62%) unsilvered.

A small mark-recapture study was conducted at Mt. Judge in 1987
(31 July, 5, 9 Aug.), in which 33 adults were marked and 16 recaptured.
Six unsilvered adults were marked (2 male, 4 female), and 3 females
recaptured, all in the mixed woods, one after 5 days. Twenty-seven
silvered adults were marked (14 male, 13 female), and 13 recaptured
(7 male, 6 female), after up to 9 days, including 5 moves completely
across the habitat, and 6 halfway across it. I conclude that silvered
adults move completely about the habitat, and females probably oviposit
in the mixed woods where host violets grow under conifers. But judging
from the restricted distribution of unsilvered adults (Table 2), these are
more local, and their restricted movement causes the habitat selection
difference. In general, unsilvered Colorado adults prefer open woods
with violets (N-facing slopes and gulch bottoms in the foothills), whereas
silvered adults also occupy more open wet valley bottoms.

Pheromones

Male odor of both forms from Mt. Judge was compared by the author.
Males of silvered and unsilvered forms smelled the same: the odor is
sweet but has a “hot” or “peppery” sensation, sweet but slightly peppery
pungent. Virtually every male had this odor, a few weaker than others.
Females lacked an odor. The description of odor is subjective, and
different observers might use different words to describe it, but it was
the same for both forms. Thus, the male pheromone is probably the
same in both forms, although the human nose certainly cannot match
the precision of laboratory instruments.

The pheromone system is complex. Males have androconial scales on
dorsal wing veins (Scott 1986b:fig. 37) which evidently produce the
pheromone odor; pheromone from these scales in the closely related
European Argynnis paphia L. causes the female to land and accept the
male (Magnus 1958). Females have a dorsal gland between abdomen
segments 7 and 8 (Scott 1986b:fig. 37). This gland in A. paphia produces

VOLUME 42, NUMBER 1 a

a pheromone that attracts males: femalelike dummies attract males but
do not elicit complete courtship, and freshly killed females are more
attractive to males than dried females (Magnus i958); virgins respond
to nearby males by exposing the abdomen gland and aiming the ab-
domen tip toward the male (Treusch 1967). Males have a paired gland
on the abdomen tip (Arnold & Fischer 1977, Scott 1986b) which, by
comparison with Heliconiini (Scott 1986b), could possibly transfer pher-
omone to the female during mating to enable mated females to produce
a third pheromone that repels males.

Courtship

Courtship of Speyeria atlantis, which is nearly identical to that of
Argynnis paphia (Magnus 1950), was described by Scott (1986b) based
mainly on unsilvered form courtships in Jefferson Co., Colorado. In
addition, a completed courtship between silvered male and female
forms was seen at Mt. Judge: female on flower when male sighted her
and landed; she fluttered her mostly spread wings with small amplitude
for 1 s, he flicked his nearly closed wings behind her for 1-2 s; she
rotated around flower top | revolution with her wings still spread while
he rotated after her and flicked his nearly closed wings once during
turn; she stopped, closed her wings, tilted forward so that her abdomen
was raised slightly but lowered from between hindwings; he spread his
wings partway; they joined.

Four courtships were seen at Mt. Judge between unsilvered males
and silvered females, as follows.

1) Male patrolled near female (prior mating status unknown) on flower, landed, flicked
wings (wingtips vibrating 0 to 1 cm apart about twice per s) for 10 s, curved abdomen
laterally to attempt joining (meanwhile female, wings closed, leaned forward with ab-
domen lowered from between closed hindwings and abdomen raised above horizontal
about 60°); wind blew them and he flew, fluttered over her for 1 s, landed, flicked beside
her 10 s, curved his abdomen but was too close and his abdomen tip missed (during his
bending she kept abdomen exposed), then he flew away. Female was evidently receptive
because she exposed her abdomen and did not perform rejection dance (fluttering wings
vigorously).

2) Male patrolled near silvered virgin (later found to have no spermatophores) on
flower, landed, flicked his wings, she crawled away with closed wings, he crawled after
her for 5 min while flicking and bending his abdomen, she stopped and spread wings
partly while he flicked and curved abdomen to attempt mating for 5 min, he flew away
(evidently she did not extrude genitalia, so he could not join). She was unreceptive even
though she did not flutter her wings, perhaps because, as judged from weak flight, she
was too young.

3) He pursued her in flight, they landed, she fluttered slightly and crawled away while
he flicked his wings and crawled behind, she got farther away, he flew up a short distance
but did not find her and flew away.

4) She raised her wings and slightly lowered and partly extruded her abdomen while
he flicked his nearly closed wings behind her, he flew away after about 30 s.

8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Data on courtship between forms are too few to be conclusive, but
no obvious courtship barriers to mating occur. Releases of reared virgins
are needed. Grey et al. (1963) inconclusively report abnormal courtship
of a few laboratory adults.

Larval Differences
(Figs. 1-14)

Color photographic slides were made of larvae and pupae from each
study site except larvae from Corwina, and some larvae and pupae
were preserved to correlate their color pattern with adult appearance.

From a distance, older larvae producing silvered adults (Figs. 5, 7-9) appear mottled
black with orangish tan spines and two middorsal white lines, whereas older larvae
producing unsilvered adults (Figs. 10-14) appear solid black with orange spines. Viewed
more closely, larvae of both forms are basically black, with a pair of middorsal whitish
lines 1 mm apart, and three rows of scoli (lateral to middorsal lines, supraspiracular, and
subspiracular) which are tan or orange with black tips. The head of both forms is black
with the dorsal half of the rear half of the head orangish.

Larvae of the silvered form (based on larvae from Tinytown, Figs. 5, 7-9) have the
middorsal whitish lines conspicuous and mostly continuous, though alternately wider and
narrower. Because Corwina pupae had less conspicuous lines than Tinytown pupae,
Corwina larvae may not have had the lines this conspicuous. Scoli of the silvered form
are orangish tan with black tips. Ground color is not as black as in the unsilvered form
so three rows of black bands with very sinuous narrowly white edges are recognizable:
along the dorsalmost scoli (edging middorsal white lines), along the supraspiracular scoli,
and in between these (Figs. 8, 9). A light gray-brown transverse band circles the rear of
each segment except middorsally, a remnant of the pale transverse stripes of Speyeria
nokomis (Edw.) larvae (Scott & Mattoon 1981).

Larvae of the unsilvered form (from O'Fallon, Critchell, Cherry Gulch, Mt. Judge) are
a little darker black and the pattern is obscured, so the black sinuous bands are unrec-
ognizable without a microscope, and the middorsal two lines are fainter and broken into
two dashed lines (Figs. 10-14). Scoli are orange with black tips. The only variation between
localities among unsilvered larvae involves the single Cherry Creek larva which had
slightly less orangish scoli. Edwards’ (1888b) description of the unsilvered form is very
similar.

The above descriptions of larvae do not correspond with descriptions of larvae of the
silvered and unsilvered forms in the Black Hills of South Dakota (Grey et al. 1963). Both
are described as identically black with orange spine shafts, the two middorsal lines grayish
white in the unsilvered form, light brown in the silvered form. Thus the two middorsal
lines are described as whiter in the unsilvered form in South Dakota, whereas they are
whiter in the silvered form in Colorado. My descriptions are based on 104 larvae and
dozens of color slides from many sites, whereas the South Dakota data are fewer.

Width of the two pale middorsal lines of the larva is apparently not closely linked to
degree of silvering of the adult; among larvae producing silvered adults, the whiteness
differed somewhat between the Tinytown and Corwina sites in Colorado as noted above,
and differed between Colorado and South Dakota adults.

Thus, both larvae and adults of the unsilvered form have more pig-
ment—more cream in adult scales, more orange on larval spines, more
black on larval body—so one can guess that the gene responsible for
the unsilvered form causes an increased deposition of some dark pig-
ment such as melanin.

Larvae and pupae of silvered ventral-hindwing S. atlantis from NE

VOLUME 42, NUMBER 1 9

Fics. 1-20. 1, First-stage larva, silvered form, Tinytown; 2, Second-stage larva, sil-
vered form, Tinytown; 3, Third-stage larva, silvered form, Tinytown; 4, Fourth-stage
larva, silvered form, Tinytown; 5, Fourth-stage larva, silvered form, Tinytown; 6, Third-
stage larva, silvered form, Tinytown; 7, Mature larva, silvered form, Tinytown; 8, Mature
larva, silvered form, Tinytown; 9, Mature larva, silvered form, Tinytown; 10, Third-
stage larva, unsilvered form, O'Fallon female C; 11, Mature larva, unsilvered form,
O'Fallon female C; 12, Mature larva, unsilvered form, Cherry Gulch; 13, Mature larva,
unsilvered form, Mt. Judge female F; 14, Mature larva, unsilvered form, Mt. Judge
female A; 15, Pupa (orange-brown wings), silvered form, Tinytown; 16, Pupa (orange-
brown wings), silvered form, Tinytown; 17, Pupa (orange-brown wings), silvered form,
Tinytown; 18, Pupa (orange-brown wings), silvered form, Tinytown; 19, Pupa (partly
orange-brown wings), unsilvered form, O'Fallon female C; 20, Pupa (black wings), un-
silvered form, Mt. Judge female F.

10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

U.S. (Edwards 1888a) are grayer than Colorado-South Dakota S. at-
lantis; larvae and pupae evidently show geographic variation as do
adults.

Pupal Differences
(Figs. 15-20)

Pupae from localities with sufficient numbers show great individual
variation, but there is no obvious important difference between silvered
and unsilvered forms. The pupa resembles S. nokomis (Scott & Mattoon
1981) in general, but is darker (orange-brown), and the posterior half
of each abdominal segment is darker because it is mottled with tiny
black dots and dashes. The anterior half of each abdominal segment is
not uniformly black as in nokomis: some pupae have a broad black
irregular band, but most have the black areas broken into spots, in-
cluding triangular spots just beside the anterior-pointing orange-brown
middorsal triangles on segments 5-7. Pupae from Tinytown have two
sinuous tan middorsal abdominal lines, but pupae from Corwina (both
sites produced silvered adults) and the other sites had weak tan mid-
dorsal lines. Pupal wing color varies from mostly black to almost wholly
orange-brown, but most are mostly orange-brown, a few black-winged.

Grey et al. (1963) describe the pupal wing cases of silvered forms as
darker with less light brown mottling than those of unsilvered forms
in the Black Hills. However, they reared only three silvered adults, so
the difference is probably due to small sample size because all Colorado
sites with large samples show considerable variation in pupal wing color.
Pupae producing silvered adults are not darker in Colorado.

DISCUSSION

There are several reasons why S. a. atlantis and S. a. hesperis could
be treated as distinct species: they often fly together, they prefer dif-
ferent microhabitats, amount of silvering seems usually linked with
ventral hindwing color, mothers usually produce offspring resembling
themselves, and their larvae differ. If scientists were aware only of
Black Hills populations, the two would certainly be treated as separate
species because they are so distinct there. Some anecdotes (coinci-
dences?) also fit the two-species theory. For instance, six unsilvered
males and one si!vered pair were found in the Mt. Judge mixed woods
28 July 1987, the silvered pair in copula.

There are several reasons why S. a. atlantis and S. a. hesperis could
be treated as one species:

|) Silvered and unsilvered forms are linked by a complete series of intermediate adults,
from slightly to partly to half to mostly silvered, although only slightly silvered inter-
mediates are common.

VOLUME 42, NUMBER 1 iat

2) Unsilvered mothers sometimes produce silvered offspring (Grey et al. 1963:146),
and half-silvered mothers often produce unsilvered offspring (Table 1).

3) In many populations, silvered forms are rare (<5%) as in the lower foothills of the
Colorado Front Range, rarity a true species might have difficulty surviving. The reverse
is also true, in which unsilvered forms are rare within silvered populations, as in the wet
center of the Black Hiils (Grey et al. 1963). However, S. coronis (Behr) is just as rare and
it survives.

4) Frequencies of the forms show clinal trends, both altitudinally in the Colorado Front
Range, and along habitat gradients. For instance, in the Black Hills (Grey et al. 1963),
atlantis is common in wet meadow habitats on poorly drained granite, and is rarer away
from these areas. Similarly, in S Colorado (Scott & Scott 1980) hesperis predominates in
the lower foothills, both forms occur in dry areas at higher altitude, and atlantis pre-
dominates in three wet meadow enclave habitats at middle altitudes: Coaldale in Arkansas
Canyon, Fremont Co.; SW of Westcliffe on Wet Mountain Valley floor, Custer Co.;
Stonewall in upper Purgatoire River valley, Las Animas Co. Such enclaves have not been
found in the Front Range W of Denver, where silvered forms are rare in the foothills
and increase in frequency with altitude until they predominate in the upper Canadian
Zone.

5) When attempts are made to divide S. atlantis into silvered and unsilvered “species ’’,
their distributions are incongruous because unsilvered forms cut an E-W swath through
the range of silvered forms, replacing them in the process (Scott 1986b).

The silvered-unsilvered division also fails to solve the problem of sympatry of S. a.
dennisi and S. a. atlantis (hollandi), both of which are silvered, in Manitoba. A species
S. dennisi could include S. atlantis ratonensis Scott from NE New Mexico and S. a. greyi
from NE Nevada, but dennisi is said to intergrade W to atlantis in Saskatchewan-Alberta,
and greyi intergrades with dodgei in S Idaho (P. C. Hammond pers. comm.), and at least
greyi seems independently evolved toward similar pallidity.

6) Other S. atlantis subspecies have polymorphisms of silvered-unsilvered adults: wa-
satchia dosP. & G. (=tetonia dosP. & G.) in W Wyoming-Utah is usually unsilvered,
chitone (Edw.) in S Utah and schellbachi Garth in N Arizona are usually silvered.

7) Other species of Speyeria have silvered-unsilvered polymorphisms: zerene (Bdv.)
in California and S Oregon, callippe (Bdv.) in N California and the Sierra Nevada, egleis
(Behr) in the Sierra and Utah, hydaspe (Bdv.) in British Columbia. These polymorphisms
are accepted by lepidopterists. Boggs (1987) hypothesized that rare unsilvered S. mor-
monia are homozygous recessives that fail to reproduce, which is dubious because S.
mormonia artonis (Edw.) are nearly always unsilvered.

8) Association between ventral hindwing color and silvering and larval color pattern
breaks down geographically. In the Black Hills and E of the continental divide in the
Colorado mountains, silvered adults have a chocolate-brown ventral hindwing (darker in
the Black Hills), and unsilvered adults usually have a reddish brown ventral hindwing.
However, in N-central New Mexico, 98% of adults (N = 60) are silvered but the ventral
hindwing varies from chocolate- to reddish brown. In SW Manitoba S. atlantis dennisi
and S. a. atlantis (a. hollandi) are 100% silvered but the ventral hindwing is usually light
brown in the former and chocolate-brown in the latter. And silvered adults have the
ventral hindwing browner in the Black Hills than in the Colorado Front Range. Larval
differences in Colorado are partially reversed in the Black Hills, and larvae are grayer
in E North America.

The conclusion that silvered and unsilvered adults are polymorphic
forms of one species seems preferable.

Paleogeography

The current geographic distribution of wing characters suggests that
the dark silvered form (S. a. atlantis) occupied the coniferous forest in
N U.S. and the Rocky Mountain foothills during the Ice Age; afterwards

12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

it moved higher in altitude and latitude. The unsilvered form with
reddish brown ventral hindwing (now S. a. hesperis, a. wasatchia, a.
irene [Bdv.]) occupied open forest in the southern Great Basin lowlands;
after the Ice Age it spread N into the mountains, E through lowland S
Wyoming to the Black Hills, and S along the Colorado mountain foot-
hills. The silvered form with narrower black lines and a pale brown
ventral hindwing (now S. atlantis dennisi and a. ratonensis) occupied
aspen parkland in the current S Great Plains or central Texas; after the
Ice Age it spread, respectively, N to Canada, and upward to a mountain
mesa. The forms became sympatric after the Ice Age.

Mechanism of Inheritance

The inheritance mechanism of silvering is unknown. Rarity of half-
silvered adults suggests dominance, but some broods with half the
offspring silvered and half unsilvered should occur but did not. If half-
silvered adults were heterozygotes, they would not produce all-unsil-
vered broods as at Critchell. If silvered is dominant, rare silvered moth-
ers would be likely to produce silvered offspring, as at Tinytown and
Corwina. O’Fallon and Cherry Gulch broods perhaps suggest modifier
genes that cause part-silvering.

Maternal inheritance seems the best guess now, and fits all the reared
broods; offspring would resemble the mother, the father having no
effect or perhaps merely modifying partly silvered offspring. Sterling
O. Mattoon (pers. comm.) states that Speyeria offspring generally re-
semble their mother very closely, although some silvered offspring have
been reared from unsilvered mothers and vice versa.

ACKNOWLEDGMENTS

I thank L. P. Grey, T. C. Emmel, and C. L. Boggs for reviewing the manuscript.

LITERATURE CITED

ARNOLD, R. A. & R. L. FISCHER. 1977. Operational mechanisms of copulation and
oviposition in Speyeria (Lepid.: Nymphalidae). Ann. Entomol. Soc. Amer. 70:455-
468. See Fig. 6.

Boccs, C. L. 1987. Demography of the unsilvered morph of Speyeria mormonia in
Colorado. J. Lepid. Soc. 41:94-97.

Epwarpbs, W. H. 1888a. Description of the preparatory stages of Argynnis atlantis,
Edw. Can. Entomol. 20:1-3.

1888b. Description of the preparatory stages of Argynnis hesperis, Edw. Can.
Entomol. 20:67-69.

GREY, L. P., A. H. MoEcK & W. H. Evans. 1963. Notes on overlapping subspecies. II.
Segregation in the Speyeria atlantis of the Black Hills (Nymphalidae). J. Lepid. Soc.
17:129-147.

Hooper, R. 1973. Butterflies of Saskatchewan. Saskatchewan Mus. Nat. Hist., Regina,
Saskatchewan. 216 pp.

Macnus, D. 1950. Beobachtungen zur Balz und Eiablage des Kaisermantels Argynnis
paphia L. (Lep., Nymphalidae). Z. Tierpsychol. 7:435-449.

VOLUME 42, NUMBER 1 13

1958. Experimentelle Untersuchungen zur Bionomie und Ethologie des Kai-
sermantels Argynnis paphia L. (Lep. Nymph.) I. Ueber optische Ausloeser von An-
fliegereaktionen und ihre Bedeutung fuer das Sichfinden der Geschlechter. Z. Tier-
psychol. 15:397—426.

ScoTT, J. A. 1986a. Larval hostplant records for butterflies and skippers (mainly from
western U.S.), with notes on their natural history. Papilio (New Series) #4. 37 pp.

1986b. The butterflies of North America, a natural history and field guide.
Stanford Univ. Press, Stanford, California. 583 pp., 64 pls.

Scott, J. A. & S.O. MATTOON. 1981. Early stages of Speyeria nokomis. J. Res. Lepid.
20:12-15.

ScoTT, J. A. & G.R. Scott. 1980. Ecology and distribution of the butterflies of southern
central Colorado. J. Res. Lepid. 17:73-128 (corrections 19:240).

TREUSCH, H. W. 1967. Bisher unbekanntes gezieltes Duftanbietin paarungsbereiter
Argynnis paphia-Weibchen. Naturwissenschaften 54:592.

Received for publication 22 April 1987; accepted 30 November 1987.

Journal of the Lepidopterists’ Society
42(1), 1988, 14-18

POPULATION FLUCTUATIONS OF
AZETA VERSICOLOR (FABRICIUS) (NOCTUIDAE)
ON GLIRICIDIA SEPIUM (JACQ.) (FABACEAE)
IN NORTHEASTERN COSTA RICA

ALLEN M. YOUNG

Invertebrate Zoology Section, Milwaukee Public Museum,
Milwaukee, Wisconsin 53233

ABSTRACT. Counts of early stages, especially caterpillars, of Azeta versicolor on the
host tree Gliricidia sepium planted as shade cover in a vanilla plantation were made
intermittently during five years. Based on field observations and rearings, the mature
caterpillar and pupa were described, noting two distinct color morphs in the former.
Tachinid parasites were also noted. Caterpillar abundance was analyzed and interpreted
in relation to monthly rainfall and leaf-flushing in the host tree, since caterpillars feed
preferentially on new (flush) leaves. Numbers of caterpillars were highly correlated with
monthly rainfall. It is concluded that population cycles of the moth are regulated by
seasonal patterns of leaf-flushing in the host.

Additional key words: immature stages, leaf flushing, population dynamics.

Impact of seasonal fluctuations in rainfall on leaf-flushing of semi or
fully deciduous host trees is a major environmental factor molding
population dynamics of noctuids and other Lepidoptera in the tropics
(Vaishampayan & Veda 1980, Blair 1982, Tucker & Pedgley 1983).
Fabaceous legume crops in the tropics are especially preferred hosts of
noctuid and pyralid defoliators, with seasonal patterns of population
outbreaks typical for several of these host species (Bradley & Carter
1982, Panchabhavi & Holihosur 1982). In many species, caterpillars
preferentially defoliate immature leaves or other most nutritious tissues
of the host, which are often only seasonally available (Futuyma &
Wasserman 1980, Bracken 1984). Here I report seasonal abundance
pattern of immature stages for the noctuid moth Azeta versicolor (Fa-
bricius) on leaves of the fabaceous legume tree Gliricidia sepium (Jacq.)
planted as shade cover in a vanilla plantation.

METHODS

Counts of life stages of Azeta versicolor were obtained on 16 dates
between March 1982 and June 1987 at “Finca La Tirimbina,” near La
Virgen (10°23'N; 84°07”W; 200 m elev.), Sarapiqui District, Heredia
Province. Within a ca. 1600 m? plot containing about 900 trees of
Gliricidia sepium planted a few years earlier to shade vanilla plants,
30 arbitrarily selected trees (canopy height ca. 3 m) were censused for
Azeta versicolor caterpillars at various times. The medium-sized (40
mm wingspan) adults and caterpillars were readily recognizable in field
censuses: adult moths are drab greenish brown with striking red ab-

VOLUME 42, NUMBER 1 15

dominal coloration, and yellowish mature caterpillars usually rest close
to the base of host trees, typically on stems and leaves of vanilla orchid
vines and other epiphytes under the trees.

On a given caterpillar census, as many as 100 samples of both mature
or immature leaves and stems on each tree (usually up to height of 1.5
m) were searched for “young” caterpillars (mixed early instars) and
eggs. Condition of canopy foliage of Gliricidia sepium was also noted
(such as presence or absence of flush leaves), providing a qualitative
picture of local timing of peak flushing periods in relation to seasonality.
A total of 80 caterpillars (later instars) were placed in clear-plastic bags
containing fresh cuttings of G. sepium and kept tightly shut for rearing.
Parasitism of caterpillars and pupae was noted from this sample.

RESULTS

Natural history. In both of two color morphs of the final stage
caterpillar, roughly equal in abundance and not sexual dimorphism,
the head is pinkish white with black dots. Thoracic and abdominal
regions of the mature caterpillar (40 mm long by 5 mm wide) have
eight lengthwise narrow bands, which, in the dark form are as follows,
dorso medial to latero ventral: (1) deep yellow; (2) faintly yellow edged
in black; (3) pale bluish streaked with tiny black lines and a single
round black dot on each segment; (4) pale bluish yellow; (5) wide pale
blue; (6) lateral (spiracular) stripe pale blue with thin black line medially
and reddish spiracle openings, each with a black dot dorsoanteriorly
and yellow dot ventroposteriorly; (7) yellow with black edging ventrally;
(8) grayish with raised black dot, one per segment. Prolegs pinkish,
each with yellow dot laterally, ringed with black. Glossy black elongate
setae on profuse raised areas of cuticle. Anal clasper faintly pinkish;
true legs reddish. In the light form, there are no black stripes bordering
other stripes.

The reddish brown pupa (20-22 mm long by 5-6 mm wide) occurs
in a loosely constructed cocoon of host leaves pulled together and an-
chored with light brown silk. Both caterpillar and pupa thrash about
vigorously when picked up. Adults are active throughout the day, and
are skittish and difficult to capture with an insect net. The spherical,
glossy yellow eggs are placed singly on the undersides of G. sepium
leaves. Of 257 eggs discovered in the field, ca. 70% were on immature
(meristem) leaves. As noted above, mature caterpillars rest on vanilla
vines and other epiphytic debris on host trunks during daytime, and
are chiefly nocturnal feeders, crawling into the G. sepium canopy to
feed. Each of 3 pupae (out of 30 reared from collected caterpillars)
yielded 1 tachinid parasite.

16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

RAINFALL (MM)

1100

1000-

800

TOTAL
CATERPILLARS

700

600 300

1 71 1 m1

400 200

100

100 #

N
\
| Cae 5 res
J FMAMJJ ASONDJ FMAMJ J ASONDJ FMAMJ J ASONDJ FMAMJJ ASONDJ FMAMJ J ASONDJ FMAMJ JASOND
1982 1983 1984 1985 1986 1987
MONTHS AND YEARS

Fic. 1. Monthly total rainfall (line), total numbers of Azeta versicolor caterpillars
(vertical bars, 5th instars represented by hatching), timing of leaf-flushing (brackets), and
periods of caterpillar absenses (x), in the 36-tree subplot of Gliricidia sepium at “Finca
La Tirimbina.” Rainfall data courtesy of Finca La Tirimbina.

Seasonal population fluctuations and leaf-flushing. Abundance of
Azeta versicolor caterpillars on sampled Gliridia sepium trees varied
greatly among census dates (Fig. 1). Aside from an occasional hesperiid
and limacodid caterpillar, I did not observe other herbivores abundant
on these trees. When the data are examined relative to rainy and dry
season periods at La Tirimbina, two patterns become apparent: (1) the
highest numbers of mature and partly grown caterpillars occurred in
the rainy season, especially June-August, approximately during the first
half of the lengthy rainy season characteristic of this locality; (2) cat-
erpillars are absent during the dry season (February—March) (Fig. 1).
A high positive correlation resulted between numbers of caterpillars
and monthly rainfall (r = 0.81, P < 0.01).

Also during July-August, as many as 500 adults were counted within
a 600 m? strip of low vegetation bordering one side of the vanilla grove
during a 2 morning census (0800-1000 h). As many as 100 eggs were
counted within the 36-tree subplot on a single day in July or August,

VOLUME 42, NUMBER 1 t7

and none were found in February or March. During dry months, host
trees are partly deciduous, and only mature leaves are present. Flow-
ering in G. sepium at La Tirimbina is most intense during March and
early April. During the first three months of the rainy season, G. sepium
exhibits intensive leaf flushing (Fig. 1).

The highest population density of Azeta versicolor at La Tirimbina
follows intense flushing of new leaves on larval host trees. The increased
availability of immature (flush) leaves during the beginning of the rainy
season provides an abundant food resource for larvae. Population build-
up can be so intense in the rainy season as to result in 80-100% defo-
liation of G. sepium on some plots. I conclude that the breeding pop-
ulation of this Neotropical noctuid fluctuates in size throughout the
year at La Tirimbina in a consistent manner, and in response to the
seasonal leaf-flushing cycle of G. sepium.

DISCUSSION

Some tropical legume crops attacked by host-specific noctuids and
other moths undergo severe defoliation at certain times of year (Singh
& Budhraja 1980). Legume tree species typically planted as a permanent
shade over perennial crops in the tropics such as cacao, coffee, and
vanilla, including G. sepium (Inostrosa & Fournier 1982), and others
such as Erythrina (Borchert 1980) undergo pronounced seasonal cycles
in leaf-flushing in direct response to water-stress and rehydration (Reich
& Borchert 1982). The complete absence of Azeta versicolor caterpillars
on Gliricidia sepium in the dry season at La Tirimbina is due to absence
of immature (newly flushed) leaves. Thus, availability of edible leaf
tissues, a consequence of seasonally regulated hostplant leaf-flushing,
determines temporal pattern of population build-up in this noctuid.
The degree to which A. versicolor exploits other larval host plants at
La Tirimbina is unknown.

Skittish behavior of the diurnally active adults, and their vivid red
abdominal colors, suggest aposematism, perhaps a consequence of larval
feeding on G. sepium, a species well known for high concentrations of
courmarin compounds in its leaves (Allen & Allen 1981). Marked build-
up of the adult population in the first half of the rainy season at La
Tirimbina suggests a population structure in which biotic regulation of
the herbivore may be minimal.

Gliricidia sepium is capable of producing a new flush of leaves
following a period of intense herbivory by (J. R. Hunter & A. M. Young
pers. obs.). The ability of G. sepium to recover rapidly from intense
defoliation may be mediated in large part by the tree’s capacity to fix
nitrogen in the soil.

18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ACKNOWLEDGMENTS

Fieldwork was supported in part by grants from the American Cocoa Research Institute.
I thank J. R. Hunter for allowing me to conduct research at his farm, Finca La Tirimbina.
I thank Jorge Mejias of La Tirimbina for field assistance, and R. W. Poole of the Systematic
Entomology Laboratory, U.S. Department of Agriculture, for identifying the moth. Voucher
specimens of the moth are deposited in the collections of the Milwaukee Public Museum.
Comments of two reviewers, and statistical advice froin R. E. Spieler were helpful in
revising the manuscript.

LITERATURE CITED

ALLEN, O. N. & E. K. ALLEN. 1981. The Leguminosae: A source book of characteristics,
uses and nodulation. Univ. Wisconsin Press, Madison, Wisconsin. 812 pp.

Bair, B. W. 1982. Seasonal abundance of Agrotis segetum (Denis & Schiff.) and A.
ipsilon (Huefn.) (Lepidoptera: Noctuidae) in Zimbabwe, and a method of forecasting
post-winter population densities. J. Entomol. Soc. South Africa 45:201-215.

BORCHERT, R. 1980. Phenology and ecophysiology of tropical trees: Erythrina peop-
pigiana O. F. Cook. Ecology 61:1065-1074.

BRACKEN, G. K. 1984. Within plant references of larvae of Mamestra configurata
(Lepidoptera: Noctuidae) feeding on oilseed rape. Can. Entomol. 116:45-—49.

BRADLEY, J. D. & D. J. CARTER. 1982. A new lyonetiid moth, a pest of winged-bean.
Syst. Entomol. 7:1-9

FutuyMa, D. J. & S. S. WASSERMAN. 1980. Resource concentration and herbivory in
oak forests. Science 210:920-922.

INostrosa, I. & L. A. FOURNIER. 1980. Efecto alelopatico de Gliricidia sepium (Jacq.)
(Madero Negro). Revista Biol. Trop. 30:35-39.

PANCHABHAVI, K. S. & S. N. HoOLrHosuR. 1982. Notes on groundnut as a new host of
Achaea janata Linn. (Lepidoptera: Noctuidae) at Dharwad, Karnataka. Ind. J. Agric.
Sci. 52:43.

REICH, P. B. & R. BORCHERT. 1982. Phenology and ecophysiology of the tropical tree,
Tabebuia neochrysantha (Bignoniaceae). Ecology 63:294-299.

SINGH, O. P. & K. BuDHRajJA. 1980. Zur Biologie des indischen Sojabohnen-Schadlings
Plusia acuta Walker (Lep., Noctuidae). Anz. Schadlingsk. Pflanzenschutz Umwelt-
schutz 53:184-185.

TUCKER, M. R. & D. E. PEDGLEy. 1983. Rainfall and outbreaks of the African army-
worm, Spodoptera exempta (Walker) (Lepidoptera: Noctuidae). Bull. Entomol. Res.
73:195-199.

VAISHAMPAYAN, S. M. & O. P. VEDA. 1980. Population dynamics of gram podborer,
Helicoverpa armigera (Hiibner) and its outbreak situation on Gram, Cicer arietinum
L. at Jabalpur. Ind. J. Entomol. 42:453-459.

Received for publication 6 August 1987; accepted 16 October 1987.

Journal of the Lepidopterists’ Society
42(1), 1988, 19-31

BUTTERFLIES OF NORTHEAST TENNESSEE

CHARLES N. WATSON JR.
Department of Entomology, Clemson University, Clemson, South Carolina 29634

AND

JOHN A. HYATT
439 Forest Hills Drive, Kingsport, Tennessee 37663

ABSTRACT. Here we give results of a 10-year survey of butterflies in a seven-county,
7000 km? area of NE Tennessee. Ninety-one species are listed and their seasonal occurrence
tabulated on a 10-day basis. Twenty-seven species are judged to be univoltine, twenty-
nine bivoltine, and twenty-one multivoltine. The remainder are thought to be migrants
or strays that do not overwinter in NE Tennessee. Comparison of our species list with
that of SW Virginia and N Georgia indicates the fauna lacks a number of lowland species
that occur in N Georgia, and some typically northern species in SW Virginia. Ten species
known to occur in both comparison areas, but not recorded here, will probably be found
in the future.

Additional key words: Appalachians, biogeography, survey, Georgia, Virginia.

There is little published information on the butterfly fauna of Ten-
nessee (Field et al. 1974). Osburn (1895a, 1895b) lists 70 species oc-
curring around Nashville. Richards (1932) provides some Tennessee
records. Watson (1946) and Snyder (1957) list some species occurring
in the Smoky Mountains. The best source for the State as a whole is
Opler (1983) which contains county distribution maps for all species
occurring in the eastern U.S.

We have collected extensively in NE Tennessee for more than 10
years. Here we summarize results of our collecting, make comparisons
with other areas in the S Appalachian region, and list additional species
likely to occur in NE Tennessee.

STUDY AREA

The area encompasses seven counties in NE Tennessee with a total area of 7000 km?
(Fig. 1). Two physiographic subdivisions of the S Appalachian region are represented.
The SE portion of the area lies within the Blue Ridge Province, the remainder in the
Ridge and Valley Province.

The peaks of the Blue Ridge are known locally as the Unaka Mountains. They are
characterized by rugged terrain and heavily forested slopes. Elevations vary from 450-
600 m in the narrow valleys to 750-1900 m on the peaks. Underlying sedimentary and
metamorphic rocks are Cambrian and Pre-Cambrian in age. Soils tend to be sandy and
acidic. Most of this portion of the area lies within the Cherokee National Forest (Miller
1974, USDA 1958, 1956, 1985).

The Ridge and Valley portion is underlain by strongly folded sedimentary rocks of
Ordovician and Cambrian age. Differential weathering has resulted in long, narrow
sandstone ridges trending NE to SW, alternating with valleys developed on less resistant
limestone and shale. The easternmost valley is broad and part of a series of connecting
valleys extending from Pennsylvania to Alabama commonly called the Great Valley

20 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

VIRGINIA

> i

SULLIVAN

-~ CARTER

15 MI
20 KM

Fic. 1. Study area in NE Tennessee, showing county boundaries. Dashed line is
approximate boundary separating Blue Ridge Province (Unaka Mts., SE) from Ridge and
Valley Province (NW).

Within it there is local relief in the form of shale knobs and entrenched streams. To the
NW, straddling the border of Greene and Hawkins counties, and extending into SW
Sullivan Co., are a group of ridges collectively called Bays Mountain. Another prominent
feature, Clinch Mountain, runs through NW Hawkins Co. Elevations average lower in
the Ridge and Valley Province, ranging from 300 m in the valleys to 600-900 m on
ridges. Ridge soils are generally sandy, shallow, and unproductive while valley soils
developed on limestone are rich and fertile (Fenneman 1938, Miller 1974, U.S. Dep.
Agric. 1953b, 1958a, 1958b, 1979, 1985).

The entire area is drained by the Holston River and its tributaries, part of the Tennessee
River drainage system. The rivers have been extensively impounded for flood control and
power generation (Hunt 1967).

Climate is characterized by mild winters and warm summers. Average annual precip-
itation is 100-150 cm except at highest elevations where it may exceed 200 cm. Topog-
raphy and altitudinal differences cause much local variation in climate. As a rule, S- and
W-facing slopes are drier than those facing N and E. Average frost-free season varies
from 190 days in NW valleys to 150 days in the Unaka mountains (Walker 1969, U.S.
Dep. Agric. 1953, 1979).

Before European settlement, the area was covered with oak-chestnut forest. Clearing
of valleys for agriculture, logging in the mountains, and chestnut blight decimated primary
forests, especially in the Ridge and Valley. Today forests are concentrated in the Unaka
Mountains and on the NW ridges. At lower elevations, oaks (Quercus spp.), hickories
(Carya spp.), yellow poplar (Liriodendron tulipfera L.) and other hardwoods are common,
often mixed with hemlock (Tsuga spp.), and several pines (Pinus spp.). The Unaka
Mountains are high enough to show altitudinal zonation. Above 900 m, northern forest
types such as sugar maple (Acer saccharum Marsh.), beech (Fagus grandifolia Ehrh.),
and yellow birch (Betula alleghaniensis Britton) are common. Above 1500 m, red spruce
(Picea rubens Sarg.), and fraser fir (Abies fraseri [Pursh.] Poir) predominate. Treeless,
dome-shaped summits called balds occur on some peaks. In the Ridge and Valley, stands
of red cedar (Juniperus virginiana L.) are common in old fields on limestone soils. Marshes
and canebrakes are rare throughout, most having been drained, cleared, or inundated by
reservoirs (Braun 1950, Walker 1969, U.S. Dep. Agric. 1953a, 1953b, 1956, 1958a, 1958b,
1979, 1985).

VOLUME 42, NUMBER 1 ya

METHODS

Most records come from collections and fields notes made by the authors from 1975
through 1986. Additional records were obtained from participants in a Southern Lepi-
dopterist Society field meeting in the area in 1980, and from collections made by students
at Sullivan (County) High School during fall 1977 and 1978. Collections at the U.S.
National Museum (USNM) and the Carnegie Museum of Natural History (CMNH) were
examined, but no additional records were found. Most specimens are retained in the
authors’ collections; others have been placed in USNM and CMNH. Some identifications
were confirmed by C. V. Covell Jr., University of Louisville, and by R. K. Robbins and
J. M. Burns (USNM). Butterfly nomenclature follows Hodges et al. (1983).

To facilitate comparison with NE Tennessee, we define SW Virginia as Giles, Mont-
gomery, and Floyd counties and those counties to the SW entirely or predominantly
within the transition zone of Clark and Clark (1951). North Georgia is defined as those
counties entirely or predominantly within the mountain region of the State as defined
by Harris (1972). Species records for these regions were obtained from Opler (1983),
Clark and Clark (1951), and Harris (1972).

RESULTS AND DISCUSSION

We recorded 91 species of butterflies and skippers from NE Tennessee
(Table 1). In addition, specimens of Celastrina ladon form neglecta-
major Tutt, considered by some to be a distinct species (Opler & Krizek
1984), have been collected in May and early June. An old sight record
for Anaea andria Scudder for which we do not have a precise date is
not included in the table but is discussed below.

The species found in NE Tennessee can be considered as falling into
two categories: residents, which overwinter in the area; and migrants
or strays, which do not normally overwinter in the area, although many
regularly occur in summer and fall.

A number of resident species are rare or local in distribution, but
only one appears limited to a particular part of the study area. Speyeria
aphrodite (F.) has been collected only in the Blue Ridge, where it is
often common at elevations above 600 m.

Analysis of flight-period data in Table 1 to determine number of
broods for resident species is complicated by the fact that the flight
period of a species at any particular locality may vary from year to
year due to climatic and biological factors. Flight periods are also
affected by elevation, beginning and ending one to three weeks later
at high elevations in the Blue Ridge than in the Ridge and Valley. For
example, summer brood Erynnis horatius (Scudder & Burgess) has been
collected at Bays Mountain Park (600 m) in Sullivan Co. from late June
through mid-August, but a fresh specimen was collected in Carter Co.
at 1200 m on 8 September.

We believe the following residents are univoltine in NE Tennessee:

Thorybes bathyllus (J. E. Smith) E. brizo (Bdv. & Leconte)
T. pylades (Scudder) E. juvenalis (F.)
Erynnis icelus (Scudder & Burgess) Wallengrenia egremet (Scudder)

22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Temporal distribution of butterfly species adults in NE Tennessee.

Species 20 31 10 20 30 10 20 31

Hesperiidae
Epargyreus clarus X xX xX xX 4
Autochton cellus xX
Achalarus lyciades xX
Thorybes bathyllus
T. pylades xX

Staphylus hayhurstii

Erynnis icelus

E. brizo

E. juvenalis

E. horatius

E. baptisiae

Pyrgus communis x

Pholisora catullus

Nastra lherminier

Ancyloxypha numitor

Thymelicus lineola

Hylephila phyleus

Polites coras x

P. themistocles

P. origenes xX

Wallengrenia egeremet

Pompeius verna x

Atalopedes campestris

Atrytone delaware

Poanes hobomok

P. zabulon xX xX

Euphyes ruricola metacomet

Amblyscirtes hegon xX

A. aesculapius

A. vialis xX X

Papilionidae

rm

~~ KX

rm xX PX
rm XK rm XK

Pr

mx MX

Battus philenor

Papilio polyxenes asterius

P. cresphontes

P. glaucus xX
P. troilus

Eurytides marcellus xX

pd dx
>> > >
x >> >
D> Dd be Dd
> dd ><

Pieridae

Pontia protodice

Artogeia virginiensis

A. rapae x
Euchloe olympia

Falcapica midea

Colias philodice xX
C. eurytheme xX
Phoebis sennae eubule

Eurema lisa

E. nicippe

ee eeEeeeeeeeeeSFSFeee

~ KKM
KKK KM
KK KKM
KKK
x Kw MM
ras

> >< >

23
ce

20

Nov
1

10

21-
3]

Oct.
1l-
20

jz

10

21-
30

Sept.
1l-
20

Continued.
=
10

21-
31

Aug.
ll-
20

i=

PABEE I
10

21-
31

July
ee Se a Oe eo Une Se Le paoIe Ie A
10 20

21-
30

June
1l-
20

1
10

VOLUME 42, NUMBER 1

» ro ba bd be
* x“ pe
> bd bd x oe
* * ree pee 5d
» ce Pa bd
x Dr >< re x x
>< ex x OX <> x DM Od re ba
mx x «Mx ex x bd > >< ><
rp Mx xx OM <r OX ra x x
rr OO KO xx xX x x
<< bd bd ><! MX bd » x < x x
>< r <>< Mx. <x <
» be <
> >< >< od ><! <
» x y < «xx » es
* rd Od » >< rd ><
y x <x xxx xX re rd >< re >< >

Species

Lycaenidae

Feniseca tarquinius
Lycaena phlaeas americana
Harkenclenus titus mopsus
Satyrium calanus falacer
S. caryaevorum

S. liparops strigosum
Calycopis cecrops

Mitoura grynea

Incisalia augustus croesioides
I. henrici

I. niphon

Parrhasius m-album
Strymon melinus

Erora laeta

Everes comyntas
Celastrina ladon

C. ebenina

Glaucopsyche lygdamus
Libytheidae

Libytheana bachmanii
Nymphalidae

Polygonia interrogationis
P. comma

Nymphalis antiopa
Vanessa virginiensis

V. cardui

V. atalanta

Junonia evarete

Euptoieta claudia

Speyeria diana

S. cybele

S. aphrodite

Clossiana bellona toddi
Phyciodes tharos
Charidryas nycteis
Euphydryas phaeton
Basilarchia arthemis astyanax
B. archippus

Apaturidae
Asterocampa celtis
A. clyton
Satyridae

Enodia anthedon
E. creola

Cyllopsis gemma
Hermeuptychia sosybius
Megisto cymela
Cercyonis pegala
Danaidae

Danaus plexippus

TABLE l.
Mar.
ie ee
20 31
x
xX
xX
x
x x
xX x

Continued.

xx KR MM

rr

KKM KM

mx

KKM MMM RK

mx

rm xX

10

rm xX

~MK MMM RK OK

mx

xx PM

rs XK

PK XS

mx KKM MX

rs XK

1]-

20

Nov.
te

10

2\-
31

Oct.
(i=
20

1
10

21-
30

Sept.
ll-
20

Continued.
l=
10

2\-
31

Aug.
ll-
20

ie

TABLE 1.
10

21-
31

July
nr ine Serre aol Ola Reo OLS. ee eas Ole.) hee don
10 20

21-
30

June
ll-
20

1
10

KKK x va
* * * a
ad ~ %X ~~ *
va ~ va

KK RRM KK KK KM ~ xX

~ xX x «KK KK ~ xX

Kr mK x KK KK ~ xX
KA KM KR KR KR KK ~ X

KKK ~ x< x «Kx

mK MK RK RRR KM
Km KKK KK KK

* x KK KKK

Kee MM KR KK KK

~ * * *

mx x Ke KR KKK KK XK

26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Poanes hobomok (Harr.) Incisalia augustus croesioides (Scudder)
Euphyes ruricola metacomet (Harr.) I. henrici (G. & R.)

Amblyscirtes hegon (Scudder) I. niphon (G. & R.)

Artogeia virginiensis (Edw.) Celastrina ebenina Clench

Euchloe olympia (Edw.) Glaucopsyche lygdamus (Doubleday)
Falcapica midea (Hbn.) Speyeria diana (Cram.)

Harkenclenus titus mopsus (Hbn.) S. aphrodite

Satyrium calanus falacer (Godt.) Euphydryas phaeton (Drury)

S. caryaevorum (McD.) Megisto cymela (Cram.)

S. liparops strigosum (Harr.) Cercyonis pegala (F.)

Speyeria cybele (F.) flies from May through September and would

appear to be multivoltine, but the long flight period is caused by stag-

gered emergence of a single brood (Opler & Krizek 1984, Scott 1986).
The following are bivoltine:

Autochon cellus (Bdv. & Leconte) Mitoura grynea (Hbn.)

Achalarus lyciades (Gey.) Nymphalis antiopa (L.)

Nastra lherminier (Latr.) Charidryas nycteis (Doubleday)
Polites coras (Cram.) Basilarchia arthemis astyanax (F.)
P. themistocles (Latr.) B. archippus (Cram.)

P. origenes (F.) Asterocampa celtis (Bdv. & Leconte)
Pompeius verna (Edw.) A. clyton (Bdv. & Leconte)
Atrytone delaware (Edw.) Enodia anthedon A. H. Clark
Poanes zabulon (Bdv. & Leconte) E. creola (Skin.)

Lycaena phleas americana (Harr.) Cyllopsis gemma (Hbn.)
Calycopis cecrops (F.) Hermeuptychia sosybius (F.)

Fresh Basilarchia archippus and B. arthemis astyanax taken in October
and early November indicate that partial third broods are produced
when mild weather persists well into fall.

Additional species are probably bivoltine, though not apparent from
our data. Erynnis horatius (Scudder & Burgess) and E. baptisae (Fbs.)
should have spring broods on the wing in April and May. They have
likely been overlooked amid large numbers of E. juvenalis flying at
that time. Pholisora catullus (F.) is also likely to have a spring brood,
and is probably more common than our records suggest. Erora laeta
(Edw.), Amblyscirtes aesculapius (F.), A. vialis (Edw.), and Staphylus
hayhurstii (Edw.) have been taken only in spring or early summer. All
four species probably have second broods in summer overlooked due
to very local occurrence.

Another group of resident species are multivoltine, with three or
more broods per year:

Epargyreus clarus (Cram.) Artogeia rapae (L:)
Ancyloxypha numitor (F.) Colias philodice Godt.
Battus philenor (L.) C. eurytheme Bdv.
Papilio polyxenes asterius Stoll Feniseca tarquinius (F.)
P. glaucus L. Strymon melinus Hbn.
P. troilus L. Everes comyntas (Godt.)

Eurytides marcellus (Cram.) Celastrina ladon (Cram.)

VOLUME 42, NUMBER 1 OF

Polygonia interrogationis (F.) V. atalanta (L.)
P. comma (Harr.) Clossiana bellona toddi (Holl.)
Vanessa virginiensis (Drury) Phyciodes tharos (Drury)

One additional species, Parrhasius m-album (Bdyv. & Leconte), is
probably multiple brooded. We have taken a worn specimen in SW
Virginia near the Tennessee line in early May, and sources indicate
that a third brood in late August-September is likely (Opler & Krizek
1984, Scott 1986).

We consider the following species to be migrants or strays:

Pyrgus communis (Grt.) E. nicippe (Cram.)

Hylephila phyleus (Drury) Libytheana bachmanii (Kirtland)
Atalopedes campestris (Bdv.) Vanessa cardui (L.)

Papilio cresphontes (Cram.) Junonia coenia (Hbn.)

Pontia protodice (Bdv. & Leconte) Euptoieta claudia (Cram.)
Phoebis sennae eubule (L.) Danaus plexippus (L.)

Eurema lisa (Bdv. & Leconte)

Most of these species overwinter in the SE coastal plain where they are
multivoltine. As their populations expand during the summer, they
move N and W,, often penetrating into the Appalachians. Although they
may reproduce during summer and fall, they generally cannot survive
winter in NE Tennessee. There are exceptions, as evidenced by an April
record for Pyrgus communis. In NE Tennessee, migrants are most likely
to be found from mid-August through October. During this period
Atalopedes campestris is one of the most common butterflies in gardens
and disturbed areas. At the other extreme, Papilio cresphontes, Pontia
protodice, and Hylephila phyleus are known from only one or two
records. Remaining species are usually present every year in varying
numbers. Libytheana bachmanii differs from the usual migrant pattern
of occurrence in that it has been found from mid-June through mid-
August. It is regularly present, but usually only as one or two individuals
at a given time and place. We include it as a migrant because we have
never collected overwintered individuals in spring.

We are not certain of the status of Thymelicus lineola (Ochs.) in NE
Tennessee. It has been taken only once, near a campground in Sullivan
Co. adjacent to a N-S interstate highway. This European species has
spread rapidly southward since it was accidently introduced into Can-
ada around 1910 (Scott 1986), and there are records from SE Kentucky
and SW Virginia (Opler 1983). If not already a resident, it is likely to
become one soon.

While walking in the late 50’s or early 60’s, the senior author saw a
single Anaea andria flying in a clover field in Sullivan Co. Without a
net he could not capture it, but followed it for a distance and was
certain of the identification. This species is resident around Center Hill

28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Lake, 130 km E of Nashville, and the junior author recently captured
several overwintered individuals in Lee Co., SW Virginia. While we
have not seen A. andria in NE Tennessee during the past 10 years, it
is somewhat migratory (Scott 1986), and should be expected on occasion.

Southwest Virginia and N Georgia have more species than NE Ten-
nessee, 120 and 108, respectively. This disparity is at least partly due
to the fact that Virginia and Georgia have been collected longer than
NE Tennessee.

Amblyscirtes aesculapius was the only species found in NE Tennessee
that has not been recorded from SW Virginia. The Clarks (1951) re-
corded it only from the coastal plain of Virginia, but there are records
from E Kentucky, and it probably occurs locally along rivers in SW
Virginia. Euchloe olympia and Clossiana bellona toddi are resident in
NE Tennessee, but are not known to occur in N Georgia. These species
are at or near the limits of their ranges in NE Tennessee.

The 39 species recorded from SW Virginia and/or N Georgia not
collected in NE Tennessee are listed in Table 2. Sixteen of these species
are known only from SW Virginia, nine from N Georgia only, and
fourteen occur in both regions.

Many species recorded from SW Virginia but not from NE Tennessee
are northern species whose ranges extend southward in the Appalachian
region. Southwest Virginia includes the entire breadth of the moun-
tainous Blue Ridge Province, and elevations in the Valley and Ridge
Province exceed 1200 m in places (Fenneman 1938). More extensive
areas of high elevation coupled with higher latitude make SW Virginia
more hospitible for some northern species than NE Tennessee.

Species recorded from N Georgia but not NE Tennessee include
Satyrium kingi (Klots & Clench), Amblyscirtes carolina (Skin.), Agrau-
lis vanillae (L.), and other species more typical of the lowland Piedmont
and Coastal Plain provinces. Relative to NE Tennessee, the Appalachian
region of N Georgia is lower in elevation and has a milder climate. In
particular, the prominent ridges that characterize the Ridge and Valley
further N are absent (Fenneman 1938). Broad valleys open onto the
Piedmont, while the oak-pine forest association and red-yellow podzolic
soils characteristic of the Piedmont extend into the Georgia portion of
the Ridge and Valley (Braun 1950, Walker 1969). These climatic and
topographic factors create favorable habitats for some lowland species,
and provide easy access for migrants.

We predict that the following species in SW Virginia and N Georgia
will eventually be found resident in NE Tennessee:

Thorybes confusis Bell Wallengrenia otho (J. E. Smith)
Erynnis martialis (Scudder) Atrytonopsis hianna (Scudder)
Hesperia metea (Scudder) Satyrium edwardsii (G. & R.)

VOLUME 42, NUMBER Il 29

TABLE 2. Butterfly species occurring in SW Virginia (VA) and N Georgia (GA) but
not recorded from NE Tennessee.

Species State
Thorybes confusis VA, GA
Erynnis martialis VA, GA
E. zarucco VA, GA
E. lucilius VA
E. persius VA
Pyrgus centaurae VA
Lerema accius VA, GA
Hesperia metea VA, GA
H. leonardus VA
H. sassacus VA
Polites mystic VA
P. vibex VA
Wallengrenia otho VA, GA
Atrytone arogos VA
Euphyes conspicua VA
E. bimacula VA
Atrytonopsis hianna VA, GA
Panoquina ocola VA, GA
Amblyscirtes carolina GA
A. alternata GA
Megathymus yuccae GA
M. harrisi GA
Zerene caesonia GA
Eurema daira VA, GA
Atlides halesus VA
Satyrium edwardsii VA, GA
S. kingi GA
Incisalia irus VA, GA
Fixenia ontario VA
Calephelis borealis VA
Agraulis vanillae GA
Charidryas gorgone GA
Speyeria idalia VA
Clossiana selene VA
Phyciodes batesii VA, GA
Polygonia progne VA
P. faunus VA, GA
Enodia portlandia GA
Satyrodes appalachia VA, GA

Incisalia irus (Godt.) Polygonia faunus (F.)
Phyciodes batesii (Reak.) Satyrodes appalachia (R. Chermock)

Hesperia leonardus (Harr.), H. sassacus (Harr.), Speyeria idalia
(Drury), and Polygonia progne (Cram.) have been recorded from bor-
dering counties in Virginia and North Carolina (Opler 1983) and also
seem likely to be found in NE Tennessee eventually.

It is possible that Amblyscirtes celia belli H. A. Freeman occurs in
NE Tennessee. We have taken it flying with Wallengrenia otho in

30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

moist woods beside an arm of Loudon Reservoir near Knoxville, Ten-
nessee, about 65 air km SW of our study area boundary. Similar habitats
should occur around reservoirs in NE Tennessee.

Additional migratory species such as Erynnis zarucco (Luc.) and
Panoquina ocola (Edw.) may eventually turn up also, but lack of direct
access from the Piedmont is a hindrance to the movement of such
species; to enter NE Tennessee, they must first pass through the rugged
North Carolina portion of the Blue Ridge, or travel a considerable
distance up valleys from Georgia.

Concentration of collecting efforts on species listed above should
increase the number of butterfly species known from NE Tennessee to
between 100 and 110.

ACKNOWLEDGMENTS

We thank P. A. Opler and an anonymous reviewer for suggesting useful changes in a
draft of this paper; also Tom Bowman of Bays Mountain Park, Sullivan Co., Tennessee,
for permitting us to collect within the Park.

LITERATURE CITED

BRAUN, L. E. 1950. Deciduous forests of eastern North America. Blakiston, Philadelphia.
596 pp.

CiarK, A. H. & L. F. CLARK. 1951. The butterflies of Virginia. Smiths. Misc. Coll. 116:
1-239.

FENNEMAN, N. M. 1938. Physiography of the eastern United States. McGraw Hill, New
York. 714 pp.

FIELD, W. D., C. F. Dos Passos & J. H. MASTERS. 1974. A bibliography of the catalogues,
lists, faunal and other papers on the butterflies of North America arranged by state
and province. Smiths. Contrib. Zool. 157. 104 pp.

Harris, L. 1972. Butterflies of Georgia. University of Oklahoma Press, Norman. 326 pp.

HopcEs, R. E. (ed.). 1983. Check list of the Lepidoptera of America north of Mexico.
E. W. Classey Ltd., London. 284 pp.

Hunt, C. B. 1967. Physiography of the United States. Freeman, San Francisco. 480 pp.

MILLER, R. A. 1974. The geologic history of Tennessee. Tenn. Dep. Cons. Div. Geol.
63 pp.

Op_LeER, P. A. 1983. County atlas of eastern United States butterflies (1840-1982). US.
Fish. Wild]. Serv. Div. Biol. Serv., Washington, D.C. 86 pp.

OpLeER, P. A. & G. O. KRIZEK. 1984. Butterflies east of the Great Plains. Johns Hopkins,
Baltimore. 294 pp.

OsBURN, W. 1895a. Rhopalocera of Tennessee. Entomol. News 6:245-248.

1895b. Rhopalocera of Tennessee—II. Entomol. News 6:281-284.

RICHARDS, A. G. 1932. Distributional studies on southeastern Rhopalocera. Bull. Brook-
lyn Entomol. Soc. 26:234-253.

ScorT, J. A. 1986. The butterflies of North America. Stanford University Press, Stanford,
California. 583 pp.

SNYDER, K. D. 1957. Checklist of insects of Great Smoky Mountains National Park.
Privately printed. 78 pp.

U.S. Dep. AGRIC. 1953a. Soil survey of Carter County, Tennessee. Series 1942, no. 4.
199 pp.

1953b. Soil survey of Sullivan County, Tennessee. Series 1944, no. 2. 199 pp.

—— 1956. Soil survey of Johnson County, Tennessee. Series 1946, no. 2. 150 pp.

1958a. Soil survey of Greene County, Tennessee. Series 1947, no. 7. 89 pp.

VOLUME 42, NUMBER 1 3]

1958b. Soil survey of Washington County, Tennessee. Series 1948, no. 5. 91 pp.

1979. Soil survey of Hawkins and Hancock counties, Tennessee. 84 pp.

1985. Soil survey of Unicoi County, Tennessee. 99 pp.

WALKER, L. C. 1969. Geography of the southern forest region. Division of Forestry,
Stephen F. Austin State University, Austin, Texas. 68 pp.

WATSON, J. R. 1946. Some August skippers of the Great Smoky Mountain National
Park and vicinity. Florida Entomol. 28:50-53.

Received for publication 2 January 1987; accepted 19 October 1987.

Journal of the Lepidopterists’ Society
42(1), 1988, 31

GENERAL NOTE

GLASSBERG, LEHMAN, AND PELLMYR COLLECTIONS
TO THE SMITHSONIAN INSTITUTION

Dr. Jeffrey S. Glassberg has donated his collection of New World butterflies to the
National Museum of Natural History (Smithsonian Institution). It consists of more than
2000 specimens, primarily Neotropical Theclinae (approximately 350 species). Dr. Glass-
berg is a molecular geneticist who lives in Chappaqua, New York, and is Vice President
for Research of Lifecodes Corp. He has a strong interest in conservation and butterfly
watching, and is currently President of the Xerces Society.

The Smithsonian Institution has received Mr. Robert Lehman’s collection of Honduran
Lepidoptera. There are 4222 meticulously spread specimens representing 1852 species,
plus about 5000 papered specimens. The Macrolepidoptera are well represented, and
there are many Pyralidae, Tortricidae, and Oecophoridae. Most of the specimens were
collected along the wet Atlantic coast of Honduras, an area that is poorly represented in
collections, and which augments the Smithsonian’s strong holdings from Mexico, Gua-
temala, Costa Rica, and Panama. Mr. Lehman has been teaching elementary school science
and, more recently, computer science, at the Mazapan School in La Ceiba, Honduras,
for 9 years, and has been collecting in Honduras since 1968.

Dr. Olle Pellmyr has donated his collection of Fennoscandian (primarily Swedish)
Lepidoptera to the Smithsonian Institution. It includes 6907 specimens of approximately
1200 species, and is rich in both Macro- and Microlepidoptera. Because so many Swedish
species are close relatives of North American ones, this collection provides important
comparative material. Dr. Pellmyr is an evolutionary biologist who works on chemical
and ecological aspects of plant-pollinator mutualism and lepidopteran courtship behavior.
He is a Swedish national, and currently a research scientist at the State University of New
York at Stony Brook.

None of the collections contains primary type specimens.

ROBERT K. ROBBINS AND GARY F. HEVEL, Department of Entomology, NHB Stop
127, Smithsonian Institution, Washington, D.C. 20560.

Journal of the Lepidopterists’ Society
42(1), 1988, 32-36

BODY WEIGHT AND WING LENGTH CHANGES IN
MINNESOTA POPULATIONS OF THE
MONARCH BUTTERFLY

WILLIAM S. HERMAN

Department of Genetics and Cell Biology, University of Minnesota,
St. Paul, Minnesota 55108-1095

ABSTRACT. Body weights and rear wing lengths were obtained from about 1900
monarch butterflies captured near Minneapolis, Minnesota, during the past decade. Mean
values for both were lowest in immigrants and highest in subsequent generations. Mean
wing length was highest in males. Mean body weights of immigrant females were higher
than those of males, but mean male body weights were higher than those of females in
subsequent generations. The data argue against the return to Minnesota of emigrants
from the previous year, and suggest that attainment of large adult size could be one
reason for monarch migration to northern regions.

Additional key words: Nymphalidae, Danaus plexippus, migration, sexual differ-
ences.

During the past several years workers in my laboratory have ex-
amined various aspects of the biology of the monarch butterfly, Danaus
plexippus L. Our studies have impressed us with the great variation
exhibited by monarch populations in our locality with respect to re-
productive status, hormone titers, behavior, and other variables (Her-
man 1985). Monarch butterflies of both sexes also exhibit predictable
changes in body weights and wing lengths during their residence in
our area, and such changes are the topic of this report.

MATERIALS AND METHODS

Animals used for this study were captured near Minneapolis, Min-
nesota, between 1976 and 1986. They were taken to the laboratory for
measurement soon after capture, usually within a few h. Whole-body
wet weights were determined to the nearest 1 mg using an analytical
balance, and rear wing maximal lengths were measured to the nearest
0.5 mm with a ruler. Immigrant butterflies rarely arrive in our locality
before 15 May, and most local monarchs emigrate by late September.
The results are therefore for animals captured 16 May to 15 September,
and data in Fig. 1 are summarized for 2-wk and 2-mo intervals during
that period. All data are presented as mean + standard error; statistical
analysis was done using Student’s t-test.

RESULTS

Mean wing lengths for both sexes were smallest during the 2-mo
period 16 May-15 July (Fig. 1). Most of these animals were presumably
immigrants from southern regions, since large numbers of monarchs

VOLUME 42, NUMBER 1 33

REAR WING LENGTH (mm)
39

$

38

ANIMAL WET WEIGHT (mg)
940
¢

l

Ce 8s ne aaal Ol 7715. 8/1 OSA

CAPTURE DATE

Fic. 1. Wing lengths and body weights of monarch butterflies captured near Min-
neapolis, Minnesota, 16 May-15 September. Data are summarized for 2-wk and 2-mo
periods. Means for 2-mo periods are shown numerically, and number of individuals given
in parentheses. Vertical lines indicate SE.

34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

do not typically emerge in our area until early July. Mean wing length
in both sexes increased significantly (P = 0.001 for both sexes) in the
2nd 2-mo period 16 July-15 September. Presumably, most of the latter
animals emerged in our locality. Mean female wing length increased
3.8% in the second 2-mo period, that of males, 3.0%. Mean male wing
lengths were significantly larger than those of females in both the Ist
(P = 0.001) and 2nd (P = 0.05) 2-mo periods. Mean wing lengths
recorded 16 July-15 September were indistinguishable from those ob-
served at emergence in monarchs reared on milkweed, Asclepias syriaca
L., in our area in July and August: 37.8 + 0.1 mm (n = 100) and 38.0 +
0.1 mm (n = 83) for females and males, respectively, on the day of
eclosion. Rear wing length varied from 29.5 to 42.0 mm in this study,
and both extremes were observed in males.

Body weights of both sexes changed in a manner similar to that of
wing lengths, with low mean values characterizing the mainly immi-
grant populations of 16 May-15 July, and significantly higher mean
values observed in monarchs that had presumably emerged in our area
16 July—15 September (Fig. 1). Mean body weights for females were
elevated 6.7% in the 2nd 2-mo period, those of males, 20.8%. Mean
female body weights were significantly larger (P = 0.001) than those
of males 16 May-15 July, principally due to higher female weights of
16 May through 15 June. Male values were significantly higher (P =
0.005) than those of females during the final 2-mo period. The lowest
mean values for both sexes were recorded in late June, when senescence
and death of immigrants is most pronounced, and the highest were
recorded in late August, when reproduction generally ceases in our
area. The increasing mean weights for both sexes from 1 July to 15
August were recorded for populations consisting principally of actively
reproducing monarchs of various ages. Mean body weights of wild-
caught butterflies never reached the mean values (680 + 32 mg [n =
26] and 652 + 11 mg [n = 109], respectively) measured on day of
eclosion for females and males reared in our area. Body weights ranged
from 195 to 836 mg during this study, and both extremes were again
found in males.

DISCUSSION

The data show that predictable variations occur in rear wing lengths
and body weights during the period that monarch butterflies reside
near Minneapolis. Small wings and low weights characterize the im-
migrant population, and both parameters increase significantly in both
sexes when monarchs that have apparently emerged in our area pre-
dominate in the local population, as they normally do after 1 July.
Causes of these variations, and their possible adaptive value, are un-

VOLUME 42, NUMBER 1 35

determined. However, the data suggest that local environmental factors
(nutrient value of foodplant, temperature, or photoperiod) during June,
July and August may provide optimal conditions for larval growth, and
thereby result in larger adults with longer wings. If so, suboptimal
conditions for larval development of the presumed immigrant gener-
ation in southern areas could account for reduced size in immigrant
butterflies. This line of reasoning implies that northward migration in
spring could be, to at least some extent, an adaptation for locating
regions that optimize adult size. Larger adults may have a greater
probability of successful southward migration, survival in the overwin-
tering colonies, or remigration.

The smaller wings of immigrants might somehow facilitate north-
ward migration, while the larger wings of animals emerging in late
August and September may be more advantageous for southward mi-
gratory flights. Perhaps larger wings are more efficient for soaring and
gliding, phenomena reported only for monarchs migrating south (Gibo
1981). Immigrant males with smaller wings might also be more suc-
cessful at mating, as reported for males in Mexican overwintering col-
onies (Van Hook 1986). James (1984) noted no significant differences
in wing lengths of Australian monarchs observed during a full year.

The data on monarch body weight generally agree with those in
other reports (Cenedella 1971, Brown & Chippendale 1974, Brower &
Glazier 1975). Other studies have reported significantly higher body
weights of males in southward migrating and overwintering monarch
populations (Tuskes & Brower 1978, Chaplin & Wells 1982). However,
others have apparently not observed periods in the monarch annual
cycle when females are significantly heavier than males, as Fig. 1
records for immigrants to our area.

Data in Fig. 1 argue against the return to our locality of monarchs
that emigrated the previous year. Our immigrants, especially females,
have intermediate weights, and, based on body weight and external
appearance, most appear to be young or middle-aged, certainly not
old. Immigrants to our area also exhibit both senescence and precipitous
weight loss (Fig. 1) within 2—4 wk after arrival, and it seems reasonable
to assume that comparable rates of aging and weight loss occur after
monarchs leave Mexican overwintering colonies. In view of these ob-
servations, it is unlikely that overwintering monarchs could leave Mex-
ican colonies in mid-March (Norman 1986), fly northward for 8-10 wk
while actively breeding, and arrive in our area with body weight and
external appearance comparable to young populations of July. Similarly,
smaller wings of our immigrants suggest they are not members of the
emigrant generation of the previous year, since emigrants have signif-
icantly larger wings. In addition, monarchs captured in Mexican col-

36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

onies in February and March 1984 had wings comparable in length to
our emigrants, and significantly larger than those of our immigrants
(Herman unpubl.). For these reasons, the data support an earlier con-
clusion (Herman 1985), and that of Malcolm et al. (1986), that most
immigrants to the northern United States are probably one generation
removed from individuals forming Mexican overwintering colonies.

ACKNOWLEDGMENTS

Some of this research was supported by the University of Minnesota Graduate School.
Ann and Alden Mikkelsen generously assisted with the capture of many animals used in
this study.

LITERATURE CITED

BROWER, L. P. & S. C. GLAZIER. 1975. Localization of heart poisons in the monarch
butterfly. Science 188:19-25.

BROWN, J. J. & G. M. CHIPPENDALE. 1974. Migration of the monarch butterfly, Danaus
plexippus: Energy sources. J. Insect Physiol. 20:1117-1130.

CHAPLIN, S. B. & P. H. WELLS. 1982. Energy reserves and metabolic expenditures of
monarch butterflies overwintering in southern California. Ecol. Entomol. 7:249-256.

CENEDELLA, R. J. 1971. The lipids of the female monarch butterfly, Danaus plexippus,
during fall migration. Insect Biochem. 1:244-247.

Giso, D. L. 1981. Altitudes attained by migrating monarch butterflies, Danaus p.
plexippus (Lepidoptera: Danaidae), as reported by glider pilots. Can. J. Zool. 59:571-
o72.

HERMAN, W. S. 1985. Hormonally mediated events in adult monarch butterflies. In
Rankin, M. A. (ed.), Migration: Mechanisms and adaptive significance. Contrib. Mar.
Sci. 27:799-815.

JAMEs, D. G. 1984. Phenology of weight, moisture and energy reserves of Australian
monarch butterflies, Danaus plexippus. Ecol. Entomol. 9:421—428.

MALCOLM, S. B., B. J. COCKRELL & L. P. BROWER. 1986. Milkweed cardenolides as
labels of the monarch’s spring remigration strategy. Abstracts from MONCON II,
Second International Conference on the Monarch Butterfly, Natural History Museum
of Los Angeles County, Los Angeles, California. P. 9.

NORMAN, C. 1986. Mexico acts to protect overwintering monarchs. Science 233:1252-
1253.

TuskEs, P. M. & L. P. BRoweErR. 1978. Overwintering ecology of the monarch butterfly,
Danaus plexippus, in California. Ecol. Entomol. 3:141-154.

VAN Hook, T. 1986. Sexual selection in monarch butterflies (Danaus plexippus L.)
overwintering in Mexico: Nonrandom mating via male choice and alternate male
mating strategies. Abstracts from MONCON II, Second International Conference on
the Monarch Butterfly, Natural History Museum of Los Angeles County, Los Angeles,
California, ‘P77,

Received for publication 4 May 1987; accepted 21 October 1987.

Journal of the Lepidopterists’ Society
42(1), 1988, 37-45

HABITAT AND RANGE OF
EUPHYDRYAS GILLETTI (NYMPHALIDAE)

ERNEST H. WILLIAMS
Department of Biology, Hamilton College, Clinton, New York 13323

ABSTRACT. Fifteen sites occupied by Euphydryas gillettii are compared according
to 10 characteristics. All sites are moist, open, mostly montane meadows, many with a
history of disturbance, commonly fire. Population size correlates with relative availability
of nectar but not with overall abundance of the usual hostplant, Lonicera involucrata.
Habitats at higher latitudes often have a southerly exposure. Reduction in hostplant size
at higher latitudes contributes to the northern range limit. Three populations likely have
become extinct since 1960, but the species range does not appear to be changing.

Additional key words: nectar, Lonicera involucrata, biogeography, extinction.

Euphydryas gillettii (Barnes), a checkerspot butterfly, occurs in dis-
crete, isolated populations (Williams et al. 1984) in the central and
northern Rocky Mountains (Ferris & Brown 1981). It is attractive and
easily caught but uncommon and not often collected. Though usually
considered a montane species (Williams et al. 1984), variation in sites
occupied by E. gillettii has not been studied, and lack of knowledge
about its habitats has led to uncertainty about its range.

Here I report characteristics of sites occupied by E. gillettii, present
range of the species, and factors influencing its distributional pattern.
This study is based on direct observation of the habitats of 15 populations
throughout the range, thus affording an uncommon view of habitat
variability in a single insect species.

METHODS

Populations of E. gillettii were located through correspondence with
collectors and researchers listed in Acknowledgments, examination of
specimen labels in collections listed in Acknowledgments, and a survey
of published reports (News Lepid. Soc., Seasonal Summaries 1960-
1986). When directions were sufficient to pinpoint locations on a to-
pographic map, I visited the sites, and assessed relative population size
and habitat characteristics.

Population size was determined by a one-day count of adults, egg
masses, and larval webs. Egg masses of E. gillettii are distinctive, easily
found, and readily counted, thus permitting quantitative comparisons
of colony size even after the flight season; in fact, egg mass counts are
better indicators of population size than adult counts because the former
are independent of weather. Eggs do not begin hatching until late in
the flight season (Williams et al. 1984), so developmental state of egg
masses at each site indicated timing of the count relative to flight season,

38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Females average one to two egg masses per individual (unpubl.); thus,
relative population size can be estimated from sum of egg masses and
adults.

‘In addition to population size, I recorded nine site characteristics,
and searched for evidence of disturbance. Observations were quantified
as much as possible for later analysis. Each site is marked on U.S.
Geological Survey and Canada Department of Energy, Mines & Re-
sources topographic maps in my possession, and latitude and elevation
were measured directly from these maps. I used a compass as well as
contour lines on the maps to determine exposure. I recorded number
of distinct shrubs or clumps of the usual hostplant, Lonicera involucrata
(Rich.) Banks (Caprifoliaceae) (Comstock 1940, Williams et al. 1984),
in open areas where egg masses and adults were found. Nectar sources
were identified (Hitchcock & Cronquist 19783), and relative nectar avail-
ability was determined by site comparison. Nearby trees were identified
and cored with a 5 mm diam increment borer for age determination.
Presence and distance to standing water and streams were recorded. I
inferred source and history of disturbance from characteristics such as
tree species and age, charring, stems gnawed by beavers, and location
in a flood plain.

RESULTS
Populations

I visited 29 localities reported as sites for Euphydryas gillettii and
found populations at 18. With my 2 previous study sites (Williams
et al. 1984), I had a total of 15 colonies throughout the geographic
distribution of the species for comparison. More than 15 egg masses
and adults were found at 7 sites (“‘large”’ populations), while fewer than
15 were found at 8 sites (“‘small” populations) (Table 1).

Habitat Characteristics

All occupied sites are wet (Table 1). Most have a small stream passing
through, though several are marshy without obvious flowing water; E.
gillettii occurs infrequently near rivers, perhaps because of flood dis-
turbance to hostplants, nectar sources, larvae, and adults. In habitat
characteristics, E. gillettii is similar to its congener E. phaeton (Drury)
(Scudder 1889). There is no observable relation between population
size and type of water present.

There appears to be a correlation between colony size and nectar
abundance (x? = 3.2, df = 1, P = 0.07). Only two sites have large
populations with low nectar availability, but these populations are mar-
ginally “large” (sites 7 & 9, Table 1). Total amount of nectar is also
important in Ewphydryas editha (Boisduval), influencing its population

VOLUME 42, NUMBER 1

39

TABLE 1. Characteristics of 15 sites occupied by Euphydryas gillettii.
Loni
Heol Nectar
Site Colony abun- availa- Nearby trees (age of largest Water
no. size! ance? bility to nearest 5 yr) (stream width) Disturbance
1 >380 >380 High Lodgepole pine (75) Stream (<1 m) Fire®
Engelmann spruce (65)
2 >380 >80 High Quaking aspen (60) Streams (<1 m) None; meadow
Subalpine fir (75) edge
3 7 10 Low Engelmann spruce (150) Stream (1-3 m) None; meadow
edge
4 2 10 Low Lodgepole pine (55) Marshy Fire; wet soil
5 4 20 Low Lodgepole pine (90) Stream (<1 m) Fire; logging
Quaking aspen (65)
6 >830 20 High Subalpine fir (155) Streams (<1 m) _ Fire’; logging
Lodgepole pine (15)
7 18 10 Low Cottonwood (40) Stream (1-3 m) Beaver activity
Lodgepole pine
Co eal 10 High Lodgepole pine (65) Stream (>5m) Flooding
9 22 20 Low Lodgepole pine (95) Marshy, Wet soil
Engelmann spruce (70) stream (<1 m)
10 8 >80 High Lodgepole pine (55) Stream (<1 m) Fire?; meadow
Engelmann spruce (50) edge
Subalpine fir (40)
die fl 5 Low. Subalpine fir (95) Marshy Fire®
_ Engelmann spruce
12 3 >80 Low Lodgepole pine Stream (1-3 m) Flooding; fire?
1s if 20 Low Engelmann spruce (195) Stream (1-3 m) Fire®
Lodgepole pine (40)
14 2 20 Low Willow (no trees) Marshy, Wet soil; graz-
stream (<1 m) ing
15 >80 5 High Lodgepole pine (75) Marshy None; meadow

edge

‘ Total number eggs and adults.
2 Approximate number Lonicera clumps in 30 x 30 m quadrat.
8 Charred tree trunks.

dynamics (Murphy et al. 1983). Nectar is supplied by a number of
genera (Table 2), mostly commonly Aster, Senecio, and Agoseris, but
each occurs conspicuously at no more than 9 of the 15 sites. Williams
et al. (1984) found the butterflies to switch nectar sources readily when
an early source senesces. Total amount of nectar thus appears more
important than particular sources.

Hostplants were considered highly abundant when there were more
than 15 distinct shrubs or clumps. In contrast to nectar availability,
hostplant abundance does not correlate directly with population size
(x2 = 0.1, df = 1, P > 0.5). Reasons are considered later.

Most sites have been disturbed (Table 1), with fire being the com-
monest natural source. Lodgepole pine, Pinus contorta Dougl., is com-

40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 2. Common nectar sources for Euphydryas gillettii at 15 study sites.

Genus Number of sites where present Genus Number of sites where present
Aster 9 Polygonum 2
Senecio 8 Antennaria It
Agoseris 7 Chrysanthemum IL
Geranium 6 Cirsium ]
Achillea 5 Geum 1
Heracleum 5 Helianthella il
Potentilla 4 Saxifraga 1
Valeriana 8 Solidago 1

mon near colonies (Table 1), indicating common disturbance history in
these areas (Pfister et al. 1977). Whatever the cause, disturbance opens
a site for growth by more hostplants and nectar sources. The few sites
not clearly showing disturbance are on edges of permanent wet mead-
ows of grasses and sedges.

At higher latitudes, occupied sites occur at lower elevations (Fig. 1,
r? = 0.49, P < 0.005). This result likely reflects colder climates and
reduced height of mountains at higher latitudes. Furthermore, impor-
tance of a minimum growing season length is shown in frequent south-
erly exposure of sites at higher latitudes, in contrast to the variable
exposure of sites at lower latitudes (Fig. 1). All large northern popu-
lations occupy sites with southern exposure, while southern sites show
no observable relation between population size and exposure. Williams
(1981) demonstrated the importance of within-habitat exposure effects;
current results suggest larger-scale influences as well.

Range

Available records of E. gillettii are mapped in Fig. 2. Sightings are
concentrated in the mountainous regions of W Wyoming, central Idaho,
NW Montana, and SW Alberta. Some regions for which there are only
older records, such as Yellowstone National Park and SW Montana,
undoubtedly support populations, but their inaccessibility makes col-
lecting sporadic. Continued existence of E. gillettii in extreme SW
Wyoming is questionable because extensive search has failed to uncover
specimens (C. F. Gillette pers. comm.). A reported record from central
Montana may be erroneous. There is also a single museum specimen
from Ontario, but improbable date as well as location suggest misla-
beling.

Sites in Alberta have smaller populations of butterflies than do those
farther south, and all northern sites have one characteristic in common:
Lonicera involucrata does not reach the large size and luxuriant growth
characteristic of Wyoming and Montana sites. In moist areas at higher

VOLUME 42, NUMBER | 4]

ELEVATION
(1000
FT- M)

42 43 44 45 46 47 48 49 50

LATITUDE (°)

Fic. 1. Elevation and latitude of fifteen Euphydryas gillettii sites. Large circles
represent “large” populations. Arrows pointing down indicate sites with southerly ex-
posure; those pointing right, easterly exposure; etc. Absence of arrow indicates site has
no obvious slope.

latitudes, willows (Salix spp.) are often taller than L. involucrata, shad-
ing them and making them less accessible to searching females; this
rarely occurs at lower latitudes. Oviposition sites are therefore scarcer
than at lower latitudes, because oviposition occurs on the highest leaves
of hostplants that are fully exposed to sunlight (Williams 1981, Williams
et al. 1984).

DISCUSSION

There appear to be four reasons for lack of correlation between
population size and abundance of L. involucrata. First, and most im-
portantly, this plant grows in moist areas regardless of amount of sun-
light, while the butterfly requires sunlit hostplants (Williams 1981). In
fact, the most luxuriant hostplants often grow in shade of conifers, but
are not used as oviposition sites. Second, an extension of the first, much
L. involucrata is over-shaded by willows at high latitudes, thus provid-
ing fewer potential oviposition sites in such areas. Third, some Euphy-

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

42

ome we
———
=
—

ALBERTA

|
|
|
Pcie pat
ee | whoa
U

en E>

SG

de
eee

wy ee

:

/

)

“AN
\

Fic. 2. Range of Euphydryas gillettii. Closed circles are sites described in this study;
open circles are locations of populations believed extinct; closed triangles are locations

VOLUME 42, NUMBER 1 43

dryas gillettii populations are mostly biennial (Williams et al. 1984),
and so may fluctuate greatly in abundance from year to year. While
most E. gillettii sites are characterized by abundant Lonicera involu-
crata, these three factors limit the size of an observed butterfly popu-
lation to less than might be expected given the total amount of Lonicera.
The fourth reason is butterfly use of alternative hostplants.

Only at one site was the colony larger than would seem possible given
the amount of nearby L. involucrata. That population (site 15, Table
1) lives where L. involucrata is uncommon, and the butterflies oviposit
extensively on two other plants, Pedicularis and another Lonicera (in
prep.). There are several possible reasons for dietary expansion in but-
terflies (Singer 1971, 1983); but whatever they may be for this popu-
lation, other study populations have not followed suit, even though all
known alternative hostplants grow throughout the Euphydryas gillettii
range. Except for site 1, where an alternative hostplant was chosen at
low frequency (less than 4% of egg masses, Williams & Bowers 1987),
I did not find eggs on or see ovipositional behavior near other plants
at the other 14 sites. Because of the known use of alternative hostplants,
I expect other E. gillettii populations use alternative hostplants as well.
The relation between population size and Lonicera involucrata abun-
dance is thus weaker than has been widely accepted.

Because its hostplants and nectar sources require wet sites, and be-
cause adults and larvae require sunlit areas for warmth, Euphydryas
gillettii most often occurs in open montane meadows. The one study
population that is not montane occupies a permanently wet, grazed
seepage area in the transition zone. Several populations were observed
along forested edges of seemingly permanent montane meadows; such
meadows may change little through time because of allelopathic in-
teractions of meadow vegetation or soil instability. More commonly,
open sites are created temporarily through disturbance. The most fre-
quent disturbance is fire, and most study sites have clearly been affected
by it. Other forms of disturbance, such as flooding, beaver activity, or
human activities like grazing and logging, also serve to open forested
areas.

Vegetational succession in disturbed areas leads to changes that make
sites less suitable through time. In particular, encroachment by sur-
rounding forest leads to greater evapotranspiration, producing a drier
site and thereby limiting growth of hostplants and nectar sources. Fur-
thermore, invasion by trees reduces the sunlight that reaches the shrub

—

where E. gillettii has been seen since 1960; open triangles are records before 1960; question
mark denotes uncertain record.

44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

and herb layer, thus eliminating warmer microsites preferred for ovi-
position (Williams 1981).

_ Life in disturbed sites suggests that E. gillettii populations are subject
to periodic extinction like E. editha (Singer & Ehrlich 1979), and such
appears to be the case. I identified with precision one site where E.
gillettii was collected in the 1960’s, but by 1983 vegetational succession
had taken place, most remaining Lonicera involucrata was shaded, and
no sign of butterflies could be found. Furthermore, human development
of recreational areas has led to loss of additional populations, one known
and one suspected.

Habitat requirements of E. gillettii, including moisture for hostplants
and nectar, and sunlight for larvae and ovipositing females, produce
the limits of its geographic distribution. Thus, plains east of the Rockies
and arid basins westward form effective biogeographic barriers to dis-
persal in either direction because of lack of water. Holdren and Ehrlich
(1981) have shown that another arid region, the Red Desert of S Wy-
oming, is the southern barrier since they successfully transplanted in-
dividuals across the barrier to central Colorado where one colony has
survived since 1977. Their transplant locales are similar to natural
habitats farther north in being wet and having an abundance of nectar
and Lonicera involucrata.

The northern range limit has been assumed to result from lower
temperatures and shorter growing season. However, all the Alberta sites
have much smaller L. involucrata, and willows dominate northern wet
sites by growing taller than other shrubs. All populations of the butterfly
at higher latitudes are smaller as well. Although no northern populations
have been found to use hostplants other than L. involucrata, alternative
hostplants used elsewhere also decline in abundance at higher latitudes.
It seems likely that competition by willows reduces size and perhaps
density of potential hostplants. Thus, fewer oviposition sites and poorer
(more shaded) ones would be found during normal hostplant searching
by females (Williams et al. 1984). I suggest that loss of oviposition sites
contributes, along with shorter growing season, to the northern limit.

Euphydryas gillettii is uacommon, but there is no evidence that its
range has been changing in recent decades. The greatest conservation
advantage this species has compared to other uncommon species is that
its habitat lies largely in mountainous areas that are not readily acces-
sible and in which there is little immediate potential for human mod-
ification. Its greatest conservation disadvantage is its occurrence through
a limited range in discrete, localized populations, which are individually
susceptible to disturbance and extinction.

VOLUME 42, NUMBER 1 45

ACKNOWLEDGMENTS

For collection records, detailed directions, maps, and correspondence, I acknowledge
contributions from the following: Karolis Bagdonas, P. J. Conway, N. S. Curtis, G. R.
DeFoliart, C. J. Durden, J. D. Eff, P. R. Ehrlich, C. D. Ferris, C. F. Gillette, L. P. Grey,
R. L. Hardesty, C. E. Holdren, F. E. Holley, W. E. Knoshaug, S. J. Kohler, N. G. Kondla,
D. D. Lawrie, J. A. Legge Jr., S. O. Mattoon, Adam Peters, F. W. Preston, J. D. Preston,
John Reichel, J. R. Slotten, R. E. Stanford, Kenneth Tidwell, W. H. Wagner Jr., and W.
D. Winter Jr. For access to collections, I thank F. H. Rindge, American Museum of
Natural History; J. D. LaFontaine, Canadian National Collection; Chen Young, Carnegie
Museum of Natural History; Richard Hoebeke, Cornell University; Leo Marnell, Glacier
National Park; R. K. Robbins, National Museum of Natural History; J. P. Donahue, Natural
History Museum of Los Angeles County; and C. A. Triplehorn, Ohio State University. I
thank T. L. Marsh, N. E. Stamp, and two anonymous reviewers for comments on a
manuscript draft, and my wife Sharon for assistance and companionship while searching
in many out-of-the-way locations. This research was funded by grants from Hamilton
College and the Brachman-Hoffman Foundation.

LITERATURE CITED

ComsTOckK, J. A. 1940. Notes on the early stages of Euphydryas gillettii Barnes. Bull.
S. Calif. Acad. Sci. 39:111-118.

FERRIS, C. D. & F. M. BROWN. 1981. Butterflies of the Rocky Mountain States. Univ.
Oklahoma Press, Norman. 442 pp.

Hitcucock, C. L. & A. CRONQUIST. 1973. Flora of the Pacific Northwest. Univ.
Washington Press, Seattle. 730 pp.

HOLDREN, C. E. & P. R. EHRLICH. 1981. Long range dispersal in checkerspot butterflies:
Transplant experiments with Euphydryas gillettii. Oecologia 50:125-129.

Murpny, D. D., A. E. LAUNER & P. R. EHRLICH. 1983. The role of adult feeding in
egg production and population dynamics of the checkerspot butterfly Euphydryas
editha. Oecologia 56:257-263.

PFISTER, R. D., B. L. KOVALCHIK, S. F. ARNO & R. C. PResBy. 1977. Forest habitat
types of Montana. U.S. Dep. Agric. Forest Service, Gen. Tech. Rep. INT-34. 178 pp.

SCUDDER, S. H. 1889. The butterflies of the eastern United States and Canada. Vol. I.
Introduction, Nymphalidae. Publ. by author, Cambridge, Massachusetts. 766 pp.

SINGER, M. C. 1971. Evolution of food plant preference in the butterfly Euphydryas
editha. Evolution 25:383-389.

1983. Determinants of multiple host use by a phytophagous insect population.
Evolution 37:389-—408.

SINGER, M. C. & P. R. EHRLICH. 1979. Population dynamics of the checkerspot butterfly
Euphydryas editha. Fortschr. Zool. 25:53-60.

WILLIAMS, E. H. 198]. Thermal influences on oviposition in the montane butterfly
Euphydryas gillettii. Oecologia 50:342-346.

WILLIAMS, E. H. & M. D. Bowers. 1987. Factors affecting host-plant use by the montane
butterfly Euphydryas gillettii (Nymphalidae). Am. Mid]. Nat. 118:153-161.

WILLIAMS, E. H., C. E. HOLDREN & P. R. EHRLICH. 1984. The life history and ecology
of Euphydryas gillettii Barnes (Nymphalidae). J. Lepid. Soc. 38:1-12.

Received for publication 27 June 1986; accepted 6 November 1987.

Journal of the Lepidopterists’ Society
42(1), 1988, 46-56

BIOLOGY OF POLYGONIA PROGNE NIGROZEPHYRUS
AND RELATED TAXA (NYMPHALIDAE)

JAMES A. SCOTT
60 Estes Street, Lakewood, Colorado 80226

ABSTRACT. The life history of Polygonia progne nigrozephyrus is compared with
that of P. gracilis zephyrus, P. faunus hylas, and P. satyrus in Colorado. Adult predator
deterrent behaviors occur: adults resemble leaves as they rest on twigs showing leaflike
undersides, roost with forewings drawn forward with antennae resting between them,
and feign death when handled. Larvae also have predator-avoidance strategies: scoli
presumably act as a physical deterrence, small larvae can drop using a silk thread, a
ventral neck gland possibly repels predators, larvae vomit on an attacker, older larvae
resemble twigs as they rest in a three-dimensional twisted-S shape, pupae resemble a
dried curled leaf or short twig. Larval host plants differ between species, with some
overlap. Identification features for the four species are presented for each stage. Despite
adult similarity of P. progne nigrozephyrus and P. gracilis zephyrus, P. g. zephyrus
larvae most resemble those of P. faunus.

Additional key words: Polygonia gracilis, P. faunus, P. satyrus, predator deterrence,
chaetotaxy.

Scott (1984) described P. progne nigrozephyrus which occurs in
Colorado-S Wyoming—Utah-SE Idaho-NE Nevada. It resembles P.
gracilis zephyrus (Edw.) on the upperside, P. p. progne (Cram.) on the
underside and in male abdominal structure, and was long confused with
zephyrus. Polygonia p. nigrozephyrus is certainly the same species as
oreas (Edw.), but some may question whether it and oreas belong to
P. progne. Early stages of nigrozephyrus are similar to those of oreas
and progne, and are distinct from zephyrus and other Polygonia; wing
undersides and abdominal structures resemble those of progne. There-
fore, nigrozephyrus does seem to be a subspecies of progne.

Since 1984, minor differences between populations of nigrozephyrus
in Colorado have been found. Adults from the E slope of the continental
divide in the Front Range usually have the dorsal hindwing darker
because the submarginal spots are the same size as those of P. g. zephyrus,
whereas adults from the W slope usually have the spots larger like those
of P. satyrus (Edw.). However, the difference is not great enough to
warrant a new name for W slope populations, and some adults from
each area resemble those from the other. The Front Range populations
may have slightly darker dorsal hindwings because of occasional im-
migration of subspecies progne, which has a very dark dorsal hindwing
margin.

An additional difference not mentioned by Scott (1984) between some
P. p. progne adults and other Polygonia, first noticed by W. H. Edwards,
involves one of the dark stripes in the ventral forewing discal cell: in
most P. p. progne the anterior stripe is unbroken, whereas in some of

VOLUME 42, NUMBER 1 47

them and in other subspecies and species the stripe is broken into two
parts.

For oviposition and larval rearing, cut host-plant sprigs were put
into water-filled vials, cotton-plugged so the water would not drain
when vials were on their sides. For older larvae, large host branches
were cut and placed in wet sand.

Adult Stage

Adults bask with wings spread (dorsal basking). In the laboratory,
some nigrozephyrus females closed the wings above the thorax and
vibrated them rapidly (up to 2 mm apart at the tips) when lights were
turned on in the morning; this is shivering behavior to raise the thorax
temperature prior to flight.

Adults of nigrozephyrus, zephyrus, and faunus (Edw.), as well as
Nymphalis milberti (God.), roost on twigs with wings closed, forewings
drawn far forward (nearly out of hindwings) and covering the head
and antennae which rest between the forewings. This posture perfects
the resemblance to a leaf on the twig by elongating the “leaf”, breaking
up its margin, and hiding antennae to avoid predation during fall,
winter, and spring. Adults frequently feign death when handled, which
would also signal a predator that the butterfly is a dead leaf.

There is evidently a circadian rhythm of oviposition, because females
laid eggs in the laboratory only during daytime, and even when lights
remained lit females begar. roosting in late afternoon. For obtaining
oviposition, fluorescent bulbs were superior to incandescent bulbs, prob-
ably because the former produce a greater and more natural amount
of ultraviolet light.

Immature Stages

Host plants. Polygonia progne nigrozephyrus feeds on gooseberry:
Ribes inerme Rydb., in Delta and Douglas counties, Colorado, R. lep-
tanthum Gray at Wiiliams Canyon, El Paso Co., Colorado. In the
laboratory, nigrozephyrus larvae accepted leaves of Ribes inerme, but
refused wax currant, R. cereum Dougl., and ate very little golden
currant, R. aureum Pursh. They ate only leaves. Additional host records
for P. p. progne, based on preserved larvae in the Smithsonian, are
gooseberry (Si. Albans, West Virginia, Monticello, New York) and cur-
rant (Centreville, Rhode Island).

Polygonia gracilis zephyrus usually eats Ribes cereum in Colorado.
However, I found an egg on R. inerme at Tinytown, Jefferson Co., on
2 June 1984, and reared it to a mature larva; and a larva under a R.
inerme leaf 5 km W Idledale, Jefferson Co., on 12 June 1984, which |
reared to an adult. In the laboratory, zephyrus larvae eat R. cereum,

48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

and do not move off its leaves to eat adjacent R. inerme leaves, although
larvae will also accept inerme. Additional host plants of zephyrus are
Ribes sanguineum (Jones 1951), and Rhododendron occidentale (larvae
reared to adults, Big Trees Park, Calaveras Co., California, 4 June 1983,
J. F. Emmel & S. O. Mattoon).

Polygonia faunus hylas (Edw.) usually eats Salix, but I found three
first-stage larvae and five eggshells under leaves of Ribes inerme at
Tinytown on 26 May 1984, and an adult emerged 20 June. In the
laboratory, fawnus larvae refused Ribes aureum leaves, but ate R. inerme
and preferred it to R. cereum.

Thus all three Polygonia will eat Ribes inerme occasionally.

The only known Colorado host of P. satyrus is Urtica dioica gracilis
(Ait.) Sel., though Humulus lupulus L. is eaten elsewhere. In the lab-
oratory, satyrus larvae accepted Humulus and Urtica leaves equally
well.

Life Cycle

Five larval instars have the following approximate head widths, re-
spectively: 0.4, 0.7, 1.2, 1.7, 2.6 mm. Stage | is easily recognized by its
black head without scoli; stage 2 has head scoli but is still black; stage
3 has head scoli but is black usually with an ochre pattern tending
toward the pattern of stages 4-5. Usual laboratory durations of nigro-
zephyrus stages at 19°C were: egg, 5-6 days; larval stages, 3, 2.5, 2, 2,
4 days, respectively; and pupa, 9-10 days; totalling 27-30 days. In the
cooler and more variable temperatures of nature, these periods are
probably nearly doubled, so that adults should appear by late July—
early August, although eggs laid in late April might produce the few
fresh late-June adults known in nature. A faunus stage 1 larva found
26 May emerged as an adult 20 June in the laboratory, even though
faunus emerges in nature only in late July and August. The laboratory
life cycle of P. p. progne is 31-32 days (Edwards 1880), of P. inter-
rogationis 28-40 days (Edwards 1882b), and of P. comma 27-88 days
(Edwards 1882a). Thus all Polygonia have similar developmental rates
indoors, and all have five larval stages. However, in Colorado P. faunus
and P. progne nigrozephyrus have only one generation per year, while
P. satyrus and P. gracilis zephyrus have two generations at low altitude
and one at high altitude; and P. interrogationis has two or three gen-
erations.

Predator-Avoidance Structures and Behavior

Stinkbugs and ants were found on R. inerme host plants and may
prey on immatures.
The scoli of stage 2-5 larvae presumably physically deter predators.

VOLUME 42, NUMBER 1 49

They slightly hurt the human skin when touched, evidently a physical
puncturing rather than an urticating chemical.

A ventral neck gland occurs on stage 2-5 larvae of all 4 Polygonia
species; it contains 2 internal transverse dark secretory pads which
perhaps produce repellent chemicals.

When grasped, the larva often bends its head arcund and vomits
green fluid onto the attacker.

Fourth- and fifth-stage larvae of nigrozephyrus grasp a twig with
the prolegs, bend the front part of the body right or left, and raise the
end of the abdomen. This “corkscrew” posture may make the larva
resemble a dead leaf or twisted twig, perhaps lessening predation by
birds. This posture also occurs in ssp. progne (Edwards 1880) and in
satyrus (C. F. Gillette pers. comm.).

Young larvae of all four species rest on the underside of a leaf, and
when older may also rest on a twig. Only older larvae of P. satyrus,
also P. comma, live in a nest. It is made by chewing the base of the
leaf on each side, thus making it droop, and silking Urtica leaf edges
down and together below the enclosed larva, which rests on the leaf
underside.

Disturbed young larvae can extrude a silk thread as they fall, then
crawl up the thread to return to the plant.

Pupae are constricted in the middle where silver spots also visually
break up the outline, making the pupa resemble a dead, shriveled leaf
or twig.

Gooseberry hosts are armed with sharp spines which act as physical
protection against vertebrates. A punctured pupa recovered completely.

Descriptions of Early Stages

Colors are based on live individuals. Immatures have been deposited in the Smithsonian
Institution. Many dozen individuals of Polygonia p. nigrozephyrus were reared from eggs
laid by females from NE of Cedaredge, Delta Co., and Nighthawk, Douglas Co. Each
stage is described, and is followed by comparisons with the other three Polygonia species
and subspecies, each of which were represented by less than 10 individuals. Segments
are named T1 for prothorax, A8 for abdominal segment 3, etc. (Fig. 3). Scoli are named
with the letter B followed by name of nearest primary seta. They are not preceded by S
because of confusion with primary seta SD1, etc.; sp is spiracle; VNG is ventral neck
gland on older larvae. Names of setae are from Hinton (1946) and Scott (1986), with
slight modifications (Scott 1988) that improve homology and make head and body setal
nomenclature different to avoid confusion.

Egg. Green, averaging 8.6 vertical ribs (Table 1), each rib steep-walled, increasing in
height to maximum at top, then disappearing; 40-50 horizontal ribs forming ladder
between adjacent vertical ribs; the day before hatching turning blackish with transparent
silvery-reflecting shell as larva becomes partly visible.

Comparison. All Polygonia eggs green. Polygonia p. progne has 8-9 ribs, P. g. zephyrus
averages 9.8, other Polygonia average 10.4-11.5 (Table 1).

First-stage larva (Figs. 1, 4, 5, 9, 11, 12, 16). Head black without pattern or horns
Body dark brown with long black setae, bumplike bases of which are chitin brown; with

50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Number of vertical ribs on eggs.

Taxon Mean SD Range N Source
Polygonia p. nigrozephyrus 8.6 0.55 8-10 40 _ this paper
P. p. progne — — 8-9 — Edwards (1880)
P. g. zephyrus 9.8 0.59 9-11 27 this paper
P. satyrus 10.4 0.54 10-12 43 this paper
P. faunus 10.5 0.63 10-12 31 __ this paper
P. interrogationis — — 8-10 — Edwards (1882b),

Pyle (1981)

P. comma (Colo.) IS 0.50 11-12 58 _ this paper
P. comma (Minn.) 10.6 0.54 10-12 388 __ this paper
Nymphalis vau-album (D. & S.) 11.0 — 9-12 — CF Gillette

pers. comm.

cream spots (Fig. 1) as follows: front half of Tl cream except for small supralateral brown
patch on some larvae; Tl cream in front of, behind, just beside black prothoracic shield;
rest of Tl brown except for 2 cream dashes extending rearward above, below spiracle.
T2-3 brown, large yellow-cream patch around D2, smaller cream patch around L1-2.
A2, A4, A6 brown, with 4 pale patches: broad cream mid-dorsal V aimed posteriorly on
anterior part of each segment; broad yellow-cream patch below D1; narrow supralateral
light brown dash; long cream sublateral dash. Al, A3, A5, A7 brown, with 4 light brown
patches on each side corresponding to pale patches on A2, A4, A6; sublateral dash cream
on A3, A5, A7, A8. A8 same as A7 but 3 upper patches slightly creamier. A9 brown,
subdorsal cream patch twice as long vertically as horizontally. A10 brown, suranal plate
black, proleg cream, proleg plate brown, large circular cream supralateral patch.

Comparison. Other Polygonia larvae very similar, with black hornless head and similar
body pattern. Polygonia satyrus same as nigrozephyrus, pale bumps cream-white, a few
creamy sublateral dashes. Polygonia g. zephyrus same as nigrozephyrus, except pale
bumps cream-white instead of yellow-white, seta D1 on T3 on whiter bump as is seta
D2, no supralateral brown patch on front of Tl though it appears on some second-stage
larvae so may be individual trait, supralateral dash on A2, A4, A6 cream, Al, A8, A5,
A7, A8 all brown except for lateral cream dash. Polygonia interrogationis similar (Ed-
wards 1882b), but P. comma “whitish-green” (Edwards 1882a). Polygonia faunus larvae
differ from all other Polygonia in having white areas expanded away from bumps: for
instance, white patch on T2, T3 includes both D1, D2 setae; on A2, A4, A6 white V
lengthened anteriorly, subdorsal white patches below D1 extend posteriorly.

Second-stage larva (Figs. 2, 6-8, 17). Head black with 2 short black spiny horns (BPA2
scoli) each with 1 long seta on tip, 5 setae on crown just below, no setae on long stalks;
bases of PA1, AG3, LH1, O2 pale, membranous; very narrow short pale line along
middorsal groove. Body reddish brown, brownish orange toward rear, similar to 1st stage
in pattern, prothorax mostly orangish yellow; orange V’s on top of A2, A4, A6, yellow-
cream areas of first stage now orange, scoli present with bases orangish. Scoli BD2 on A2,
A4, A6 ochre on some larvae, mostly brown on most, other scoli black. BD2 scoli on T2,
T3, A2, A4, A6 rest on large orange bumps making segments conspicuously paler, other
scoli rest on small orangish bumps. Body has weak cream mid-dorsal, subdorsal spots
which help form abdominal V’s; remaining segments have thin wavy lateral cream line
between BL1 scoli, thin wavy supralateral cream line between BSD1 scoli. Tiny pale
subdorsal transverse dashes present. Ventral neck gland present.

Comparison. Polygonia g. zephyrus has slightly shorter horns, body undergoes less
color change from first stage: color pattern the same, pale patches still white, though BD2
on A2, A4, A6 yellow-cream, in some larvae blackish, making segments still paler on top,
other scoli black. Only BD2 on T2-8, A2, A4, A6 rest on yellow-cream bumps; other scoli
rest on small whitish bumps. Tiny cream transverse dashes occur behind, before BD2 on
A2, A4, A6 to help form V’s as in P. satyrus; middorsal, subdorsal, supralateral, lateral

VOLUME 42, NUMBER 1 51

D1 D2

XD1 aN FI RST a STAGE vt ih DL

iJ

S)

I

ACVAS TEA TAST AG Ar “AG AS AiO

Fics. 1-8. Setal maps of Polygonia progne nigrozephyrus larvae. 1, First stage. Color
pattern shown on some segments, except that plates at base of setae, including prothoracic
shield and suranal plate, are dark brown; T2 and T8 patterns similar; Al, A8, A5, A7
patterns similar; A2, A4, A6, A8 patterns similar except that A8 darker; 2, Second stage;
3, Fifth (mature) stage. L inside circle is true leg; P inside circle is proleg; S inside circle
is scolus. Hundreds of small setae not shown. See text for further explanation.

white spots present. Polygonia satyrus resembles |st-stage zephyrus, thus head black, T1
mostly white except for black prothoracic shield. BD2 on A2, A4, A6 also yellow-cream;
other scoli black, except BD1 on A6 whitish, BD1 on A4 partly whitish, BL1 on A4, A6,
A7, A8 mostly white. BD2 on T2, T3, A2, A4, A6 rest on large yellow-cream bumps;
other scoli rest on small tan hills, though BD1 on A2, A4, A6, BL1’s rest on fairly white
bumps. Polygonia faunus has enlarged white areas compared to other species, on at least
1 larva BD1 and BD2 on A2, A4, A6 pale. Ventral neck gland occurs in all 3 Polygonia.

Third-stage larva (Figs. 8, 13, 18). Head black with black scoli, following structures
ochre: Mid-dorsal notch, adfrontal cleavage line (lateral to frontoclypeus), lower fronto-
clypeus, head just above antennae, bases of all major setae except black horns; but some
individuals have head mostly black, nearly devoid of pattern. Head setae AG3, PA], LH1,
O2 on long ochre stalks. Body dark brown, with long mostly orange scoli: BL1, BSD1
mostly black; BD2 mostly orange; BD2 on T3, A2, A4, A6 strongly orange; scoli on T2,
A10 mostly black. Body pattern similar to stages 4—5.

Comparison. Polygonia g. zephyrus larvae have BD1, BD2 more whitish cream on
abdomen. Polygonia satyrus differs greatly: head black with cream notch on top running

52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

iA OSS

Fics. 4-15. Leg and cranial setae of Polygonia progne nigrozephyrus. 4, Ventral-
medial view of first-stage larval thoracic leg, showing setae typical of butterflies; 5, Setae
and olfactory pores of first-stage larval head; 6, Head of second-stage larva. Head horn
derived from, or incorporates, seta PA2. Setae PA], AG8, LH1, O2 arise from small cones
on transparent circles of exoskeleton. X’s show positions of setae present on some larvae;
7, Head horn of second-stage larva, includes PA2 seta of first stage; 8, Ventral neck gland
of larval stages 2-5 partly everted. It appears slitlike when retracted, is fully everted in
some preserved larvae; 9, Mandible of first-stage larva; 10, Head setae of fifth-stage
(mature) larva with primary setae whose origin is traceable to first-stage larval seta lettered;
11, Labrum of first-stage larval head, anterior view showing one olfactory pore; 12,
Labrum of first-stage larval head, posterior view showing two olfactory pores, three
spatulate setae; 13, Labrum of third-stage larval head, anterior view; 14, Labrum of
fourth-stage larval head, anterior view; 15, Labrum of fifth-stage (mature) larval head,
anterior view.

forward to inverted cream V on face, head horns, setae mostly black, some setae on sides
and lower face white; body has lateral cream band with cream BL1; top of body cream
with cream scoli, black dashes in shape of V without point angling forward from each
BD1.

Fourth-stage larva (Figs. 8, 14). Head as in mature larva. Body similar to mature
larva, but scoli more orangish, BD1, BD2 on A8, A5, A7 with dark brown ring around
each above base, whereas other scoli and all scoli on mature larva, lack brown ring.

Comparison. The other species also resemble mature larva.

Mature larva (Figs. 3, 8, 10, 15, 19, 20, color photo on pl. 3 of Scott 1986). Head
black, horns dark brown, orangish cream notch on top, orange-red W on front consisting
of streak along upper part of each adfrontal cleavage line plus streak angling down from
base of each horn, lower 3rd of frontoclypeus orange-brown, orange-red patch surrounding
eye cluster, orangish mottling beside neck. Some setae everywhere on head including
AG3, PA1, LH1, O2 orange-red, on long orange stalks; AG2, some dorsal setae beside
neck, about 3 lateral setae beside neck on smaller orange stalks. Body scoli ochre, only
needle tips orange, except: BD2, BSD1 on T2 black with some orange branches; BD2 on
T3 mostly black, orangish on basal 5th, BSD1 ochre; BD2 on A8 partly black, BSD1
mostly black, BD1, BL1 ochre; BD2 on AQ partly black; BD2 on A10 black. Body blackish
brown in ground color, with complex pattern. Tl brown with mid-dorsal, subdorsal,
supraspiracular, subspiracular orangish lines, some small mostly orange spinelike setae;
mid-dorsal ochre band extending from head to T1, narrowing on T2, very narrow on TS.
A few ochre transverse dorsal lines between T1, T2, between A8-10. Body joints between

VOLUME 42, NUMBER 1 53

T2, A8 have 5 ochre joint lines, line 2 grayish, lines 1, 3 widest, separated by 4 black
joint lines, most posterior very narrow. Segments T2, T3 ochre on top, with paired short
black grooves on either side of black mid-dorsal line. Segments Al, A2 similar but paired
dark grooves form brown transverse streak behind BD1. A1, especially A2, begin to show
dorsal black rearward-aimed V’s characteristic of all Polygonia on A3-8. Tip of V blunt,
wide, corresponding to brown transverse streak on Al-—2 just behind BD1, each arm of
V thickest in middle anterodorsal to BD2 where V becomes orangish black, outlined by
ochre bands as thick as V itself. Three more black spots posterior to point of each V that
continue point: black transverse mid-dorsal dash formed by 2 interruptions in 1st black
joint line circling segment, narrower dash formed by narrower interruptions in next joint
line, mid-dorsal black triangular spot on anterior edge of posterior segment. Ochre joint
lines stop at 2 wavy lateral lines characteristic of all Polygonia. Upper wavy lateral line
orange, on each segment obliquely extending from BSD1, which is ochre with orange
base, up, forward then down; behind BSD1 obliquely extending down, backward then
down, forward, resembling orange staple aimed down, forward, centered on BSD1. Upper
line interrupted between segments by last 3 ochre joint lines which splinter into about 5
ochre wavy narrow lines that stop just above lower wavy lateral line. Lower wavy lateral
line ochre, extending from each BL1 obliquely up, forward, then straight forward, then
angling down toward BL1 of preceding segment. Beneath this line a vague ochre line
above prolegs. Prolegs, underside blackish brown, ochre ventral bands running along
abdomen on each side of mid-ventral line. Ventral neck gland present.

Of more than 50 larvae, a few slightly paler (dorsal areas yellow anteriorly, cream
behind). Early 5th stage slightly more pinkish violet as orange-red scoli of 4th stage change
to ochre.

Comparison. California P. p. oreas, based on preserved larvae, photos, same as ni-
grozephyrus, except that top front of former oranger, yellowish orange vs. orangish yellow
on top of thorax, Al-2; BSD1 on orange upper wavy lateral band more orangish than
nigrozephyrus, ochre in latter with only base orangish. Based on 50-year-old preserved
larvae in Smithsonian, ssp. progne similar to nigrozephyrus in structures, all pattern
elements seem present, though impossibie to discern true colors; dorsal V-marks, transverse
lines between segments present. Edwards (1880) described progne color as buff (ochre),
dorsal area “reddish” (probably orangish ochre) around black V’s; he described T2-3,
A9-10 scoli as black, others ochre as in nigrozephyrus; described BSD1 as black, but
contradicted on the preserved larvae, these being pale also. T1 collar described as yellow
in progne, and is pale in the preserved larvae, whereas it is black except for mid-dorsal
line in nigrozephyrus, other 3 species. Head seems to have larger black areas in nigro-
zephyrus than ssp. progne. Evidently ssp. progne larva does not change color from front
to rear as much as western subspecies, and dark brown areas of former are smaller.

Mature larvae of other Polygonia species differ greatly. All 3 have black V's on top of
abdomen slightly narrower than nigrozephyrus, point of each V less strongly connected.
All 3 have wavy lower lateral lines as in nigrozephyrus, but these are slightly reddish
cream in zephyrus, red-orange in faunus, orangish cream in satyrus. Polygonia g. zephyrus
(Fig. 27, color photo on pl. 2 of Scott 1986) much more 2-toned, top of segments T2-3,
Al-2 red-orange, especially T3, A2); top of A3-8 whitish, especially A4, A6 which are
yellowish white. Basic pattern elements of zephyrus same as in nigrozephyrus, but wavy
lateral lines weak, slightly reddish, scoli black except BL1 along lower wavy line whitish
in some larvae, BD1 orange within orange areas, white within white areas. Head of
zephyrus also mostly black, except for white mid-dorsal notch, sometimes thin orange
inverted V on front, scattered small white seta bases. Some zephyrus larvae have T3, A2
orangest on top, A4, A6 whitest, whereas in others T2-3, Al-2 equally red-orange on
top, A3-8 equally white on top. Latter characteristic of P. fawnus, which has top of body
orange in front, white behind as in photo 14 of faunus (=silvius) in Pyle (1981). Polygonia
faunus has both wavy lateral lines red-orange, BL1 on lower line white, head black with
some cream setae, cream dorsal notch, orange W on front. Thus mature larvae of P. g.
zephyrus, P. faunus are similar. Polygonia c-album L. larvae resemble faunus closely
(photos in Pyle 1981, Whalley 1979:19, Brooks & Knight 1985:79).

Polygonia satyrus mature larvae differ greatly from other Polygonia (Fig. 25, color

54 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

photo on pl. 2 of Scott 1986). Top near-uniform yellow, same pattern elements present:
head black, inverted cream V on front, mid-dorsal cream notch, some small cream setae;
middorsal line cream on thorax, transverse rings between segments, dorsal V’s present.
However, entire top of body greenish yellow, T2-3, Al-2 ochre-yellow in some larvae,
lower wavy line thick, pale yellow, orangish between segments in some larvae, yellow
BL1’s; upper wavy line nearly absent, with black BSD1’s or line thin, orange, with cream
BSD1’s in some larvae.

Polygonia interrogationis (Fab.) mature larvae are also very different from other
Polygonia (Pyle 1981:photo 15, Edwards 1882b). Polygonia comma (Harr.) mature larvae
vary (Edwards 1882a).

Ventral neck gland present in stages 2-5 of P. g. zephyrus, P. faunus, P. satyrus.

Chaetotaxy (Figs. 1-7, 9-15). Head of lst-stage larva has only primary setae. Second-
stage head has many secondary setae, scoli BPA2; each horn incorporates seta PA2 of Ist
stage because scolus in same position as lst-stage PA2, other dorsal primary setae rec-
ognizable on 2nd-stage head by large size, position. Each 2nd-stage horn has long PA2
seta on tip, 5 setae on crown below tip. Head setae, horns on stages 3-5 like those of stage
2 except for proliferation of small setae, primary setae recognizable on mature larval
head by larger size, horn still including only 1 primary seta, PA2. Setae on labrum constant
at 6 on each side, 3 spatulate setae on posterior oral surface, during larval stages 1-5,
setae on other mouthparts also constant, except mandible setae which rise from 2 on each
stage 1 mandible to about 10 on stage 5.

Proprioceptor setae, those that detect cuticular folds telescoping over, on head, body
same as present in other Lepidoptera.

Body of Ist-stage larvae has mainly primary setae, also some secondary L2 setae, present
on all individuals examined, on T3, Al—8; on A3-6 of some larvae 4th L seta present
near L2. On 2nd-stage body, many secondary setae, scoli appear. Body scoli of 2nd-stage
larva not homologous with Ist-stage larval primary setae because primary setae of Ist
stage occur on 2nd stage, sometimes slightly moved in position, with scoli. Thus A10 of
lst stage has paler spot where BSD1 appears on 2nd stage, yet both stages have same
dorsal primary setae on A10; on T2-3, 2nd stage retains same SD, L setae of stage 1 adds
BLI1; on A8, 2nd stage retains same D1-2, SD1 setae of stage 1, adds BD1, BD2. Body
scoli add small setae between stages 2-5, otherwise change little. Small SD plate on T2-
3 of stage 2 disappears, only 1 or 2 setae remain on stage 5. Body setae multiply between
stages, hundreds of which are not shown on stage 5 setal map (Fig. 3). Crochets typical
of butterflies: 14 of anterior 8 prolegs forming circle in stage 1, medial crescent in mature
larvae; 12 anal crochets form anteromedial crescent in all stages. Each true leg has 5, 2,
6, 2 tactile setae plus 3, 1, 0, 2 proprioceptor setae on Ist 4 leg segments of stage 1, the
usual number in lst-stage butterflies, additional setae joining these on mature larvae, Ist
segment having about 8 setae, for instance, on mature larvae. No anal comb present on
any stage.

Comparison. Setae, scoli of all larval stages same in 4 Polygonia compared, also in
mature P. interrogationis larvae based on preserved specimens, Petersen (1965) showing
drawing of mature interrogationis larva: thus secondary Ist-stage L2 seta occurs in all
species, L1 on A3-6 in some zephyrus individuals splitting into 3 instead of 2 setae,
making 4 L’s instead of the normal 8, head horn on stages 2-5 incorporating primary
seta PA2, consisting of 1 terminal setae, crown of 5 main setae below. Secondary L2 seta
on |st-stage T3-A8 distinguishes Polygonia from Nymphalis, Vanessa.

Pupa (Figs. 21-24, color photo on pl. 5 of Scott 1986). Usually pinkish tan, sometimes
paler, rarely blackish gray. Segments T3, Al, A2 have silver or gold subdorsal spot, usually
silver on T3, Al, often gold on A2 because of reddish tan A2 top, making 6 in all, mid-
dorsal silver streak sometimes on Al. Segment A2, to lesser extent A3, reddish tan on top.
Four abdominal bands: lateral tan-edged brown band, mid-ventral tan-edged brown
band, mid-dorsal brown-edged tan line. Basal half of each tibia brown. Sliver of hindwing
just above forewing brown. Light-brown V’s on A4-7, weakly on A8, on both sides of tan
mid-dorsal line, 1 arm of each V ending at each subdorsal cone. Broad brown, often
greenish brown, band crosses wing from tornus to mid-costa, short brown subapical band
parallel to it. Many cones, bumps usually at larval scoli positions: very small mid-dorsal

VOLUME 42, NUMBER 1 ap

Fics. 16-28. Polygonia larvae and pupae. 16-24 P. progne nigrozephyrus from Delta
Co., Colorado; 25-28 other taxa as noted from Jefferson Co., Colorado. 16, First-stage
larva, dorsal view; 17, Second-stage larva, dorsolateral view; 18, Third-stage larva, dorsal
view; 19, Fifth-stage larva, dorsal view; 20, Fifth-stage larva, lateral view; 21, Pupa,
dorsal view; 22, Pupa, lateral view; 23, 24, Pupae, lateral views showing variation; 25,
P. satyrus mature larva, lateral view; 26, P. satyrus pupa, lateral view; 27, P. gracilis
zephyrus mature larva, dorsal view; 28, P. faunus hylas pupa, lateral view.

bump on A2-8; large subdorsal cone on T2-3, Al—8; supralateral bump on A3-7; lateral
bump on A4-8, lateral bump on each head horn; large bump on wing base; bump on
lower basal corner of wing; subventral bump on A5-6, another on head, 1 on each tibia;
2 stout cones (horns) projecting forward from each side of head; mid-dorsal keel on T2.

Silk pad spun by pupating larva bright pink.

Comparison. All Polygonia pupae have similar silver or gold spots in saddle, similar
cones, keels, horns, dark bands on abdomen, wings. Species differ in overall color, shape,
size of cones, horns. Polygonia p. oreas resembles nigrozephyrus, but 2 oreas pupae seen
were brown, not pinkish tan. Polygonia p. progne pupa (Edwards 1880) also pinkish
brown like nigrozephyrus, with similar markings; head, thorax sometimes greenish brown.
Polygonia g. zephyrus like nigrozephyrus in shape, but most individuals light brown,
some creamy gray or tinged with green, rarely blackish gray, abdomen more mottled,
subdorsal area on A4 lighter than on other segments, on A5—A7 a paler streak angling
forward, down from each subdorsal cone. Few zephyrus pupae resemble nigrozephyrus
in overall color, yet reddish tan top of A2 of nigrozephyrus identifies most. P. faunus
pupa (Fig. 28, color photo 14 of Pyle 1981, as silvius) light brown (often with reddish

56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

flush on top of A2-8 as in nigrozephyrus) or dark gray, easily identified by elongate shape,
>10% longer, long head horns, twice as long as other Polygonia. P. satyrus pupa (Fig.
26, color photo on pl. 5 of Scott 1986) paler, tan or straw, sometimes yellowish dorsally,
rarely brown all over, easily identified by mid-dorsal T2 keel being twice as high as other
species, subdorsal abdomen cones about twice as large. P. interrogationis similar in color
to some nigrozephyrus, faunus, with similar sized bumps, but its T2 keel very large (color
photo 15 of Pyle 1981, Edwards 1882b). P. comma pupa quite variable (Edwards 1882a).
Polygonia c-album pupa brown, resembling nigrozephyrus in shape but T2 keel larger
as in satyrus (Brooks & Knight 1985:79).

Oddly, silk cremaster pad spun by pupating larvae colored differently in other species:
bright pink in nigrozephyrus, also interrogationis (photo in Pyle 1981); pale pink in
zephyrus, faunus; yellowish white, rarely faintly pink, in satyrus.

ACKNOWLEDGMENTS

I thank J. R. Heitzman for larvae of P. interrogationis, R. K. Robbins for loans of
Polygonia immatures from the Smithsonian Institution, P. A. Opler for photos of P.
progne oreas immatures, and C. F. Gillette for reviewing the manuscript.

LITERATURE CITED

Brooks, M. & C. KNIGHT. 1985. A complete pocket guide to British butterflies. Jonathan
Cape, London. 159 pp.

EDWARDS, W. H. 1880. Description of the preparatory stages of Grapta progne. Can.
Entomol. 12:9-16.

1882a. Description of the preparatory stages of Grapta comma. Can. Entomol.

14:189-196.

1882b. Description of the preparatory stages of Grapta interrogationis. Can.
Entomol. 14:200-208.

HINTON, H. E. 1946. On the homology and nomenclature of the setae of lepidopterous
larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Roy. Entomol.
Soc. London 97:1-37.

JONEs, L. 1951. An annotated checklist of the Macrolepidoptera of British Columbia.
Entomol. Soc. British Columbia Occ. Pap. 1:1-148.

PETERSEN, A. 1965. Larvae of insects. Part I. Lepidoptera and plant-infesting Hyme-
noptera. Publ. by author, Columbus, Ohio. 315 pp.

PyLE, R. M. 1981. The Audubon Society field guide to North American butterflies.
Knopf, New York. 916 pp.

ScCoTT, J. A. 1984. A review of Polygonia progne (oreas) and P. gracilis (zephyrus)
(Nymphalidae), including a new subspecies from the southern Rocky Mountains. J.
Res. Lepid. 23:197-210.

1986. The butterflies of North America. A natural history and field guide.

Stanford Univ. Press, Stanford, California. 583 pp.

1988. The small forest: Chaetotaxy of first-stage butterfly larvae. In preparation.

WHALLEY, P. 1979. Butterflies. Hamlyn nature guides. Hamlyn Pub. Group, London.
128 pp.

Received for publication 22 April 1987; accepted 2 October 1987.

Journal of the Lepidopterists’ Society
42(1), 1988, 57-58

GENERAL NOTE

DIFFERING OVIPOSITION AND LARVAL FEEDING STRATEGIES IN
TWO COLOTIS BUTTERFLIES SHARING THE SAME FOOD PLANT

Additional key words: Pieridae, Colotis amatus, C. vestalis, eggs, Salvadoraceae.

There is much interest in the habit of certain butterfly species laying eggs in clusters.
It is generally agreed that cluster-laying is a derived trait, the ancestral butterfly having
laid single eggs. Cluster-laying has evolved independently several times in all butterfly
families. Its significance has been subject to a variety of interpretations. The purpose of
this paper is to present oviposition data for two closely related species of Colotis in New
Delhi, India.

The species in question are Colotis amatus F., whose geographic distribution covers
most of Africa, Arabia, India, and Sri Lanka; and C. vestalis Butler, found in NW India,
Pakistan, and East Africa, but unaccountably absent from Arabia (Larsen, T. B. 1983,
Fauna of Saudi Arabia 5:333-478). Together with C. phisadia Godart, C. amatus and C.
vestalis form a small section of the genus that feed on Salvadoraceae rather than on the
more usual Capparidaceae.

In New Delhi both butterflies feed on Salvadora persica L. and S. oleoides Decaisne.
Usually both are phenologically synchronous, and occur on the same trees or bushes. In
size and behavior they are very similar and were not the ground colours salmon and
white, respectively, they would be difficult to tell apart on the wing. M. A. Wynther-
Blyth (1957, Butterflies of the Indian Region, Bombay Natural History Society, Bombay,
523 pp.) even suggests they interbreed, interspecific copula having been observed.

Given the overall similarity, the difference in oviposition behavior is startling. Colotis
amatus lays clusters averaging ca. 30 eggs on upper surfaces of fresh leaves at outer
extremities of the host plant (Table 1). Eggs are evenly spaced within each clutch. Colotis
vestalis lays single eggs deep inside the host plant, usually on a twig or a branch, rarely
on an old leaf. I observed eggs being laid as far as 90 cm from the nearest leaf, a
considerable distance for a small, freshly hatched larva to travel. Larvae of C. amatus
feed gregariously on fresh foliage, but group cohesion weakens in final instars. Those of
C. vestalis feed singly on old leaves, usually deep inside the bush or tree. I never found
both species on the same leaf.

The egg of C. vestalis is chalk white with 20-22 keels extending from the micropyle
to the base. It is covered in fine hairs, best visible when the egg is submerged in fluid.
Egg volume appeared 15-20% greater than that of C. amatus. The latter’s eggs are yellow,
have only 14-16 keels, lack hairs, and unlike those of C. vestalis are covered with a sticky
substance. Midges and mosquitoes were often found trapped on egg clutches.

S. Courtney (1984, Am. Nat. 123:276-281) mentions that Aporia crataegi L. in Morocco
may adjust egg-clutch size to food plant quality. The data are given in more detail by S.
Courtney (1986, Adv. Ecol. Res. 15:51-131). Colotis amatus clutch-size on the broad-
leaved Salvadora persica averaged 28.7 eggs (n = 106), and on the narrow-leaved S.
oleoides, 22.7 (n = 17) in my Delhi sample; the difference is not statistically significant.

Although these two common butterflies are synchronous and share foodplants, they
seem to be noncompetitive. I never saw complete defoliation of food plants. There are a
number of potential pathways for two such butterflies to evolve different ovipositing
strategies, but data to support any specific hypothesis are not available. Probably no single
causal factor underlies all egg clustering. However, available data do not support the
hypothesis of R. A. Fisher (1930, The genetical theory of natural selection, Clarendon
Press, Oxford, 272 pp.) that egg clustering leads to aposematism; if anything C. vestalis,
which feeds on old leaves, should be the more aposematic of the two. I masticated a
number of specimens without finding the least pungency or emetic response, although I
found other aposematic butterflies emetic (Larsen, T. B. 1983, Entomol. Ree. J. Var. 95:
66-67).

The closest parallel I have seen to the two Colotis species is that of Eurema hecabe L.
and E. blanda Boisduval in Papua New Guinea and S India. The former lays single eggs,

58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Number of eggs in 123 clutches laid in the wild by Colotis amatus in New
Delhi, India (autumn 1986).

No. eggs in clutch No. clutches No. eggs in clutch No. clutches

1-5 0 41-45 3)
6-10 2 46-50 3
11-15 10 51-55 0
16-20 24 56-60 2
21-25 25 61-65 il
26-30 19 66-70 1
31-35 17 71-75 1
36-40 13 76+ 0

Average 27.9 eggs per clutch.

the latter clutches. However, in both places the two show more ecological and spatial
segregation than Colotis; they can feed on the same plants but usually do not do so in
the same locality. In Yemen I noticed that Capparidaceae-feeding Colotis tend towards
local food plant specialisation.

The Urtica feeding members of the Vanessini in the Palaearctic fall into two groups.
Vanessa lay single eggs, Aglais lay clutches. Members of both genera are often found on
the same batch of nettles, but as in Colotis complete defoliation is rare.

This paper was prepared under a general research grant for 1987 kindly provided by
the Carlsberg Foundation of Denmark.

TORBEN B. LARSEN, Snoghoj alle 29C, DK 2770 Kastrup, Denmark.

Received for publication 23 july 1987; accepted 26 October 1987.

Journal of the Lepidopterists’ Society
42(1), 1988, 59

BOOK REVIEW

THE LIVES OF BUTTERFLIES, by Matthew M. Douglas. 1986. xv + 241 pp. 16 pp. color
photographs. University of Michigan Press, Ann Arbor, Michigan, U.S.A. Hard cover.
$45.00.

This attractive book is the product of a scientist and teacher whose enthusiasm is
contagious. Its strengths include substantive explanations of many aspects of work on
butterfly biology, its discussion of experimental and other evidence for scientific conclu-
sions, and its emphasis on scientific literature. The book is rich in clear, often detailed,
explanations of work in several major areas: anatomy, development, and evolution of
morphological features of life stages; biophysical, physiological, and ecological constraints
on life stages and community structure; behavioral, biochemical and ecological aspects
of speciation and coevolution with plants. This exposition is accompanied by many black-
and-white diagrams (often from published original drawings or photographs), a section
of color photographs illustrating activities and morphological characteristics of life stages,
a glossary, several appendices, and a useful index. This combination makes the book an
engaging, accessible, self-contained store of information.

In addition, the author enhances the book’s informational content in two ways. First,
he places specific examples in a conceptual context by discussing considerations that
underlie specific hypotheses. Explanations of how observations and experimental data are
collected contribute to a clear sense of how scientific questions are raised and examined,
and why “answers may be open to alternative interpretations. This aspect of the book
reflects the author’s experience as a university professor; many of his discussions would
make good lecture notes for an advanced undergraduate course. This bold focus on
processes of scientific research may be the book’s most important contribution to its
educational goals. Second, the book’s emphasis on recent research literature provides a
resource for further study.

The question of readership presents problems for the book. While ostensibly written
for an audience that has some background in biology, its attempt to appeal to both lay
and professional audiences sometimes creates disconcerting inconsistencies. For example,
the author describes “‘sex-limited mimicry’ as a special case of Batesian mimicry in which
one sex mimics unpalatable models; he includes a definition of this term in the glossary.
Ten pages later, however, “sex-limited”’ is used colloquially to describe distribution of a
trait whose pattern of inheritance is sex-linked. This colloquial use of a term that has
specific meaning in genetics is confusing. Similarly, the author emphasizes his personal
research experience in a way likely to engage the interest of lay readers. To a professional
readership, however, such emphasis is likely to seem egotistical and annoying.

This book thus attempts the dual challenges of engaging and educating a lay readership
as well as concisely reviewing recent literature for a professional audience. This is a rarely
attempted goal, and the author presents us with a unique solution. The book’s value to
its potential professional audience lies in its conciseness and timely review of much recent
literature. Its appeal to this audience is uncertain, because professors whose students study
these research topics in class may assign the original literature rather than this book.
However, the author’s contribution to explicating butterfly biology and scientific research
for a lay audience is a noteworthy success.

F. S. CHEW, Department of Biology, Tufts University, Medford, Massachusetts 02155.

Journal of the Lepidopterists’ Society
42(1), 1988, 60-61

OBITUARY

ABNER ALEXANDER TOWERS (1916-1987): A Tribute

The Lepidopterists’ Society lost one of its charter members with the passing of Abner
A. Towers, who was well-known to collectors in the Southeast and to participants in the
first of the collecting expeditions to Ecuador organized by Thomas C. Emmel and Gio-
vanna Holbrook.

Abner Alexander Towers

Born in Gadsden, Alabama, 28 January 1916, Abner Towers developed an interest in
wildlife as a boy, particularly his lifelong fascination with butterflies and moths, birds,
and other flying creatures. He grew up in Gadsden, completing his primary schooling
there, then attended the Kent School, in Kent, Connecticut, during which time he began
to collect and study Lepidoptera seriously. At the age of eighteen, in 1934, he took his
first trip to the Florida Keys expressly to observe and collect butterflies and moths. He
attended the Massachusetts Institute of Technology as a general science major, and, after
earning the Bachelor of Science in 1939, served as an officer in the U.S. Army Corps of
Engineers. He spent most of World War II in the Aleutian and Philippine Islands.
Following the war he settled in Georgia, the state he would call home for the rest of his
life. Abner Towers married, raised a family, and built a career as an engineer and chemist,
and was often described in both capacities by co-workers and peers as “brilliant.” In
August 1972, he cofounded A-Jay Chemical Company, in Powder Springs, Georgia, an
industrial chemical firm he continued to administer until his terminal illness.

In the 1950’s Abner resumed his study of the Lepidoptera of the region, focusing his
attention almost entirely on the butterflies of Georgia and Florida, and he steadily built
an impressive collection containing substantial series of virtually all the species recorded
from the two states. He established a strong friendship with Lucien Harris Jr., and his
contributions to Harris's The Butterflies of Georgia (University of Oklahoma Press, 1972)
were significant, and included numerous state records and field observations. Abner’s

VOLUME 42, NUMBER 1 61

persistent and dedicated collecting subsequently added several species to the Georgia
butterfly fauna, including Mitoura hesseli Rawson & Ziegler, and, in 1981, he participated
in the discovery of a new geometrid, described as Narraga georgiana Covell, Finkelstein
& Towers (J. Res. Lepid. 23:161-168, 1984). Occasionally, when his other responsibilities
allowed, Abner traveled and collected outside the country; most notable were his collecting
trips to Ecuador in 1980 and Jamaica in 1982.

Abner Towers died 18 March 1987, after a bravely fought three-year battle with
leukemia. He is survived by his wife, Margaret Le Craw Towers, his children John A.
Towers, Marsha Towers Endictor, and Andrea Towers Rohaly, and his sister Harriet
Towers Bjelouvucic. His friends and co-workers remember him as “a man of warmth...
who always took time to inquire of people’s families, discuss their hobbies, jobs or personal
interests and give advice, if asked, with a sincerity derived from a love of people. He
was totally unselfish with his time, his knowledge and his abilities.” (From a eulogy by
Alan Shipp and Polly Buford.)

His collection was donated to the University of Florida in 1985 and deposited in the
Florida State Collection of Arthropods, Gainesville. In addition to the Lepidopterists’
Society, Abner was a charter member of the Southern Lepidopterists, a group he served
since its founding in 1978 as Georgia zone coordinator.

“Abner Towers was a gentle man, and a gentleman. He will be missed.” (Shipp and

Buford. )

IRVING L. FINKELSTEIN, 425 Springdale Drive N.E., Atlanta, Georgia 30305.

Date of Issue (Vol. 42, No. 1): 16 March 1988

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EDITORIAL STAFF OF THE JOURNAL
WILLIAM E. MILLER, Editor

Dept. of Entomology
University of Minnesota
St. Paul, Minnesota 55108 U.S.A.

Associate Editors:
M. DEANE BOWERS, BOYCE A. DRUMMOND III, DOUGLAS C. FERGUSON,
ROBERT C. LEDERHOUSE, THEODORE D. SARGENT, ROBERT K. ROBBINS

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SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London.
209 pp.

196la. Some contributions to population genetics resulting from the study of

the Lepidoptera. Adv. Genet. 10:165-216.

In General Notes and Technical Comments, references should be shortened and given
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M., 1961, Sym. R. Entomol. Soc. London 1:23-30) without underlining.

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CONTENTS

SPEYERIA ATLANTIS IN COLORADO: REARING STUDIES CONCERNING
THE RELATION BETWEEN SILVERED AND UNSILVERED FORMS.
James A. Scott: 0.0 ee

POPULATION FLUCTUATIONS OF AZETA VERSICOLOR (FABRICIUS)
(NOCTUIDAE) ON GLIRICIDIA SEPIUM (JACQ.) (FABACEAE) IN

NORTHEASTERN Costa Rica. Allen M. Young WW
BUTTERFLIES OF NORTHEAST TENNESSEE. Charles N. Watson Jr.
i> John A. Hyatt, 200

BODY WEIGHT AND WING LENGTH CHANGES IN MINNESOTA
POPULATIONS OF THE MONARCH BUTTERFLY. William S.
Herman .cc

HABITAT AND RANGE OF EUPHYDRYAS GILLETTI (NYMPHALIDAE).
Ernest H, Williams uu ee

BIOLOGY OF POLYGONIA PROGNE NIGROZEPHYRUS AND RELATED
TAXA (NYMPHALIDAE). James A. Scott 23a :

GENERAL NOTES

Glassberg, Lehman, and Pellmyr collections to the Smithsonian Institution.
Robert K. Robbins <> Gary F. Hevel 2. ee

Differing oviposition and larval feeding strategies in two Colotis butterflies
sharing the same food plant... Torben B. Larsen _.____.__._

Book REVIEW
The Lives of Butterflies. °F.S. Chew i220) ee

OBITUARY
Abner Alexander Towers (1916-1987): a tribute. Irving L. Finkelstein

46

31

57

59

Volume 42 1988 Number 2

ISSN 0024-0966

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES
- Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

24 May 1988

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

JERRY A. POWELL, President JEAN-FRANCOIS LANDRY, Vice
DouGLas C. FERGUSON, Immediate Past President

President ATUHIRO SIBATANI, Vice
JACQUELINE Y. MILLER, Vice President President
RICHARD A. ARNOLD, Secretary JAMES P. TUTTLE, Treasurer

Members at large:

MIRNA M. CASAGRANDE M. DEANE BOWERS JULIAN P. DONAHUE
EDWARD C. KNUDSON RICHARD L. BROWN JOHN E. RAWLINS
FREDERICK W. STEHR PAUL A. OPLER Jo BREWER

The object of the Lepidopterists’ Society, which was formed in May 1947 and for-
mally constituted in December 1950, is “to promote the science of lepidopterology in
all its branches, .... to issue a periodical and other publications on Lepidoptera, to fa-
cilitate the exchange of specimens and ideas by both the professional worker and the
amateur in the field; to secure cooperation in all measures’ directed towards these aims.

Membership in the Society is open to all persons interested in the study of Lepi-
doptera. All members receive the Journal and the News of the Lepidopterists Society.
Institutions may subscribe to the Journal but may not become members. Prospective
members should send to the Treasurer full dues for the current year, together with their
full name, address, and special lepidopterological interests. In alternate years a list of
members of the Society is issued, with addresses and special interests. There are four
numbers in each volume of the Journal, scheduled for February, May, August and
November, and six numbers of the News each year.

Active members—annual dues $25.00
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Life members—single sum $350.00
Institutional subscriptions—annual $40.00

Send remittances, payable to The Lepidopterists’ Society, to: James P. Tuttle, Treasurer,
3838 Fernleigh Ave., Troy, Michigan 48083-5715, U.S.A.; and address changes to: Julian
P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, California
90007-4057 U.S.A. For information about the Society, contact: Richard A. Arnold, Sec-
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Cover illustration: Mature larva of Papilio polyxenes asterius Stoll on wild carrot,
Daucus carota L. Submitted by John V. Calhoun.

JOURNAL OF

THe LEPIDOPTERISTS’ SOCIETY

Volume 42 1988 Number 2

Journal of the Lepidopterists’ Society
42(2), 1988, 63-98

IMPACT OF OUTDOOR LIGHTING ON MOTHS:
AN ASSESSMENT

KENNETH D. FRANK
2508 Pine St., Philadelphia, Pennsylvania 19103

ABSTRACT. Outdoor lighting has sharply increased over the last four decades. Lep-
idopterists have blamed it for causing declines in populations of moths. How outdoor
lighting affects moths, however, has never been comprehensively assessed. The current
study makes such an assessment on the basis of published literature. Outdoor lighting
disturbs flight, navigation, vision, migration, dispersal, oviposition, mating, feeding and
crypsis in some moths. In addition it may disturb circadian rhythms and photoperiodism.
It exposes moths to inereased predation by birds, bats, spiders, and other predators.
However, destruction of vast numbers of moths in light traps has not eradicated moth
populations. Diverse species of moths have been found in illuminated urban environments,
and extinctions due to electric lighting have not been documented. Outdoor lighting does
not appear to affect flight or other activities of many moths, and counterbalancing eco-
logical forces may reduce or negate those disturbances which do occur. Despite these
observations outdoor lighting may influence some populations of moths. The result may
be evolutionary modification of moth behavior, or disruption or elimination of moth
populations. The impact of lighting may increase in the future as outdoor lighting expands
into new areas and illuminates moth populations threatened by other disturbances. Re-
ducing exposure to lighting may help protect moths in small, endangered habitats. Low-
pressure sodium lamps are less likely than are other lamps to elicit flight-to-light behavior,
and to shift circadian rhythms. They may be used to reduce adverse effects of lighting.

Additional key words: conservation, evolution, flight, urban ecology, light pollution.

Since the invention of the incandescent lamp over a hundred years
ago, outdoor lighting has progressively increased. The growth has been
characterized by expansion into new geographic areas, development of
new lamps with new spectral characteristics, and increases in total
amount of light and radiant energy (Riegel 1973, Hendry 1984, Sullivan
1984). Outdoor lighting has transformed the nocturnal face of the earth
(Croft 1978). However, despite universal awareness that electric light
disturbs behavior of nocturnal insects, the ecological impact of outdoor
lighting has never been comprehensively assessed.

The possibility that outdoor lighting may adversely affect our fauna

64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

is well recognized. Lepidopterists have blamed outdoor lighting for
declines in populations of North American moths, especially saturniids
in the northeastern United States (Holland 1908, Ferguson 1971, Hessel
1976, Muller 1979, Worth & Muller 1979, Krivda 1980, Pyle et al.
1981). This view assumes a direct causal link between lamps and faunal
change. Fundamental questions about such a link, however, have never
been closely examined: What mechanisms might link lamps with changes
in populations of moths? If lamps cause populations of moths to change,
specifically what might the changes be? How important are effects of
lighting compared to effects of other environmental disturbances? This
study examines each of these questions. It investigates the hypothesis
that outdoor lighting influences populations of moths.

The investigation is based on a review of literature. The presentation
is organized into three sections. The first section describes distribution,
growth, energy, and spectral composition of outdoor lighting. The sec-
ond describes how lamps affect behavior, life functions and survival of
individual moths. The third explores how such effects may disturb moth
populations; it also discusses measures to reduce disturbances caused
by lighting. Citations are deliberately extensive to facilitate retrieval
of source material which is widely scattered among different disciplines.

LIGHTING

Nocturnal images of earth viewed from orbiting satellites show the
distribution of outdoor lighting (Fig. 1). In the United States this dis-
tribution coincides with that of the country’s population (Croft 1978).
Nocturnal illumination is clustered around all large metropolitan areas,
with greatest concentration in the Northeast corridor. Viewed from an
airplane, nocturnal lighting delineates a web of interconnecting road-
ways lined with illumination from houses, parking lots, billboards, and
other landmarks. Such aerial observation suggests that lighting forms
an illuminated web that envelops the nocturnal environment of Lepi-
doptera. The web’s density varies with human population density, and
its distribution is continental.

The magnitude of lighting in a major metropolitan area is illustrated
by Philadelphia’s streetlighting (Table 1). Philadelphia has 100,000 high-
pressure sodium streetlamps at a density of almost 300 lamps/km?. The
energy they radiate equals more than 10 kilowatts/km?, an order of
magnitude greater than the energy density of moonlight at full moon
(Agee 1969). During the last 4 decades, lamp size (lumens) increased
7-fold, number of lamps tripled, and type of lamp changed from tung-
sten filament and mercury to high-pressure sodium (Figs. 2 & 3) (Wain-
wright 1961, C. A. Oerkvitz pers. comm.). Nationwide per capita con-
sumption of electrical power for streetlighting is similar to that of

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66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Streetlamps in Philadelphia, 1983. Total lamps, lumens, and demand (watts)
from C. A. Oerkvitz (pers. comm.). Radiant energy calculated from GTE Products Corp.
(Sylvania) (1977b). Demographic data from World Almanac (1986).

Number
Streetlamp parameter Total Per capita Per km?
Lamps 1.0 x 10° 3.8 X 107 2.8 x 10?
Lumens 1.8 x 10° Le <Oe 5.0 x 10°
Radiant energy (watts) emitted for
wavelengths 350-700 nm 5.6 x 10° 3.3 1.6 x 10!
Electric power demand (watts) 2.2 x 10° 3y x NO 6.1 x 10!

Philadelphia, and growth in lumens has been comparable or higher
(Riegel 1973, Edison Electric Institute 1971, 1985, Sullivan 1984).

Conversion from mercury to high-pressure sodium lamps reduces
radiant energy at the short-wavelength end of the spectrum. However,
high-pressure sodium light is spectrally broad and does include radiant
energy in the blue spectral region (Fig. 2B).

In contrast to high-pressure sodium light, low-pressure sodium light
is spectrally narrow. It excludes practically all energy in the ultraviolet,
blue, and green regions of the spectrum (Fig. 2A). Viewed through a
spectroscope, its spectrum contains a bright yellow-orange line (actually
2 spectral lines very close together) near 589 nm. Because the human
eye is particularly sensitive to light in the 589 nm region, low-pressure
sodium lamps can provide bright illumination with comparatively little
radiant energy (Finch 1978). Compared to other lamps used for outdoor
lighting, low-pressure sodium lamps minimize environmental exposure
to radiant energy both in number of wavelengths and number of watts.
These lamps are used for streetlighting and other outdoor lighting, but
much less frequently than are high-pressure sodium lamps.

Conversion of streetlamps from mercury to high-pressure sodium has
changed the spectral distribution of outdoor lighting, but it has not
changed it as much or as clearly as one might suppose. Mercury lamps,
for example, are still used for residential and commercial lighting in
Philadelphia, and for streetlighting in neighboring areas. Tungsten fil-
ament (Fig. 3), low-pressure sodium, metal halide (Fig. 2C) and flu-
orescent lamps (Sorcar 1982) all contribute to spectral diversity of out-
door lighting in the city. While density and distribution of outdoor
lighting have increased, spectral composition has diversified.

EFFECTS ON INDIVIDUAL MOTHS
Vision
Bright light can lower sensitivity of moth eyes 1000-fold (Bernhard
& Ottoson 1960a, Hoglund & Struwe 1970, Agee 1972, 1973, Eguchi

VOLUME 42, NUMBER 2

Energy emitted (watts per 10 nm)

A

350 400 450 500 550 600 650 700
Wavelength (nm)

Electrical input = 135 watts
Luminous output = 20,000 lumens
Manufacturer: North American Philips Lighting Corp.

Low-pressure sodium

Energy emitted (watts per 10 nm)

Wavelength (nm)

Electrical input = 400 watts
Luminous output = 34,000 lumens
Manufacturer: GTE Products Corp. (Sylvania)

Metal halide

Ultraviolet Violet

350 400 500

Energy emitted (watts per 10 nm)

350 400 450 500 550 600 650

Wavelength (nm)
Electrical input = 400 watts

Luminous output = 50,000 lumens
Manufacturer: GTE Products Corp. (Sylvania)

High-pressure sodium

Energy emitted (watts per 10 nm)

D

350 400 450 500 550 600
Wavelength (nm)

Electrical input = 400 watts
Luminous output = 20,500 lumens
Manufacturer: GTE Products Corp. (Sylvania)

Clear mercury

550

Wavelength (nm)

ie 42:

67

700

Spectral energy distribution of vapor discharge lamps. Sources for A: Judd

1951, Finch 1978, Illuminating Engineering Society 1981, North American Philips Light-
ing Corp. 1982. Sources for B, C, and D: GTE Products Corporation (Sylvania) 1977a,

1977b, 1979.

& Horikoshi 1984). Electroretinographic studies suggest what happens
to the visual sensitivity of a moth that flies to a lamp. If the moth
remains at the lamp and then flies away, full visual sensitivity may not
return for 30 min or longer (Bernhard & Ottoson 1960a, 1960b, Agee
1972). This effect requires exposure to the lamp over a period of time,
probably 10 min or longer (Day 1941, Héglund 1963, Yagi & Koyama

68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Relative energy

100

Wavelength (nm)

Electrical input: 1000 watts
Luminous output: 23,100 lumens
Color temperature: 3030° Kelvin

Fic. 3. Spectral energy distribution of tungsten filament (“incandescent ’’) lamp. Sources:
GTE Products Corporation 1972, 1974.

1963). A moth flying away from a lamp into relative darkness on a
cloudy, moonless night may be functionally blind until enough time
has elapsed for it to become fully dark-adapted.

Continuous exposure to bright electric lamps could in theory “dazzle”
moths. This means it could stimulate the moth retina so intensely that
the retina could not respond to additional increases in light. The result
would be functional blindness so long as the moth remained exposed
close to the lamp. Electroretinographic evidence, however, suggests that
lamps do not dazzle moths (Eguchi & Horikoshi 1984).

Net effects of electric lighting on moth vision may vary according
to local conditions as well as moth behavior. Urban lighting increases
background illumination which in turn may help moths see. Electric
lighting in some areas has increased nocturnal sky brightness as much
as 20-fold (Hendry 1984). However, the spectral composition, polariza-
tion and spatial distribution of outdoor lighting varies widely in different
settings. In some locations they may differ so much from that of natural
nocturnal light that they create visual artifacts and distortions. One

VOLUME 42, NUMBER 2 69

outcome of disturbed vision is flight to outdoor lamps, but many dis-
turbances in visual function and behavior are possible.

The suggestion that urban lighting influences nocturnal vision of
moths may appear paradoxical. Municipal light sources have shifted
away from mercury lamps and toward high-pressure sodium lamps.
One might suppose that moth retinas are insensitive to the relatively
long wavelengths which characterize most of the energy contained in
high-pressure sodium light (Fig. 2B). Moths, for example, do not fly to
the 589 nm light of low-pressure sodium lamps (Fig. 2A), or do so rarely
(Robinson 1952). Such a supposition, however, is incorrect: electroreti-
nograms of moths consistently demonstrate sensitivity to light in the
589 nm region, and most studies have found maximum sensitivity in
the green rather than ultraviolet part of the spectrum (Jahn & Crescitelli
1939, Héglund & Struwe 1970, Hsiao 1972, Mikkola 1972, Agee 1978,
MacFarlane & Eaton 1978, Langer et al. 1979, Mitchell & Agee 1981,
Eguchi et al. 1982). Retinal sensitivity extends farther into the long-
wavelength end of the spectrum than flight-to-light behavior typically
would suggest (Mikkola 1972, MacFarlane & Eaton 1973, Mitchell &
Agee 1981).

Navigation

Diversion to lamps. Three hundred fifty-six species of Macrolepi-
doptera, or about a third of those species found in all of Great Britain,
were collected at a single light trap in England (Williams 1939). Com-
parable findings have been reported in Britain and North America
(Dirks 1937, Robinson & Robinson 1950a, Beebe 1958, Bretherton 1954.
Moore 1955, Langmaid 1959, Hosny 1959, Holzman 1961, Moulding
& Madenjian 1979). Tens of thousands of moths have flown to a single
lamp in a single evening (Robinson & Robinson 1950a), and huge
swarms of moths have aggregated around urban light sources (Howe
1959). On the other hand, some species of nocturnal moths rarely fly
to lamps even though large populations of them may be flying nearby
(Bretherton 1954, Taylor & Carter 1961, Janzen 1983). A variety of
physiologic, behavioral and environmental factors may determine which
species of moths fly to light and when (Geier 1960, Gehring & Madsen
19638, Milyanovskii 1975, Mazokhin-Porshnyakov 1975, Janzen 1983,
1984).

Large numbers of moths flying to lamps may give a false impression
that lamps divert moths from great distances. Effective radius of a 125-
watt mercury vapor light trap was initially reported to be 91 m, but
later estimates reduced the figure to 17 m, and the most recent analysis
cut the distance to 3 m (Robinson & Robinson 1950a, Robinson 1960,
Baker & Sadovy 1978). Other studies have shown flight-to-light dis-

70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

tances of 10 m or less (Stanley 1932, Hamilton & Steiner 1939, Hartstack
et al. 1971, Plaut 1971). Long-distance estimates ranging up to half a
kilometer represent either extrapolation, artificial conditions or both
(Graham et al. 1961, Hsiao 1972, Agee 1972, Stewart et al. 1969, Plaut
1971, Bowden & Morris 1975).

If the mechanism by which a lamp disturbs moths depends on di-
version of flight paths to the lamp, then the moths disturbed must be
limited to those flying in the geographic area immediately adjacent to
the lamp. In this sense any direct effects of a particular lamp would
tend to be local, except when topography (Beebe 1949, Beebe & Fleming
1951), foodplants, pheromones, or other factors concentrate moths near
the lamp. Only in urban regions would density and distribution of
lamps be great enough to influence large populations of moths over
broad geographical areas.

Effects of electric lamps in urban areas, however, may be much
smaller than one might expect. Robinson & Robinson (1950a) noted
that lamps in isolated phone booths appear to be much more effective
in eliciting flight-to-light behavior than are clusters of bright urban
lamps located immediately adjacent to areas with large populations of
moths. They demonstrated that lamps interfere with each other’s ca-
pacity to elicit flight-to-light behavior, and the closer together the lamps,
the greater the interference. The high density which characterizes dis-
tribution of urban lamps suppresses flight-to-light behavior.

Urban lighting may suppress flight to light for a number of reasons.
Light trap collections vary with the lunar cycle and are lowest at full
moon (Williams et al. 1956, Agee et al. 1972, Nemec 1971, Dufay 1964,
Bowden & Church 1978, Janzen 1983, Stradling et al. 1983). A similar
correlation with moonlight cannot be demonstrated when nocturnal
flight is measured by suction traps (Williams et al. 1956, Danthana-
rayana 1986), pheromone-baited traps (Saario et al. 1970, Janzen 1984)
or radar (Schaefer 1976). Moths active at dusk typically appear in
suction traps before they appear in light traps (Taylor & Carter 1961).
Eye pigment must be in a position of dark adaptation before moths
will fly to light (Collins 1934), and even relatively dim background
light can cause the pigment to move away from this position (Bernhard
& Ottoson 1964). Diffuse urban light, like moonlight and twilight,
reduces the darkness essential for flight-to-light behavior.

The moon not only increases background lighting but also constitutes
a concentrated source of light by which insects may be able to orient
(Sotthibandhu & Baker 1979). Moths flying by lunar navigation may
bypass lamps (Baker & Sadovy 1978). Lamps may provide navigational
cues which suppress flight to other lamps.

Light sources that emit large amounts of ultraviolet energy are gen-

VOLUME 42, NUMBER 2 7

erally most effective in eliciting flight-to-light behavior (Williams et al.
1955, Glick & Hollingsworth 1955, Klyuchko 1957, Deay et al. 1965,
Mazokhin-Porshnyakov 1969, 1975, Mikkola 1972, Sargent 1976, Mitch-
ell & Agee 1981). Conversion of mercury streetlamps to high-pressure
sodium and metal halide streetlamps has undoubtedly tended to reduce
flight to streetlamps. On the other hand, moths do fly to high-pressure
sodium and metal halide lamps, and a small minority of species may
fly preferentially to lamps with little or no ultraviolet emission (Klyuch-
ko 1957, Mikkola 1972). Unlike high-pressure sodium lamps, however,
low-pressure sodium lamps rarely elicit flight-to-light behavior (Rob-
inson 1952).

In summary, increases in electric lighting do not necessarily impair
nocturnal vision and navigation. Under some conditions they may im-
prove moths’ nocturnal vision and suppress flight-to-light behavior.

Diversion away from lamps. Electric lamps may also divert moths
away from them (Robinson & Robinson 1950a, Robinson 1952, Herms
1929, 19382, Nomura 1969, Nemec 1969, Hsiao 1972). These effects
may depend in part on spectral output of the lamp (Mazokhin-Porsh-
nyakov 1969, 1975, Nomura 1969). Several theories attempt to explain
this behavior (Hsiao 1972), but none accounts for diversity of flight
paths at lamps (Janzen 1984): while some moths make spiral or circular
flights around lamps and land several meters away, others make a
beeline straight to lamps and crash into them. Flight paths approaching
lamps may zig-zag or be totally chaotic (Holzman 1961, Mazokhin-
Porshnyakov 1969, Janzen 1984). Diversion away from lamps has been
debated (Bretherton 1950, Robinson & Robinson 1950b). Evidence that
moths avoid large illuminated areas (Herms 1929, 1932, Nomura 1969,
Nemec 1969) is inconclusive, but this behavior is more difficult to
demonstrate than flight to lamps.

Lamps suppress flight of moths that fly to them. Moths approaching
lamps may land near them and remain quiescent for a moment or for
the entire night. Lamps suppress flight of some species more than others
(Blest 1963, Graham et al. 1964). In some cases lamps do not appear
to suppress flight; in other cases they excite quiescent moths into flight
(Collins 1984, Hsiao 1972). Diurnal moths occasionally fly at night to
lamps (Engelhardt 1946, Janzen 1983), but here it is unclear whether
the lamps help to initiate nocturnal flight.

Diversion and suppression of flight may impair orientation and nav-
igation based on lunar, stellar or other visual celestial cues (Mazokhin-
Porshnyakov 1969, Sotthibandhu & Baker 1979, Wehner 1984) includ-
ing polarization of celestial light (Danthanarayana & Dashper 1986).
It also may impair navigation and orientation based on geomagnetic,
gravitational, barometric, aerodynamic, inertial, olfactory, acoustic or

12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

visual terrestrial cues (Baker & Kuenen 1982, Baker & Mather 1982,
Janzen 1984, Schone 1984, Riley & Reynolds 1986). How much electric
lighting disturbs use of particular cues may be expected to vary in part
according to which cues the moth happens to be using at the moment
it encounters the lamp.

Migration and Dispersal

Light sources divert moths engaged in migratory or dispersal flights
(Cockerell 1914, Williams 1937, Beebe & Fleming 1951, Wolf et al.
1986). Urban lighting surrounds habitats isolated by urban sprawl, so
that moths may have to traverse dozens of kilometers of densely illu-
minated territory to arrive at potential breeding sites. Moths flying high
(Glick 1965) may fly to urban light sources on tall buildings (Stanley
1932, Glick 1961). Because location of natural flyways is poorly doc-
umented for North American moths, one cannot determine the extent
urban lighting may intersect long-range natural migration routes here.
In Venezuela, vast numbers of migrating moths aggregated around
lamps near a narrow mountain pass which functions as a natural flyway
(Beebe 1949, Beebe & Fleming 1951). Lighting along roads following
topographical features such as valleys, rivers, and coastlines might se-
lectively interfere with North American moth migrations (Fig. 1).

Oviposition

Electric lighting can disturb oviposition. Light-trap surveys have
shown that the vast majority of females collected at lamps are gravid
(Dirks 1937, Ficht et al. 1940, Glick & Hollingsworth 1954, Geier 1960,
Gehring & Madsen 1963) although males usually outnumber them
(Dirks 1937, Williams 1939, Sargent 1976, Worth & Muller 1979, Janzen
1984). Flight to light can shift oviposition to sites located near the lamp
(Ficht et al. 1940, Martin & Houser 1941, Pfrimmer & Lukefahr 1955,
Beaty et al. 1951, Nemec 1969, Brown 1984). Eggs may be deposited
on lampposts, window screens, buildings, and other unsuitable sites near
lamps. Egg densities may be several-fold higher on plants near lamps
(Martin & Houser 1941). The result may be larval overcrowding and
increased susceptibility to starvation, microbial infection, and preda-
tion.

Lamps shift the distribution of oviposition sites toward them probably
by diverting ovipositing females and not by stimulating oviposition. In
cornfields, Ostrinia nubilalis (Hbn.) (Pyralidae) tends to oviposit near
lamps (Ficht et al. 1940, Beaty et al. 1951), but in the laboratory
nocturnal illumination suppresses O. nubilalis oviposition (Skopik &
Takeda 1980). Similar observations have been reported in Pectinophora
gossypiella (Saund.) (Gelechiidae) (Pfrimmer & Lukefahr 1955, Lu-

VOLUME 42, NUMBER 2 Fle

kefahr & Griffin 1957, Henneberry and Leal 1979). Outdoor lighting
may decrease oviposition by Cydia pomonella (L.) (Tortricidae) and
Heliothis spp. (Noctuidae), although the mechanism is unclear (Herms
1929, 1982, Nemec 1969).

Mating

Outdoor lighting does not prevent mating in certain Saturniidae:
male Hyalophora cecropia (L.) and Samia cynthia (Drury) complete
long-distance mating flights to virgin females at night across illuminated
urban territory, and breed in urban habitats (Rau & Rau 1929, Pyle
1975, Sternburg et al. 1981, Waldbauer & Sternburg 1982). Most freshly
emerged female saturniids do not fly at all until they have emitted
pheromone and mated (Blest 1963, Nassig & Peigler 1984, Waldbauer
& Sternburg 1979). Male sphingids and saturniids fly to virgin females
before they fly to nearby electric lamps (Allen & Hodge 1955, Worth
& Muller 1979, Janzen 1984). Almost all female Cydia pomonella col-
lected at black lights have already mated (Gehring & Madsen 1963).
Although more males than females typically fly to lamps, the capacity
of males to mate with more than one female (Rau & Rau 1929, Allen
& Hodge 1955, Lukefahr & Griffin 1957, Vail et al. 1968) may moderate
the reproductive impact of disproportionate harm to males.

In contrast, electric lighting may have a major effect on mating in
certain Noctuidae. Heliothis zea (Boddie) is an example. The peak time
of night during which H. zea flies to light traps coincides with the
period of copulation (Graham et al. 1964, Stewart et al. 1967). Only a
third to a half of female H. zea collected at light sources have mated
(Gentry et al. 1971, Vail et al. 1968). In the laboratory, H. zea will not
mate unless its eyes are in a state of dark adaptation, as indicated by
the presence of eye glow. Light intensity must be below 0.015 wW/
cm?, the intensity of light of a quarter-moon (Agee 1969). The suggestion
is that H. zea females fly to light sources whose radiant energy suppresses
mating.

A criticism of this scenario is that unmated H. zea females that fly
to light may be migrating (Raulston et al. 1986) and therefore sexually
immature (Johnson 1969). Female H. zea in the laboratory do not mate
for 30-60 h after eclosion (Agee 1969). However, even if unmated
females at lamps were sexually immature migrants, the lamps could
disrupt reproductively important behavior, such as flight to locations
where courtship and mating would be likely to occur. Furthermore,
outdoor lighting may interfere with H. zea mating regardless of flight
to light. Levels of light that suppress mating in the laboratory (Agee
1969) are well below ambient levels of light in electrically illuminated
environments outdoors. Low levels of incandescent light (Nemec 1969)

74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

and moonlight (Nemec 1971) have influenced activities of Heliothis
spp. in the field.

Other evidence suggests that lighting may interfere with mating.
Unmated females of four other noctuid species fly to lamps (Vail et al.
1968). Male sphingids caught in light traps baited with virgin females
do not seek out the females (Hoffman et al. 1966). In the laboratory,
even dim electric light (0.3 lux) suppresses female Trichoplusia ni
(Hbn.) (Noctuidae) pheromone release and male response to pheromone
(Shorey & Gaston 1964, 1965, Sower et al. 1970). Electric light also
suppresses female pheromone release and male response to pheromone
in Dioryctria abietivorella (Grt.) (Pyralidae) (Fatzinger 1979). Mating
by Pectinophora gossypiella requires a period of relative darkness last-
ing at least 7 h (Lukefahr & Griffin 1957).

Feeding

Moths may feed in illuminated environments. Sphingids and noctuids
visit food sources in full view of electric lamps located sometimes less
than a few meters away, or they fly to electric light sources after they
have completed feeding (Bretherton 1954, Milyanovskii 1975, Mazo-
khin-Porshnyakov 1975, Janzen 1983, 1984). I have observed Buddleja
(Gentianaceae) blossoms covered with noctuids at night (2300 h) vir-
tually directly under a tungsten filament street lamp illuminating a
heavily traveled road in Quisset, Massachusetts. Light from automobile
headlamps and from a flashlight did not alter the moths’ activities.

Electric lamps, however, may interfere with feeding. Orchard illu-
mination has reduced the number of Cydia pomonella feeding at bait
(Herms 1932). In Japan, orchard illumination has been used to protect
fruit from damage by fruit-piercing noctuids (Nomura 1969). Light
has disturbed nectaring sphingids (Brown 1976). Diversion of moths
away from light may explain why lamps interfere with feeding.
Suppression in feeding is moot for the large number of moth adults
that never feed (Norris 1936).

Electric lighting theoretically could injure larval foodplants. Sodium
vapor lighting may harm plants by disrupting photoperiodic regulation
of growth and development (Sinnadurai 1981, Cathey & Campbell
1975, Shropshire 1977), but such effects are apparently greater indoors
in greenhouses than outdoors on the street (Andresen 1978).

Time Keeping
Electric lighting can delay or advance vital activities of moths and
their larvae, and these shifts could affect the insects as much as changes
in the activities themselves (Beck 1980, Saunders 1982). This possibility
has been the basis for proposals to exploit biological clocks for purposes

VOLUME 42, NUMBER 2 vis)

of pest control (Barker et al. 1964, Nelson 1967). In a field trial, however,
light exposure failed to prevent diapause in larvae of Adoxophyes orana
(F.R.) (Tortricidae) (Berlinger & Ankersmit 1976). The trial suggests
that it is easier to manipulate biological clocks indoors than outdoors
where temperature and other factors cannot be controlled.

Biological clocks of flying insects, however, may be much more sus-
ceptible to outdoor electric lighting than those of larvae. This is because
flight to light increases exposure to radiant energy. Exposure to a pulse
of light lasting only 15 min is sufficient to attenuate a circadian rhythm
in Drosophila; light 10° times more intense produces the same effect
after only 10 sec; light 10° times more intense does it after an exposure
of less than 0.1 sec (Chandrashekaran & Engelmann 1976). Energy for
even a minute fraction of a second (photoflash) can disturb photope-
riodic clocks in larvae of Lepidoptera (Barker et al. 1964). The an-
thropomorphic observation that quiescent moths adjacent to a lamp are
“asleep because they think it is daytime’? may be close to the truth.

Shifts in timing of nocturnal behavior of moths at lamps do not
necessarily imply shifts in phase of endogenous rhythms. Changes in
timing of behavior could represent other responses to light, or they
could represent complex mixtures of responses. Regardless of these
possibilities, magnitude and character of responses may vary according
to when in the circadian cycle exposure to light occurs (Pittendrigh &
Minis 1971, Skopik & Takeda 1980). Responses may also vary depending
on spectral output of the lamp. For example, Pectinophora gossypiella
has two light-sensitive clocks, only one of which responds to the 589
nm light emitted by low-pressure sodium lamps (Bruce & Minis 1969,
Pittendrigh et al. 1970).

Theoretical Effects

To what extent nocturnal flight to light affects timing of nocturnal
behavior has never been formally investigated. For example, if a moth
flies to a light source, receives intense irradiation for 15 min, and flies
away, how will its activities during the rest of the night be affected?
If a male, will its mating period still coincide with that of females not
exposed to light? If a female, will pheromone release still occur during
the flight period of males? Shifts in mating times could cause sympatric,
closely related species to attempt to mate with each other; such species
normally do not mate with each other in part because their different
mating periods keep them temporally segregated (Tuttle 1985).

Synchronization of activities with lunar rhythms may help moths
navigate, mate, and avoid predators (Danthanarayana 1986). Lamps
may disturb oviposition synchronized to lunar rhythms (Nemec 1969,
1971). To what extent moth activity synchronizes with lunar rhythms,

76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

and to what extent electric lighting may disturb such synchrony war-
rants investigation.

Predation

Bats, birds, skunks, toads, and spiders hunt moths flying to lamps
(Stanley 1932, Thaxter 1957, Holzman 1961, Krivda 1980, Covell 1985,
Brower 1986). Lamps increase predation by clumping prey, and directly
exposing them to attack (Turnbull 1964). Concentrated experience with
particular species may help birds learn to defeat defenses based on
surprise, novelty, or deceit (Blest 1957, Wickler 1968, Coppinger 1970,
Sargent 1973b, Pietrewicz & Kamil 1979). Lamps also can destroy
defensive behavior, such as that required for crypsis (Sargent & Keiper
1969, Sargent 1973a, 1976). The outcome is exemplified by a dark,
bark-colored moth conspicuously resting on a white wall near a lamp
at dawn. Lamps may help birds learn to recognize unpalatable species,
but moths unpalatable to some birds may be acceptable to others (L6hrl
1979). Lamps may enable different birds to pick and choose among
different possible prey. Because moths often land before they arrive at
lamps, lamps may provide predators with far more prey than one might
expect from the moths immediately adjacent to the lamp (Hartstack et
al. 1968).

Parasitoids of Lepidoptera fly to electric light sources (Collins &
Nixon 1930, Cline et al. 1983). Electric lighting could reduce predation
on Lepidoptera by suppressing populations of parasitoids (Worth &
Muller 1979). It may divert parasitoids used for biological control of
pest Lepidoptera in warehouses (Cline et al. 1983). Even brief exposure
to intense sources of radiant energy (photoflash) may sterilize minute
hymenopterous parasites which survive the radiation (Riordan 1964).
Theoretically, lighting could affect secondary parasites, thus potentially
disturbing the food chain at three levels, and producing changes in
populations which would be difficult to predict (Frank 1986).

EFFECTS ON MOTH POPULATIONS
Evidence Against Effects

Migration and dispersal. Even though lamps may contribute to the
destruction of vast numbers of moths, the impact on moth populations
may be negligible. For example, more than 10000 Autographa (Plusia)
gamma (L.) (Noctuidae) were collected in a light trap in one season in
England (Robinson & Robinson 1950a). In England the population of
A. gamma is maintained almost entirely by immigration in spring from
southern Europe (Ford 1972). A particular light source in England
should have a negligible influence on the breeding stock which annually

VOLUME 42, NUMBER 2 rol

replenishes the population of A. gamma around it. Seasonal movement
of moths over long distances is not rare (Williams et al. 1942, Williams
1958, Johnson 1969, Ford 1972) and may be sustained by wind trans-
porting moths at altitudes sometimes hundreds of meters above most
electric light sources (Glick 1965, Mikkola 1986, Raulston et al. 1986,
Wolf et al. 1986).

Failure to suppress agricultural pests and other species. One might
expect that light traps could substantially reduce or eliminate some
moth populations. However, elaborate efforts to exploit such traps for
pest control have failed, and successes could not be consistently repli-
cated (Cantelo 1974, Hienton 1974). The failure has been attributed to
influx of moths from outlying areas, but light trapping may fail to
control insect populations even on small islands. On St. Croix, United
States Virgin Islands, 250 black-light traps were deployed during a
period of 43 months. The island is 208 km? in area. Although decreases
in light-trap collections suggested that traps were depleting the island’s
sphingids (Cantelo et al. 1972a, 1972b), other studies using the same
traps at the same time found similar decreases in collections of Heliothis
zea even though traps collected only a minute fraction of the island’s
H. zea population (Cantelo et al. 1973, 1974, Snow et al. 1969). Fur-
thermore, light-trap collections of sphingids were beginning to increase
at the time the study was terminated. Meteorologic and density-de-
pendent ecological forces may determine the size of moth populations
exposed to lighting, even on isolated islands.

Failure of light traps to reduce insect populations extends beyond
species of agricultural interest. Williams (1939) examined 150 species
of Noctuidae and Geometridae collected in his stationary light trap
during a 4-year period in Rothamsted. Comparison of numbers of
individuals of each species collected from year to year provided no
evidence of any consistent declines in populations, except possibly in
the case of one geometrid. More recent observations at Rothamsted
extended Williams’ studies. Taylor et al. (1978) tabulated annual num-
ber of species and number of specimens of each trapped at Rothamsted
from 1966 to 1975, and also calculated an index of diversity for each
year. No downward trends are apparent, despite wide fluctuations from
year to year.

Prevalence of urban moths. The above studies did not simulate urban
conditions where lighting is dense and widespread. However, large
numbers of species have been collected in urban areas in Britain and
the United States (Langmaid 1959, Lutz 1941). Collections based on a
nationwide network of 172 light traps in Britain suggest that moth
populations in areas undergoing urban changes can substantially recover
despite electric lighting (Taylor et al. 1978). In North America, some

78 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

saturniid species not only tolerate urban lighting but may actually thrive
better in urban than in rural habitats. Hyalophora cecropia and Samia
cynthia are two examples. The ecology of both species is complex, and
numerous factors other than lighting can account for changes in their
abundance in illuminated environments (Sternburg et al. 1981, Frank
1986). In New England, eight species of Catocala (Noctuidae) thrive
in illuminated urban or suburban areas. Seven of these species can be
found within a mile of downtown New Haven, and one occurs in
downtown Boston. Several depend almost entirely on urban-suburban
shade trees (D. F. Schweitzer pers. comm.).

Extinctions unrelated to lighting. Most declines and extinctions in
moth populations can be linked to specific circumstances unrelated to
lighting (Bretherton 1951, Ford 1972, Heath 1974). These include de-
forestation, agriculture, and draining of fens. Destruction of habitats
as a cause of widespread declines in Lepidoptera populations has been
described in detail for European butterflies (Kudrna 1986). In Britain,
many species of moths became scarce around the middle of the last
century, but after World War I the situation reversed, probably because
of favorable climatic changes (Heath 1974). Declines in numbers of
Malacosoma americanum (F.) (Lasiocampidae) in Winnipeg, Mani-
toba, have been attributed to English sparrows (Passer domesticus L.,
Passeridae) eating the moths at lamps (Krivda 1980), but M. ameri-
canum populations fluctuate at intervals independent of changes in
lighting. Interval duration is about 10 years (Johnson & Lyon 1976).
Attacks by microbial and parasitic agents probably account for periodic
reductions in populations of this species (Lutz 1941).

Saturniid populations in the northeastern United States declined in
the 1950’s. This observation is supported by dates of last capture for
species represented in regional collections, and by surveys of collectors
(Ferguson 1971, Hessel 1976, D. F. Schweitzer pers. comm.). Popula-
tions of some saturniid species have since shown signs of recovery,
whereas other saturniids, especially the two Citheronia species native
to the area, have failed to recover in several states (D. F. Schweitzer
pers. comm.). Declines that occurred in the 1950’s coincided with wide-
spread aerial spraying against gypsy moth, and recoveries coincided
with drastic curtailment of this spraying (D. F. Schweitzer pers. comm.,
Gerardi & Grimm 1979). Whether pesticides can account for changes
in saturniid populations is unclear. However, changes in populations of
saturniids as a group correlate poorly with changes in outdoor lighting.

Evidence for Effects

Small colonies exposed to lighting. Evidence that outdoor electric
lighting has the capacity to affect populations of moths is illustrated by

VOLUME 42, NUMBER 2 79

Hydraecia petasitis Doubleday (Noctuidae) in Finland. Only three or
four isolated colonies are known to exist in the country. The isolation
is not due to urbanization but rather to the fact that the species in
Finland is at the extreme tip of its range. Two small colonies were
studied, one covering 700 m?, the other 800 m?. A mark-recapture
experiment conducted during 48 days in one colony demonstrated that
a trap equipped with an 80-watt mercury lamp captured 53% of males
in the colony and 30% of females at least once. The colony was estimated
to consist of 218 individuals. These and other observations suggest that
continuous light trapping could destroy this population. The authors
point out that this species is only mildly attracted to light, and that the
effect of light trapping might be more severe for other Lepidoptera
(Vaisanen & Hublin 1983). The number of moths the authors trapped
probably underestimated the number that flew to the lamps (Hartstack
et al. 1968).

The Finnish light-trap study demonstrates that a substantial propor-
tion of individual moths within a geographically small colony may fly
to an electric lamp. It is conceivable that disturbances in oviposition,
mating, feeding, vision, navigation, dispersal, crypsis, circadian rhythms
or photoperiodism would be sufficient to disrupt an already shaky
population or to impede establishment of a new one. Disruptive effects
would be even greater when caused by lamps in special conditions.
These include lamps in traps equipped with electrocuting grids (“bug
zappers’) and lamps near bird feeders and bird houses. Lamps may
incinerate or desiccate moths trapped inside poorly constructed or bro-
ken luminaires. Lamps near hostplants may disturb females attracted
to the plants, or they may disturb males attracted to the females. Lamps
in open garages and pavilions may direct moths into areas from which
they cannot escape. Automobile headlamps and streetlamps divert moths
into the paths of moving vehicles.

Urbanization and fragmentation of habitats. The same urban changes
that increase outdoor electric lighting also tend to fragment habitats
(MacArthur & Wilson 1967). The result is creation of small colonies
exposed to electric illumination. Man has made many species of British
moths in effect relict faunas, remnants of a bygone era when their
habitats were much more widespread (Bretherton 1951, Ford 1972).
Three species of noctuids once plentiful in southern California have
been reduced to small, isolated colonies, in one instance in the vicinity
of the Los Angeles International Airport (Hessel 1976). Urban gardens
and parks now function as important faunal reservoirs (Frankie & Ehler
1978, Davis 1978, 1982, Owen 1978, Schaefer 1982). Urbanization in-
creases both vulnerability and exposure of moth populations to lamps.

Lighting as a selective force. Outdoor lighting may act as a selective

80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

force against particular individuals within a population. For example,
it may select against individuals that tend most strongly to exhibit flight-
to-light behavior. In the Finnish light-trap study, such individuals would
include those that flew into traps most frequently. Industrial melanism
demonstrates that urban change may cause evolutionary change in
populations of moths, and that disturbances in crypsis can generate the
selective forces needed to produce such evolution (Kettlewell 1973,
Cook et al. 1986). Electric lighting disturbs crypsis, but also a multitude
of other functions. That some species of noctuids and other nocturnal
moths do not fly to nearby light sources, or do so only rarely (Bretherton
1954, Taylor & Carter 1961, Janzen 1983), suggests that evolutionary
modification of flight-to-light behavior has already occurred, although
the causes are unknown.

Responses to selective pressures produced by lighting may be diverse.
For species active at dusk, natural selection could favor individuals that
fly at the beginning of the population’s flight period, rather than at the
end when flight to light occurs. The evolutionary response would be a
shift in flight period rather than a specific change in flight-to-light
behavior. Biological clocks are in part genetically controlled, and clock
mutants affecting time of eclosion and locomotor activity have been
identified in Drosophila (Konopka & Benzer 1971, Yu et al. 1987). In
moths, different races or strains of a single species exhibit different
photoperiodic behavior (Gardiner 1982, Ankersmit & Adkisson 1967),
and selective pressures can account for such differences (Tauber &
Tauber 1978, Hoy 1978, Waldbauer 1978). On the other hand, ad-
vancing or delaying flight times could disturb species segregation me-
diated through allochronic flight periods (Tuttle 1985), or it could expose
moths to increased predation by birds or bats that fly only at certain
times. Any evolutionary response to selective pressures generated by
electric lighting would have to represent a net response to opposing
selective pressures.

The diversity of moth behavior around lamps suggests a multitude
of possible mechanisms for reducing adverse effects of electric light.
The degree to which moths of different species fly to lamps may depend
on the degree to which they respond to alternative navigational cues
that compete with the lamps (Janzen 1984). Suppression of flight-to-
light behavior could take the form of increasing responsiveness to com-
peting stimuli such as olfactory, geomagnetic, aerodynamic, gravita-
tional and inertial cues, plus alternative visual cues (Baker & Kuenen
1982, Baker & Mather 1982, Schone 1984, Janzen 1984, Riley & Reyn-
olds 1986). Within a population of moths, variation exists not only in
tendency of different individuals to fly to light, but also in tendency to
linger at the light or fly past it. Variation may also exist in tendencies

VOLUME 42, NUMBER 2 81

to avoid lamps or oviposit near them. Evolutionary changes in response
to electric lighting may be complex.

Forces opposing evolutionary reduction of flight-to-light behavior,
however, are difficult to understand and assess in individual cases.
Studies have employed suction traps to measure aerial densities of moth
populations and at the same time light traps to measure flight to light.
These studies suggest that Xestia (Amathes) c-nigrum (L.) (Noctuidae)
is 5000 times as likely to fly to light as Amphipyra tragopoginis (Cl.)
(Noctuidae) (Taylor & Carter 1961). Why these two noctuids behave
so differently around lamps is a mystery. Failure to evolve seemingly
advantageous adaptations has been well described in Lepidoptera (Ehr-
lich 1984). Populations of moths may resist strong selective pressures
to evolve defenses against adverse effects of electric light.

Fewer moths at urban lamps. Evolutionary changes in wing color-
ation can be documented by inspection of collections of moths obtained
over a period of time (Kettlewell 1973). Evolutionary changes in flight-
to-light behavior cannot be documented in this way. Observations a
century ago, however, are worth noting. Riley (1892: 51) advises col-
lectors where to look for moths: “‘. .. nowadays the electric lights in all
large cities furnish the best collecting places, and hundreds of species
may be taken in almost any desired quantity.”’ Denton (1900:35) was
more explicit:

While employed in Washington, D.C., I made a splendid collection of the moths of
that region simply by going the rounds of a number of electric lights every evening.
The lamps about the Treasury Building were sometimes very productive of fine spec-
imens and the broad stone steps and pillars were frequently littered with moths, May
flies beetles, etc., where one could stand and pick out his desiderata with little difficulty.
I captured several of the Regal Walnut moths (Citheronia regalis) and a number of
our largest and handsomest sphinxes. Besides making the acquaintance of a number
of insects new to me, I met several entomologists who, like myself, had been attracted
to the lights by the abundance of specimens.

Today lamps in big cities such as Washington, D.C., Philadelphia, and
Boston rank among the worst places to collect moths or meet ento-
mologists. Reductions in numbers of moths flying to lamps have been
noted in other locations (Hessel 1976, Muller 1979, Janzen 1983). De-
creases in moths at urban lamps can be explained by many factors,
including declines in moth populations, dilution of moths among thou-
sands of city light sources, and suppression of flight-to-light behavior
as a result of diffuse background light. However, reductions in numbers
of moths flying to urban lamps are what one would expect if urban
moths today were genetically less inclined to fly to lamps than were
those a century ago.

In densely illuminated urban environments, lighting may have fa-
vored species that either fly during the day, do not fly to lamps, or do

82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

not fly at all. Urban pests exemplify such species. These include sesiids
(Engelhardt 1946) and domestic tineids (Ebeling 1978). Species with
flightless females include the bagworm moth, Thyridopteryx ephem-
eraeformis (Haw.) (Psychidae), gypsy moth, Lymantria dispar (L.),
(Lymantriidae), white-marked tussock moth, Orgyia leucostigma (J. E.
Smith) (Lymantriidae), and fall cankerworm, Alsophila pometaria
(Harris) (Geometridae) (Lutz 1941, Drooz 1985). The two urban sa-
turniids, Hyalophora cecropia and Samia cynthia, do not commonly
fly to urban light sources (G. P. Waldbauer pers. comm., Covell 1984).
The extent to which lighting may have influenced the kinds of moths
inhabiting densely illuminated urban environments is unclear.

METHODS TO REDUCE DISTURBANCES

Low-pressure sodium lamps may be used to reduce disturbances
caused by lighting. Low-pressure sodium lamps elicit flight-to-light
behavior less frequently than do other lamps (Robinson 1952). They
do not disturb certain circadian rhythms of Lepidoptera and other
insects (Frank & Zimmerman 1969, Bruce & Minis 1969, Pittendrigh
et al. 1970, Truman 1976). The low-pressure sodium lamp radiates less
energy than does any other kind of lamp of equal illuminance (Finch
1978).

A variety of measures may protect moths from adverse effects of
outdoor lighting. Lamp-free reserves such as sheltered hollows shielded
from lighting have been suggested to save the glow worm, Lampyris
noctiluca L.. (Coleoptera: Lampyridae), a species whose survival in
Britain may be threatened by outdoor lighting (Crowson 1981). To
reduce lighting impact in habitats already exposed to lamps, the most
effective action is to turn off the lamps. Low-pressure sodium lamps
may replace other lamps when illumination is essential. Filters to block
ultraviolet light may be installed over mercury vapor lamps, and shields
may be placed around lamps to block stray light. Low-watt orange-
colored incandescent lamps (“bug lights’) may replace ordinary in-
candescent lamps, but some moths fly to these lamps. Bird feeders may
be removed from windowsills, lampposts, and other sites close to light
sources. “Bug zappers” should be turned off. Natural light-traps such
as open garages may be closed to prevent entry of insects. Operators
of nearby commercial light sources such as illuminated billboards may
be contacted and invited to save money and moths by turning lamps
off during those hours of night and early morning when billboards are
rarely seen.

Although the feasibility of such changes may be questioned, several
North American cities have taken similar steps to reduce light pollution.
Light pollution interferes with astronomical work at observatories (Hen-

VOLUME 42, NUMBER 2 83

dry 1984). These cities have converted streetlamps to low-pressure so-
dium, required ultraviolet-blocking filters over mercury lamps, imposed
curfews on the use of commercial lighting, and mandated shielding of
luminaires (Hendry 1984). Low-pressure sodium lighting, however, has
provoked political controversy on aesthetic and other grounds (San Jose
Committee of the Whole 1980).

CONCLUSION

Effects of outdoor lighting may be divergent. They vary according
to species, lamps, and habitats. Improved levels of illumination may
increase nocturnal vision, but creation of visual artifacts may disturb
vision. Increased numbers of lamps may promote flight-to-light behav-
ior, but high levels of background light may suppress this behavior.
Expansion of streetlighting may increase flight to streetlamps, but shifts
from mercury to sodium lamps may decrease it. Diversion of moths to
lamps may increase numbers of moths in illuminated areas, but diver-
sion of moths away from lamps may decrease numbers. Lamps may
suppress oviposition in the laboratory, but oviposition may increase or
decrease near lamps in the field. Clumping of moths near lamps may
increase predation by birds and bats, but destruction of parasitic wasps
and flies at lamps may decrease predation. Disturbances such as habitat
destruction and urbanization may further confound effects of outdoor
lighting.

Several conclusions emerge from the observations on lighting. Out-
door lighting may destroy vast numbers of individual moths without
apparently suppressing populations of moths. However, it disturbs some
populations more than others, and it disturbs some individuals more
than others in the same population. It generates selective pressures
favoring adaptations for protection against adverse effects of lamps.
The result may be evolutionary changes in behavior, or changes in the
kinds of moths inhabiting illuminated environments. These changes
may increase through time as urban expansion fragments habitats and
exposes smaller moth populations to electric illumination.

Conservation efforts need to consider adverse effects of outdoor light-
ing. If one wishes to protect Lepidoptera in small, endangered habitats
exposed to outdoor lighting, reducing or changing exposure may be
helpful. In such habitats light traps including “bug zappers’” may de-
plete populations of moths. Some cities have attempted to reduce light
pollution to protect astronomical observatories. Whether similar large-
scale restrictions on lighting might help to conserve Lepidoptera has
yet to be demonstrated.

Future research could help clarify lighting impact. Despite abundant
evidence that outdoor lighting affects individual moths, few studies

84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

have attempted to quantify lighting effects on moth populations. Evi-
dence that lighting has suppressed populations of particular moths such
as saturniids is weak. Studies similar to those on the effects of illumi-
nation of orchards and cotton fields (Herms 1929, 1932, Nomura 1969,
Nemec 1969) could be extended to other settings and species. Faunal
surveys, life history studies, and ecological studies could examine Lep-
idoptera in differently illuminated environments. Behavioral and phys-
iological studies could investigate the possible evolution of tolerance to
adverse effects of lighting. The method might include comparison of
Lepidoptera sampled from large geographic regions that possess dif-
ferent levels or kinds of outdoor illumination.

ACKNOWLEDGMENTS

D. F. Schweitzer, D. C. Ferguson, and anonymous reviewers provided valuable infor-
mation and criticism. C. A. Oerkvitz of the City of Philadelphia Department of Streets
provided data on street lighting in Philadelphia. R. L. Edwards introduced me to astro-
nomical literature on light pollution. The National Oceanic and Atmospheric Adminis-
tration provided nocturnal satellite images of the United States. GTE Corporation granted
permission to reproduce spectral distribution graphs for Sylvania lamps. The Illuminating
Engineering Society granted permission to reproduce the spectral distribution graph for
the low-pressure sodium lamp. R. H. Frank and S. E. Frank edited the manuscript. E.
F. Hoeber, T. R. Hoeber, and C. A. Baylor provided special assistance. This paper was
presented in part at a meeting of the American Entomological Society on 18 February
1987.

LITERATURE CITED

AGEE, H. R. 1969. Mating behavior of bollworm moths. Ann. Entomol. Soc. Am. 62:
1120-1122.

1972. Sensory response of the compound eye of adult Heliothis zea and H.

virescens to ultraviolet stimuli. Ann. Entomol. Soc. Am. 65:701—705.

1973. Spectral sensitivity of the compound eyes of field-collected adult bollworms
and tobacco budworms. Ann. Entomol. Soc. Am. 66:613-615.

AGEE, H. R., J. C. WEBB & H. M. Tarr. 1972. Activity of bollworm moths influenced
by the full moon. Environ. Entomol. 1:384-385.

ALLEN, N. & C. R. HODGE. 1955. Mating habits of the tobacco hornworm. J. Econ.
Entomol. 48:526-528.

ANDRESEN, J. W. 1978. Do streetlights turn city trees into late-growing insomniacs?
Am. For. 84:12.

ANKERSMIT, G. W. & P. L. ADKISSON. 1967. Photoperiodic responses of certain geo-
graphic strains of Pectinophora gossypiella (Lepidoptera). J. Insect Physiol. 13:553-
064.

BAKER, R. R. & J. G. MATHER. 1982. Magnetic compass sense in the large yellow
underwing moth, Noctua pronuba L. Anim. Behav. 30:543-548.

BAKER, R. R. & Y. SAaDOvy. 1978. The distance and nature of the light trap response
of moths. Nature 276:818-821.

BAKER, T. C. & L. P.S. KUENEN. 1982. Pheromone source location by flying moths: A
supplementary non-anemotactic mechanism. Science 216:424—427.

BARKER, R. J., C. F. COHEN & A. MAYER. 1964. Photoflashes: A potential new tool for
control of insect populations. Science 145:1195-1197.

BEATY, H. H., J. H. Litty & D. L. CALDERWOOD. 1951. Use of radiant energy for corn
borer control. Agr. Engin. 32:421-422, 426, 429.

VOLUME 42, NUMBER 2 85

BEcK, S. D. 1980. Insect photoperiodism. 2nd ed. Academic Press, New York. 387 pp.

BEEBE, R. 1953. Sampling Michigan Lepidoptera by the fixed light trap. Lepid. News
20:28.

BEEBE, W. 1949. Insect migration at Rancho Grande in north-central Venezuela. General
account. Zoologica 34:107-145.

BEEBE, W. & H. FLEMING. 1951. Migration of day-flying moths through Portachuelo
Pass, Rancho Grande, north-central Venezuela. Zoologica 36:243-266.

BERLINGER, M. J. & G. W. ANKERSMIT. 1976. Manipulation with the photoperiod as a
method of control of Adoxophyes orana (Lepidoptera: Tortricidae). Entomol. Exp.
Appl. 19:96-107.

BERNHARD, C. G. & D. OTTOSON. 1960a. Comparative studies on dark adaptation in
the compound eyes of nocturnal and diurnal Lepidoptera. J. Gen. Physiol. 44:195-
208.

1960b. Studies on the relation between the pigment migration and the sensitivity

changes during dark adaptation in diurnal and nocturnal Lepidoptera. J. Gen. Physiol.

44:205-215.

1964. Quantitative studies on pigment migration and light sensitivity in the
compound eye at different light intensitities. J. Gen. Physiol. 47:465-—478.

BLEsT, A. D. 1957. The function of eyespot patterns in the Lepidoptera. Behavior 11:
209-256.

1968. Longevity, palatability and natural selection in five species of New World
saturniid moth. Nature 197:1183-1186.

BOWDEN, J. & B. M. CHURCH. 1973. The influence of moonlight on catches of insects
in light-traps in Africa. Part I]. The effect of moon phase on light-trap catches. Bull.
Entomol. Res. 63:129-142.

BOWDEN, J. & M. G. Morris. 1975. The influence of moonlight on catches of insects
in light traps in Africa. III. The effective radius of a mercury-vapor light-trap and
the analysis of catches using effective radius. Bull. Entomol. Res. 63:303-348.

BRETHERTON, R. F. 1950. The behavior of moths at light traps: A comment. Entomol.
Gaz. 1:102-104.

1951. Our lost butterflies and moths. Entomol. Gaz. 2:211-240.

1954. Moth traps and their lamps: An attempt at comparative analysis. Entomol.
Gaz. 5:145-154.

BROWER, A. E. 1986. Predation on Catocala moths (Noctuidae). J. Lepid. Soc. 39:280-
283.

Brown, C. H. 1976. A survey of the Sphingidae of Sanibel Island, Florida. J. Lepid.
Soc. 30:230-233.

Brown, L. N. 1984. Population outbreak of pandora moths (Coloradia pandora Blake)
on the Kaibab Plateau, Arizona (Saturniidae). J. Lepid. Soc. 38:65.

BRUCE, V. C. & D. H. MINIs. 1969. Circadian clock action spectrum in a photoperiodic
moth. Science 163:183-1835.

CANTELO, W. W. 1974. Blacklight traps as control agents: An appraisal. Bull. Entomol.
Soc. Am. 20:279-282.

CANTELO, W. W., J. L. GOODENOUGH, A. H. BAUMHOVER, J. S. SMITH JR., J. M. STANLEY
& T. J. HENNEBERRY. 1974. Mass trapping with blacklight: Effects on isolated
populations of insects. Environ. Entomol. 3:389-395.

CANTELO, W. W., J. S. SMITH JR., A. H. BAUMHOVER, J. M. STANLEY & T. J. HENNEBERRY.
1972a. Suppression of an isolated population of the tobacco hornworm with blacklight
traps unbaited or baited with virgin female moths. Environ. Entomol. 1:253-258.

CANTELO, W. W., J. S. SMITH JR., A. H. BAUMHOVER, J. M. STANLEY, T. J. HENNEBERRY
& M. B. PEACE. 1972b. The suppression of isolated populations of sphingids by
blacklight traps. Environ. Entomol. 1:753-759.

1973. Changes in the population levels of 17 insect species during a 32-year
blacklight trapping program. Environ. Entomol. 2:1033-1038.

CaTHEY, H. M. & L. E. CAMPBELL. 1975. Effectiveness of five vision-lighting sources
on photo-regulation of 22 species of ornamental plants. J. Am. Soc. Hort. Sci. 100:
65-71.

86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

CHANDRASHEKARAN, M. K. & W. ENGELMANN. 1976. Amplitude attenuation of the
circadian rhythm in Drosophila with light pulses of varying irradiance and duration.
Intern. J. Chronobiol. 3:221-240.

CLINE, L. D., B. R. FLAHERTY & J. W. PRESS. 1983. Response of selected parasitoids
and predators of stored-grain-product insects to whitelight or blacklight traps. J.
Econ. Entomol. 76:298-301.

COCKERELL, T. D. A. 1914. The cotton-worm moth in Colorado. J. Econ. Entomol. 7:
405.

Co.Luins, D. 1934. Iris-pigment migration and its relation to behavior in the codling
moth. J. Exper. Zool. 69:165-197.

CoLLins, D. & M. W. Nixon. 1930. Responses to light of the bud moth and leaf roller.
New York Agr. Exper. Sta. Geneva Bull. 583. 32 pp.

Cook, L. M.,G.S. MANI & M. E. VARLEY. 1986. Postindustrial melanism in the peppered
moth. Science 231:611-623.

CopPINGER, R. P. 1970. The effect of experience and novelty on avian feeding behavior
with reference to the evolution of warning coloration in butterflies. II. Reactions of
naive birds to novel insects. Am. Nat. 104:323-335.

CovELL, C. V. 1984. A field guide to the moths of eastern North America. Houghton
Mifflin, Boston. 496 pp.

1985. News and notes. Kentucky Lepid. 11(4):8.

Crot, T. A. 1978. Nighttime images of the earth from space. Sci. Am. July:86-98.

Crowson, R. A. 1981. The biology of Coleoptera. Academic Press, New York. 802 pp.

DANTHANARAYANA, W. 1986. Lunar periodicity of insect flight and migration, pp. 88-
119. In Danthanarayana, W. (ed.), Insect flight: Dispersal and migration. Springer-
Verlag, New York. 289 pp.

DANTHANARAYANA, W. & S. DASHPER. 1986. Response of some night-flying insects to
polarized light, pp. 120-127. In Danthanarayana, W. (ed.), Insect flight: Dispersal
and migration. Springer-Verlag, New York. 289 pp.

Davis, B. N. K. 1978. Urbanisation and the diversity of insects, pp. 126-138. In Mound,
L. A. and N. Waloff (eds.), Diversity of insect faunas (Symp. Roy. Entomol. Soc.
London, No. 9). Blackwell Scientific Pub., London.

1982. Habitat diversity and invertebrates in urban areas, pp. 49-63. In Born-
kamm, J.A. Lee, and M.R.D. Seaward (ed.), Urban Ecology (2nd European Ecological
Symposium). Blackwell Scientific Pub., London, 370 pp.

Day, M. F. 1941. Pigment migration in the eyes of the moth, Ephestia kuehniella
Zeller. Biol. Bull. 53:275-291.

DEAY, H. O., J. R. BARRET & J. G. HARSTACK. 1965. Field studies of flight response of
Heliothis zea to electric light traps, including radiation characteristics of lamps used.
Proc. North Central Br. Entomol. Soc. Am. 20:109-116.

DENTON, S. F. 1900. Moths and butterflies of the United States east of the Rocky
Mountains. Vol. 1. The moths. Bradlee Whidden, Boston. 161 pp.

Dirks, C.O. 1937. Biological studies of Maine moths by light trap methods. Bull. Maine
Agr. Exper. Sta. 389. 162 pp.

Drooz, A. T. (ed.). 1985. Insects of eastern forests. U.S. Dept. Agr. Misc. Publ. 1426.
608 pp.

Duray, C. 1964. Contribution a |’étude du phototropisme des lépidoptéres noctuides.
Ann. Sci. Nat. Zool. Paris (12 Sér.) 1:281-406.

EBELING, W. 1978. Urban entomology. University of California, Berkeley. 695 pp.

EDISON ELECTRIC INSTITUTE. 1967. A report on street and highway lighting throughout
the United States. Street and Highway Lighting Committee, Edison Electric Institute,
New York. 29 pp.

1971. Statistical yearbook of the electric utility industry, 1970. Edison Electric

Institute, New York.

1985. Statistical yearbook of the electric utility industry, 1984. Edison Electric
Institute, Washington, D.C. 102 pp.

EGUCHI, E. & T. HoRIkOsHI. 1984. Comparison of stimulus-response (V-log I) functions

VOLUME 42, NUMBER 2 87

in five types of Lepidopteran compound eyes (46 species). J. Comp. Physiol. A 154:
3-12.

EGucuI, E., K. WATANABE, T. HARIYAMA & K. YAMAMOTO. 1982. A comparison of
electrophysiologically determined spectral responses in 35 species of Lepidoptera. J.
Insect Physiol. 28:675-682.

EHRLICH, P. R. 1984. The structure and dynamics of butterfly populations, pp. 24—40.
In Vane-Wright, R. I. and P. R. Ackery (eds.), The biology of butterflies. Academic
Press, London. 429 pp.

ENGELHARDT, G. P. 1946. The North American clear-wing moths of the family Ae-
geriidae. U.S. Nat. Mus. Bull. 190. 222 pp.

FATZINGER, C. W. 1979. Circadian rhythmicity of sex pheromone release by Dioryctria
abietella (Lepidoptera: Pyralidae (Phycitinae)) and the effect of a diel light cycle on
its precopulatory behavior. Ann. Entomol. Soc. Am. 66:1147-1153.

FERGUSON, D. C. 1971. The moths of North America. Fasc. 20.2A. Bombycoidea.
Saturniidae (in part). E. W. Classey, London. 153 pp.

FICHT, G. A., T. E. HIENTON & J. M. Fore. 1940. The use of electric traps in the
control of the European corn borer. Agr. Engin. 21:87-89.

FINCH, D. M. 1978. Atmospheric light pollution. J. lum. Engin. Soc. January:105-117.

ForD, E. B. 1972. Moths. 3rd ed. Collins, London. 266 pp.

FRANK, K. D. 1986. History of the ailanthus silk moth (Lepidoptera: Saturniidae) in
Philadelphia: A case study in urban ecology. Entomol. News 97:41-51.

FRANK, K. D. & W. F. ZIMMERMAN. 1969. Action spectrum for phase shifts of a circadian
rhythm in Drosophila. Science 163:688-689.

FRANKIE, G. W. & L. E. EHLER. 1978. Ecology of insects in urban environments. Ann.
Rev. Entomol. 23:367-387.

GARDINER, B. O. 1982. A silkmoth rearer’s handbook. 3rd ed. Amateur Entomologists’
Society, Hanworth, Middlesex, England. 255 pp.

GEHRING, R. D. & H. F- MADSEN. 1963. Some aspects of the mating and oviposition
behavior of the codling moth, Carpocapsa pomonella. J. Econ. Entomol. 56:140-143.

GEIER, P. W. 1960. Physiologic age of codling moth females (Cydia pomonella L.)
caught in bait and light traps. Nature 185(4714):709.

GENTRY, C. R., W. A. DICKERSON JR., & J. M. STANLEY. 1971. Populations and mating
of adult tobacco budworms and corn earworms in northwest Florida indicated by
traps. J. Econ. Entomol. 64:335-338.

GERARDI, M. H. & J. K. Grimm. 1979. The history, biology, damage, and control of
the gypsy moth Porthetria dispar (L.). Fairleigh Dickinson Univ. Press, Rutherford,
New Jersey. 233 pp.

GLick, P. A. 1961. Light traps for detection, pp. 43-47. In Response of insects to
induced light. U.S. Dept. Agr., Agr. Research Service, ARS 20-10, 66 pp.

1965. Review of collections of Lepidoptera by airplane. J. Lepid. Soc. 19:130-

137.

GLIckK, P. A. & J. P. HOLLINGSWoRTH. 1954. Response of the pink bollworm moth to
certain ultraviolet and visible radiation. J. Econ. Entomol. 47:81-86.

1955. Response of the pink bollworm and other cotton insects to certain ultra-
violet and visible radiation. J. Econ. Entomol. 48:173-177.

GRAHAM, H. M., P. A. GLICK & J. P. HOLLINGSWORTH. 1961. Effective range of argon
glow lamp survey traps for pink bollworm adults. J. Econ. Entomol. 54:788-799.

GRAHAM, H. M., P. A. GLICK & D. F. MarTIN. 1964. Nocturnal activity of adults of
six lepidopterous pests of cotton as indicated by light trap collections. Ann. Entomol.
Soc. Am. 57:328-332.

GTE PRODUCTS CORPORATION (SYLVANIA). 1972. Tungsten halogen lamps. Sylvania
Lighting Center, Danvers, Massachusetts. Engin. Bull. 0-349, 12 pp.

1974. Sylvania incandescent lamps. Sylvania Lighting Center, Danvers, Mas-

sachusetts. Engin. Bull. 0-324, 18 pp.

1977a. Sylvania high intensity discharge lamps; mercury lamps. Sylvania Light-

ing Center, Danvers, Massachusetts. Engin. Bull. 0-346, 20 pp.

88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1977b. High pressure sodium, lumalux, and unalux lamps. Sylvania Lighting

Center, Danvers, Massachusetts. Engin. Bull. 0-348, 16 pp.

1979. Sylvania high intensity discharge lamps. Sylvania Lighting Center, Dan-
vers, Massachusetts. Engin. Bull. 0-344, 22 pp.

HAMILTON, D. W. & L. F. STEINER. 1939. Light traps and codling moth control. J.
Econ. Entomol. 32:867-872.

HARTSTACK, A. W., J. P. HOLLINGSWORTH & D. A. LINDQUIST. 1968. A technique for
measuring trapping efficiency of electric light traps. J. Econ. Entomol. 61:546-552.

HARTSTACK, A. W., J. P. HOLLINGSWorTH, R. L. RipGway & H. H. Hunt. 1971.
Determination of trap spacings required to control an insect population. J. Econ.
Entomol. 64:1090-1100.

HEATH, J. 1974. A century of change in the Lepidoptera, pp. 275-292. In Hawksworth,
L. (ed.), The changing flora and fauna of Britain. Academic Press, London. 461 pp.

HENDRY, A. 1984. Light pollution: A status report. Sky and Telescope June:504—507.

HENNEBERRY, T. J. & M. P. LEAL. 1979. Pink bollworm: Effects of temperature, pho-
toperiod and light intensity, moth age, and mating frequency on oviposition and egg
viability. J. Econ. Entomol. 72:489-492.

Heros, W. B. 1929. A field test of the effect of artificial light on the behavior of the
codling moth Carpocapsa pomonella Linn. J. Econ. Entomol. 22:78-88.

1932. Deterrent effect of artificial light on the codling moth. Hilgardia 7:263-

280.

HESSEL, S. A. 1976. A preliminary scan of rare and endangered Nearctic moths. Atala
4:19-21.

HIENTON, T. E. 1974. Summary of investigations of electric insect traps. U.S. Dept.
Agr. Tech. Bull. 1498. 136 pp.

HOFFMAN, J. D., F. R. LAwson & B. PEACE. 1966. Attraction of blacklight traps baited
with virgin female tobacco hornworm moths. J. Econ. Entomol. 59:809-811.

HOGLUND, G. 1963. Glow, sensitivity changes and pigment migration in the compound
eye of nocturnal Lepidoptera. Life Sciences No. 4, Pergamon Press. Pp. 275-280.

HOGLUND, G. & G. STRUWE. 1970. Pigment migration and spectral sensitivity in the
compound eye of moths. Z. Vergl. Physiol. 67:229-237.

HOLLAND, W. J. 1903. The moth book. Doubleday, New York. 479 pp.

HOLZMAN, R. 1961. Collecting Sphingidae with a mercury vapor lamp. J. Lepid. Soc.
15:191-194.

Hosny, M. M. 1959. Review of results and a complete list of Macrolepidoptera caught
in two ultra-violet light traps during 24 moths at Rothamsted, Hertfordshire. Entomol.
Monthly Mag. 95:226-237.

Howe, W. H. 1959. A swarm of noctuid moths in southeastern Kansas. J. Lepid. Soc.
13:26.

Hoy, M. A. 1978. Variability in diapause attributes of insects and mites: Some evolu-
tionary and practical implications, pp. 101-126. In Dingle, H (ed.), Evolution of
migration and diapause. Springer-Verlag, New York. 284 pp.

Hsiao, H. S. 1972. Attraction of moths to light and infrared radiation. San Francisco
Press, San Francisco. 89 pp.

[ILLUMINATING ENGINEERING SOCIETY OF NORTH AMERICA. 1981. IES lighting hand-
book. 1981. 2 vols. Illuminating Engineering Society of North America, New York.

JAHN, T. L. & F. CRESCITELLI. 1939. The electrical response of the cecropia moth eye.
J. Cell. Comp. Physiol, 13:118-119,

JANZEN, D. H. 1983. Insects, pp. 619-645. In Janzen, D. H. (ed.), Costa Rican natural
history. Univ. Chicago Press, Chicago. 816 pp.

1984. Two ways to be a tropical big moth: Santa Rosa saturniids and sphingids.
Oxford Surveys in Evolutionary Biology I:85-140.

JOHNSON, C. G. 1969. Migration and dispersal of insects by flight. Methuen, London.
763 pp.

JOHNSON, W. T. & H. H. Lyon. 1976. Insects that feed on trees and shrubs. Comstock
Publishing Assoc., Cornell Univ. Press, Ithaca, New York. 463 pp.

Jupp, D. B. 1951. Basic correlates of the visual stimulus, pp. 811-867. In Stevens, S. S.
(ed.), Handbook of experimental psychology. John Wiley, New York. 1436 pp.

VOLUME 42, NUMBER 2 89

KETTLEWELL, B. 1973. The evolution of melanism. Clarendon Press, Oxford. 423 pp.

KLyucHko, Z. F. 1957. On the flight of noctuids (Lepidoptera, Noctuidae) towards
different light sources. Dokl. Biol. Sci. Sec., Proc. Acad. Sci. U.S.S.R. 117:951-954.
(Russian.) Publ. in English by Amer. Inst. Bio. Sci.

Konopka, R. J. & S. BENZER. 1971. Clock mutants of Drosophila melanogaster. Proc.
Nat. Acad. Sci. 68:2112-2116.

Krivpa, W. V. 1980. English sparrows feeding on adult Malacosama americana. News
Lepid. Soc. No. 3, p. 41.

KUDRNA, O. 1986. Butterflies of Europe. Vol. 8. Aspects of the conservation of butterflies
in Europe. AULA-Verlag, Wiesbaden, Federal Republic of Germany. 323 pp.
LANGER, H., B. HAMANN & C. C. MEINECKE. 1979. Tetrachromatic visual system in

the moth Spodoptera exempta (Insecta: Noctuidae). J. Comp. Physiol. 129:235-239.

LANGMAID, J. R. 1959. Moths of a Portsmouth garden—A four-year appreciation. Ento-
mol. Gaz. 10:159-164.

LOHRL, H. 1979. Zur Nahrungsauswahl insektenfressender Vogel. Vogelwelt 100(1/2):
48-54.

LUKEFAHR, M. & J. GRIFFIN. 1957. Mating and oviposition habits of the pink bollworm
moth. J. Econ. Entomol. 50:487-490.

Lutz, F. E. 1941. A lot of insects. Entomology in a suburban garden. Putnam, New
York. 303 pp.

MACARTHUR, R. H. & E.O. WILSON. 1967. The theory of island biogeography. Princeton
Univ. Press, Princeton, New Jersey. 203 pp.

MACFARLANE, J. H. & J. L. EATON. 1973. Comparison of electroretinogram and elec-
tromyogram responses to radiant energy stimulation in the moth Trichoplusia ni. J.
Insect Physiol. 19:811-822.

MARTIN, C. H. & J. S. Houser. 1941. Numbers of Heliothis armigera (Hbn.) and two
other moths captured at light traps. J. Econ. Entomol. 34:555-559.

MAZOKHIN-PORSHNYAKOV, G. A. 1969. Insect vision. Plenum Press, New York. 306 pp.

1975. The attractive effect of light and choice of optimum transmitter for light
traps, pp. 106-111. In Pristavko, V. P. (ed.), Insect behavior as a basis for developing
control measures against pests of field crops and forests. (Russian.) Publ. in English
in 1981 by Amerind Publ. Co., New Delhi, India, for U.S. Dept. Agr. and National
Science Foundation, Washington, D.C.

MIKKOLA, K. 1972. Behavioral and electrophysiological responses of night-flying insects,
especially Lepidoptera, to near-ultraviolet and visible light. Ann. Zool. Fenn. 9:225-
254.

1986. Direction of insect migrations in relation to the wind, pp. 152-171. In
Danthanarayana, W. (ed.), Insect flight: Dispersal and migration. Springer-Verlag,
New York. 289 pp.

MILYANOVSKI, E. S. 1975. Some characteristic features of the attraction of light traps
for Lepidoptera, pp. 117-119. In Pristavko, V. P. (ed.), Insect behavior as a basis for
developing control measures against pests of field crops and forests. (Russian.) Publ.
in English in 1981 by Amerind Publ. Co., New Delhi, India, for U.S. Dept. Agr. and
National Science Foundation, Washington, D.C.

MITCHELL, E. R. & H. R. AGEE. 1981. Response of beet and fall armyworm moths to
different colored lamps in the laboratory and field. J. Environ. Sci. Health A 16:387-
396.

Moore, S. 1955. An annotated list of the moths of Michigan, exclusive of Tineoidea
(Lepidoptera). Mus. Zool. Univ. Mich. Misc. Publ. 88, 87 pp.

MOULDING, J. D. & J. J. MADENJIAN. 1979. Macrolepidopteran moths light-trapped in
a New Jersey oak forest (Lepidoptera). Proc. Entomol. Soc. Wash. 81:135-144.
MULLER, J. 1979. Fourth addition to the supplemental list of Macrolepidoptera of New

Jersey. J. Lepid. Soc. 33:174-178.

NAssic, W. & R. S. PEIGLER. 1984. The life-history of Actias maenas (Saturniidae). J.
Lepid. Soc. 38:114-123.

NELSON, S. O. 1967. Electromagnetic energy, pp. 89-145. In Kilgore, W. W. and R.
L. Doutt (eds.), Pest control: Biologic, physical and selected chemical methods. Ac-
ademic Press, New York.

90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

NEMEC, S. J. 1969. Use of artificial lighting to reduce Heliothis spp. populations in
cotton fields. J. Econ. Entomol. 62:1138-1140.

1971. Effects of lunar phases on light-trap collections and populations of boll-
worm moths. J. Econ. Entomol. 64:860-863.

Nomura, K. 1969. Studies on orchard illumination against fruit-piercing moths. Rev.
Plant Prot. Res. (Tokyo) 2:122-124.

Norris, M. J. 1936. The feeding-habits of the adult Lepidoptera Heteroneura. Trans.
Roy. Entomol. Soc. London 85:61-90.

NorTH AMERICAN PHILIPS LIGHTING CORPORATION. 1982. Technical bulletin on SOX
low pressure sodium lighting. Hightsown, New Jersey. 6 pp.

OwEN, D. F. 1978. Insect diversity in an English suburban garden, pp. 13-30. In
Frankie, G. W. and C. S. Koehler (eds.), Perspectives in urban entomology. Academic
Press, New York. 417 pp.

PFRIMMER, T. R. & M. J. LUKEFAHR. 1955. Experiments with light traps for control of
the pink bollworm. U.S. Dept. of Agr., Agr. Res. Serv. ARS 33-6, 9 pp.

PreTREwIcz, A. T. & A. C. KAmIL. 1979. Search image formation in the blue jay
(Cyanocitta cristata). Science 204:1332-1333.

PITTENDRIGH, C. S., J. H. EICHHORN, D. H. MINIS & V. G. BRUCE. 1970. Circadian
systems VI. Photoperiodic time measurement in Pectinophora gossypiella. Proc. Nat.
Acad. Sci. USA 66:758-764.

PITTENDRIGH, C. S. & D. H. MINIs. 1971. The photoperiodic time measurement in
Pectinophora gossypiella and its relation to the circadian system in that species, pp.
212-250. In Menaker, M. (ed.), Biochronometry. Natl. Acad. Sci., Washington, D.C.
662 pp.

PLaut, H. N. 1971. Distance of attraction of moths of Spodoptera littoralis to BL
radiation, and recapture of moths released at different distances of an ESA blacklight
standard trap. J. Econ. Entomol. 64:1402-1404.

PYLE, R. M. 1975. Silk moth of the railroad yard. Nat. Hist. 84(5):45-51.

PYLE, R. M., M. BENTZIEN & P. OPLER. 1981. Insect conservation. Ann. Rev. Entomol.
26:233-258.

Rau, P & N. Rau. 1929. The sex attraction and rhythmic periodicity in the giant
saturniid moths. Trans. St. Louis Acad. Sci. 26:83-221.

RAULSTON, J. R., S. D. PAIR, F. A. PEDRAZA MARTINEZ, J. K. WESTBROOK, A. N. SPARKS
& V. M. SANCHEZ VALDEZ. 1986. Ecological studies indicating the migration of
Heliothis zea, Spodoptera frugiperda, and Heliothis virescens from Northeastern
Mexico and Texas, pp. 204-220. In Danthanarayana, W. (ed.), Insect flight: Dispersal
and migration. Springer-Verlag, New York. 289 pp.

RIEGEL, K. W. 1973. Light pollution. Science 179:1285-1291.

RILEY, C. V. 1892. Directions for collecting and preserving insects. U.S. Nat. Mus. Bull.
39:51.

RILEY, J. R. & D. R. REYNOLDs. 1986. Orientation at night by high-flying insects, pp.
71-87. In Danthanarayana, W. (ed.), Insect flight: Dispersal and migration. Springer-
Verlag, New York. 289 pp.

RIORDIN, D. F. 1964. High-intensity flash discharge as a source of radiant energy for
sterilizing insects. Nature 204:1332.

ROBINSON, H. S. 1952. On the behaviour of night-flying insects in the neighbourhood
of a bright source of light. Proc. Roy. Entomol. Soc. London (A) 27:13-21.

ROBINSON, H. S. & P. J. M. ROBINSON. 1950a. Some notes on the observed behavior of
Lepidoptera in flight in the vicinity of light sources together with a description of a
light trap designed to take entomological samples. Entomol. Gaz. 1:3-20.

1950b. Reply to Mr. F. Bretherton’s observations. Entomol. Gaz. 1:104—107.

ROBINSON, P. J. M. 1960. An experiment with moths on the effectiveness of a mercury
vapour light trap. Entomol. Gaz. 11:119-132.

SAARIO, C. A., H. H. SHOREY & L. K. GASTON. 1970. Sex pheromones of noctuid moths
XIX. Effect of environmental and seasonal factors on captures of males of Trichoplusia
ni in pheromone-baited traps. Ann. Entomol. Soc. Am. 63:667-672.

SAN JOSE COMMITTEE OF THE WHOLE. 1980. San Jose study and report on low pressure

VOLUME 42, NUMBER 2 91

sodium lighting. Submitted by A. R. Turturici, Director of Public Works, to City of
San Jose, San Jose, California. 84 pp.

SARGENT, T. D. 1973a. Behavioral adaptations of cryptic moths VI. Further experimental
studies on bark-like species. J. Lepid. Soc. 27:8-12.

1973b. Studies on the Catocala (Noctuidae) of southern New England IV. A

preliminary analysis of beak-damaged specimens, with discussion of anomaly as a

potential anti-predator function of hindwing diversity. J. Lepid. Soc. 27:175-192.

1976. Legion of night; the underwing moths. Univ. Massachusetts Press, Amherst,
Massachusetts. 222 pp.

SARGENT, T. D. & R. R. KeEIPER. 1969. Behavioral adaptations of cryptic moths I.
Preliminary studies on bark-like species. J. Lepid. Soc. 23:1-9.

SAUNDERS, D. S. 1982. Insect clocks. 2nd ed. Pergamon Press, Oxford. 409 pp.
SCHAEFER, G. W. 1976. Radar observations of insect flight, pp. 157-197. In Rainey, R.
C. (ed.), Insect flight. Symp. Roy. Entomol. Soc. London 7. Wiley, New York.
SCHAEFER, M. 1982. Studies of the arthropod fauna of green urban ecosystems, pp. 65—
74. In Bornkamm, R., J. A. Lee, and M. R. D. Seaward (eds.), Urban ecology. 2nd

European Ecological Symposium. Blackwell, London. 370 pp.

SCHONE, H. 1984. Spatial orientation. The spatial control of behavior in animals and
man. Princeton Univ. Press, Princeton, New Jersey. 346 pp.

SHOREY, H. H. & L. K. GASTON. 1964. Sex pheromones of noctuid moths III. Inhibition
of male response to the sex pheromone in Trichoplusia ni (Lepidoptera: Noctuidae).
Ann. Entomol. Soc. Am. 57:775-779.

1965. Sex pheromones of noctuid moths VIII. Orientation to light by pheromone-
stimulated males of Trichoplusia ni (Lepidoptera: Noctuidae). Ann. Entomol. Soc.
Am. 58:833-836.

SHROPSHIRE, W. 1977. Photomorphogenesis, pp. 281-312. In Smith, K. C. (ed.), The
science of photobiology. Plenum, New York. 430 pp.

SINNADURAI, S. 1981. High pressure sodium street lights affect crops in Ghana. World
Crops (Nov/Dec):120-122.

SKOPIK, S. D. & M. TAKEDA. 1980. Circadian control of oviposition activity in Ostrinia
nubilalis. Am. J. Physiol. 239:R259-R264.

SNow, J. W., W. W. CANTELO & M. C. BOWMAN. 1969. Distribution of the corn earworm
on St. Croix, U.S. Virgin Islands, and its relation to suppression programs. J. Econ.
Entomol. 62:606-611.

SoRCcAR, P. C. 1982. Energy saving lighting systems. Van Nostrand Reinhold, New
York. 346 pp.

SOTTHIBANDHU, S. & R. R. BAKER. 1979. Celestial orientation by the large yellow
underwing moth, Noctua pronuba L. Anim. Behav. 27:786-800.

SOWER, L. I., H. H. SHoREY & L. K. GAsTon. 1970. Sex pheromones of noctuid moths
XXI. Light : dark cycle regulation and inhibition of sex pheromone release by females
of Trichoplusia ni. Ann. Entomol. Soc. Am. 63:1090-1092.

STANLEY, W. W. 1932. Observations on the flight of noctuid moths. Ann. Entomol. Soc.
Am. 25:366-368.

STERNBURG, J. G., G. P. WALDBAUER & A. G. SCARBROUGH. 1981. Distribution of
cecropia moth (Saturniidae) in central Illinois: A study in urban ecology. J. Lepid.
Soc. 35:304-320.

STEWART, P. A., J. J. LAM & J. L. BLYTHE. 1969. Influence of distance on attraction of
tobacco hornworm and corn earworm moths to radiations of a blacklight lamp. J.
Econ. Entomol. 62:58-61.

STEWART, P. A., J. J. LAM & J. D. HOFFMAN. 1967. Activity of tobacco hornworm and
corn earworm moths as determined by traps equipped with blacklight lamps. J. Econ.
Entomol. 60:1520-1522.

STRADLING, D. J., C. J. Lecc & F. D. BENNETT. 1983. Observations on the Sphingidae
(Lepidoptera) of Trinidad. Bull. Entomol. Res. 73:201-232.

SULLIVAN, W. T. 1984. Our endangered night skies. Sky and Telescope May:412-414.

TAUBER, M. J. & C. A. TAUBER. 1978. Evolution of phenological strategies in insects:
A comparative approach with ecophysiological and genetic implications, pp. 49-71.

92 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

In Dingle, H. (ed.), Evolution of migration and diapause. Springer-Verlag, New York.
284 pp.

TayLor, L. R. & C. I. CarTER. 1961. The analysis of numbers and distribution in an
aerial population of Macrolepidoptera. Trans. Roy. Entomol. Soc. London 113:369-
386.

TayLor, L. R., R. A. FRENCH & I. P. Worwop. 1978. The Rothamsted insect survey
and the urbanization of land in Great Britain, pp. 31-66. In Frankie, G. W. and C.
S. Koehler (eds.), Perspectives in urban entomology. Academic Press, New York. 417

pp.

THAXTER, E. L. 1957. A southern visitor comes north (Sphingidae). Lepid. News 11:
43.

TRUMAN, J. W. 1976. Extraretinal photoreception in insects. Photochem. Photobiol. 23:
215-225.

TURNBULL, A. L. 1964. The search for prey by a web-building spider Achaearanea
tepidariorum (C. L. Koch) (Araneae, Theridiidae). Can. Entomol. 96:568-579.
TUTTLE, J. P. 1985. Maintaining species integrity between sympatric populations of
Hyalophora cecropia and Hyalophora columbia (Saturniidae) in Central Michigan.

J. Lepid. Soc. 39:65-84.

VaiL, P. V., A. F. HOWLAND & T. J. HENNEBERRY. 1968. Seasonal distribution and
mating of female noctuid moths in blacklight trapping studies. Ann. Entomol. Soc.
Am. 61:405-411.

VAISANEN, R. & C. HUBLIN. 1983. The effect of continuous light trapping on moth
populations. A mark-recapture experiment on Hydraecia petasitis (Lepidoptera, Noc-
tuidae). Notulae Entomol. 63:187-191.

WAINWRIGHT, N. B. 1961. History of the Philadelphia Electric Company 1881-1961.
Philadelphia Electric Company, Philadelphia. 416 pp.

WALDBAUER, G. P. 1978. Phenological adaptation and the polymodal emergence pat-
terns of insects, pp. 127-144. In Dingle, H. (ed.), Evolution of migration and diapause.
Springer-Verlag, New York. 284 pp.

WALDBAUER, G. P. & J. G. STERNBURG. 1979. Inbreeding depression and a behavioral
mechanism for its avoidance in Hyalophora cecropia. Am. Mid]. Nat. 102:204—208.

1982. Long mating flights by male Hyalophora cecropia (L.) (Saturniidae). J.
Lepid. Soc. 36:154-155.

WEHNER, R. 1984. Astronavigation in insects. Ann. Rev. Entomol. 29:277-298.

WICKLER, W. 1968. Mimicry in plants and animals. McGraw Hill, New York. 255 pp.

WILLIAMS, C. B. 1937. Butterfly travelers; some varieties migrate thousands of miles.
Nat. Geogr. Mag. 61:568-585.

1939. An analysis of four years captures of insects in a light trap. Part 1. General

survey; sex proportion; phenology; and time of flight. Trans. Roy. Entomol. Soc.

London 86:79-132.

1958. Insect migration. Collins, London. 235 pp.

WILLIAMS, C. B., G. F. COCKBILL & M. E. Gipss. 1942. Studies on the migration of
Lepidoptera. Trans. Roy. Entomol. Soc. Lond. 92:102-283.

WILLIAMS, C. B., R. A. FRENCH & M. M. Hosni. 1955. A second experiment on testing
the relative efficiency of insect traps. Bull. Entomol. Res. 46:193-204.

WILLIAMS, C. B., B. P. SINGH & S. EL ZIADY. 1956. An investigation into the possible
effects of moonlight on the activity of insects in the field. Proc. Roy. Entomol. Soc.
London (A) 31:135-144.

Wo Lr, W. W., A. N. Sparks, S. D. Pair, J. K. WESTBROOK & F. M. TRUESDALE. 1986.
Radar observations and collections of insects in the Gulf of Mexico, pp. 221-234. In
Danthanarayana, W. (ed.), Insect flight: Dispersal and migration. Springer-Verlag,
New York. 289 pp.

WORLD ALMANAC. 1986. Newspaper Enterprise Association, Inc., New York. 928 pp.

Worth, C. B. & J. MULLER. 1979. Captures of large moths by an ultraviolet light trap.
J. Lepid. Soc. 33:261-265.

YAGI, N. & N. KOYAMA. 1963. The compound eye of Lepidoptera. Shinkyo Press, Tokyo.
319 pp.

VOLUME 42, NUMBER 2 93

Yu, Q., A. C. JACQuiIER, Y. CITRI, M. HAMBLEN, J. C. HALL & M. RosBasH. 1987.
Molecular mapping of point mutations in the period gene that stop or speed up
biological clocks in Drosophila melanogaster. Proc. Nat. Acad. Sci. USA 84:784-788.

Received for publication 23 February 1987; accepted 24 December 1987.

Journal of the Lepidopterists’ Society
42(2), 1988, 94-102

HYBRIDIZATION BETWEEN TWO SPECIES OF
SWALLOWTAILS, MEIOSIS MECHANISM,
AND THE GENESIS OF GYNANDROMORPHS

ROBERT BLANCHARD
150 rue des Carmes, 46000 CAHORS, France

AND

HENRI DESCIMON!

Laboratoire de Systématique Evolutive, Université de Provence,
3 place Victor Hugo, 13331 MARSEILLE CEDEX 3, France

ABSTRACT. Hybridization between Papilio machaon L. and P. polyxenes asterius
Stoll was carried out over four generations by backcrossing black female F, and further
hybrids with wild machaon males. A bilateral gynandromorph (symmetrical mosaic for
the black-yellow phenotype) was obtained. In the fourth generation, one brood from a
single female had negligible mortality but yielded an abnormal sex ratio opposite that
predicted by Haldane’s Rule (45 males/86 females, ca. 1:2). The black-yellow character
followed a perfect 1:1 segregation. Reexamination of previous data suggests that meiosis
in Lepidoptera follows an unusual pattern: the sister chromatids segregate during the first
division, and crossing over is frequently absent in females. Bilateral gynandromorphs are
generally due to fertilization of binucleate oocytes. Segregation during the first meiotic
division also can explain the patterns of gynandromorphs arising as autosomal mosaics,
such as those described here.

Additional key words: Papilio machaon, P. polyxenes asterius, Papilionidae, sex ratio.

Hybridization between Papilio machaon L. from Europe and Japan
and P. polyxenes asterius Stoll from the United States was first under-
taken more than 20 years ago (Clarke & Sheppard 1953, 1955, Ae 1966).
Commercial availability of pupae of both species has allowed many
breeders to easily carry out this cross since then. We recently performed
such crosses, and our results are peculiar. Moreover, a spectacular gyn-
andromorph emerged from one of our broods, and we compare this
specimen with other examples recently described by Clarke and Clarke
(1983).

MATERIALS AND METHODS

Papilio polyxenes asterius stocks were established from diapausing
pupae obtained from Chicago, Cook Co., Illinois, and we obtained P.
machaon from Cahors, Lot, France. The insects were hand-paired using
the technique of Clarke (1952). Mated females were fed with a honey-
water mixture (1:10), and allowed to oviposit in a gauze cage on carrot
(Daucus) leaves. Either sunshine or light of a 60 W bulb at a distance
of 20 cm was used to activate the insect. Number of ova laid per female

' Correspondence should be addressed to the second author.

VOLUME 42, NUMBER 2 95

was between 50 and 100, sometimes more. Larvae were reared on carrot
leaves, or occasionally on other Umbelliferae. While no pathology was
noticed in former broods, it has inhibited breeding in recent years. This
fact may be related to extensive use in our neighborhood of the insec-
ticide “Bactospéine”’, which contains strains of Bacillus thuringiensis
Berl. Many diseased pupae and adults showed teratological atrophies
comparable to those induced by toxin of this bacterium (Burgerjon &
Biache 1967). Similar abnormalities also have been observed in pure
strains of Papilio polyxenes asterius bred in the United States (Carter
& Feeny 1985).

We encountered difficulties obtaining functional males in our breed-
ing stocks, even among non-hybrids. Thus, hybrid females were always
used, while the males were pure machaon from wild stocks. Crosses
were performed over four generations.

RESULTS

Several broods comprised the F, generation (2 asterius X 6 machaon),
and gave the same results as those of Clarke and Sheppard’s (1958,
1956) experiments: each offspring was as melanic as asterius and the
anal eye-spot was intermediate. F, backcrosses (? F, x 6 machaon) gave
the expected 1:1 segregation between “black” and “yellow”. In one
brood, a remarkable gynandromorph was obtained (Figs. 1, 2). It is
bilateral, with all of the left side being female with a “black” phenotype,
and the eye-spot very close to machaon. The underside, although me-
lanic, shows a strong machaon influence in distal parts of the wing.
These features are characteristic of this kind of backcross. Markedly
smaller, the right side is mainly male, and extremely machaon-like.
However, on the hindwing, a melanic patch is present in the anal part.
Its shape is complex, and its anterior border coincides with a com-
partment limit (Sibatani 1983) in the middle of the cell. The body is
conspicuously halved in “black” and “yellow”.

The third and fourth generations were obtained by pairing melanic
females from the previous backcross with wild male machaon. As noted
by Clarke and Sheppard (1956), fertility gradually increased. In the
fourth generation, we were fortunate to obtain a large, healthy brood
from a single female: 131 adults from 185 ova. Among them, 65 were
of the “‘yellow” phenotype (18 males, 48 females) and 66 of the “black”
(27 males, 38 females). Therefore, if the “yellow” /“black”’ ratio of 65/
66 is truly 1:1, the sex ratio is strongly distorted (45/86, x? = 12.88,
P < 0.001). The latter proportion is close to a 1:2 ratio. The cross presents
another intriguing feature: the excess of females is more marked in the
“yellow” phenotype than in the “black”, where it does not even reach
a significant level (27/38, x? = 1.86, P < 0.2). Analysis of these data

96 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fics. 1, 2. P. machaon x P. polyxenes asterias F, hybrid, bipartite mosaic and
gynandromorph. 1, Dorsal surface; 2, Ventral surface. Shown at *% natural size.

through a contingency table indicates that this abnormality of distri-
bution is on the borderline of significance (x? = 2.955, 0.10 > P >
0.05).

A second gynandromorph arose in another brood of the fourth gen-
eration; it is a “mosaic’’, with the same “black” phenotype throughout.
Gynandromorphism is apparent only in the parts where the male differs
from the female. This specimen resembles closely those described by
Clarke et al. (1977). No element of symmetry could be observed in this
individual.

DISCUSSION

Some gynandromorphs of Papilio have already been described. A
discussion once arose in the News of the Lepidopterists Society about
interpretation of gynandromorphs of Papilio glaucus L. (Walsten 1977,
Silberglied 1977); a reanalysis of these examples was provided by Clarke
and Clarke (1983). We next review some problems raised by sex genetics
and the origin of gynandromorphs in Lepidoptera.

Morgan and Bridges (1919) showed that, in Drosophila, gynandro-
morphism is due to an irregular disjunction of sex chromosomes, leading
to the loss of an “X” in one of the daughter cells. So, one-half of the
organism would bear an “XX” set and would be female, and the other
an “XO” and would be male (the Y chromosome is considered to bear
very little information in this insect). In only one case in Lepidoptera
has this mechanism been conclusively demonstrated, in the moth Abrax-
as grossulariata (Morgan & Bridges 1919), but Clarke and Clarke (1983)
consider it a very likely explanation in some other cases. In most other
examples, another mechanism seems to be involved: fertilization of a
binucleate oocyte, as explained by Goldschmidt (1931). During meiosis,

VOLUME 42, NUMBER 2 97

Fic. 8. Schematic presentation of the mechanism producing gynandromorphs from
a binucleate oocyte. Z, W: sex chromosomes; A: autosomal stock. pb: polar bodies from
the former division; spz: the two spermatozoa which will fertilize both female pronuclei.

the two successive divisions lead to four haploid nuclei. Normally, three
of them are eliminated. In some cases, which appear to be scarce, but
whose frequency may be increased by certain mutations, two nuclei
remain in the central zone of the oocyte, and both become fertilized.
There follows a juxtaposition of the two eggs, which may have a genetic
composition as different as any combination of two brothers and sisters.
Figure 8 illustrates this phenomenon; we use “Z”’ and “W’”’ for hetero-
chromosomes, ZZ being male and ZW female. This phenomenon has
been observed and photographed by Goldschmidt and Katsuki (1927).

As Robinson (1971) pointed out, this mechanism raises problems
related to chromosome segregation in meiosis. Meiosis may proceed in
two ways:

1) Sister chromatids issuing from the same single parental one may
separate in the first mitosis of meiosis; the second mitosis therefore
dissociates mother- and father-issuing homologous chromosomes.

2) The first mitosis separates mother- and father-issuing sets of chro-
mosomes, and the second one, the sister chromatids.

The second way is considered normal in animals and plants. Of
course, division of the centromere is expected to play a key role in this
phenomenon. Actually, it is very difficult to observe the process cyto-
logically and to demonstrate it genetically. It is only in oocytes that
daughter cells undergo such a dissimilar fate.

The study of gynandromorphs and mosaics originating from binu-
cleate oocytes may provide a clue to the precise order of chromatid
segregation. When such abnormalities arise, they are most likely due
to the two pronuclei issuing from the second mitosis remaining in the

98 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

middle of the oocyte. Gynandromorphs and bipartite mosaics can arise
only if chromosome sets present in each symmetrical fertile pronuclei
are genetically different. This implies that meiosis follows the first way
above.

Cockayne (1935) clearly showed that there are two types of respective
segregation in autosomes and sex chromosomes. 1) In Bombyx mori,
gynandromorphs and uni- or bisexual mosaics are observed with the
same frequency (Goldschmidt & Katsuki 1927). This shows that the
two fertile pronuclei may either be both “Z” or both “W’’, or one “Z”
and one “W’’, and it means that the chromosomes are segregating at
random, likely due to an achiasmatic meiosis. 2) In Argynnis paphia,
Goldschmidt and Fischer (1927) studied a strain where gynandro-
morphism occurred regularly, probably because of a mutation produc-
ing abnormal meiosis. In some cases, the autosomal and sex-conditioned
mutant “valesina”’ was involved in the crosses. In contrast with Bombyx,
“valesina’’-normal mosaics are only observed when there is also gyn-
andromorphism, and no unisexual mosaic occurs in this strain. The
unambiguous conclusion (not stated by Goldschmidt and Fischer or
Cockayne) is that always, when a binucleate oocyte is formed, one
pronucleus bears a “Z” and the other a “W’’—a strong argument in
favor of meiosis with preliminary separation of sister chromatids. In
the sphingid Laothoe populi, and in many other instances, things appear
identical. An illustrative example was recently provided by Platt (1983),
in artificial hybrids of Limenitis arthemis and L. lorquini; he also
interpreted the bipartite mosaic-gynandromorph he obtained by the
“double egg” theory. However, we are reminded of a halved “alba’’-
orange female of Colias croceus figured by Frohawk (1938); since
various kinds of gynandromorphs have been described in this species
(including “alba” female-orange male), this case might rather corre-
spond to the silkworm type; however, mosaics may arise from various
causes and, isolated, this record remains inconclusive.

Previous paragraphs deal only with bipartite gynandromorphs and
mosaics; however, most sexual mosaics are asymmetrical, which can be
explained in two ways: either, in the case of binucleate oocytes, one
pronucleus becomes shifted from the central region of the oocyte, or
sex chromosomes segregate abnormally during further division of em-
bryonic cells. This latter event most likely explains the minute patches
which characterize the bulk of so-called gynandromorphs.

The first gynandromorph described in this study fits perfectly with
the double-oocyte theory, and supports the assumption that meiosis
obeys the first-named way in Papilio. In the double oocyte that gave
rise to this individual, the left pronucleus was ‘““W” and “black”? and
the right one “Z’’ and “yellow” (Fig. 4). The only puzzling point comes

VOLUME 42, NUMBER 2 99

€ BLACK
(| YELLOW

G3\G)) 9

Fic. 4. Scheme of the mechanism leading to the gynandromorph of Figs. 1, 2. Only
the color character-bearing autosome pair is figured.

from the small melanic patch on the right hindwing. It is inadmissible
that it arose from a chromosome loss, since “black”? is dominant and
could not be present in the genetic stock of the right half. Thus it should
be due either to presence of an uneliminated third pronucleus, remnant
from the first division of meiosis, or to an erratic cell coming from the
left half. The second gynandromorph may be best explained by an
atypical segregation of sex chromosomes during embryogenesis. If it
arose from a binucleate oocyte, this would imply that achiasmatic mei-
osis could occur in Papilio as in Bombyx.

According to Suomalainen (1965), the first-named type of meiosis is
determined by the holocentric nature of centromeres, which he has
indeed observed in Lepidoptera; he states also that, in this order, no
crossing over occurs in the female sex. These assumptions have suffered
controversy from Robinson (1971) and White (1978) as remaining un-
demonstrated in the whole of Lepidoptera, but they have been firmly
ascertained for some species, such as Bombyx mori (Tazima 1964),
Heliconius (Turner & Sheppard 1975), Anagasta kuehniella (Traut
1977).

The abnormal sex ratio observed in one brood of the fourth generation

100 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

(which is corroborated by other less quantitative observations made at
the same time) is most difficult to interpret. We have no definitive
explanation to propose; we only offer remarks that may help future
investigations. The distortion goes against Haldane’s (1922) Rule. Here,
it is the heterogametic sex that is favored; the 6/@ ratio is close to 1:2.
It is not possible to explain this discrepancy by the death of one-half
of the males, since the mortality from egg to adult was very low. We
must invoke a possible abnormality during meiosis.

A non-disjunction of “Z’’ chromosomes during oogenesis could pro-
duce a sex ratio distortion in the direction observed. It should give rise
to a proportion of 2 (W, A) oocytes, 1 (ZZ, A) and 1 (O, A). Hence,
following fertilization, we should have 1/2 ZW, 2A normal females,
1/4 ZO, 2A females (such a formula is usually considered to correspond
to females) and 1/4 ZZZ, 2A males. But should the two latter types of
individuals display a viable phenotype? We could not detect any ab-
normality in offspring of the concerned brood. Moreover, if generalized,
this mechanism should give rise to 3/4 females, while we obseryed 45/
86 (x? = 6.5, P < 0.01); therefore, non-segregation acted only partially.

Meiotic drive is another phenomenon which could lead to sex ratio
distortion. This is a preferential segregation of certain chromosomes in
functional gametes (detailed review in Zimmering et al. 1970). Recent
data indicate that this phenomenon is rather widespread, and involves
a higher frequency sex and “B” (heterochromatine) chromosomes. Sex
chromosomes differ in both of our species by a heterochromatic segment
present only in machaon. Moreover, the “W” from asterius does not
pair perfectly with the “Z’’ from either species (Clarke et al. 1977). Do
these peculiarities trigger meiotic drive preferentially directing the “Z”’
towards a polar body? Such a phenomenon would produce normal
karyotypes; being unaware of the problem, we did not check karyotypes.

Moreover, we should consider that sex ratio distortion perhaps affects
the “yellow” phenotype somewhat more; this could mean that there is
“attraction” between the asterius-originated ““W” and the color-con-
trolling autosome which comes from machaon. One possible explanation
is that these two chromosomes possess certain sequences in common,
and that they could pair, at least partly, during meiotic prophase. This
should obviously affect further segregation, the color-bearing autosomes
being the “drivers’’, as is indicated by their overall 1:1 proportion. Both
species should therefore differ by a translocation between sex chro-
mosomes and color-bearing autosomes. This hypothesis is not as fancy
as it may appear at first sight, since in related American species such
as Papilio glaucus, the color-controlling segment itself is carried on the
“W”’ chromosome (Clarke & Clarke 1983).

VOLUME 42, NUMBER 2 101

CONCLUSION

Reexamination of previous data and analysis of the experiments pre-
sented here allow us to conclude that:

1) In butterflies, at least in the vast majority, meiosis obeys a rather
unusual pattern where chromatid segregation follows an order opposite
the normal one. This is also probably true for moths (the Silkworm case
being the most extreme, since in its meiosis it is achiasmatic).

2) Bilateral gynandromorphs arise most often from fertilization of a
double oocyte. The determinism of mosaic gynandromorphs is more
complex and may result from completely different causes.

3) In Papilio, one can carry interspecific crosses over a large number
of generations by using backcrosses, in a kind of “monitored introgres-
sion’.

4) Even in Papilio, however, a residual amount of genetic incom-
patibility occurs. Sex chromosomes are the most sensitive to disturbances
resulting from this incompatibility.

We surmise that some kind of abnormal chromosome segregation
takes place in interspecific crosses, and we hope our findings will stim-
ulate further research on these questions.

ACKNOWLEDGMENT

We thank A. P. Platt who greatly helped us improve redaction of this paper.

LITERATURE CITED

AE, S. A. 1966. A further study of hybrids in the Papilio machaon group. Bull. Jap.
Entomol. Acad. 2:19-30.

BURGERJON, A. & G. BIACHE. 1967. Effets tératologiques chez les nymphes et les adultes
d’insectes dont les larves ont ingéré des doses sublétales de toxine thermostable. C.R.
Acad. Sci. Paris 264:2423-2425.

CarTER, M. & P. FEENY. 1985. Techniques for maintaining a culture of the black
swallowtail butterfly, Papilio polyxenes asterius Stoll (Papilionidae). J. Lepid. Soc.
39:125-133.

CLARKE, C. A. 1952. Hand pairing of Papilio machaon in February. Entomol. Rec. J.
Var. 64:98-100.

CLARKE, C. A. & F. M. M. CLARKE. 1983. Abnormalities of wing pattern in the eastern
tiger swallowtail butterfly, Papilio glaucus. Syst. Entomol. 8:25-28.

CLARKE, C. A., U. MITTwocH & W. TrauT. 1977. Linkage and cytogenetic studies in
the swallowtail butterflies Papilio polyxenes Fab. and Papilio machaon L. and their
hybrids. Proc. Roy. Soc. Lond. B 198:385-399.

CLARKE, C. A. & P. M. SHEPPARD. 1953. Further observations on hybrid swallowtails.
Entomol. Rec. J. Var. Suppl. 65:1-12.

1955. A preliminary report on the genetics of the machaon group of swallowtail

butterflies. Evolution 9:182-201.

1956. A further report on the genetics of the machaon group of swallowtail
butterflies. Evolution 10:66-73.

CockayNnE, E. A. 1935. The origin of gynandromorphs in the Lepidoptera from bi-
nucleate ova. Trans. Roy. Entomol. Soc. Lond. 83:509-521.

102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

FROHAWK, F. G. 1938. Varieties of British butterflies. Ward & Lock, London, 200 pp.

GOLDSCHMIDT, R. B. 1931. Die sexuellen Zwischenstufen. Springer, Berlin. 528 pp.

GOLDSCHMIDT, R. B. & E. FISCHER. 1927. Erblicher Gynandromorphismus bei Schmet-
terlingen. Wilhelm Roux’ Arch. Entw. Mech. Org. 109:1-18.

GOLDSCHMIDT, R. B. & K. KATSUKI. 1927. Erblicher Gynandromorphismus und so-
matische Mosaik Bildung von Bombyx mori. Biol. Zbl. 47:45-54.

HALDANE, J. B. S. 1922. Sex ratio and unisexual sterility in hybrid animals. J. Genet.
12:101-109.

MorcGan, T. H. & C. B. BRIDGES. 1919. The origin of gynandromorphs. Carnegie Inst.
Wash. Publ. 278:1-122.

PLATT, A. P. 1983. Evolution of North American admiral butterflies. Bull. Entomol.
Soc, Amy 29322;

ROBINSON, R. 1971. Lepidoptera genetics. Pergamon Press, Oxford. 687 pp.

SIBATANI, A. 1983. A compilation of data on wing Homeosis in Lepidoptera. J. Res.
Lepid. 22:1—46.

SILBERGLIED, R. 1978. Letter to the editor. News Lepid. Soc. Jan/Feb 1978: 4.

SUOMALAINEN, E. 1965. On the chromosomes of the geometrid moth genus Cidaria.
Chromosoma 16:166-184.

TAZIMA, Y. 1964. The genetics of the silkworm. Logos Press, London. 253 pp.

TRAUT, W. 1977. A study of recombination, formation of chiasmata and synaptonemal
complexes in female and male meiosis of Ephestia kuhniella (Lepidoptera). Genetica
47:135-142.

TURNER, J. R. G. & P. M. SHEPPARD. 1975. Absence of crossing-over in female butterflies
(Heliconius). Heredity 34:265-269.

WALSTEN, D. M. 1977. Letter to the editor. News Lepid. Soc. Sept/Oct 1977: 6.

WuitE, M. J. D. 1973. Animal cytology and evolution. 8rd ed. Cambridge Univ. Press,
Cambridge. 961 pp.

ZIMMERING, S., L. SANDLER & B. NICOLETTI. 1970. Mechanisms of meiotic drive. Ann.
Rev. Genet. 4:409-436.

Received for publication 12 December 1986; accepted 26 January 1988.

Journal of the Lepidopterists’ Society
42(2), 1988, 103-115

EXTERNAL GENITALIC MORPHOLOGY AND COPULATORY
MECHANISM OF CYANOTRICHA NECYBIA (FELDER)
(DIOPTIDAE)

JAMES S. MILLER

Curatorial Fellow, Department of Entomology,
American Museum of Natural History,
Central Park West at 79th Street, New York, New York 10024

ABSTRACT. External genitalia of Cyanotricha necyria (Felder) exhibit characters
that occur in the Notodontidae and Dioptidae. These provide further evidence that the
two groups are closely related. Dissection of two C. necyria pairs in copulo revealed two
features unique among copulatory mechanisms described in Lepidoptera. First, only the
male vesica, rather than the aedoeagus and vesica, are inserted into the female. Secondly,
during copulation the female is pulled into the male abdomen, and his eighth segment
applies dorsoventral pressure on the female’s seventh abdominal segment. This mechanism
is facilitated by a long membrane between the male eighth and ninth abdominal segments.
The first trait is probably restricted to only some dioptid species, while the second may
represent a synapomorphy for a larger group that would include all dioptids, and all or
some notodontids.

Additional key words: Noctuoidea, Notodontidae, Josiinae, functional morphology.

Genitalic structure has been one of the most important sources of
character information in Lepidoptera systematics. Taxonomists often
use differences in genitalic morphology to separate species, and ho-
mologous similarities have provided characters for defining higher cat-
egories in Lepidoptera classification (Mehta 1933, Mutuura 1972, Dug-
dale 1974, Common 1975). Unfortunately, we know little concerning
functional morphology of genitalia. A knowledge of function may aid
in determining homology of genitalic structures, something that has
proved to be extremely difficult and controversial. In addition, a func-
tional approach can provide important new characters for understand-
ing phylogenetic relations. For example, Stekolnikov and Kuznetsov
(1982) used functional morphology of male genitalia to provide char-
acters for higher classification of ennomine geometrids, and Stekolnikov
(1967a) contributed new data concerning familial relations among but-
terflies. In this paper I describe the external genitalia and mechanism
of copulation in a dioptid moth, Cyanotricha necyria (Felder).

Forbes (1939) was among the first to examine musculature of male
genitalia in Lepidoptera, and his study provided the basis for subsequent
research (Birket-Smith 1974). Several workers have described muscu-
lature of male and female butterfly genitalia (Shirozu & Yamamoto
1953, Hannemann 1954a, 1954b; Ehrlich & Davidson 1961, Stekolnikov
1967a), while there have been fewer such studies on moths (Hannemann
1957, Stekolnikov 1967b).

Studies of copulatory mechanisms in Lepidoptera are rare. Perhaps

104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

the first was that of Chapman (1916a, 1916b), who attempted to de-
termine the mechanism of copulation in lycaenids. However, he was
unable to adequately preserve specimens in copulo. Arnold and Fischer
(1977) analyzed genitalic muscle attachments and the method of cop-
ulation in three Speyeria species (Nymphalidae), and De Jong (1978)
described the copulatory mechanism in Carcharodus boeticus Reverdin
(Hesperiidae). Stekolnikov (1965) compared copulatory mechanisms of
four moth species, Spilosoma menthastri Esper (Arctiidae), Acronicta
rumicis L. (Noctuidae), Antheraea pernyi Guérin (Saturniidae), and
Dendrolimus pini L. (Lasiocampidae). In a remarkable series of papers,
Callahan (1958, 1960), Callahan and Chapin (1960), and Callahan and
Cascio (1963) presented a detailed analysis of copulation, spermato-
phore production, and egg formation in Noctuidae. They examined 11]
noctuid species, including Helicoverpa zea (Boddie), Pseudaletia uni-
puncta (Haworth), Peridroma saucia (Hubner), and 8 members of
Plusiinae. Their methods included serial dissection of moth pairs at
various stages during copulation.

The study described here is the first on moths related to notodontids,
and illustrates some unique features concerning their genitalia and
mechanism of copulation. Cyanotricha necyria is a member of Diop-
tidae, a group comprising approximately 400 species of diurnal, Neo-
tropical moths (Bryk 1930, Hering 1925). Although it is acknowledged
that they are closely related to Notodontidae (Franclemont 1970), their
precise phylogenetic position remains unresolved; the group may ul-
timately be reclassified as a notodontid tribe (Minet 1983, Miller 1987,
S. Weller unpubl.). The genus Cyanotricha Prout, which contains only
two species, C. necyria and C. bellona (Druce), was placed by Kiriakoff
(1950) in the dioptid subfamily Josiinae, a well-defined monophyletic
group of approximately 100 species (J. Miller unpubl.). Cyanotricha
necyria (Figs. 1 & 2) is an iridescent blue-green moth with an orange-
brown dash at the forewing base between veins Sc and Rs, and a
forewing length between 15 and 18 mm. It is found from central Peru
N to southern Colombia, whereas the other Cyanotricha species, C.
bellona, which is less common in museum collections, has been recorded
only in central Peru at elevations up to 4200 m. Like many other
members of Josiinae, C. necyria larvae feed on Passiflora (Passiflora-
ceae), and the moth is currently being tested as an agent to control the
spread of P. mollissima (HBK) Bailey, a forest weed in Hawaii (Markin
et al. in press).

METHODS

Two pairs of pinned Cyanotricha necyria, preserved in copulo, were
found in the collection at the United States National Museum. Each

VOLUME 42, NUMBER 2 105

Fics. 1-8. Cyanotricha necyria (Felder) in dorsal view. 1, Male; 2, Female; 3, In
copulo, male at left.

had been prepared by putting a pin through the male thorax, and the
wings of the male and female had been left folded (Fig. 3). According
to label data, both pairs were from the Dognin collection and had been
collected in the “Environs de Loja’, Ecuador, by Abbé Gaujon, one
pair in 1885 and the other in 1886.

For both pairs I used the same dissecting technique. The abdomens
were broken from the male and female thoraces and placed, still joined,
in 10% KOH for 12 h. They were then moved to 70% ethanol, cleaned
of scales and soft tissues, and drawn using a camera lucida attached to
a dissecting microscope. Drawings were made at two points during
dissection: (1) with abdominal segments 1-6 of the male and female
removed; and (2) with abdominal segments 7 and 8 and the left valve

106 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 4. Male terminalia of C. necyria in lateral view, anterior at left. A, aedoeagus;
s8, sternite 8; Su, uncus; T, tegumen; t8, tergite 8; V, valve; Vn, vinculum.

of the male removed, and abdominal segment 7 of the female removed.
These drawings were overlaid to produce Fig. 11.

To better understand genitalic morphology in C. necyria, I dissected
three additional males and three additional females, all from the same
locality and collector as the pairs taken in copulo. The unpaired spec-
imens were prepared and stained using techniques previously described
(Miller 1987). All preparations are in the U.S. National Museum, Wash-
ington, D.C.

Morphological terminology follows Klots (1970), Sibatani (1972), and
Ogata et al. (1957). Rather than follow the recommendation of Ogata
et al. and Sibatani, who proposed the term sociuncus, I use two terms,
socii and uncus, following Klots.

RESULTS AND DISCUSSION
General Features of Cyanotricha necyria Genitalia

External genitalia of C. necyria exhibit features unique to notodontids
and dioptids. These strengthen the argument that the two groups are
closely related. In C. necyria there is a long membrane between segment
8 and the tegumen + vinculum (Fig. 4). The latter are collectively
termed the ring, which is thought to be homologous with abdominal
segment 9 (Snodgrass 1935, Klots 1970). Genitalia in this species are
normally enveloped within the abdomen. In Speyeria, where there is
also extrusion of male genitalia during mating, movement is effected
by protractor and retractor muscles, aided by hemolymph pressure
(Arnold & Fischer 1977). An extremely long membrane between seg-
ments 8 and 9, combined with ability to withdraw genitalia inside the

VOLUME 42, NUMBER 2 107

j<— 1.0 mm —>|

Fics. 5, 6. Male eighth abdominal segment of C. necyria. 5, Tergite (dorsal view);
6, Sternite (ventral view).

abdomen, is typical of dioptids, but is also found throughout Notodon-
tidae (Mehta 1933). This trait may represent a synapomorphy for the
notodontid-related groups.

The male eighth abdominal segment in dioptids and notodontids is
usually modified. In C. necyria there are excavations along the posterior
margins of the tergite and sternite, and apodemes on their anterior
margins (Figs. 5 & 6). In many dioptids and notodontids the posterior
margin of the sternite and tergite is heavily sclerotized, sometimes
bearing spines (J. Miller unpubl.). There is also much variation in shape
of the apodemes on the anterior margin of sternite 8; they are frequently
much longer than in C. necyria.

The sacculus of the valve in C. necyria is large with numerous pleats
(Fig. 7), and the rest of the valve, except for the costa, is membranous.
The pleated sacculus was described by Barth (1955) for Hemiceras
(Notodontidae), but is another feature found frequently in dioptids and
notodontids (Forbes 1942, Holloway 1988, Miller 1987). The pleats
enclose androconia, and probably unfold during courtship, extruding
the scales, which then presumably disseminate male scent. Mehta (1933)
characterized notodontids as lacking the saccus, an internal extension
of the vinculum. Male genitalia of C. necyria illustrate that the saccus
is absent in some dioptids as well. The slender uncus and socii are
hinged on the tegumen.

The aedoeagus of C. necyria (Fig. 8) is typical in shape for members
of Josiinae, being short, deep dorsoventrally, and large relative to the
rest of the genitalia. A row of cornuti on the vesica, terminating in a
set of robust, spinelike cornuti, is also common in the group (J. Miller
unpubl.).

108 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

C

Fics. 7, 8. Male genitalia of C. necyria. 7, Genitalia in posterior view with aedoeagus
removed; 8, Aedoeagus in lateral view (anterior at left). At, anal tube; C, cornuti; Cs,
costa of valve; D, diaphragma; E, opening of vesica; Sc, saccus; Sl, sacculus; U, uncus; T,
tegumen; Th, ventral tooth of aedoeagus; Tr, transtilla; Vn, vinculum; Vs, vesica.

In female genitalia of C. necyria (Fig. 9), tergite 8 is membranous
dorsally. The ostium is surrounded by postvaginal and antevaginal plates,
which hinge on a point dorsal to the opening. There are small spines
inside the proximal portion of the corpus bursae. A feature found in
C. necyria and only a few other dioptids is the large, convoluted,
sclerotized band which wraps around the corpus bursae. In noctuids,
large muscles attach to the corpus (Callahan & Cascio 1963). Once the
male has deposited the spermatophore in the corpus bursae, these mus-

VOLUME 42, NUMBER 2 109

~— 1.0 mm —>|

Fic. 9. Female genitalia of C. necyria in lateral view, anterior at right. Aa, anterior
apophyses; Ap, posterior apophyses; Av, antevaginal plate; B, sclerotized band; Cb, corpus
bursae; Db, ductus bursae; Ds, ductus seminalis; P, papillae anales; Pv, postvaginal plate;
S, signum; Sp, basal spines of corpus bursae.

cles are thought to squeeze seminal fluid and sperm into the ductus
seminalis. The sclerotized band of C. necyria, in conjunction with these
muscles, may serve to break up the spermatophore. In C. necyria the
ductus seminalis is located laterally on the corpus bursae, whereas in
most dioptids it is located on the ductus bursae (J. Miller unpubl.). The
signum is composed of a group of long spines protruding into the corpus
bursae from a concave sclerotized region. Petersen (1907) and Callahan
(1958) suggested that the signum functions to hold the spermatophore
in place. It is a site of muscle attachment in Helicoverpa zea (Callahan
& Cascio 1968).

Copulatory Mechanism of Cyanotricha necyria

The interrelations of male and female genitalia during copulation
are shown in Figs. 10 and 11. Between the papillae anales of the female,
a membranous invagination allows for insertion of the male uncus,
which is reflexed downward. In these preparations the uncus almost
engages the postvaginal plate of the female. In freshly preserved ma-
terial with the musculature intact, it most likely would do so. Stekolnikov
(1965) found that the uncus engages the postvaginal plate in Spilosoma
menthastri; the configuration he described is almost identical with that
of C. necyria. Stekolnikov stated that the uncus in Acronicta rumicis
engages the female’s eighth sternite, but his illustration suggests that
the ‘eighth sternite’ in A. rwmicis is the same structure as the postvaginal
plate in Spilosoma and Cyanotricha.

110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Mt7 Mts Ft7 11

Ms8&

Fics. 10, 11. Lateral view of male and female C. necyria in copulo, anterior of male
at left, anterior of female at right. 10, Pair #1 with abdominal segments 1-8 and left
valve of male removed, and abdominal segments 1-7 of female removed; 11, Pair #2
with abdominal segments 1-6 and left valve of male removed, and abdominal segments
1-6 of female removed. A, aedoeagus; C, cornuti; Fs7, female sternite 7; Ft7, female
tergite 7; Ms7, male sternite 7; Ms8, male sternite 8; Mt7, male tergite 7; Mt8, male tergite
8; O, ostium bursae of female; Si, socii; Tr, transtilla; U, uncus; V, valve; Vs, vesica.

VOLUME 42, NUMBER 2 Lyd

In C. necyria the socii rest on top of the papillae anales during
copulation and would seem to apply downward pressure on them (Figs.
10 & 11). A large muscle (“#1” in Forbes 1939) has its origin on the
tegumen and its insertion at the base of the socii. This muscle has been
observed in all Lepidoptera studied, and acts to flex the socii and uncus
(Stekolnikov 1965, Arnold & Fischer 1977).

Judging from their position, the valvae of C. necyria apply lateral
pressure on the female terminal segments. The sacculus is elongate and
fairly rigid in most Lepidoptera. Muscles originate on the sacculus and
insert on the clasper of the valve. When these are flexed, the claspers
squeeze the female laterally (Forbes 1939, Arnold & Fischer 1977). The
valve of C. necyria has a membranous sacculus and lacks a clasper (Fig.
7). It may be that only the valval costa provides traction during cop-
ulation.

Eversion of the vesica is effected by the combined forces of aerostatic
pressure and muscle action (Callahan 1958). In Noctuidae the cornuti
appear to serve two functions (Callahan 1958, Callahan & Chapin 1960):
First, while the vesica is being everted, the cornuti, which at this time
point inward, help drag the formed collum of the spermatophore into
the ductus bursae. Secondly, when the vesica is fully everted and the
cornuti point outward, they help manipulate the spermatophore so that
it properly orients in the corpus bursae. Shape and orientation of the
spermatophore is extremely specific in lepidopteran species (Williams
1940, 1941, Callahan 1960). Usually its aperture is placed in close
proximity to the opening of the female’s ductus seminalis. In addition,
the movements of the vesica within the corpus bursae can be extremely
complex. Callahan and Chapin (1960) argued that there is a “lock and
key” mechanism at work during copulation that serves to inhibit mating
between species. However, their research convinced them that it is not
the relative shapes of the male valvae and female genitalia that is
critical, as most previous authors had proposed, but is instead the con-
figuration of the everted vesica and its ability to correctly place the
spermatophore.

Unlike Noctuidae (Callahan & Chapin 1960, Takeuchi & Miyashita
1975) and Arctiidae (Stekolnikov 1965), the aedoeagus of C. necyria
does not actually enter the female, but a small ventral tooth on the
aedoeagus (Fig. 8) appears to insert into the antevaginal plate (Figs. 10
& 11). The diaphragma of C. necyria holds the aedoeagus tightly in
place, whereas in many other Lepidoptera it is loose and allows the
phallus to penetrate the female when the aedoeagus protractor muscles
are activated (Forbes 1939, Stekolnikov 1965, Arnold & Fischer 1977).
Opposing muscles insert on the saccus, and lack of movement of the
aedoeagus in C. necyria may account for absence of the saccus. Judging

112 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

from the morphology of the diaphragma and aedoeagus in other diop-
tids, the characteristic of having only the vesica enter the female may
define a restricted group of species. In copulating C. necyria, cornuti
of the everted vesica were in apposition with basal spines of the corpus
bursae (Figs. 10 & 11). This seemed to hold male and female genitalia
together even after the left valve of the male had been removed. The
two sets of spines may become entangled.

During copulation, the male genitalia of C. necyria are withdrawn
into the abdomen to a point approximately even with segment 7 (Fig.
11). This is facilitated by the long intersegmental membrane between
segment 8 and the ring (Fig. 4). It would be useful to know which
muscles pull the genitalia in. Their morphology may prove to be another
unique feature of dioptids and notodontids. In C. necyria the tergite
and sternite of male segment 8 have an important holding function;
when the female is pulled into the male abdomen, they appear to apply
dorsoventral pressure on her seventh segment. Highly modified male
eighth tergites and sternites are found in many notodontid and dioptid
species (Franclemont 1970, Holloway 1983), which suggests that a hold-
ing function is typical for the group, and possibly represents a synap-
omorphy for the entire lineage.

CONCLUSIONS

One feature of copulation seems common to all lepidopterans studied:
the male uncus is inserted between the papillae anales and applies
pressure on the dorsal surface of the female’s postvaginal plate. Other
aspects are unique to each species. The female of Speyeria is held at
three points: the uncus secures the tergum of segment 8, valvae apply
lateral pressure on the papillae anales, and the base of the valve secures
sternite 7 of the female (Arnold & Fischer 1977). In Carcharodus the
intersegmental membrane between segments 7 and 8 of the female is
expanded. The uncus engages the postvaginal plate, and valvae grip
the female’s intersegmental membrane (De Jong 1978). There are two
points of contact in Spilosoma and Acronicta: the uncus secures the
female postvaginal plate, and valvae apply lateral pressure at the base
of the ductus bursae (Stekolnikov 1965). My study has shown that the
female of Cyanotricha necyyria is held in three places: the uncus engages
the female postvaginal plate, valvae grasp her terminal segments lat-
erally, and the male eighth abdominal segment applies dorsoventral
pressure on female segment 7. A fourth possible point is the cornuti of
the vesica, which seem to become entangled with spines located at the
base of the corpus bursae, but dissection of freshly preserved material
is required to confirm this.

Copulation in C. necyria is unique among Lepidoptera so far de-

VOLUME 42, NUMBER 2 113

scribed in that an exceptionally long membrane between abdominal
segments 8 and 9 of the male allows the female to be pulled into the
abdomen during copulation. The male eighth segment then aids in
grasping the female, and may provide the majority of force for holding
the pair together. This trait could prove to be another synapomorphy
for the dioptid-notodontid lineage, but its distribution among species
has not been adequately documented. Such information may be crucial
in clarifying phylogenetic relations among these taxa.

ACKNOWLEDGMENTS

Support was provided by a Kalbfleisch Curatorial Fellowship from the American Mu-
seum of Natural History, and by a Smithsonian postdoctoral fellowship. I thank F. H.
Rindge, J. G. Franclemont, Susan Weller, Ian Kitching, and an anonymous reviewer for
comments on the manuscript. Marcus Matthews (British Museum) and George Markin
(U.S. Dept. Agr., Hawaii) kindly provided locality data for C. necyria and C. bellona.
Adult photographs were taken by Victor Krantz of the U.S. National Museum. Study of
specimens at the National Museum was made possible through the generosity of Robert
Robbins and Robert Poole.

LITERATURE CITED

ARNOLD, R. A. & R. L. FISCHER. 1977. Operational mechanisms of copulation and
oviposition in Speyeria (Lepidoptera: Nymphalidae). Ann. Entomol. Soc. Am. 70:
455-468.

BARTH, R. 1955. Maennliche Duftorgane brasilianischer Lepidopteren. 10. Mitteilung:
Hemiceras proximata Dogn. (Notodontidae). An. Acad. Bras. Ciéne. 27:539-544.

BIRKET-SMITH, S. J. R. 1974. Morphology of the male genitalia of Lepidoptera. I.
Ditrysia. Entomol. Scand. 5:1-22.

BryYk, F. 1930. Dioptidae. In Strand, E. (ed.), Lepidopterorum Catalogus (42). Junk,
Berlin. 65 pp.

CALLAHAN, P. S. 1958. Serial morphology as a technique for determination of repro-
ductive patterns in the corn earworm, Heliothis zea (Boddie). Ann. Entomol. Soc.
Am. 51:413-428.

1960. A morphological study of spermatophore placement and mating in the
subfamily Plusiinae (Noctuidae, Lepidoptera). Proc. XI Int. Congr. Entomol. (Vienna)
1:339-345.

CALLAHAN, P. S. & T. Cascio. 1963. Histology of the reproductive tracts and trans-
mission of sperm in the corn earworm, Heliothis zea. Ann. Entomol. Soc. Am. 56:
535-556.

CALLAHAN, P. S. & J. B. CHAPIN. 1960. Morphology of the reproductive systems and
mating in two representative members of the family Noctuidae, Pseudaletia uni-
puncta and Peridroma margaritosa, with comparison to Heliothis zea. Ann. Entomol.
Soc. Am. 53:763-782.

CHAPMAN, T. A. 1916a. On the pairing of the Plebeiid blue butterflies (Lycaenidae,
tribe Plebiidi). Trans. Entomol. Soc. London 1916:156-180.

1916b. The rein-sheath in Plebeiid blues: A correction of and addition to paper
VI. Trans. Entomol. Soc. London 1916:297-300.

Common, I. F. B. 1975. Evolution and classification of the Lepidoptera. Ann. Rev.
Entomol. 20:183-208.

DUGDALE, J. S. 1974. Female genital configuration in the classification of Lepidoptera.
N.Z. J. Zool. 1:127-146.

EHRLICH, P. R. & S.E. Davipson. 1961. The internal anatomy of the Monarch butterfly
(Danaus plexippus) L. (Lepidoptera: Nymphalidae). Microentomol. 24:85-133.
FORBES, W. T. M. 1939. The muscles of the lepidopterous male genitalia. Ann. Entomol.

Soc. Am. 32:1-10.

114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1942. Family Notodontidae, pp. 235-318. The Lepidoptera of Barro Colorado
_ Island, Panama. Bull. Mus. Comp. Zool. 85:99-322.

FRANCLEMONT, J. G. 1970. Dioptidae, pp. 9-11. In Dominick, R. B. et al. (eds.),
Prospectus of the moths of America north of Mexico. E. W. Classey Ltd. and Wedge
Entomological Research Foundation, London.

HANNEMANN, H. J. 1954a. Zur funktionellen Anatomie des mannlichen Kopulations-
apparates von Argynnis paphia L. Zool. Anz. 152:266-274.

1954b. Zur Muskelfunktion der weiblichen Genitalsegmente von Argynnis pa-

phia L. Zool. Anz. 153:149-154.

1957. Die mannlichen Terminalia von Micropteryx calthella L. (Lep. Microp-
terygidae). Deutch. Entomol. Z. 4:209-222.

HERING, E.M. 1925. Dioptidae, pp. 501-534. In Seitz, A. (ed.), Die Gross-schmetterlinge
der Erde. Vol. 6. Alfred Kernen, Stuttgart.

HoLioway, J. D. 1983. The moths of Borneo: Family Notodontidae. Malay. Nat. J. 37:
1-107.

Jonc, R. DE. 1978. Functional morphology of the genitalia of Carcharodus boeticus
stauderi Rev. (Lepidoptera, Hesperiidae). Neth. J. Zool. 28:206-212.

KIRIAKOFF, S. G. 1950. Recherches sur les organes tympanique des Lépidoptéres en
rapport avec la classification. III. Dioptidae. Bull. Ann. Soc. Entomol. Belg. 86:67-
86

K.Lots, A. B. 1970. Lepidoptera, pp. 115-130. In Tuxen, S. L. (ed.), Taxonomist’s glossary
of genitalia in insects. Munksgaard, Copenhagen. 359 pp.

MARKIN, G. P., R. F. NAGATA, M. RojJAS DE HERNANDEZ & E. E. TRUJILLO. 1988.
Biology and behavior of the South American moth, Cyanotricha necyria (Felder)
(Lepidoptera: Notodontidae), a potential biocontrol agent in Hawaii of the forest
weed, Passiflora mollissima (HBK) Bailey. Proc. Hawaiian Entomol. Soc. In press.

MEHTA, D. R. 1933. Comparative morphology of the male genitalia in Lepidoptera.
Rec. Indian Mus. (Calcutta) 35:197-266.

MILLER, J. S. 1987. A revision of the genus Phryganidia Packard, with description of
a new species (Lepidoptera: Dioptidae). Proc. Entomol. Soc. Wash. 89:303-321.
MINET, J. 1983. Eléments sur la systematique des Notodontidae et nouvelles données
concernant leur étude faunistique 4 Madagascar (Lepidoptera: Noctuoidea). Bull.

Soc. Entomol. Fr. 87:354—870.

Muruura, A. 1972. Morphology of the female terminalia in Lepidoptera, and its
taxonomic significance. Can. Entomol. 104:1055-1071.

OcaTA, M., Y. OKADA, H. OKAGAKI & A. SIBATANI. 1957. Male genitalia of Lepidoptera:
Morphology and nomenclature. III. Appendages pertaining to the tenth somite. Ann.
Entomol. Soc. Amer. 50:237-244.

PETERSEN, W. 1907. Uber die Spermatophoren der Schmetterlinge. Z. Wiss. Zool. 88:
117-130.

SHIROZU, T. & H. YAMAMOTO. 1953. Morphology of the male genital organs of Argy-
ronome laodice japonica Ménétriés (Lepidoptera: Nymphalidae). Sieboldia (Acta
Biol., Fukuoka) 1:161-168.

SIBATANI, A. 1972. Male genitalia of Lepidoptera: Morphology and nomenclature IV.
Notes on Tuxen’s “Taxonomist’s Glossary of Genitalia in Insects’: Second enlarged
edition. J. Lepid. Soc. 26:117-122.

SNODGRASS, R. E. 1935. Principles of insect morphology. McGraw-Hill, New York. 667
PP.

STEKOLNIKOV, A. A. 1965. Functional morphology of the copulatory apparatus in some
Lepidoptera. Entomol. Rev. 44:143-149.

1967a. Phylogenetic relationships within the Rhopalocera on the basis of the

functional morphology of the genital apparatus. Entomol. Rev. 46:1-11.

1967b. Functional morphology of the copulatory apparatus in the primitive
Lepidoptera and general evolutionary trends in the genitalia of the Lepidoptera.
Entomol. Rev. 46:400-409.

STEKOLNIKOV, A. A. & V. I. KUZNETSOV. 1982. Functional morphology of the male

VOLUME 42, NUMBER 2 ES

genitalia and a new tribal division of the Ennominae (Lepidoptera, Geometridae).
Entomol. Rev. 61:92-122.
TAKEUCHI, S. & K. MIyASHITA. 1975. The process of spermatophore transfer during
the mating of Spodoptera litura F. Jap. J. Appl. Entomol. Zool. 19:41-46.
WILLIAMS, J. L. 1940. The anatomy of the internal genitalia and the mating behavior
of some lasiocampid moths. J. Morphol. 67:411-438.
1941. The relations of the spermatophore to the female reproductive ducts in
Lepidoptera. Entomol. News 52:61-65.

Received for publication 27 September 1987; accepted 8 February 1988.

Journal of the Lepidopterists’ Society
42(2), 1988, 116-119

A NEW SPECIES OF CATOCALA FROM THE
SOUTHEAST UNITED STATES

VERNON A. BROU JR.
137 Jack Loyd Rd., Abita Springs, Louisiana 70420

ABSTRACT. Catocala charlottae is described from Louisiana and Florida, the type
series consisting of 100 males and 64 females. The new species is differentiated from its
most similar ally, C. alabamae Grote, mainly by genitalic characters. Adults and genitalia
of both sexes of both species are illustrated.

Additional key words: Noctuidae, Catocala charlottae, C. alabamae, taxonomy, un-
derwings.

The small-bodied underwing described here, Catocala charlottae,
closely resembles C. alabamae Grote both superficially and in male
genitalia. Catocala charlottae has been taken at the type locality in
Louisiana, and at several Florida localities.

Louisiana C. alabamae are similar to those occurring through most
of its known range. A lifesize color photo of the C. alabamae holotype
in the British Museum (Natural History) was examined and it precisely
matched Louisiana C. alabamae.

Catocala charlottae Brou, new species

(Figs. 1, 2, 5, 6)

Forewing length of males averaging 19.5 mm (18.5-21.2 mm, N = 54); of females,
20.9 mm (19.7—21.8 mm, N = 32). Forewing slate gray with distinct bold black antemedial
line and anal dash. Most specimens have a bold medium brown broad line paralleling
basal side of antemedial line, absent above R,. Same brown coloring evident between
postmedial and subterminal line and especially noticeable as a distinct brown patch below
anal dash. Brown spot at middle of costal margin above vein R,. Reniform and subreniform
present, sometimes diffuse and indistinct. Forewing underside exhibiting a pale yellow
postmedial band bordered on both sides with dark brown bands. Fringe dark with darker
brown bars. Basal half of forewing stronger orange-yellow than outer half with fine black
line on Cu,.

Hindwing above with black inner band and outer marginal band with connecting
black-barred off-white fringe. Underside with yellow postmedial band bordered on both
sides with dark brown bands. Yellow on costal half of hindwing pale, while that half
along inner margin is bolder orange-yellow.

Male genitalia (Fig. 5) (N = 12). Cucular areas along costal margin of valva sickle
shaped; mid-costal edge minimally squared, terminal edge finely serrated. Uncus semi-
circular and acuminate.

Female genitalia (Fig. 6) (N = 10). Papillae anales elongated, strongly sclerotized.
Posterior edge of lamella antevaginalis straight, abruptly angled inwardly to ostium bursae
in a long narrow V-shape.

_ Flight period. At the type locality, specimens were taken at light and fermented bait
from 30 April to 23 June, with peak occurrence on 22 May (N = 177). Specimens taken
after the fourth week usually were worn and tattered.

Discussion. In Louisiana, C. charlottae appears on the wing about two weeks earlier
than C. alabamae. In Louisiana, adult C. alabamae were taken from 13 May to 16 June
(N = 38), with peak occurrence on 2 June.

Both Louisiana and Florida populations of C. charlottae are consistent in maculation

VOLUME 42, NUMBER 2 TAF

Fics. 1-4. Catocala adults from the C. charlottae type locality. 1, C. charlottae, 6
holotype; 2, C. charlottae, ? allotype; 3, C. alabamae, é coll. 13 May 1985; 4, C. alabamae,
2 coll. 10 June 1985.

and size; the only noticeable exterior difference is the slightly darker appearance of
Florida specimens. Forewing lengths of male Louisiana C. charlottae (N = 108) average
7% larger than those of male Louisiana alabamae which average 18.2 mm (16.6-19.5
mm, N = 20). Forewing lengths of female Louisiana C. charlottae (N = 32) average 9%
larger than those of female Louisiana C. alabamae which average 19.1 mm (17.9-20.1
mm, N = 14). The upper forewings of C. charlottae lack the overall blue-green suffusion
present on C. alabamae. Occasionally, fresh C. charlottae exhibit a few diffuse greenish
scales around the reniform, but these are sometimes evident only with magnification.

Male genitalia of C. charlottae are similar to C. alabamae (N = 8) except that the mid-
costal margin is squared to a lesser degree in the former (Figs. 5, 7). Female genitalia of
C. charlottae differ more noticeably from those of C. alabamae (N = 6). In the latter,
the posterior edge of the lamella antevaginalis is angled caudally approaching the midline
and abruptly angled inwardly to ostium bursae in a wide V-shape (Fig. 8).

Since C. charlottae has been collected with typical C. alabamae in both Louisiana and
Florida localities, a distance of 660 miles (1062 km), it does not seem likely that the
former is part of a phenotypic cline of the latter.

More than 30 Catocala species have been collected at the C. charlottae type locality.
This habitat is a longleaf pine region, a gently rolling hilly area interspersed with flatwoods
and sloughs. It is rich in diverse natural vegetation, the secondary growth being so dense
that it is impenetrable except in slough areas. The C. charlottae larva may be a Rosaceae
feeder. Four possible host species common at the type locality are Prunus serotina Ehrh.,
Malus angustifolia (Ait.), Crataegus marshallii Ellgeston, and Aronia arbutifolia (L.).

118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 5-8. Catocala genitalia from the C. charlottae type locality. 5, C. charlottae,

8 coll. 10 May 1986; 6, C. charlottae, 2 coll. 15 May 1986; 7, C. alabamae, 6 coll. 2 June
1986; 8, C. alabamae, ¢ coll. 3 June 1986.

Catocala alabamae has the broader geographic range, occurring in Missouri, Texas,
the southwestern States, the Gulf States, Tennessee, South Carolina, and Florida (Barnes
& McDunnough 1918, Holland 1903:269, Sargent 1976:70, Covell 1984:315).

Types. Holotype 4 (Fig. 1) 4.2 miles (6.7 km) NE Abita Springs, sec. 24,T6SR12E, St.
Tammany Parish, Lousiana, 7 May 1985, V. A. Brou Jr.; allotype 2 (Fig. 2) same data,
30 April 1985; Both in United States National Museum, Washington, D.C. Paratypes:
same locality, 96 4, 56 2, 30 April to 23 June 1983-87; Jacksonville, Duval Co., Florida,

VOLUME 42, NUMBER 2 119

3 6,5 2, 15 May to 7 June 1977-85; Seminole Jr. College, Seminole Co., Florida, 1 2, 21
May 1974; Sanford, Seminole Co., Florida, 1 2, 13 May 1985. Paratypes are deposited in
the Florida State Collection of Arthropods, Gainesville; Louisiana State University, Baton
Rouge; and the author's collection.

ACKNOWLEDGMENTS

I thank the following for assistance: H. D. Baggett, Gainesville, Florida; D. C. Ferguson,
Systematic Entomology Laboratory, Agricultural Research Service, Washington, D.C.; H.
A. Freeman, Garland, Texas; R. M. Gilmore, Winter Springs, Florida; E. L. Quinter,
American Museum of Natural History, New York, New York; J. R. Slotten, Jacksonville,
Florida; G. Strickland, Baton Rouge, Louisiana.

LITERATURE CITED

BARNES, W. & J. MCDUNNOUGH. 1918. Illustrations of the North American species of
the genus Catocala. Mem. Am. Mus. Nat. Hist. (New Ser.) 3(1):1-47.

COVELL, C. V. 1985. A field guide to the moths of eastern North America. Houghton
Mifflin, Boston. 496 pp.

HOLLAND, W. J. 1903. The moth book. Doubleday, Page & Co., New York. Reprinted
1968, Dover, New York. 479 pp.

SARGENT, T. D. 1976. Legion of night: The underwing moths. Univ. Massachusetts
Press, Amherst. 222 pp.

Received for publication 26 January 1987; accepted 10 February 1988.

Journal of the Lepidopterists’ Society
42(2), 1988, 120-131

BIOLOGY OF THE BLUEBERRY LEAFTIER
CROESIA CURVALANA (KEARFOTT) (TORTRICIDAE):
A FIELD AND LABORATORY STUDY

B. M. PONDER AND W. D. SEABROOK

Department of Biology, University of New Brunswick,
Fredericton, New Brunswick E3B 5A8, Canada

ABSTRACT. Biology of Croesia curvalana (Kearfott) is described for the first time.
Laboratory-laid eggs were white, later brown, 0.6 mm in diam., and were deposited
singly under blueberry branches. Seventy-five percent hatched when given a 7-day chilling
at 6°C followed by a 24-week cold treatment at 0°C. Four instars occurred during the
21-day larval development period at 21°C. Male pupal stage was 9 days, 2 days less than
females. Field eggs were laid on surface litter under blueberry plants in July and August,
and eggs overwintered. Flower buds were invaded by emerging larvae in the last part
of April and early May, and pupation occurred during the first half of June. During four
years of study, the flight season began at Blackville, New Brunswick, during the first
week of July, and later at Pouch Cove, Newfoundland. Larvae from Pouch Cove were
parasitized by Chorinaeus excessorius (Davies). In trapping experiments, virgin female
Croesia curvalana attracted the largest proportion of males between 2200 h and 2400 h.
Male Croesia curvalana were attracted to sex attractant lures and virgin Choristoneura
fumiferana (Clem.) females between 2000 h and 0400 h.

Additional key words: Tortricidae, eggs, diel periodicity, trapping, Vaccinium an-
gustifolium.

Croesia curvalana (Kearfott), commonly called blueberry leaftier
(BBLT), are responsible for serious crop losses in Newfoundland, where
lowbush blueberries, Vaccinium angustifolium (Ait.), are a two-mil-
lion-pound export crop (Morris 1981). The insect was first recorded in
Newfoundland in 1979, and subsequently reported to infest up to 30
percent of blueberry buds in 12 fields sampled in New Brunswick (G.
Wood pers. comm.). Incidence of infestation is increasing due to change
in blueberry cultivation practices: field burning every two years has
been replaced by mowing because of rising oil prices and soil damage.

Kearfott (1907) described Croesia curvalana, as one of four “vari-
eties” of Tortrix albicomana (Clemens) feeding an oak, rose, and huck-
leberry. MacKay (1962) described larval morphology based on a prob-
able last instar. She placed it in the tribe Tortricini, genus Argyrotoza.
Larval host plants were said to be Vacciniaceae, with distribution from
Nova Scotia to British Columbia. Subsequently Powell (1964) and Hodges
et al. (1983) listed the insect as Croesia curvalana.

In 1979 we discovered that adult male C. curvalana were attracted
to virgin spruce budworm, Choristoneura fumiferana (Clemens), (SBW)
adult females. Trapped moths were identified by the Forest Insect and
Disease Survey (FIDS) of Environment Canada, and the Biosystematic
Research Centre, Agriculture Canada. Initial trap capture of C. cur-
valana occurred in traps hung at 1.5 m above ground in balsam fir

VOLUME 42, NUMBER 2 IPAl

stands. This finding suggested that some components of spruce bud-
worm sex pheromone were also BBLT sex attractants. Sanders and
Weatherston (1976) identified the primary components of SBW sex
pheromone as (E) and (Z)-11-tetradecenal (96:4), and Silk et al. (1980)
found traces of tetradecanal and E-11-tetradecenyl-acetate in the ef-
fluvia.

Little is known of BBLT biology. This paper describes laboratory
studies from 1980 to 1983 concerning duration of egg diapause, larval,
and pupal development; suitability of oviposition substrates, and fe-
cundity. Field studies are also presented which explore the BBLT life
cycle and examine diel periodicity of male attraction to calling females.

MATERIALS AND METHODS
Laboratory

Egg treatment during diapause. Eggs collected in August 1981 failed
to hatch when held in the laboratory for 4 months at 21°C with tem-
perature variations of +2°C. A cold treatment was therefore provided
for egg collections made the next year. Sequencing of temperature and
photoperiod throughout diapause in the laboratory was similar to that
used for SBW (Stehr 1954). Field-collected eggs laid on dead leaves
under blueberry plants were stored in Petri dishes lined with dampened
filter paper, sealed with parafilm, and held at 21 + 2°C for 18 to 37
days. One batch of 1144 eggs was chilled at 6 + 1°C for 7 days in a
dark refrigerator. A 2nd batch of eggs was exposed to cold treatment
of 0 + 1°C in a freezer with no chilling. Samples from both batches
were removed from the freezer after 18, 21 and 24 weeks. All eggs
were placed in a refrigerator at 6 + 1°C for 2 days before being exposed
to a constant temperature of 2] + 2°C in a 17L:7D photoperiod. Time
required for eggs to hatch after removal from the freezer and per-
centage hatch were measured.

Larval and pupal rearing. After hatch, 994 larvae were transferred
using a mohair brush to artificial-leaf-meal diet in plastic creamer cups
(4 per cup) or to young foliage, and reared at 21 + 2°C in a 17L:7D
cycle at 70% RH. Diet was that developed for SBW (McMorran 1965)
as modified by Grisdale (1973), to which was added dried blueberry
leaf meal (50% v/v). The meal was produced by drying and grinding
previously frozen leaves from June collections. The diet was allowed
to dry for a day at room temperature, which made it draw away from
cup sides and provide niches for larvae. Twenty larvae were reared
singly from hatch to adult emergence to determine number of larval
instars and time required for maturation. Exuvial head capsules were
collected and widths measured.

122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Oviposition substrates. When neonate larvae were transferred from
leaves to diet, mold spores also transferred immediately contaminated
the diet. Therefore, a study was undertaken to examine suitability of
other oviposition substrates. In 1983, 10 newly emerged virgin females
were placed with males at a ratio of 1:1.5 in each of 8 screened cages
measuring 30 cm’. Moths were provided with a live blueberry branch,
a 10% sucrose source, and a selection of oviposition substrates: parafilm,
waxed paper, aluminum foil, filter paper strips, all 3cm x 15 cm, and
glass (2 bottles each of 21 cm surface area) which were placed on cage
bottoms. In two cages, dried leaves were also offered as an oviposition
substrate. Numbers and viability of eggs deposited on each substrate
were determined. Cages were maintained in a 17L:7D light cycle at
corresponding temperatures of 21 and 17°C with 70% RH for 3 weeks,
after which eggs were counted. Eggs on the various substrates were
given the cold treatment found effective in the initial diapause study,
and emerging larvae counted.

Fecundity. Numbers of eggs produced by virgin and mated female
moths under laboratory conditions were investigated. Single virgin fe-
males and male-female pairs were reared in 12-ml vials containing a
10% sucrose source. Seventy-one virgins 0 to 11 days old were dissected
and their eggs counted. Females were dissected in a 5.5-cm Petri dish
to expose the reproductive system. One or more drops of Shaeffer Script
permanent blue-black ink diluted 50% with water was added to the
preparation. Ovarioles were separated and eggs counted using a base-
lit dissecting microscope at 160-400 magnification. Eggs were regarded
as mature when they reached ca. 0.8 mm diam., the size at oviposition.
Eggs, unfertilized and fertilized, were also counted from 31 mated 5-
to 10-day-old females.

Field

Areas studied. Two geographically distinct regions were selected for
study. The Pouch Cove, Newfoundland, blueberry barrens, which were
used in 1984 and 1985, were rocky and windswept. An area near
Blackville, New Brunswick, which was used from 1980 to 1984, had
less open terrain, and blueberry plants were often interspersed with
ferns, small trees, and bushes.

Life history. In late August 1981, two weeks after the flight season,
whole live blueberry plants, surrounding vegetation, and surface litter
were collected at Blackville. All materials were examined in the labo-
ratory for BBLT eggs. Samples of leaf litter were again collected from
the same area in October to check for egg hatch.

Timing of insect development at Blackville was investigated in 1981.
First invasion of flower buds by larvae was monitored by microscopic

VOLUME 42, NUMBER 2 123

1F

5.0mm;

Fic. 1. Croesia curvalana life stages and injury to host. A, Fertile (arrow) and infertile
eggs; B, Fully mature egg two days before eclosion; C, Newly emerged larva with empty
egg; D, First-instar entry holes in blueberry buds; E, Fourth instar; F, Male and female
pupae (arrows indicate moveable abdominal segments).

10mm

124 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Effect of chilling at 6 + 1°C and duration in freezer at 0 + 1°C on timing
and success of Croesia curvalana egg hatch. All eggs received a post-freezer period of 2
days in refrigerator at 6 + 1°C before being incubated at 21 + 2°C. N = 2100.

Percent hatch

No.chilling Percent of hatching eggs after removal from freezer of eggs sub-
days in ING: wees) «kee a ee iccicdno
refrigerator in freezer 5-11 days 12-18 days 19-25 days 26-32 days treatment

4 24 45 44 9 2 75

if 21 4 66 28 2, ay

7 18 10) 64 39 i! DO

0 24 8 Do 31 6 49

0 “il 6) DO 45 0 20

6) 18 0) 36 A7 ies 31

examination of blueberry plant clippings beginning in early April. Fo-
liage was subsequently clipped for examination at two-week intervals
for observations of larval and pupal development.

Parasites. In 1984, 528 late instars collected at Blackville and 102
from Pouch Cove were reared singly in the laboratory on foliage to
determine incidence of parasitization.

Trap height. Height of male flight and trap height for optimal male
capture were investigated at Blackville in mid-August 1980. Initial
capture of male BBLT in SBW-baited traps had occurred at a height
of 1.5 m. Four Pherocon® 1C sticky traps were each baited with 2
spruce budworm virgin females 0 to 24 h old in small screen cages,
and 4 Pherocon® traps were left empty. Two moth-baited traps, and
two empty Pherocon® 1C traps were hung 1.5 m above the blueberry
canopy and 2 of each at 10 cm above the canopy. Traps were 30 m
apart. Captured moths were counted each day for 10 days and the
SBW virgins were replaced every 2 days.

Flight season. Onset and duration of the flight season was studied
during four seasons in Blackville and two seasons at Pouch Cove by
sweep netting and by capturing males in sticky traps. Traps were placed
at canopy level in advance of the flight season and monitored every 48
h. They were baited with polyvinyl chloride (PVC) lures (Fitzgerald
et al. 1973) formulated by G. Lonergan, Department of Chemistry,
University of New Brunswick, to release (E) and (Z)-11-tetradecenal
(95:5) with small amounts of (E)-11-tetradecanyl-acetate (0.2) at the
rate of 1 SBW equivalent (Sanders 1981). These had attracted leaftier
males in previous experiments (Ponder unpubl.).

Periodicity of sexual activity. Diel periodicity of sexual activity un-
der field conditions was observed at Blackville between 13 and 16 July
1982 in a 96-h trapping study. Pherocon® 1C traps were each baited
with one of the following: two virgin SBW females, two virgin BBLT

VOLUME 42, NUMBER 2 125

females, a PVC lure releasing a sex attractant at the rate of one SBW
equivalent, or a blank PVC formulated without attractants. Each trap
type was replicated three times within the array. Virgin SBW females
were included in this experiment as lures because of their proven success
in the capture of male leaftiers. Traps were separated by 30 m and
their initial positions in the 12-placement array were selected by random
numbers. Trapped moths were counted and traps were moved foward
by 1 position every 2 h because the population was not uniformly
distributed.

RESULTS AND DISCUSSION
Laboratory

Egg treatment during diapause. Three to 4 days after removal from
18 to 24 weeks in the freezer, and 2 to 3 days before hatch, a black
head and larval outline could be observed inside eggs (Fig. 1B).

Significant decreases in egg mortality occurred with acclimation.
Chilling eggs at 6°C in the refrigerator for one week before putting
them in the freezer enhanced hatch (Table 1). The longer period of 24
weeks in the freezer resulted in significantly increased egg hatch (2-
way ANOVA w/o replication, P < 0.05). Seventy-five percent of eggs
hatched if given a 24-week freezer treatment after 7 days of chilling
(Table 1).

Larval and pupal rearing. Larvae matured through 4 instars to pu-
pation in 21 (SD + 3, N = 20) days at 21 + 2°C. Mean head capsule
widths (mm + SD) progressing through instars were 0.25 + 0.03, 0.85 +
0.04, 0.57 + 0.05, and 1.22 + 0.04. Hatchlings were 1.2 mm long,
cream colored, with a dark thoracic shield and black head (Fig. 1C).
Second and third instars remained cream colored, had black heads and
thoracic shields, and dark anal shields. Fourth instars (Fig. 1E) became
yellow, and the head changed to cinnamon brown; the thoracic shield
was cinnamon brown medially, shading to dark brown laterally. Male
gonads in the fifth abdominal segment were maroon, simplifying larval
sexing. Exuvial head capsules appeared slightly lighter in color than
the head in the last two instars.

Mortality was high in the 994 larvae fed leaf-meal diet; only 22%
survived compared with 50% on fresh foliage under the same conditions,
though maturation time was approximately equal. Mortality of diet-
fed larvae could be attributed in part to mold transferred with hatch-
lings from leaves to diet. No attempt was made to surface-sterilize eggs.
Larvae did not feed on previously frozen blueberry foliage unless mixed
with SBW diet. An attempt had been made in 1980 to rear 230 larvae
on SBW diet without addition of blueberry meal. Three larvae survived

126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Mean no. mature eggs per female

! sauause Virgin
60 = Mated

0 1 2 3 4 +) 6 q 8 9 10 11

Moth age (days)

Fic. 2. Numbers of mature eggs in mated (N = 31) and virgin (N = 71) female
Croesia curvalana in relation to age. Vertical bars indicate SD.

to pupation, and larval development time was 75 to 80 days. Newly
hatched larvae seldom burrowed into flower buds (Fig. 1D) on diet,
but rather spun nests between the diet and creamer cup wall.

Pupation occurred either in spun nests between diet and creamer
cup wall, or under the lid. Males pupated before females, and mean
male pupal duration was 9.2 (SD + 0.8, N = 20) days which was shorter
by 2 days than mean female pupal duration of 11.5 (SD + 1.8, N =
20) days. Male pupae could be distinguished by presence of a fourth
moveable abdominal segment (Fig. 1F), females having only three
moveable segments.

Oviposition substrates. Few eggs were deposited by BBLT females
in the laboratory. Such eggs were scattered singly on materials placed
in cage bottoms. Eggs were white, flattened, convex, 0.6 mm in diam.,

VOLUME 42, NUMBER 2 127

and had a clear pebbled surface. Within a week they changed to reddish
brown as the enclosed embryo matured (Fig. 1A). In total, 1061 eggs
were collected in 1983 on synthetic substrates, 44% on waxed paper,
37% on parafilm, 8% on aluminum foil, 7% on filter paper, and 4% on
glass. No preference was shown for dried leaves as an oviposition sub-
strate. Egg maturation and hatch after diapause was 66% on waxed
paper, and 64% on filter paper, but only 32% on parafilm. Other sub-
strates resulted in less hatch.

Fecundity. No mature eggs were found when 5 newly emerged
virgins were dissected; however, a mean of 90.8 (SD + 11.4) immature
eggs were counted. In mated females, ca. % of eggs matured by the
Ath day (Fig. 2), and up to50% matured by the 7th day after emergence.
Twenty-six percent matured by the 7th day in virgins. Both mated and
virgin females laid few eggs in vials. A total of 4 unfertilized eggs laid
by the 71 virgins remained white. Fifty-five fertile eggs which changed
from white to brown were laid by the 31 mated females.

Decrease in numbers of eggs in virgin females between days 8 and
11 (Fig. 2) suggested that their eggs were resorbed. Most moths (79%)
died between 11 and 12 days after emergence; however, some lived 15
days.

Many factors contributed to the difficulty of rearing this insect on
artificial-meal diet. Fecundity was low. The preferred oviposition sub-
strate was ill-defined, and many substrates proved unsuitable for com-
plete egg maturation. Hatchlings were small, delicate, and difficult to
locate even with a microscope. Year-round rearing of BBLT using fresh
vegetation would be impractical without light- and temperature-con-
trolled greenhouse and refrigeration for continuous propagation of blue-
berry plants.

Field

Life history. Eggs were laid singly on dried leaf litter under blueberry
plants, and were difficult to locate even with a microscope. No eggs
were located on living plant leaves, stems, or branches. October col-
lections of eggs indicated that hatch had not yet occurred. Seventy-five
percent of field-collected eggs hatched successfully in the laboratory
when given a treatment of 1 week at 6°C and 24 weeks at 0°C. These
findings indicate that eggs overwinter. Blueberry cultivation practices
that incorporate field mowing rather than burning may therefore result
in increased leaftier populations.

Foliage clipped weekly from April to mid-June at Blackville indicated
that infestation of flower buds by first-instars occurred during the last
two weeks of April (Table 2). Larvae burrowed into closed flower buds
leaving a round hole marked by an accumulation of yellow frass. Fre-

128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 2. Development of Croesia curvalana larvae, Blackville, N.B., 1981.

No. larvae
Date Stage of blueberry foliage collected Instar no. or stage
18 April Closed buds 0
24 April Closed buds 6 1
6 May Expanded flower buds 327 1
12 May Expanded leaf buds 12] 1
129 2
22 May Young leaves 62 2
102 3
2 June Expanded leaves and flowers 247 3
153 4
12 July Flowers and immature fruit 2 4
187 pupae
15 pupal cases

quently, two larvae were found feeding in the same bud. Larvae sub-
sequently fed on swelling leaf buds. Numbers of buds infested with
larvae increased to mid-May, indicating a three-week period of egg
hatch. Visible plant damage peaked just before larvae pupated. At this
time, terminal leaf growth was webbed and eaten, and larger leaves
were folded or webbed together to form shelters. Increased numbers
of abandoned shelters during late-instar development suggested that
larvae moved frequently.

First appearance of pupae in the field at Blackville ranged from the
first to third weeks in June. Males pupated before females. The dark
brown pupae could be found sandwiched in shelters or occasionally
hanging freely by the cremaster from blueberry twigs.

The moths were 5 to 7 mm long, and were of a yellowish hue with
forewing markings of rust and yellow. Toward the end of the flight
season, spent moths lost many wing scales, which made them appear
cream colored. First male moth emergence at Blackville, as established
by trap capture, occurred during the first week of July in all four years
of study. It occurred thus regardless of differences in weather during
the larval and pupal stages. Sweep-net collections indicated that, as in
the laboratory, males emerged before females. Sweep-net collections
were achieved two to three days after first male trap capture. First
sweep-net collections had a male: female ratio per sweep of 0.116:0.044,
which changed to 0.009:0.008 by the end of the flight season. At Pouch
Cove where the population was higher, the ratio changed from
0.23:0.02 to 0.22:0.16 by the end of the flight season. Moth location
may have had a bearing on results. At Blackville where mean day time
temperatures were 5°C higher than Pouch Cove, moths preferred shad-

VOLUME 42, NUMBER 2 129

TABLE 3. Trap capture of male Croesia curvalana in Pherocon® 1C traps hung at
different heights in 10 days, Blackville, N.B., 1980. Number of traps = 8.

Trap height above Mean 24-h catch per trap by Mean 24-h catch per
lueberry canopy virgin spruce budworm unbaited trap
oe 6.0 0.1
10 cm 17.6 0.7

ed areas, while at Pouch Cove where the barrens had mean wind
velocities of 21.6 km/h and mean RH of 82%, moths were located in
sheltered areas of deep vegetation. Female moths may have been at a
lower stratum or beneath vegetation during oviposition, making sam-
pling for females by sweep-net unreliable.

Length of flight season in the 4 years of study at Blackville ranged
from 30 to 47 days; at Pouch Cove, it began in 1984 on 12 July and
lasted 35 days in 1984, and in 1985 on 19 July and lasted 28 days.

Parasites. Larvae collected at Blackville were parasitized 10% by
tachinid flies which emerged as larvae from their hosts. Tachinid pupar-
ia were held in the laboratory for eight months without a cold period.
One fly emerged in too poor condition to identify. No tachinids were
found in larvae from Pouch Cove. Two ichneumonids emerged as adults
from pupae collected as larvae at Pouch Cove. These were identified
by the Biosystematic Research Centre, Agricultre Canada, as Chori-
naeus excessorius Davies. This parasite has not been reported previously
from BBLT.

Trap height. Significantly more male BBLT were captured in traps
hung at 10 cm than at 1.5 m above the foliage in both unbaited and
virgin SBW baited traps x?, = 109.3, P < 0.001, df = 1) (Table 3)
which confirmed visual observations that moths flew immediately above
the foliage.

Croesia semipurpurana (Kft.), a species morphologically similar to
C. curvalana, is attracted to traps hung at 1.5 m baited with components
(Grant et al. 1981) which are also part of the spruce budworm sex
pheromone bouquet.

Periodicity of sexual activity. The largest proportion of BBLT males
trapped by all bait types was between 2200 and 0200 h (Table 4). Virgin
BBLT females captured the largest proportion of BBLT males from
2200 to 2400 h (P = 0.05, Chi square for multiple proportion, Zar 1984)
indicating that female sex pheromone release (calling) took place during
these hours. PVCs which released attractant continuously over a 24-h
period attracted BBLT males consistently between 2000 and 0400 h,
suggesting that male flight period and attraction to lures may extend

130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 4. Proportion of 96-h trap capture of male Croesia curvalana by 2-h intervals
at Blackville, N.B., 1982. Baits in Pherocon® 1C traps were replicated 3 times (N =
1513). No moths were captured between 1000 and 1400 h.

Trapping interval (h) (AST)
1400— 1600—- 1800- 2000- 2200- 2400—- 0200—- 0400- 0600- 0800-

Bait 1600 1800 2000 2200 2400 0200 0400 0600 0800 1000
Two virgin BBLT 0.00 0.00 0.00 0.08 055 0.27 0.02 0.02 0.02 0.05
Blank PVC 0.11 000 000 004 044 033 0.04 0.00 0.04 0.00
PVC sex 0.01 0.01 001 026 030 030 011 001 0.01 0.00

Two virgin SBW 0.00 0.00 0.00 0.23 0.57 0.16 0.03 0.00 0.00 0.01

on either side of the virgin female BBLT calling period. This was
confirmed by the finding that virgin female SBW, which would have
been calling from 2000 h to 2400 h (Palaniswamy & Seabrook 1985),
also attracted BBLT males before the BBLT female calling period.

ACKNOWLEDGMENTS

We thank Agriculture Canada in St. John’s, Newfoundland for providing laboratory
space and personnel, and George Wood and R. F. Morris for advice and encouragement.
We are indebted to the Government of Newfoundland and Labrador for providing a
truck for fieldwork in 1984 and 1985, and to Paul Hendrickson, Small Fruit Specialist,
Government of Newfoundland, for aid in selecting infested areas. We thank L. J. Dyer
and D. C. Eidt for helpful criticisms of the manuscript, and L. R. Kipp for assistance
with numerical testing. Financial support for this study was received by W. D. Seabrook
from Agriculture Canada and the Natural Sciences and Engineering Research Council.

LITERATURE CITED

FITZGERALD, T. D., A. D. ST. Ciair, G. E. DATERMAN & R. G. SMITH. 1973. Slow
release plastic formulation of the cabbage looper pheromone cis-7-dodeceny] acetate:
Release rate and biological activity. Environ. Entomol. 2:607—610.

GRANT, G. G., D. FRECH, L. MACDONALD & B. DOYLE. 1981. Effect of additional
components on a sex attractant for the oak leaf shredder, Croesia semipurpurana
(Lepidoptera: Tortricidae). Can. Entomol. 13:449-451.

GRISDALE, D. 1973. Large volume preparation and processing of a synthetic diet for
insect rearing. Can. Entomol. 105:1553-1557.

HopcEs, R. W., T. DoMINIck, D. R. Davis, D. C. FERGUSON, J. C. FRANCLEMONT, E.
G. MUNROE & J. A. POWELL (eds.). 1983. Check list of the Lepidoptera of America
north of Mexico including Greenland. E. W. Classey Ltd., Wedge Entomological
Research Foundation, London. 284 pp.

KEARFOTT, W. D. 1907. New North American Tortricidae. Trans. Am. Entomol. Soc.
33:73.

MacKay, M. 1962. Larvae of the North American Tortricidae (Lepidoptera: Tortrici-
dae). Can. Entomol. Suppl. 28. 182 pp.

McMorran, A. 1965. A synthetic diet for spruce budworm Choristoneura fumiferana
(Clem.) (Lepidoptera: Tortricidae). Can. Entomol. 97:58-62.

Morris, R. 1981. Fighting blueberry pests in Newfoundland. News and Features,
‘Agriculture Canada. No. 1938. 50 pp.

PALANISWAMY, P. & W. D. SEABROOK. 1985. The alteration of calling behaviour by
female Choristoneura fumiferana when exposed to synthetic sex pheromone. Ento-
mol. Exp. Appl. 37:13-16.

VOLUME 42, NUMBER 2 ee

POWELL, J. A. 1964. Biological and taxonomic studies on tortricine moths, with reference
to the species in California. Univ. Calif. Publ. Entomol. 32, 307 pp.

SANDERS, C. J. 1981. Release rates and attraction of PVC lures containing synthetic sex
attractant of the spruce budworm, Choristoneura fumiferana (Lepidoptera: Tortrici-
dae). Can. Entomol. 113:103-111.

SANDERS, C. J. & J. WEATHERSTON. 1976. Sex pheromone of the eastern spruce budworm
(Lepidoptera: Tortricidae) optimum blend of trans-and-cis-11-tetradecenal. Can.
Entomol. 108:1285-1290.

SILK, P. J.,S. H. TAN, C. J. WEISNER, R. J. Ross & G. C. LONERGAN. 1980. Sex pheromone
chemistry of the eastern spruce budworm, Choristoneura fumiferana. Environ. Ento-
mol. 9:640-644.

STEHR, G. 1954. A laboratory method for rearing the spruce budworm, Choristoneura
fumiferana (Clem.) (Lepidoptera: Tortricidae). Can. Entomol. 86:423-428.

ZAR, J. H. 1984. Biostatistical analysis. Prentice-Hall, Englewood Cliffs, New Jersey.
718 pp.

Received for publication 28 August 1987; accepted 27 January 1988.

Journal of the Lepidopterists’ Society
42(2), 1988, 1382-137

HYLESIA ACUTA (SATURNIDAE) AND ITS
AGGREGATE LARVAL AND PUPAL POUCH

KIRBY L. WOLFE
Rt. 5, Box 169-C, Escondido, California 92025

ABSTRACT. Developmental stages of Hylesia acuta are described and illustrated,
and two examples of aggregate larval and pupal pouches are reported. Larval development
from eggs gathered in Chiapas, Mexico, required ca. 80 days at 25-32°C. Late instars
constructed a tough, silk pouch where they remained by day, emerging to feed at night.
The first instar is pale yellowish tan with a black head; the mature larva is dark saffron
mottled with green. All instars possess urticating spines, but not adults. Larvae constructed
individual cocoons before pupating together in the pouch, in which they remained for
eight months before emergence. A wild pupal pouch found in Veracruz, Mexico, contained
46 pupae from which 42 adults emerged. A dissected female yielded 374 ova. The species
appears to be univoltine, adults emerging during the tropical wet season.

Additional key words: Mexico, silkmoths, immature stages.

The American genus Hylesia contains about 100 species of small
silkmoths (C. Lemaire pers. comm.), the biology of most unknown. The
genus has achieved notoriety in parts of South America because of
urticating abdominal hairs of some females (Lamy & Lemaire 1983)
used to cover egg masses (Gardner 1982). Hairs in some species cause
severe dermatitis in man (Pesce & Delgado 1971, and others).

In the Central American Hylesia lineata Druce, ova pass the dry
season in a felt nest (Janzen 1984). Though H. nigricans Berg, whose
immature stages and behavior were illustrated by Lampe (1986), also
has overwintering eggs, at least several other species do not (Gardner
1982). Pupation is solitary in most known species.

Aggregate pupation above ground in a silk pouch is unusual among
Saturniidae. A “communal cocoon” of an unidentified species of Neo-
diphthera (Saturniinae) from New Guinea contains about a dozen co-
coons (R. S. Peigler pers. comm.). Stoll (1791) illustrated a “gregarious
cocoon” ascribed to Phalaena Bombyx bibiana, which Bouvier (1925)
believed was a Hylesia (Hemileucinae) species. Beutelspacher (1985)
found Hylesia frigida Schaus gregarious larvae and pupae in loose silk
pouches in Mexico. Cockerell, in Packard (1914), quoted Dyar as having
a specimen of Hylesia tapabex Dyar “‘bred from a ‘gregarious podlike
cocoon.” Bouvier (1924a, 1924b, 1925) described two aggregate pupal
pouches of this species from Venezuela. A nest of H. tapabex is preserved
in the Muséum national d’Histoire naturelle, Paris (C. Lemaire pers.
comm.). The present study establishes that Hylesia acuta Druce, closely
related to H. tapabex, also pupates in a shared pouch.

Hylesia acuta is a small moth (forewing length 2.5-3.1 em) with
marked sexual dimorphism, the female resembling many Hylesia species,

VOLUME 42, NUMBER 2 133

while the male is distinctive (Fig. 3). Described by Druce (1886) from
“North Mexico,” its known range extends along the eastern and western
lowlands of central Mexico S into Guatemala and E into Yucatan and
British Honduras (Schiissler 1934, Hoffmann 1942, C. Lemaire pers.
comm.). Its biology and aggregate pupation behavior have not been
previously reported.

REARED MATERIAL

A wild female captured near Huixtla, Chiapas, Mexico, oviposited
in a paper bag on 10 July 1983. The eggs were white, upright ovals
placed in a single-layered dense cluster, partially hidden with brown
abdominal hairs (Fig. 1). They numbered ca. 100. Refrigerated at ca.
50°C for 8 days to delay hatching, the eggs were then kept in a covered
plastic petri dish under natural daylight at temperatures varying be-
tween 25 and 32°C. They hatched in 25 days, and were transferred to
a tight styrene box 11 x 11 x 4cm. During the first week they refused
to feed on any plants offered, instead eating eggshells. Occasionally
they wandered in single file, returning to the nest without feeding.
Eventually most did accept Brazilian pepper tree, Schinus terebinthi-
folius Raddi (Anacardiaceae). Plants refused included plum, Robinia
pseudoacacia L., Rhus laurina Nutt., Quercus agrifolia Née, and others.
Larvae were then placed on small pepper tree branches in a container
of water in a screened cage in a humid greenhouse at 25-32°C, with
foliage replaced daily. From the earliest stages they spun a loose silken
platform to which they returned after group wanderings or feeding.

Six weeks after hatching, the half-grown larvae spun a silk tent at
the end of a branch. It appeared as a small, broad cone (ca. 7 cm diam.),
of dense silk tipped on its side. A hole near the vertex allowed larval
access; frass fell through a slit at the bottom.

When the tent was completed, the larvae became nocturnal, not
appearing until at least 1 h after dark when they emerged and traveled
in single file, stopping to feed in densely packed rows on mature (darker)
leaves. Such tandem movement in early instars is typical among hemi-
leucines as noted in Hemileuca oliviae Cockerell (Capinera 1980), Hy-
lesia lineata (Janzen 1984), and other species (Lemaire 1971).

The earliest instars were pale yellowish tan, with typical hemileucine
spination retained by all subsequent instars (Figs. 2, 4). This color
deepened, and after the fourth instar was dark saffron indistinctly
mottled with green. One dorsal and two subdorsal longitudinal stripes
were straw colored. Mature larvae were plump (Fig. 2). Instar duration
and number were not determined.

Half-grown larvae became diurnally active during two days as they
spun a larger tent (ca. 15 cm diam.). Incorporating material from the

134 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

CM I 2 3 4

i 1G ssorsineeanccmmenseenieaaens ae
5 6 7 ~ = “Beek © fio acetone

Fics. 1-6. Hylesia acuta. 1, Eggs partially hidden by female abdominal hairs; 2,
Mature larva; 3, Adult female (upper) and male (lower); 4, Early instars feeding on

Schinus terebinthifolius; 5, Aggregate larval and pupal “nest” pouch of captive reared
larvae; 6, Pouch containing wild pupae.

first tent, its walls resembled thin leather, shiny on the inside, and white
on the outside (Fig. 5).

Larvae molted within the tent, and the cast-off skins fell through the
bottom slit. Shortly before pupation larval mortality increased. Five
survivors ceased feeding at 10 weeks, reaching a length of ca. 50 mm.
A flashlight beamed through the tent at night revealed their silhouettes,
which became progressively less visible as they worked during three
weeks to fill it with firm, woolly white silk. Four small holes or “escape”’

VOLUME 42, NUMBER 2 135

tunnels to the outside were made, two by larvae chewing through the
cage’s fiber glass screen where it adhered to the pouch.

By 1 December noticeable larval activity had ceased. Several weeks
later, the pouch was opened, revealing a small mass of cocoons imbed-
ded in silk. From hatching to pupation was ca. 80 days. The next year
in the first week of August, five imagines emerged.

WILD MATERIAL

A cordiform pupal pouch of H. acuta with viable pupae was found
near Papantla, Veracruz, Mexico, in a small, dead tree ca. 4 m above
ground on 27 July 1986. Surrounded by a fresh growth of tall grasses,
the tree appeared to have been Bursera simaruba (L.) Sarg. (Bursera-
ceae). Imprints of fresh leaves were imbedded on the surface of the
pouch. The pouch measured 13.0 x 9.5 x 5.0 cm, and contained 46
pupae. It had been constructed during the previous wet season, since
a new wet season was just beginning.

This pouch was angular (Fig. 6), and possessed three widely spaced
holes on its upper surface, providing access for feeding larvae and exits
for emerging imagines.

Inside, pupae were arranged in double-walled, fusiform cocoons,
tightly fit and adhering to one another. Cocoons were parallel in a band
three layers thick which wrapped around the supporting branch within
the pouch. Exit vents were oriented upward, and opened’ on several
smooth-lined corridors through the dense silk wool to the outside open-
ings of the pouch.

The pupae conform to Bouvier’s (1924a, 1924b, 1925) description of
Hylesia tapabex. He pointed out that while H. tapabex, known to pupate
aggregately, does not possess a cremaster, solitary pupating Hylesia
species do possess one. This is further evidenced in the solitary pupae
of H. nigricans (Lampe 1986) and H. lineata (pers. obs.) which possess
cremasters.

Emergence of imagines began in September. From 46 pupae, 19
males and 23 females emerged during three weeks. Average emergence
was at 1715 h PDT (SD = 1554-1836 h PDT, n = 9). Notably, no
parasitism was found. Numerous attempts were made to achieve mat-
ings between emerged siblings without success.

Observations of captive females suggest that H. acuta oviposits three
or more clusters. Dissection of a newly emerged female yielded 374
ova. Females produced no stinging when their abdomens were rubbed
on the author’s skin.

Voucher imagines are in the San Diego Museum of Natural History,
and collections of C. Lemaire, S. Stone, M. Smith, D. Herbin, and the
author.

136 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

DISCUSSION

Hylesia are among the smallest American saturniids. Those that live
under extreme seasonal conditions have evolved strategies to survive
the harsh conditions of a dry season (Janzen 1984). In H. acuta, a
protective nest has evolved in which pupae survive the dry season. The
tough, leathery pouch filled with silk wool protects them against adverse
weather and perhaps parasites. Its light silvery gray color reflects heat,
and its resemblance to a wasp nest may discourage avian predation.

Published records (Beutelspacher 1978), correspondence (C. Lemaire
pers. comm.), and labels of wild collected specimens indicate dates of
capture predominately from June through August, with records as late
as September, during the season of heaviest rains.

ACKNOWLEDGMENTS

I thank Claude Lemaire for encouragement and assistance. He and R. S. Peigler
provided key literature and commented on the manuscript. I also thank C. R. Beutel-
spacher B. for comments, and D. K. Faulkner for technical suggestions; also Daniel Herbin,
Claude Lemaire, M. J. Smith, and Steve Stone for emergence observations; and Marvin
Valverde for assistance in collecting the material studied.

LITERATURE CITED

BEUTELSPACHER B., C. R. 1978. Familias Sphingidae y Saturniidae (Lepidoptera) de
Las Minas, Veracruz, México. An. Inst. Biol. Univ. Nal. Auton. México 49 (Zoolo-
gia)(1):219-230.

1985. Ciclo de la vida de Hylesia frigida Schaus (Lepidoptera: Saturniidae),
una plaga forestal en Chiapas. An. Inst. Biol. Univ. Nal. Auton. Mexico 56 (Zoolo-
gia)(2):465—476.

Bouvier, E.-L. 1924a. Sur la nidification et les métamorphoses de quelques Saturniens
hemileucides. Compt. Rend. Acad. Sci. Paris (Biology) 179:858-861.

1924b. Contribution a |’étude des Saturniens. Ann. Sci. Nat. (Zool.), Série ioeme

7:138-178.

1925. Contributions a la connaissance des métamorphoses chez les Saturniens
hemileucides. Encyclop. Entomol. B. II(1):3-10.

CAPINERA, J. L. 1980. A trail pheromone from silk produced by larvae of the range
caterpillar Hemileuca oliviae (Lepidoptera: Saturniidae) and observations on aggre-
gation behavior. J. Chem. Ecol. 6:655-664.

Druce, H. 1886. Insecta. Lepidoptera—Heterocera. In Godman, F. and O. Salvin,
Biologia Centrali-Americana, I. Taylor & Francis, London. 197 pp.

GARDINER, B.O.C. 1982. A silkmoth rearer’s handbook. 3rd ed. (Amateur Entomologist
12) 255 pp.

HOFFMANN, C. C. 1942. Catalogo sistematico y zoogeografico de los lepiddpteros me-
xicanos. Parte 3. Sphingoidea y Saturnioidea. An. Inst. Biol. Mexico 13(1):213-256.

JANZEN, D. H. 1984. Natural history of Hylesia lineata (Saturniidae: Hemileucinae) in
Santa Rosa National Park, Costa Rica. J. Kansas Entomol. Soc. 57:490-514.

LAMPE, R. E. J. 1986. Die Praimaginalstadien von Hylesia nigricans Berg 1875 (Lep::
Saturniidae). Entomol. Z. 96(1-2):7-12.

LAMY, M. & C. LEMarrE. 1983. Contribution 4 la systématique des Hylesia: étude au
microscope électronique 4 balayage des fléchettes urticantes Lep.: Saturniidae. Bull.
Soc. Entomol. France 88:175-192.

LEMAIRE, C. 1971. Révision de genre Automeris Hiibner et des genres voisins. Biogéo-
graphie, éthologie, morphologie, taxonomie. Mém. Mus. Nat. Hist. Natl. N. Sér., Sér.
A, Zool. 68(92):1-232.

VOLUME 42, NUMBER 2 137

PACKARD, A. S. 1914. Monograph of the bombycine moths of North America, pt. 3.
Mem. Natl. Acad. Sci. 12: 1-276, 5038-516.

PrEscE, H. & A. DELGADO. 1971. Poisoning from adult moths and caterpillars, pp. 119-
156. In Bucherl, W. and E. B. Buckley (eds.), Venomous animals and their venoms.
Vol. III. Venomous invertebrates. Academic Press, New York.

SCHUSSLER, H. 1934. Saturniidae, in E. Strand, Lepidopterorum Catalogus, pars 58, pp.
325-484. W. Junk, Berlin.

STOLL, C. [1787-1790] 1791. A Anhangsel van het werk de uitlandsche Kapellen
voorkomende in de drie Waereld-Deelen Asia, Africa en America, door den Heere
Pieter Cramer ... (1791). S. J. Baalde, Amsterdam, B. Wild, Utrecht. Pp. 1-184, pls.

1-92.

Received for publication 16 April 1987; accepted 9 February 1988.

Journal of the Lepidopterists’ Society
42(2), 1988, 138-1438

EUROPEAN CORN BORER REPRODUCTION: EFFECTS OF
HONEY IN IMBIBED WATER

WILLIAM E. MILLER

Department of Entomology, University of Minnesota,
St. Paul, Minnesota 55108

ABSTRACT. European corn borer adults are well known to imbibe water, without
which their reproduction is greatly decreased. Whether their reproduction is enhanced
by sugars in imbibed water has long been unresolved. Two groups totalling more than
50 captive fertile pairs, one group receiving 15% honey-water to imbibe, the other plain
water, were compared with respect to 10 reproductive attributes. Honey in imbibed water
significantly improved performance in four attributes, resulting in heavier eggs, more
females maintaining or increasing egg weight during the oviposition period, fewer females
with immature oocytes at death, and more unlaid eggs that were mature. Adult nutritional
ecology seems a potential factor in the dynamics of populations.

Additional key words: Ostrinia nubilalis, Pyralidae, Pyraustinae, fecundity, egg weight.

Imbibing of water by adults of the European corn borer, Ostrinia
nubilalis (Hiibner), is well documented (Vance 1949). Imbibed water
greatly increases adult lifespan and fecundity; without it, reproduction
is severely depressed (Barlow & Mutchmor 1963, Kira et al. 1969).
Concerning sugars in imbibed water, reported effects are contradictory,
one paper claiming no further improvement in reproduction (Caffrey
& Worthley 1927), and another claiming the opposite (Kozhantshikov
1938). Unfortunately, the first paper offers no supporting data, and the
second enumerates data insufficiently for independent assessment. Al-
though this contradiction has never been addressed, the prevailing view,
as reflected in mass rearing practice (Reed et al. 1972), is that sugars
in imbibed water do not enhance reproduction. Clarification seems
desirable for at least two reasons. First, European corn borer adults
aggregate in grass and weedy vegetation bordering host fields (Showers
et al. 1976), where potential adult food sources such as nectaries and
honeydew-producing insects may occur. Second, if sugars do enhance
reproduction, mass rearing programs might thereby increase production
with little extra effort.

Here I report how European corn borer adults receiving honey-water
and plain water performed in the 10 reproductive attributes listed in
Table 1.

MATERIALS AND METHODS

One experiment was done. In it, 78 single female-male pairs of pupae
were numbered, and the 39 even-numbered ones assigned to the treat-
ment group, and the 39 odd-numbered ones to the control group. After
adults eclosed, treatment pairs received honey-water to imbibe, control

VOLUME 42, NUMBER 2 139

pairs plain water. After all moths had expired, reproduction data gath-
ered from both groups were compared.

Pupae were obtained from a culture at the University of Minnesota
originating in Jowa and maintained according to standard European
corn borer production methods (Reed et al. 1972). The pupae were
sexed, and pairs placed in 1-pint (0.47 liter) cardboard ice cream con-
tainers capped with Petri dish lids. Lids were lined with waxed paper
to ensure a surface suitable for oviposition. Containers were kept in a
walk-in environmental chamber programmed for 16 h light at 27°C
and 8 h dark at 17°C, both at 60% RH.

Each container had a 35 cm® foam-latex sponge that dispensed dis-
tilled water in the control group and 15% (by volume) honey-water in
the treatment group. Sponges and liquids were renewed every second
day. Honey was used as the sugar source because in composition (White
1975) it conveniently simulates hexose-rich shallow-flower nectar (Baker
& Baker 1983) and insect honeydew (Auclair 1968).

Reproduction data were gathered once daily near mid-day. Mating
and fertilization success was ascertained by holding one or several early
egg masses for a week or until the dark larval heads showed through
chorions. Preoviposition period was measured from female eclosion to
first oviposition; such data were used only when the male of the pair
eclosed no later than one day after the female because late male eclosion
prolonged the period.

“Early eggs” refers to eggs laid on the first or second day of ovipo-
sition, “late eggs’ to eggs laid on the fourth to eighth day of oviposition.
Mean egg weight was determined from one or more masses totalling
17 to 184 eggs removed intact from the waxed paper and weighed to
the nearest 0.5 mg. Mature unlaid eggs were counted in excised ovaries
at stereomicroscope magnifications up to 65x. Maturity of unlaid eggs
was judged by size and chorionation. Eggs were deemed chorionated
if they did not readily absorb 0.3% aqueous methylene blue after 3 min
exposure (Jennings 1974). Immature oocytes in expired females could
not be counted accurately, so only their presence or absence was re-
corded. Only data from fertile pairs were analyzed because some re-
productive attributes are atypical in the absence of insemination.

Referral of species mentioned in this paper to Pyraustinae is based
on the classification of Fletcher and Nye (1984).

RESULTS AND DISCUSSION

Of the 78 pupal pairings, 79% resulted in fertile eggs, a level believed
high enough to provide representative adult performance. Data were
analyzed from fertile pairs numbering 25 and 27 in the plain-water
and honey-water groups, respectively, these numbers also reflecting

140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

losses from mishaps like moth escapes. The fewest number of obser-
vations on any attribute in either group was 19 for preoviposition period
in the water-imbibing control, reflecting a further loss of data on this
attribute resulting from late male eclosions.

Four differences between the two imbibing groups were significant
(Table 1). Thus, in the honey-water group, a greater proportion of
unlaid eggs was mature, fewer expired females contained immature
oocytes, late eggs were heavier, and more females maintained or in-
creased egg weight during the oviposition period. The first and second
differences suggest that honey-water imbibers approached full repro-
ductive potential more closely than plain-water imbibers. The third
and fourth differences presumably reflect conversion of glucose and
fructose in honey to lipid that became incorporated into oocytes during
egg maturation (Kozhantshikov 1938, Downer & Matthews 1976). Con-
sequences of differing egg weights have not been investigated in the
European corn borer, but in other moth species, heavier eggs produce
larvae more likely to survive (Barbosa & Capinera 1978, Harvey 1985).
Hence, sugars in imbibed water might enhance European corn borer
fitness if females live long enough to lay the heavier eggs. Some do live
long enough (Elliott et al. 1982): in 9 of 14 samples taken from June
to September, % of wild mated females were 4 or more days old, the
onset age for heavier eggs in the present study.

Although 6 of the 10 reproductive attributes did not differ signifi-
cantly between imbibing groups (P = 0.06, one-tailed Student t-tests),
all attributes except female lifespan show differences of 1 to 150% in
favor of the honey-water group (Table 1). Shorter female lifespan in
the honey-water group seems anomalous, but no cause was evident.
Despite this attribute, the honey-water group outperformed the plain-
water group in an overall comparison of reproduction as follows. Of
eight independent attributes (omitting number of unlaid mature eggs
and percentage females maintaining or increasing egg weight, which
are facets of other attributes), seven show gains resulting from honey
imbibing whether individually significant or not, and such an outcome
is not likely due to chance (P < 0.05, one-tailed sign test).

Because European corn borer fecundity varies directly with body
size (Vance 1949), the possibility that body-size differences between
imbibing groups caused attribute differences was examined. Vance
(1949) did not express the relation mathematically; he tabulated six
class means for number of eggs laid (y) and corresponding initial adult
female weight (mg) (x). Based on retrospective frequency-weighted
analysis of class means, the relation can be quantified as y = 8.5x +
153 (66n, r? = 0.88, P < 0.001). The correlation coefficient is overes-
timated because it is derived from means rather than individual values,

VOLUME 42, NUMBER 2 14]

TABLE 1. Reproductive performance of European corn borer adults receiving plain
water and 15% honey-water. Means and percentages are based on 19 to 27 observations
per treatment group.

Mean (+SD) or percentage PSepanee
Attribute Plain water Honey-water hanes
Lifespan, days
Female Dior st jor kGiata= 5A =*
Male 16.8 + 4.0 18.2. +.4.5 8
Preoviposition period, days 285-13 Sieh 2 —18
No. mature oocytes
Laid GOK 191 623 + 237 4
Unlaid 28 + 38 a0: 053* 78
Total 630 + 170 673 + 209 i
% females containing immature
oocytes at death 86 o3* =38
Egg weight, mg
Early eggs 0.0638 + 0.0048 0.0643 + 0.0060 1
Late eggs 0.0618 + 0.0066 0.0652 + 0.0070* 6
% females maintaining or
increasing egg weight 24 60* 150

* Significantly different (P < 0.05, based on one-tailed Student t-tests for means; 2 x 2 contingency tables and ad-
justed-G tests for numbers underlying percentages).

but the relation is nevertheless striking. Such a relation could not have
affected attribute differences in the present study for two reasons. First,
length of one forewing, a surrogate for body weight (Miller 1977),
averaged 13.5 (SD + 0.6) mm and 13.4 (SD + 0.8) mm in females of
the plain-water and honey-water groups, respectively. These means are
identical statistically, the difference, 0.1 mm, being less than 1% of
either. Second, in neither imbibing group did any egg attribute correlate
significantly with female forewing length (r? < 0.08, P > 0.20).

In virgin European corn borer females, 80 egg follicles have been
seen in one ovariole (Drecktrah & Brindley 1967). This number trans-
lates to 640 per female, near the average total number of mature eggs
per female in the present study (Table 1). Number of oocytes already
mature at female eclosion averaged 92 (SD + 16, 4n) in the present
study, as determined for females 0-4 h old averaging 14.0 mm in
forewing length. Subtracting 92 from total mature eggs in the plain-
water and honey-water groups leaves 538 and 581 eggs, respectively.
The latter numbers suggest that more than 80% of oocytes mature after
females eclose (538/630 = 0.85; 581/673 = 0.86), and that ample op-
portunity exists for adult nutrition to enhance oogenesis.

Although European corn borer adults have not been reported to

142 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

imbibe sugary liquids in the wild, it is possible they do so opportunistical-
ly because adults of other Pyraustinae use such liquids. Both sexes of
Pyrausta orphisalis Walker have been seen taking nectar from flowers
(Campbell & Pike 1985). In Cnaphalocrocis medinalis (Guenée), im-
bibed sugars and planthopper honeydew greatly increased adult life-
span and fecundity (Waldbauer et al. 1980). Moreover, both sexes of
European corn borer have well developed proboscides: females with
forewing length (w) averaging 14.0 mm had proboscides with uncoiled
length (p) averaging 7.3 mm (4n), forming a p/w ratio of 0.52. This
length of proboscis could provide access to floral nectar of many kinds
of plants, and the ratio is well within the range signifying flower vis-
itation in other lepidopterans (Opler & Krizek 1984:31).

In conclusion, some differences in reproductive attributes between
treatment groups are subtle. They nevertheless bring one step nearer
resolution the old uncertainty whether sugars in imbibed water affect
European corn borer reproduction. The adult’s use of sugary liquids in
nature remains to be shown. If it is shown, adult nutritional ecology
could be a factor in the dynamics of populations, with sugar consump-
tion perhaps elevating population quality and crop damage. Such an
outcome could lead to removal of natural sources of sugar as a supple-
mental management technique for the insect.

ACKNOWLEDGMENTS

This study was supported by a U.S. Dept. Agr. Cooperative State Research Service
grant. I thank K. V. Bui for laboratory assistance, D. A. Andow, K. R. Ostlie, T. C. Schenk
and three anonymous readers for useful manuscript reviews, and B. A. Drummond for
serving as special Journal editor for this paper. This is paper 15868, Scientific Journal
Series, Minnesota Agricultural Experiment Station, University of Minnesota, St. Paul,
Minnesota 55108.

LITERATURE CITED

AUCLAIR, J. L. 1963. Aphid feeding and nutrition. Ann. Rev. Entomol. 8:439-490.

BAKER, H. G. & I. BAKER. 1983. Floral nectar sugar constituents in relation to pollinator
type, pp. 117-141. In Jones, C. E. and R. J. Little (eds.), Handbook of experimental
pollination biology. Scientific & Academic Editions, New York. 558 pp.

BARBOSA, P. & J. L. CAPINERA. 1978. Population quality, dispersal and numerical change
in the gypsy moth, Lymantria dispar (L.). Oecologia (Berl.) 36:203-209.

BARLOw, C. A. & J. A. MuTCHMoR. 1963. Some effects of rainfall on the population
dynamics of the European corn borer, Ostrinia nubilalis (Hbn.) (Pyraustidae: Lep-
idoptera). Entomol. Expt. Appl. 6:21-36.

CAFFREY, D. J. & L. H. WoRTHLEY. 1927. A progress report on the investigations of
the European corn borer. USDA Bull. 1476, 154 pp.

CAMPBELL, C. L. & K. S. PIKE. 1985. Life history and biology of Pyrausta orphisalis
Walker (Lepidoptera: Pyralidae) on mint in Washington. Pan-Pac. Entomol. 61:42-
47.

DOWNER, R. G. H. & J. R. MATTHEWS. 1976. Patterns of lipid distribution and utilisation
in insects. Am. Zool. 16:733-745.

DRECKTRAH, H. G. & T. A. BRINDLEY. 1967. Morphology of the internal reproductive
systems of the European corn borer. Iowa State J. Sci. 41:467-480.

VOLUME 42, NUMBER 2 143

ELLIOTT, W. M., J. D. RICHARDSON & J. FOUNK. 1982. The age of female European
corn borer moths, Ostrinia nubilalis (Lepidoptera: Pyralidae), in the field and tests
of its use in forecasting damage to green peppers. Can. Entomol. 114:769-774.

FLETCHER, D. S. & I. W. B. NyE. 1984. The generic names of moths of the world. Vol.
5. Pyraloidea. British Museum (Natural History), London. 185 pp.

Harvey, G. T. 1985. Egg weight as a factor in the overwintering survival of spruce
budworm (Lepidoptera: Tortricidae) larvae. Can. Entomol. 117:1451-1461.

JENNINGS, D. T. 1974. Potential fecundity of Rhyacionia neomexicana (Dyar) (Olethreu-
tidae) related to pupal size. J. Lepid. Soc. 28:131-136.

Kira, M. T., W. D. GUTHRIE & J. L. HuGGANS. 1969. Effect of drinking water on
production of eggs by the European corn borer. J. Econ. Entomol. 62:1366-1368.

KOZHANTSHIKOV, I. W. 1938. Carbohydrate and fat metabolism in adult Lepidoptera.
Bull. Entomol. Res. 29:103-114.

MILLER, W. E. 1977. Wing measure as a size index in Lepidoptera: The family Olethreu-
tidae. Ann. Entomol. Soc. Am. 70:253-256.

Oper, P. A. & G. O. KRIZEK. 1984. Butterflies east of the Great Plains. Johns Hopkins,
Baltimore. 294 pp.

REED, G. L., W. B. SHOWERS, J. L. HUGGANS & S. W. CARTER. 1972. Improved procedure
for mass rearing the European corn borer. J. Econ. Entomol. 65:1472-1476.

SHOWERS, W. B., G. L. REED, J. F. ROBINSON & M. B. DEROzARI. 1976. Flight and
sexual activity of the European corn borer. Environ. Entomol. 5:1099-1104.

VANCE, A. M. 1949. Some physiological relationships of the female European corn borer
moth in controlled environments. J. Econ. Entomol. 42:474—484.

WALDBAUER, G. P., A. P. MARCIANO & P. K. PATHAK. 1980. Life-span and fecundity
of adult rice leaf folders, Cnaphalocrocis medinalis (Guenée) (Lepidoptera: Pyrali-
dae), on sugar sources, including honeydew from the brown planthopper, Nilparvata
lugens (Stal) (Hemiptera: Delphacidae). Bull. Entomol. Res. 70:65-71.

WuitE, J. W. 1975. Composition of honey, pp. 157-206. In Crane, E. (ed.), Honey: A
comprehensive survey. Heinemann, London. 608 pp.

Received for publication 13 July 1987; accepted 8 January 1988.

Journal of the Lepidopterists’ Society
42(2), 1988, 144-145

GENERAL NOTE

PARASITOID AND HOSTPLANT RECORDS FOR GENUS SCHINIA
(NOCTUIDAE) IN TEXAS

- Additional key words: Schinia bina, S. arcigera, S. chrysella, Tachinidae, Hyme-
noptera.

In an earlier report, R. S. Peigler and S. B. Vinson (1984, Southw. Entomol. 9:48-51)
listed 24 species of Schinia Hubner collected in Brazos Co., Texas, and commented on
abundance of the adults. The present paper deals with some observations on the immature
stages. The life-cycle for most species is as follows: eggs are deposited in autumn into
flowers of composites (Asteraceae), most species specializing on one or a few host genera.
Larvae mature in less than one month, pupation is below ground, and adults emerge the
following autumn. Peak abundance of larvae as well as adults is in fall.

We experienced difficulty in associating field-collected larvae with adults because dia-
pause was not terminated by various treatments, and pupae often died before adults
emerged. Reared material emerged at the next normal flight time for some species, but
this was the exception. A few pupae produced adults only after being held for two or
more years. Some parasitoids were equally delayed in emerging as adults. Consequently,
some records below are cited as “Schinia sp.’ where we were unable to associate larvae
with adults. Interspecifically, Schinia larvae are as variable as adults in color and pattern
(Covell, C. V. 1984, A field guide to the moths of eastern North America, Houghton
Mifflin, Boston, Pl. 29), but larvae can confidently be assigned to this genus based on
general appearance.

Previously published records of parasitism in Schinia are few. P. H. Arnaud (1978, A
host-parasite catalog of North American Tachinidae (Diptera), U.S. Dept. Agr. Misc. Publ.
1319, 860 pp.) listed only one record for a tachinid attacking Schinia: Winthemia quad-
ripustalata (Fabricius) parasitizing Schinia septentrionalis Walker (=S. brevis Grote).
Only one record for a hymenopterous parasitoid attacking Schinia was cited by P. M.
Marsh (in Krombein, K. V., P. D. Hurd, D. R. Smith, B. D. Burks (eds.), 1979, Catalog
of Hymenoptera in America north of Mexico, vol. 1:268, Smithsonian Press, Washington,
D.C.): the braconid Cardiochiles magnus Mao in Schinia sp. Another braconid, Microplitis
croceipes (Cresson) (det. by P. M. Marsh) was reared from Schinia olivacea J. B. Smith
collected in Live Oak Co., Texas, and another tachinid, Gymnoclytia unicolor Brooks
(det. C. W. Sabrosky) from Schinia olivacea in Bexar Co., Texas (R. O. Kendall pers.
comm. ).

Our larvae were collected in or on composite inflorescences. They were kept individually
in the laboratory on artificial diet (Vanderzant, E. S., C. D. Richardson & S. W. Fort
1962, J. Econ. Entomol. 55:140) in plastic shell vials plugged with cotton. It was necessary
to isolate larvae to prevent cannibalism, a problem also noted by D. F. Hardwick (1958,
Can. Entomol. Suppl. 6:1-116). For hostplants, we follow nomenclature of D. S. Correll
and M. C. Johnston (1970, Manual of the vascular plants of Texas, Texas Research
Foundation, Renner, Texas, 1881 pp.). All records below are from Brazos Co., in E-central
Texas. Species of Schinia most commonly collected by us in the larval stage were S. bina
(Guenée), S. arcigera (Guenée), S. chrysella Grote, and S. bifascia Hiibner. Also, many
larvae of S. nundina (Drury) were collected from flowers of goldenrod (Solidago spp.)
in October, but few adults were because they are rarely phototactic.

The following parasitoids were reared:

Diptera
Tachinidae

Plagiomima similis (Townsend). One specimen reared from larva of Schinia bina
collected in fall. Puparium formed outside host and overwintered before emerging.
Eucelatoria sp. (armigera Coquillet of authors). One specimen reared from Schinia
sp. Puparium formed outside host, adult emerged in fall without diapausing.
Winthemia rufopicta (Bigot). One specimen reared from Schinia sp.

VOLUME 42, NUMBER 2 145

Hymenoptera
Ichneumonidae

Ophion sp. (det. R. S. Peigler using I. D. Gauld & P. A. Mitchell, 1981, The taxonomy,
distribution and host preferences of Indo-Papuan parasitic wasps of the subfamily
Ophioninae (Hymenoptera: Ichneumonidae), Commonwealth Agric. Bur., Slough, 611
pp.). One reared from Schinia sp.

Campoletis sonorensis (Cameron). A few reared from 8rd instar Schinia bina and
S. chrysella. White cocoons formed alongside dried host remains.

Pristomerus spinator (Fabricius). One reared from 2nd instar Schinia bina.

Braconidae

Cardiochiles abdominalis (Cresson). Thirty parasitoids reared from Schinia bina and
S. arcigera. Larvae of both host species were collected on Aster spinosus Benth.

Microplitis croceipes (Cresson). Two reared from larvae of Schinia chrysella collected
on Xanthocephalum dracunuloides (DC) Shinners.

Cotesia marginiventris (Cresson). From larvae of Schinia chrysella collected on
Xanthocephalum dracunuloides we reared 58 parasitoids. From larvae of Schinia
bifascia collected on Ambrosia trifida L. we reared 18 parasitoids.

Meteoris sp., probably laphygmae Viereck. One specimen reared from Schinia sp.

Schinia belongs to the same subfamily as Heliothis virescens (Fabricius) and H. zea
(Boddie), two important agricultural pests. Entomologists working on Heliothis would be
well advised to determine which species of Schinia occur in their region and at what
population levels, since these could be significant alternate hosts for Heliothis parasitoids.
Most of the parasitoids listed here attack Heliothis (Krombein et al., above).

We thank P. M. Marsh, C. W. Sabrosky, and R. W. Carlson (Systematic Entomology
Laboratory, U.S. Department of Agriculture, Beltsville, Maryland) for identifying para-
sitoids. Robert Wyatt (University of Georgia, formerly Texas A&M University) identified
hostplants. Roy O. Kendall kindly offered his previously unpublished records. Adult
Schinia were determined by D. F. Hardwick (Biosystematics Research Institute, Agri-
culture Canada, Ottawa). Voucher specimens of parasitoids are in the National Museum
of Natural History (Washington, D.C.) and Texas A&M University Entomology Depart-
ment collections. Paper approved as TA-23149 by Director, Texas Agricultural Experi-
ment Station.

RICHARD S. PEIGLER AND S. BRADLEIGH VINSON, Department of Entomology, Texas
A&M University, College Station, Texas 77848.

Received for publication 3 November 1987; accepted 5 February 1988.

Journal of the Lepidopterists’ Society
42(2), 1988, 146-147

BOOK REVIEWS

ANIMAL EVOLUTION IN CHANGING ENVIRONMENTS WITH SPECIAL REFERENCE TO ABNOR-
MAL METAMORPHOSIS, by Ryuichi Matsuda. 1987. John Wiley & Sons, New York. 355
pp. $44.95.

Truth happens to an idea. It becomes true, is made true by events. Its verity is in fact
an event, a process: the process namely of its verifying itself, its verification. Its validity
is the process of its validation.—William James, The Varieties of Religious Experience
(1902)

What is truth? Specifically, what is truth in evolutionary biology? Neo-Darwinism
remains constantly under attack; Fundamentalist Christians may be its most conspicuous
antagonists, but neither Darwinism nor the neo-Darwinian synthesis has ever sat well
among secular philosophers and humanists of various persuasions, and their objections to
them surface and resurface periodically—the proverbial old wine in new bottles. The
inheritance of acquired characteristics is an idea hallowed by time if not by recent
consensus; it was a familiar theme in 19th- and early 20th-century lepidopterology, which
in those days was at the frontier of evolutionary science. Its revival and embrace in Stalin’s
Soviet Union, with the concomitant suppression of Mendelian genetics for decades, added
to its discredit elsewhere. But the idea of Lamarckian inheritance survives, and not only
among nostalgic old Reds. It has a certain appeal to idealistic young radicals with no ties
to Stalinism but with a faith in the perfectability of mankind through struggle, shared
by old Lamarckists like Paul Kammerer. It also survives apart from politics among those
who cannot accept an undirected (“random,’ but this word is always misused in such
literature) process which leads to adaptive results. This position leads to some kind of
vitalism. Animal Evolution in Changing Environments has links to the vitalist tradition.
It is an exercise in wish-fulfillment: neo-Lamarckism must be true, therefore it is. Such
declarations, alas, have no bearing on truth itself, only on our perception of what constitutes
persuasive evidence pro or con. For lepidopterists this book is a window on an acrimonious
argument which is an important part of our tradition, and is once again prominent in
the broader sphere of evolutionary biology.

The book is in two parts. Part I is a polemic in favor of the notion that radical novelty
in evolution is generated by genetic assimilation acting on components of the process of
development, particularly on metamorphosis as expressed in stressful environments. Es-
sentially the entire argument was advanced by Matsuda in an article in the Canadian
Journal of Zoology in 1982, which can be seen as a précis of the book. It is summed up
even more concisely by fig. 6 of the present volume (p. 244). Part I occupies the first 53
pages, concluding with a “proposal of pan-environmentalism”’: “Environment consists of
both morphogenetic and selective factors ... the former induces, by response of the
genotype, variation upon which the selective factor(s) works...” and, graciously, “Neo-
Darwinism may be retained as a method of analysis of the evolutionary process where
the effect of environmental change or development is minor or negligible” (pp. 52-53).
Part II occupies pages 57-355 and is a comprehensive and detailed bibliographic catalogue
of cases of abnormal metamorphosis, neoteny, etc. judged by the author to be evolution-
arily significant, arranged by taxa. (It also contains, in the aforementioned fig. 6 and
accompanying text, the clearest statement of what the author’s model is.) This is a
remarkable achievement which would be of great value to theoreticians (who in these
intellectually impoverished times in the English-speaking world know little comparative
zoology as a rule)—if only they would read it. It does not read like a novel. It reads more
like the telephone book. Matsuda is no Darwin or Gould or Dawkins, and the book suffers
from disorganization and choppiness as well as a remarkably dull style for so fervent on
advocate. And it must be read critically; like most compilers (the eccentric biogeographer
Leon Croizat is a very good comparison), Matsuda himself accepts too much at face value
and is prone to wish-fulfilling interpretation. As a student of genetic assimilation myself,
however, I confess that about half of Matsuda’s bibliography was new to me.

Because I have worked on phenotypic plasticity and genetic assimilation in butterflies

VOLUME 42, NUMBER 2 147

for some 20 years, Matsuda and IJ maintained a correspondence for some time which
ultimately led to shared frustration. It was frustrating for Matsuda because he interpreted
my results differently than I did, but was unable to convince me that he was right; it was
frustrating for me because he seemed so plainly an enthusiast who was after verification
of his ideas, which he equated with truth. (To be fair, clearly he saw me as unduly
wedded to conventional neo-Darwinism.) More recently I had a somewhat similar inter-
action with Mae-Wan Ho, of Ho and Saunders, Beyond Neo-Darwinism; interestingly,
Matsuda and Ho never did agree on the mechanism of genetic assimilation, although
both professed a post-Darwinian, neo-Lamarckian viewpoint. A sociology-of-science ap-
proach to genetic assimilation as a problem has been undertaken by an American student,
and his work should be forthcoming soon. It may clarify some of the issues, but its author
has expressed the desire to avoid ideology as a factor. I think this is a mistake.

I am unhappy with Matsuda’s handling of my own work and of butterfly polyphenism
generally. This is no trivial matter. Historically, butterfly work informed and shaped the
opinions not only of specialists like Standfuss and Fischer, but of generalizers and theo-
reticians who inspired much work and controversy—people like Kammerer, Weismann,
Schmalhausen, and Goldschmidt, to name a very mixed bunch. I am especially unhappy
because I think Matsuda was really on to something, and his premature declaration of
victory will turn so many readers off that what is valid and important in this book will
once again be relegated to oblivion. Matsuda, a morphologist by trade, had a fair grasp
of both vertebrate and invertebrate endocrinology, but his model depends on his repeated
invocation of “the mechanism of gene control,” and this does not ring true. It is akin to
the promiscuous use of similar language by paleontologists—macroevolutionists. One such,
a friend of mine in fact, invoked “reverse transcriptase” in a seminar and was asked in
all innocence by a paleontology grad student if he could explain what that was and how
it worked; of course he could not. Neither could Matsuda, and he stopped short even of
citing relevant literature, including references I gave him. Literature searching ended in
1983, but a lot of highly relevant stuff was already available by then. One searches in
vain for the real quasi-Lamarckian literature here—exciting stuff such as Gorezynski and
Steele on the immune system, John Campbell on gene automodulation, Spergel and others
on heritable drug-induced metabolic defects and hormone problems, Cullis on genotrophy
in flax—none of which would prove Matsuda’s case, but which might at least render it
more plausible. As it is, Matsuda clearly did not grasp this literature, and his death shortly
before the book went to the publisher denied him the opportunity to make a case to
impress any but the already-convinced.

Studies of wing-pattern modification in butterflies may or may not ultimately help to
unravel the Lamarckian problem, but we may continue working with the knowledge that
this book does not close the matter. Perhaps someday someone will be able to make the
assertions Matsuda made in this book, and back them up with a solid case rather than a
lot of arm-waving. Then and only then will truth “happen to” the neo-Lamarckian idea.

ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal-
ifornia 95616.

Journal of the Lepidopterists’ Society
42(2), 1988, 147-149

THE BUTTERFLY GARDEN, by Mathew Tekulsky, introd. by Robert Michael Pyle, illus.
by Susanah Brown. 1985. Harvard Common Press, Boston. x + 144 pp. $8.95 (paper),
$16.95 (cloth).

THE BUTTERFLY GARDENER, by Miriam Rothschild and Clive Farrell, illus. by Elisabeth
Luard. 1983. Michael Joseph Ltd. and Rainbird Publishing Group Ltd., London. 128 pp.
UK £7.95 (hardbound).

148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

When L. Hugh Newman and Moira Savonius published their classic book Create a
Butterfly Garden in 1967 (John Baker, London), they crystallized and explored for the
first time the theory of gardening to encourage butterflies. Native butterflies must be
tempted into gardens by their favorite nectar flowers, they asserted, and some may
establish breeding populations in gardens if the proper larval foodplants are grown.

Occasional articles on the subject have since been published in horticultural, lepidop-
terological, and environmental journals, but Rothschild and Farrell's The Butterfly Gar-
dener and Tekulsky’s The Butterfly Garden are the first book-length treatments of this
special type of horticulture to appear in almost 20 years. Both are welcome, helpful
additions to the limited and scattered literature on butterfly gardening.

The Butterfly Gardener is a strange marriage of what might better have been two
separate volumes. The first part, “The Outdoor Butterfly Gardener,” is by Miriam Roths-
child of the renowned family of British naturalists (daughter of Charles, niece of Walter),
an eminent, even awesome entomologist, naturalist, and conservationist in her own right.
The second part, “The Indoor Butterfly Gardener,” is by Clive Farrell who designed and
established the famous London Butterfly House at Syon Park. Rothschild further treats
the theme begun by Newman and Savonius, while Farrell explores a very different
subject—the butterfly zoo, wherein breeding populations of tropical butterflies are main-
tained inside a special greenhouse containing their caterpillar hostplants and nectar flow-
ers.

“Flowers and insects have travelled down the ages together, bound up in a kaleidoscopic
rainbow relationship of mutual benefit and mutual exploitation,’ writes Rothschild. Her
large garden is divided into three sections, a stone-walled kitchen garden, the grounds
surrounding the house and courtyard, and an acre of flowering hayfield in which she has
established more than 100 species of wild plants. Her seasonal approach is practical,
emphasizing cultivation, conservation, and management of grasses, shrubs, and wildflow-
ers that serve as larval foodplants and adult nectar sources for butterflies. The book includes
directions for gathering and sowing wildflower seeds, techniques for mowing fields to
minimize disturbance to butterflies in all life stages, and a butterfly garden design.
Likewise, much useful information on British butterflies is enmeshed in anecdotes and
in historical, cultural, and literary allusions that are a pleasure to wander through, just
like a butterfly-filled garden itself. Her chapter on “Grass” is one of the most original,
inspired, and delightful short essays I have read. The book deals less with butterflies and
gardening than with the author’s unique appreciation of them. Her wit and erudition,
child-like curiosity, sensitivity, and humility, as well as her love of gardens, plants, but-
terflies, and people and her understanding of how they interact, shine in every line. This
is a piece of great literature. Like Carl Sagan, she is able to distill the spirit of her subject
in popular prose.

Farrell's chapters detail the indoor culture of exotic butterflies and their foodplants as
a display for public education and enjoyment. Warmth, light, humidity, and ventilation
are important considerations in a greenhouse managed for insects as well as plants. Also,
an enclosure of this type must be very tight to prevent escape of butterflies and entry of
parasites and predators, and no pesticides can be used. In richly informative, straight-
forward prose, Farrell treats each aspect of indoor butterfly culture, concentrating on
easily reared tropical species. His level of detail is thoughtful, helpful, and indicates vast
experience and a real talent to communicate. His directions for breeding captive butterflies
are among the best available. Farrell’s contribution is unique in the literature.

_ Mathew Tekulsky’s The Butterfly Garden is the first comprehensive textbook on the
subject, and the first butterfly gardening book slanted to the United States and Canada.
The author basically reviews existing butterfly gardening lore, giving detailed abstracts
of longer original treatments. He includes three chapters on garden setup and plants, plus
information on feeders for butterflies like those for hummingbirds, hibernation boxes for
adult anglewing butterflies, bait traps, and educational activities for the butterfly garden.
Although the author lives in California, he included examples of plants and butterflies
from all parts of the continent north of Mexico. Tekulsky is an excellent writer (his chapter
transitions are especially well done), but I have a vague sense of disappointment at the

VOLUME 42, NUMBER 2 149

lack of original material. Even so, I expect the book to become a classic because it is so
thorough. It proved a useful text for a butterfly gardening class I taught in 1987.

Both books are well illustrated. Rothschild and Farrell’s has eight exquisite color pho-
tographs by Kazuo Unno, Carl Wallace, and Tony Evans, a color dust jacket, and 21 pen-
and-ink drawings and decorations by Elisabeth Luard. Tekulsky’s has 43 lovely pencil
drawings and a beautiful color cover by Susanah Brown. Luard’s designs are often very
fine (especially the frontispiece), and her drawings do successfully communicate concepts,
but poor technique frequently shows in an irritating overuse of stipple-dots. Brown’s
pencil drawings are wonderful. Both artists have depicted plants and butterflies in lifelike
poses.

Each book contains appendices on garden butterflies; wild and cultivated nectar flowers;
commercial sources of seeds, plants, butterflies, and equipment; organizations and pub-
lications dealing with horticulture, Lepidoptera, wildflowers, and conservation; and ref-
erences. Tekulsky’s section on “Further Reading” is the most complete bibliography of
the subject I have seen. The appendices are the most practical sections of the works.

For those who derive great pleasure from seeing the living, moving color of wild
butterflies among their blooms, these books are a must!

ROBERT Diric, L. H. Bailey Hortorium Herbarium, Division of Biological Sciences,
467 Mann Library, Cornell University, Ithaca, New York 14858.

Journal of the Lepidopterists’ Society
42(2), 1988, 149-151

SPHINGIDAE MUNDI. Hawk Moths of the World, by Bernard D’Abrera. E. W. Classey,
Faringdon, England. 226 pp. 79 plates. 250 x 340 mm, hard cover. £97.50 (ca. $145.00).

Somebody once remarked to Dr. O. Niemeyer, the architect who designed most of the
government buildings in Brasilia, the modern capital of Brazil, “your architecture is
beautiful, but not always functional”. Niemeyer kindly replied: “beauty is a function”.
D Abrera’s book is a beautiful book, and, in Dr. Niemeyer’s concept, this book fulfills
that function perfectly. It is artistically designed, and the plates are magnificent. The
colors of specimens, especially those of Neotropical species, are well-balanced. Except for
species represented by old, faded, and descaled specimens, the creatures would not be
ashamed of their portraits.

To help the reader understand viewpoints to follow, we provide some background
information. The first author met D’Abrera in 1979 at the British Museum (Natural
History). They frequently spent long hours discussing work, dreams, and difficulties.
D’Abrera does not regard himself a professional entomologist. He is, above all, an artist
whose main interest is to express his talents through butterflies and moths, and at the
same time to produce something beautiful and useful to others. Also, he is not supported
by taxpayers, so has to work under great pressures, especially economic pressures. It is
difficult to write books on butterflies and moths for a livelihood and to finance publication.
This includes the cost of travelling more than 12,000 miles (19,300 km) from his home
to the British Museum (Natural History), where he has to do his work, and production
financing which includes preparing plates, writing text, designing, type setting, color
separation, printing and binding, and export!

We offer this background for several reasons. First, it is important to recognize the
motivation and personal sacrifice behind D’Abrera’s books. Second, previous reviews of
D’Abrera’s books may have been unfair. We do not deny there are mistakes, but are they
solely the author’s fault, or do they reflect the chaotic state of lepidopteran taxonomy?
D Abrera clearly says that the main objective of this book “. . . is to provide, in a synoptic
form, a modern illustrated systematic list of the known species of the Hawk Moths

150 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

(Sphingidae) of the World. It is not a revision.” Is it the responsibility of authors who
produce such books to solve all the taxonomic problems before publishing something
useful? The task would take many years and involve many workers. Finally, it is possible
that some previous reviewers did not, or did not try to, understand the audience to which
D Abrera’s books are directed. Professional entomologists might feel that D’Abrera’s books
only add to the confusion already accumulated. But, what about the people who do not
have access to good collections, who cannot travel to museums around the world to examine
material, who do not have access to an extensive library? Should they wait another century
until the taxonomic chaos is sorted out? In this case, it is preferable to have his work with
errors than nothing at all.

This book is divided into four sections: introduction, text and plates, bibliography, and
index. The introduction consists of a Foreword explaining objectives, conditions under
which the work was carried out, and the style followed throughout. It is followed by an
Acknowledgements section expressing gratitude to the late Alan Hayes. The book is based
on a check list prepared by Hayes, and on the collection he curated for more than 20
years at the British Museum (Natural History). The Introduction is elegantly written, and
describes previous works and general sphingid biology. A useful two-page section entitled
“Notes for the Guidance of the Reader’ explains abbreviations and symbols used in text
and plates, and includes three figures showing important features of wings, pretarsal joint,
and head. A Systematic Catalogue of Genera, listing genera in the same order as the text,
ends the introduction.

The body of the book—the text and plates—is organized practically. The book was
designed to present text alternating with plates so that when the book is opened, text is
on the left-hand side, and figures are on the opposite page. This allows the reader to
consult the book without having to turn pages back and forth to compare text and figures.
This format could not be followed consistently, however, because when many small
specimens are on a plate, the accompanying text cannot be included on a single page.
The text provides useful information such as variation in color patterns, diagnostic char-
acters, and larval host plants. The plates show entire moths in actual size, and in several
cases the underside is also shown to distinguish similar species. The Appendix consists of
4 plates which illustrate 39 type specimens of species not represented in the British Museum
(Natural History). They vary in quality: some are good; others are poor, but useful, since
most have never been figured. Selected References omits some important works such as
M. Draudt (1931. In A. Seitz, Macrolepidoptera of the World); W. Rothschild & K.
Jordan (1907. Genera Insectorum); and H. Wagner (1913-1919. Lepidopterorum Ca-
talogus). Finally, there are two indexes, one for genera and one for species.

This book has long been needed. The last comprehensive treatment of Neotropical
Sphingidae was that of Draudt (cited above). D’Abrera recognizes approximately 1050
species, and illustrates more than 1000. Draudt listed only around 480 names and figured
260 Neotropical specimens. Further, Draudt’s work had many faults, specifically with
regard to combinations, that are corrected in D’Abrera’s book. But we disagree with the
sinking of Neococytius, and the transfer of N. cluentius to Cocytius; several cases pointed
out by D’Abrera, such as Dolba and Dolbogene, should have been synonymized following
the same criteria. D’Abrera himself regards the latter differences as “trivial.” It would
have been better to use the same criteria throughout, or to leave genera alone.

The Neotropical species were checked, and no misidentifications found. However, the
book has a few mistakes that should have been corrected by the editorial panel. The most
serious is authorship and date of family Sphingidae, which is actually Latreille 1802, not
Samouelle 1819; and the nominate subfamily name should bear the same authorship and
date. Other mistakes include incongruences between numbers of species given by the
author and numbers treated in genera. For example, under Paonias it is stated that there
are two species, but three are actually treated. The same occurs with Hemeroplanes,
where the numbers are four and five, respectively. In the plates, Callionima neivai and
Eumorpha adamsi are identified as “neavei” and “damasi.” A figure of Protaleuron
rhodogaster is stated to be in the Appendix, but no figure was found.

By examining this book, it becomes evident that Sphingidae are in desperate need of
a revision of higher classification. Several groups such as Xylophanes, Theretra and

VOLUME 42, NUMBER 2 Se |

Cechena have species that are difficult to tell apart superficially but are kept in separate
genera because they occur in different faunistic regions. Because the groups were too
large, or because of difficulties in communication, each fauna was studied separately. The
result is that each faunistic region has an independent set of genera. It would be useful
to subject the genera to a rigorous character analysis. Cladists, here is a good subject to
study!

D’Abrera did a good job on this book. For those who want to start or to organize a
collection, and to study the group, it is a good starting reference. We recognize that for
many who live in the Poor World, £97.50 is a lot of money. However, when we consider
the quality of this book, the prices of books of lesser quality, and the fact that this is a
book one would buy only once in a lifetime, it is a bargain.

We thank R. W. Hodges for critically commenting on the review.

VITOR O. BECKER, Centro de Pesquisa Agropecuaria dos Cerrados, P.O. Box 70-0023,
73300 Planaltina, DF Brazil, AND M. ALMA SOLIS, Maryland Center for Systematic
Entomology, % National Museum of Natural History, Smithsonian Institution, NHB
127, Washington, D.C. 20560.

Journal of the Lepidopterists’ Society
42(2), 1988, 152-153

MANUSCRIPT REVIEWERS, 1987

The merit of a scientific journal depends on the quality of its reviewers as well as of
its authors, but the former are usually unknown to readers. The Journal acknowledges
with gratitude the services of the people listed below from whom the editor received
manuscript reviews in 1987.

Suzanne Y. Allyson, Ottawa, Ontario, Canada
David A. Andow, St. Paul, MN

R. Robin Baker, Manchester, UK

Carol L. Boggs, Stanford, CA

Susan S. Borkin, Milwaukee, WI

M. Deane Bowers, Cambridge, MA

G. K. Bracken, Winnipeg, Manitoba, Canada
Lincoln P. Brower, Gainesville, FL

Keith S. Brown Jr., Campinas, Sao Paulo, Brazil
Richard L. Brown, Mississippi State, MS

John M. Burns, Washington, DC

J. F. Gates Clarke, Washington, DC
John M. Coffman, Timberville, VA
Steven P. Courtney, Eugene, OR
Charles V. Covell Jr., Louisville, KY
Joseph D. Culin, Clemson, SC

P. T. Dang, Ottawa, Ontario, Canada
Donald R. Davis, Washington, DC

John C. Downey, Cedar Falls, IA

Boyce A. Drummond III, Florissant, CO

John N. Eliot, Taunton, Somerset, UK
Thomas C. Emmel, Gainesville, FL

Douglas C. Ferguson, Washington, DC
Clifford D. Ferris, Laramie, WY

Lawrence F. Gall, New Haven, CT
Clyde F. Gillette, Salt Lake City, UT
Edward H. Glass, Geneva, NY
George L. Godfrey, Champaign, IL

George T. Harvey, Sault Ste. Marie, Ontario, Canada
Ronald W. Hodges, Washington, DC

Dale W. Jenkins, Sarasota, FL

A. B. S. King, Patancheru, Andhra Pradesh, India
R. L. Kitching, Armidale, NSW, Australia
Ronald M. Knaus, Baton Rouge, LA

Loke T. Kok, Blacksburg, VA

Edward N. Lambremont, Baton Rouge, LA
Robert C. Lederhouse, East Lansing, MI
Normal C. Leppla, Gainesville, FL

C. Don MacNeill, Oakland, CA

Lee D. Miller, Sarasota, FL

Thomas A. Miller, Frederick, MD
Charles W. Mitter, College Park, MD
Dennis D. Murphy, Stanford, CA

VOLUME 42, NUMBER 2

S. S. Nicolay, Virginia Beach, VA
Paul A. Opler, Fort Collins, CO

Daniel R. Papaj, Amherst, MA
Richard S. Peigler, Lakewood, CO
Daniel A. Potter, Lexington, KY
Jerry A. Powell, Berkeley, CA

John E. Rawlins, Pittsburgh, PA
Robert K. Robbins, Washington, DC
George C. Rock, Raleigh, NC

Theodore D. Sargent, Amherst, MA
Dale F. Schweitzer, Boston, MA
James A. Scott, Lakewood, CO
Oakley Shields, Mariposa, CA
William B. Showers, Ankeny, IA
Steven R. Sims, St. Louis, MO
Michael W. Stimmann, Davis, CA

Norman B. Tindale, Palo Alto, CA
M. R. Tucker, London, UK
Paul M. Tuskes, San Diego, CA

Gilbert P. Waldbauer, Urbana, IL
Susan J. Weller, Washington, DC
David A. West, Blacksburg, VA
Christer Wiklund, Stockholm, Sweden

Date of Issue (Vol. 42, No. 2): 24 May 1988

153

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EDITORIAL STAFF OF THE JOURNAL
WILLIAM E. MILLER, Editor

Dept. of Entomology
University of Minnesota
St. Paul, Minnesota 55108 U.S.A.

Associate Editors and Editorial Committee:
M. DEANE BOWERS, BOYCE A. DRUMMOND III, DOUGLAS C. FERGUSON,
LAWRENCE F. GALL, ROBERT C. LEDERHOUSE, THOMAS A. MILLER,
THEODORE D. SARGENT, ROBERT K. ROBBINS

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories
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SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London.
209 pp.

196la. Some contributions to population genetics resulting from the study of

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In General Notes and Technical Comments, references should be shortened and given
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CONTENTS

IMPACT OF OUTDOOR LIGHTING ON MOTHS: AN _ ASSESS-
MENT. Kenneth D: Frank 000
HYBRIDIZATION BETWEEN TWO SPECIES OF SWALLOWTAILS, MEIO-
SIS MECHANISM, AND THE GENESIS OF GYNANDRO-
MORPHS. Robert Blanchard t+ Henri D€SCiMON -ecccccccecnccenceee

EXTERNAL GENITALIC MORPHOLOGY AND COPULATORY MECHA-
NISM OF CYANOTRICHA NECYRIA (FELDER) (DIOPTI-
DAE)... James S..Miller 000

A NEW SPECIES OF CATOCALA FROM THE SOUTHEAST UNITED
STATES. Vernon A. Brow Jr) uu.)

BIOLOGY OF THE BLUEBERRY LEAFTIER CROESIA CURVALANA
(KEARFOTT) (TORTRICIDAE): A FIELD AND LABORATORY
sTuDY. B.M. Ponder & W. D. Seabrook 22.

HYLESIA ACUTA (SATURNIIDAE) AND ITS AGGREGATE LARVAL AND
PUPAL POUCH. Kirby L. Wolfe

EUROPEAN CORN BORER REPRODUCTION: EFFECTS OF HONEY IN
IMBIBED WATER. William E. Miller 0. OS

GENERAL NOTE

Parasitoid and hostplant records for genus Schinia (Noctuidae) in Tex-
as. Richard S. Peigler:¢7'S. Bradleigh Vinson’...

Book REVIEWS
Animal Evolution in Changing Environments with Special Reference to
Abnormal Metamorphosis. Arthur M. Shagprir0 o.ccccccecceoccsocceneeescersnececeseeeeeeee
The Butterfly Garden and The Butterfly Gardener. Robert Dirig

Sphingidae Mundi. Hawk Moths of the World. Vitor O. Becker & M. Alma
Solis

63

116

144

Volume 42 1988 Number 3

ISSN 0024-0966

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

30 August 1988

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

JULIAN P. DONAHUE, President GEORGE W. GIBBS, Vice
JERRY A. POWELL, Immediate Past President

President RONALD LEUSCHNER, Vice
JOHN N. ELIOT, Vice President | President
RICHARD A. ARNOLD, Secretary JAMES P. TUTTLE, Treasurer

Members at large:

JOHN W. BROWN M. DEANE BOWERS FREDERICK W. STEHR
MOGENS C. NIELSEN RICHARD L. BROWN JOHN E. RAWLINS
FLOYD W. PRESTON PAUL A. OPLER Jo BREWER

The object of the Lepidopterists’ Society, which was formed in May 1947 and for-
mally constituted in December 1950, is “‘to promote the science of lepidopterology in
all its branches, .... to issue a periodical and other publications on Lepidoptera, to fa-
cilitate the exchange of specimens and ideas by both the professional worker and the
amateur in the field; to secure cooperation in all measures’ directed towards these aims.

Membership in the Society is open to all persons interested in the study of Lepi-
doptera. All members receive the Journal and the News of the Lepidopterists Society.
Institutions may subscribe to the Journal but may not become members. Prospective
members should send to the Treasurer full dues for the current year, together with their
full name, address, and special lepidopterological interests. In alternate years a list of
members of the Society is issued, with addresses and special interests. There are four
numbers in each volume of the Journal, scheduled for February, May, August and
November, and six numbers of the News each year.

Active members—annual dues $25.00
Student members—annual dues $15.00
Sustaining members—annual dues $35.00
Life members—single sum $350.00
Institutional subscriptions—annual $40.00

Send remittances, payable to The Lepidopterists’ Society, to: James P. Tuttle, Treasurer,
3838 Fernleigh Ave., Troy, Michigan 48083-5715, U.S.A.; and address changes to: Julian
P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, California
90007-4057 U.S.A. For information about the Society, contact: Richard A. Arnold, Sec-
retary, 50 Cleaveland Rd., #38, Pleasant Hill, California 94523-3765, U.S.A.

To obtain:

Back issues of the Journal and News (write for availability and prices); The Com-
memorative Volume ($10.00; $6.00 to members, postpaid); A Catalogue/Checklist of
the Butterflies of America North of Mexico (clothbound $17.00, $10.00 to members;
paperbound $8.50, $5.00 to members): order (make remittance payable to “The Lepi-
dopterists’ Society”) from the Publications Coordinator, Ronald Leuschner, 1900 John
St., Manhattan Beach, California 90266-2608, U.S.A.

Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for
$40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’
Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los
Angeles, California 90007-4057. Second-class postage paid at Los Angeles, California and
additional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’
Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, California 90007-
4057.

Cover illustration: Male walnut sphinx, Laothoe juglandis (J. E. Smith), in a typical

day resting posture (1% life size). Submitted by Gerald P. Wykes, 2569 Reinhardt Road,
Monroe, Michigan 48161.

JOURNAL OF

Tue LepriporprTrerists’ SOCIETY

Volume 42 1988 Number 3

Journal of the Lepidopterists’ Society
42(3), 1988, 155-163

CYRIL FRANKLIN DOS PASSOS (1887-1986)

RONALD S. WILKINSON
228 Ninth Street, N.E., Washington, D.C. 20002

Cyril Franklin dos Passos, eminent student of Nearctic Rhopalocera,
and a Charter and Honorary Life Member of the Lepidopterists’ So-
ciety, died on 29 October 1986, only a few months before he would
have celebrated his 100th birthday.

Many who are aware of Cyril’s entomological contributions may not
know that during an extraordinarily long and full life he had two quite
distinct and successive careers. The second is of more concern to us,
but as Cyril did not begin his work on butterflies until he was past the
age of 40, something must be said of the first.

Cyril was always aware of the Portuguese heritage of his family, a
fact that this writer recalls most vividly because of the excellent Mad-
eiran wines served at the dos Passos table. Cyril’s paternal grandfather
Manoel (later Manuel) dos Passos emigrated to the United States from
Ponta do Sol, Madeira, in 1830, becoming a cobbler and later a shoe-
maker, and finally settling in Philadelphia where he married Lucinda
Cattell. There were six children, including Cyril’s father Benjamin
Franklin Dos Passos (the American family had capitalized the Portu-
guese lower case d) and an older brother, John Randolph, who would
also be of considerable importance in Cyril’s life.

John Randolph studied law and became an eminent and affluent New
York City attorney, specializing in brokerage and corporation law. He
defended trusts, opposed regulation of business by government, and
wrote extensively on these and other subjects. He took Benjamin Frank-
lin into his law firm, which became Dos Passos Brothers. Cyril, an only
child, was born in New York on 7 February 1887. His mother, Isabel
Kirker Strong, was of English descent. His father died in 1898 when
Cyril was eleven.

By his own account of his education, Cyril attended several private

156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

schools, finally spending eight years at Cutler School in New York City,
and graduating in 1905. He read law for two years at Dos Passos
Brothers, and entered New York Law School in 1907. Cyril attended
classes while working half days in his uncle’s Broad Street offices. After
receiving his LL. B. cum laude in 1909, he was admitted to the New
York bar and began to practice in the family firm, becoming a partner.

John Randolph Dos Passos represented railroad interests, and played
a large part in the reorganization of various lines. His protege Cyril
eventually entered the railroad business, becoming president and a
director of the Kansas, Oklahoma & Gulf Railway Company, which
ran from Joplin, Missouri, to Denison, Texas. When a competing line
forced Cyril’s company into receivership, he incorporated and became
secretary, treasurer, and later president of a brokerage firm, the New
York and Hanseatic Corporation. During these business activities Cyril
maintained his place in the law firm. When John Randolph died in
1917, Cyril inherited the “good will, name and business of Dos Passos
Brothers” and his uncle’s extensive law library. He was, however, Cyril
dos Passos, because his mother had encouraged him to use the Portu-
guese form. (Despite an interest in his own Portuguese heritage, John
Randolph’s son and Cyril’s cousin, the well-known literary figure John
Roderigo Dos Passos, chose to retain the Americanized capitalization.)

On 3 August 1927 Cyril married Viola Harriet Van Hise, who would
direct his interests toward entomology. She was the youngest daughter
of Anthony H. Van Hise and Harriet Louise Archer, and was born at
Newark, New Jersey, on 24 November 1891. Having earned a com-
fortable fortune, Cyril was able in 1928 to retire from law and business,
and devote his time to leisure pursuits. The couple lived in Ridgewood,
New Jersey, for several years, and enjoyed their summers at the Range-
ley, Maine “camp” which Cyril had built before the marriage for
hunting, fishing, and other recreation. Their son Manuel, who survives
him, was born on 4 February 1929.

Thus was the stage set for Cyril dos Passos’ second career. In later
life he enjoyed telling the story of his discussions with Viola about
taking up an instructive and useful pursuit. Cyril suggested art. Viola,
who had been reading copies of W. J. Holland’s butterfly and moth
books, voted for entomology. She won the day. They decided that she
would collect and study Nearctic moths, and he would devote his at-
tention to butterflies.

Collecting began in earnest at the Rangeley camp in 1929. The two
set out a sugaring trail, a line of Rummel bait traps, and eventually
had a large light trap constructed. A neighbor suggested that Cyril visit
the Department of Entomology of the American Museum of Natural
History (AMNH), and he did so, making the friendship of Frank E.

VOLUME 42, NUMBER 3 V7

Cyril F. dos Passos and friend at Quimby Pond Camps, Rangeley, Maine, 1973

Watson, who helped identify his and Viola’s captures. Cyril assisted
Watson in disinfecting and otherwise caring for the AMNH collection,
and was soon an unofficial “regular,” enjoying the encouragement of
the department curator, Frank E. Lutz.

Meanwhile, planning was under way for the magnificent French
Provincial house which would become well known to Cyril’s scientific
friends. He had always been partial to things French; his mother (who
spoke French fluently and had a French maid) had taken him to France
a number of times. Ideas for the house and grounds were assembled

158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

from books on minor chateaux and turned over to an architect. Con-
struction began in spring 1931 on ninety partially wooded acres in
Mendham, New Jersey. Cyril and Viola wished to move in at the earliest
opportunity, and were able to do so by winter.

At the Mendham “chateau” part of the second story was devoted to
specimen storage and a rapidly growing entomological library. Viola
placed her moths in traditional cabinets, separated by a partition from
Cyril’s collection. He adopted the use of Watson-Comstock store boxes,
designed by his AMNH friends Watson and William P. Comstock, and
constructed at Cyril’s expense by the museum carpenter. Cyril described
the box in his 1957 obituary of Comstock as “‘quite an ingenious, light
and inexpensive affair with top and bottom of heavy cardboard and
sides of wood. The bottom is lined with pressed cork.” As the collections
grew, Viola’s moths were moved to another spacious room, and Cyril’s
specimens filled the original area.

His earliest entomological publications were undertaken with a young
correspondent and collecting companion, L. Paul Grey, who has con-
tributed a memoir of his own to this Journal issue. The 1934 dos Passos-
Grey annotated list of Maine butterflies and its supplement were natural
beginnings for the initially Maine-oriented pair who would later revise
the Nearctic “argynnids.” Cyril gradually became a respected taxon-
omist, but during his career also published investigations of life histories
of imperfectly known butterflies. He was an avid field collector, and
because of his financial resources was able to augment his own efforts
greatly by hiring collectors to work for him in some remote areas. For
example, the first of his many papers with descriptions of new subspecies
was based on material sent to him from Newfoundland by Hugh Mclsaac.

In 1986 Cyril was appointed Research Associate in the Department
of Entomology, AMNH, through Lutz’s recommendation. The appoint-
ment was regularly renewed until the year before Cyril’s death, so that
he served a full half-century on the scientific staff. He was instrumental
in acquiring the first really large collection of North American butter-
flies to be added to the AMNH’s then relatively modest holdings, being
a substantial contributor to the purchase of Jeane D. Gunder’s 27,000
specimens in 1937. Cyril later published a catalogue of the Gunder
types; he also obtained Gunder’s library and added it to his own. As
time passed, Cyril was able to buy a number of established collections
of significance and integrate them into his previous holdings. One of
his purchases was the Alberta and Illinois material of Thomas E. Bean,
a correspondent of William Henry Edwards who supplied considerable
data used in the third volume of The Butterflies of North America;
among others were the collections of Max Rothke, E. H. Blackmore,
R. F. Sternitzky, Owen Bryant and Louis Doerfel. Types went to AMNH,

VOLUME 42, NUMBER 3 159

although Cyril retained most paratypes. When his purchases included
moths, these were placed in Viola’s cabinets.

Cyril’s concern about the significance of types led him to devise an
improved method of photographing type specimens and their labels.
An apparatus for the purpose had been described by Gunder in 1930,
but it had defects, which Cyril remedied. His folding device, utilizing
a Leica Model F camera and adjustable floodlights, could adequately
record the insect and the many labels often found on types, and could
be placed in a suitcase for travel. Cyril visited American and European
museums with his camera, and although he restricted his activities
chiefly to recording types of North American Rhopalocera, he hoped
that through cooperation all remaining type specimens of Lepidoptera
could be photographed; while many types might be lost to science over
the years, Cyril argued that photographs would create a record that
could last indefinitely. By 1945, when he published a description of his
apparatus, he had recorded as many as 1200 types. The project was
continued, and Cyril’s photographs are now in AMNH. Some have been
reproduced in his own papers and those of other workers. His original
idea still has merit.

Cyril’s early taxonomic work chiefly concerned Lycaenidae and Sa-
tyridae, although he also published on nymphalids. His first synonymic
catalogue, which appeared in 1939, was of the North American Satyr-
idae, part of a proposed but ill-fated catalogue with references to orig-
inal descriptions of all Rhopalocera north of the Mexican border, edited
by F. Martin Brown and R. W. L. Potts, which failed from lack of
funding.

During World War II Cyril suffered a great loss. Viola, who had
continued to collect moths, had a heart attack in 1939, and her activities
were restricted. She died at Rangeley on 29 August 1944. Later in that
year Cyril donated her collection, which included over 12,000 speci-
mens, to AMNH.

The collaboration with L. Paul Grey on the Argynninae, discussed
in the accompanying memoir, began to bear fruit during the war years.
Their first three joint papers appeared in 1942 and 1945; the third was
one of three independent genitalic studies (the others by B. C. S. Warren
and F. A. T. Reuss) which led to a new scheme of classification of the
subfamily, restricting the genera Argynnis and Brenthis to the Pale-
arctic region, leaving Boloria as Holarctic, and Speyeria and Euptoieta
as Nearctic genera. The dos Passos-Grey systematic catalogue of Spey-
eria was published in 1947. They concluded that although 109 published
names attributable to Speyeria were valid, only 13 species were in-
volved.

Reviewing the revision in The Lepidopterists News, Charles L.

160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Remington noted that before the work of dos Passos and Grey, “different
authors accepted a widely varying number of distinct species in the
group, many supposed affinities were entirely wrong, and uncorrelated
new ‘races’ continued to be described. The challenge of ordering the
chaos was grasped at that time by L. Paul Grey, who disposed of his
excellent collection of North American Lepidoptera to devote all his
time and space to the ‘Args.’ He was fortunate to be joined by C. F.
dos Passos, who had the means, the equipment, and the methodical
mind to scour the scattered literature, visit a number of museums to
examine carefully the types, and study the numerous nomenclatorial
problems.” The 1947 paper has recently been mentioned by Scott in
The Butterflies of North America (1986) as helping to turn the trend
of “splitting” into the more sophisticated concept of species we have
today.

When the Lepidopterists’ Society was formally constituted in May
1947, Cyril was a charter member. He served on the very early Board
of Specialists (which identified specimens for Society members) for the
family Satyridae. For the first two and a half years of its existence the
Society operated under “articles of organization,’ published in the first
issue of The Lepidopterists News. In 1950 editor Remington asked
Cyril to prepare a formal constitution and by-laws. He did so, and
served as chairman of an international committee to study and approve
the draft, which was ratified by members at the first annual meeting.
Cyril’s committee appointed temporary officers to serve the Society
until the first election by the membership, and it was due to the dos
Passos committee’s good judgment that the Society’s first president was
a lepidopterist of the very highest reputation, Cyril’s friend James H.
McDunnough, whom he had met while photographing types at the
Canadian National Collection in the 1980’s. Cyril served on the Karl
Jordan Medal Awards Committee, and was elected an Honorary Life
Member in 1978.

He attended the International Congresses of Zoology at Paris (1948),
Copenhagen (1953), and London (1958), participating in the prior col-
loquia, sections, and other activities devoted to nomenclature. He read
papers on nomenclature at Copenhagen and London, and frequently
during the decade (as well as occasionally afterwards) contributed to
the Bulletin of Zoological Nomenclature, proposing and commenting
on decisions of the International Commission, and discussing and sug-
gesting amendments to the Régles Internationales, later the Interna-
tional Code of Zoological Nomenclature. He also traveled to a number
of International Congresses of Entomology, and during these and other
journeys out of the country, he made many scientific friendships and

VOLUME 42, NUMBER 8 161

added considerable material to his entomological holdings. He espe-
cially enjoyed collecting in Europe, and did so widely.

During this active period, Cyril was appointed Research Associate
by the Carnegie Museum (1952). He continued to publish on Satyridae
and on topics as diverse as the eye colors of Colias and the ethics of
scientific criticism. On 3 September 1959, Cyril married Maria Amalia
Pita Pestana Reis, who survives him. She is the daughter of Maria Pita
de Macedo and Miguel Pestana dos Reis and was born in Ponta do Sol,
the birthplace of Cyril’s paternal grandfather. Maria AmAélia brought
Cyril much happiness, and the great success of his second marriage was
evident to his friends.

The result of a project of some years’ length appeared in 1964 as A
Synonymic List of the Nearctic Rhopalocera, this Society’s Memoir
No. 1 and, with its supplements, Cyril’s most significant and useful
contribution as single author. Much of his time in later years was devoted
to the full catalogue of Nearctic butterflies announced as forthcoming
in the introduction to his 1964 checklist. The typescript eventually grew
to seven volumes, but the work was discontinued due to the impending
appearance of Miller and Brown’s A Catalogue/Checklist of the But-
terflies of America North of Mexico (1981).

Work on such tasks as the checklist and catalogue was made easier
because Cyril had built one of the most extensive private entomological
libraries in America. When he wished to search the literature he seldom
had to leave his home, for most of the works in which North American
butterflies were described, from the 18th century onward, were there,
not only monographs but runs of journals. For an historian and bib-
liographer of entomology, the most exciting part of a visit to the dos
Passos chateau was the time spent in the library. One example will
suffice; during research on John Abbot, I was examining varying wa-
termarks in copies of Smith and Abbot’s The Natural History of the
Rarer Lepidopterous Insects of Georgia (1797) to determine the length
of its publishing history. Cyril was able to show me not one copy but
two, the second being a volume of the plates issued later with a pub-
lisher’s imprint I have never seen elsewhere than in the great library
which was donated to Wittenberg University, Springfield, Ohio, during
Cyril’s last years.

His concern with books and libraries, and his devotion to AMNH led
Cyril to give considerable assistance to the Museum’s library. He pub-
lished a number of bibliographical papers and (with William D. Field
and John H. Masters) a very useful volume, A Bibliography of the
Catalogs, Lists, Faunal and Other Papers on the Butterflies of North
America North of Mexico Arranged by State and Province (1974).

162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Some of Cyril’s bibliographical writings dealt with the actual dates of
publication of literature containing descriptions of insects, which of
course are important in determining priority. Cyril’s frequent work
with descriptions quite naturally led to an interest in the history of
entomology, to which his major contribution was his edition of William
Henry Edwards’ entomological reminiscences (1951); the manuscript
was loaned to him for the purpose by Edwards’ granddaughter.

Among Cyril’s later publications were two substantial studies (and
models for emulation) co-authored with Alexander B. Klots. The first
(1969) concerned the pierid Anthocharis midea (Hubner). As explained
in their introduction, for many years they had recognized the need for
a detailed investigation of midea to clarify such problems as geographic
variation and nomenclature, and had been accumulating data and spec-
imens toward that end. The resulting paper, which also treated life
history, foodplants, and parasites, was an exemplary discussion of a
species and its subspecies. They had also been gathering data about the
lycaenid Erora laeta (W. H. Edwards), and had jointly and severally
conducted field studies between 1934 and 1968. Their thorough paper
on the genus Erora (1982) examined early stages, ethology, ecology,
and geographic distribution of laeta and E. quaderna sanfordi dos
Passos, and provided synonymies. An extensive taxonomic study of the
satyrids Lethe portlandia (Fabricius) and L. anthedon (Clark) was
published by J. Richard Heitzman and Cyril (1974); incidentally, these
three papers provide excellent examples of the use of Cyril’s photo-
graphs of type specimens. On his own, Cyril produced his usual variety
of publications nearly to the end, although his last appearance in print
was as co-author with Clifford D. Ferris, James A. Ebner, and J. Donald
Lafontaine of an annotated list of Yukon butterflies (1983). It was
appropriate that Cyril’s final paper concerned the far-northern fauna
he loved.

Cyril donated his entomological collection to AMNH in 1980. At that
time the butterflies were contained in over 1250 store boxes. Announcing
the event in the Journal, curator Frederick H. Rindge stated that the
collection was undoubtedly “the single largest and most complete one
of North American butterflies ever made by one individual.’ He noted
that the gift included over 65,000 specimens; of those which were
mounted and identified, 57,870 were North American and 6182 were
European. There were 464 paratypes (Cyril had consistently deposited
holotypes and allotypes in AMNH) and 617 slides, chiefly of genitalia
but also of venation. Cyril intended that his correspondence and other
manuscript materials should go to AMNH, and the transfer was made
by Maria Amalia in 1987.

Cyril was a member of a number of scientific societies, and a Fellow

VOLUME 42, NUMBER 3 163

of the Royal Entomological Society of London (1950-). He was also a
Fellow of the Linnean Society of London (1977-), where many years
previously he had photographed Linnaeus’ types of North American
butterflies. Wittenberg University awarded him an honorary D.Sc. in
1965, and McDunnough named the copper Lycaena epixanthe dos-
passosi after him in 1940.

Those who knew Cyril were aware that his interests were by no
means limited to entomology. They ranged over the entire field of
natural history, including ornithology, geology and paleontology, and
extended to archaeology. He assembled extensive and valuable holdings
of classic postage stamps and covers of the United States, including
proofs, and also acquired the stamps of Nepal, Tibet, Heligoland, the
British Commonwealth, and France. He was a contributor to philatelic
journals.

Cyril was a man of many parts. He gave to entomology an unrivaled
private collection of Nearctic Rhopalocera, many examples of financial
generosity, and 50 years of publications of high professional quality.
Much has been written by historians about the professionalization of
science, a relatively recent transition from a past in which scientific
foundations were laid by workers educated in other areas. In some cases
the process has led to too rigid a distinction between professional and
amateur. It is still possible to make important and lasting contributions
to entomology without earning a graduate degree in the subject or a
related discipline; witness Cyril F. dos Passos, an amateur who made
our science his profession and served it very well.

Journal of the Lepidopterists’ Society
42(3), 1988, 164-167

MEMORIES OF CYRIL F. DOS PASSOS (1887-1986)

LIONEL PAUL GREY
Rte. 1, Box 1925, Lincoln, Maine 04457

The magnitude of “d.P.’s’” accomplishments, as chronicled in the
preceding article by R. S. Wilkinson, will attest that he was an unusual
individual. Some of my memories of him, and experiences with him
as a correspondent, collaborator and friend, perhaps will add to the
picture of the kind of person who could live so full a life, with out-
standing success in so many varied endeavors.

Quite inevitably he had to be an energetic worker, but in addition
he carried methodical procedure and time budgeting almost to the
status of an art form. When he was in his prime, few if any random
intrusions crept into schedules determined days or weeks in advance.
Usually there was a brief nap after lunch, followed by a half-hour’s
walk; aside from this, very little else in the way of relaxation except at
meals. Those were ceremonious, especially at dinner, and served at the
precise times appointed. He had uncompromising ideas about how
things should be done, including the conduct of his own life and the
running of his household. When you dined with him you wore a coat
and tie. I used to describe him (jokingly, and never within his hearing)
as “the last of the barons’. Indeed he was more the Old World aristocrat
(in the best sense) than the American businessman.

My first contact with him came shortly after he had begun studies
at the American Museum. He wrote to me for information on collecting
Maine butterflies. I learned that he had a summer camp at Rangeley,
where he came to escape his perennial troubles with hay fever. This
lead to meetings, discussions, and the beginning of a lifelong friendship.
Our first meeting probably left a lingering impression, to put it deli-
cately, since I was on a manure cart at the time, spreading richness on
my father’s farm. Anyhow, he never forgot that I was his “very first
entomological correspondent”, and he came to be my closest associate
among the amateur lepidopterists who, at that time, in the very early
30’s, comprised a rather small fraternity.

On one of our earlier outings we collected Oeneis katahdin New-
comb. Cyril had reserved a cabin for us at a sporting camp on Daicey
Pond, reached by a long hike from where we had to leave the car. The
next day we paddled across the lake and struck out through the woods
with only occasional glimpses of the distant mountain. The region then
was almost as wild as in Thoreau’s time, with few trails. We finally
came out in a clearing where a major campground of Baxter State Park
now is located; here we picked up Katahdin Stream which we followed

VOLUME 42, NUMBER 3 165

up to the steep slopes of the spruce belt, and from thence to timberline
and up over the rocks to the tableland. I still remember how amazed
I was that a city lawyer could find his way through the woods as well
as a country native, and could endure the long day’s ordeal without
apparent discomfort. Worse yet, he caught more katahdin than I did,
using the slow stalking approach while I was dashing hither and yon
making wild swoops at anything arising from the tundra. Even the
day’s end had a lesson for me, when, in the evening back at our cabin,
he spent a few minutes writing in a diary, advising me to consider the
uses of such. For him, in future years, there would be no doubt con-
cerning what he had accomplished during this particular day of his
life, nor would there be any lack of details should he ever wish to refresh
his memory about his series of katahdin butterflies and the place where
he took them.

Subsequent occasions bore out the conclusion that Cyril was a tough
physical specimen, which never would have been guessed in view of
his small stature and rather delicate frame. But then, this seems to have
been a family heritage, judging from stories told of his cousin, John,
the well-known author. Apparently the latter had lead a wild life,
soaking up “local color” in some of the most dangerous places on earth,
a midget holding his own among giants. As one commentator put it,
“John actually did the things Hemingway bragged about doing’’. This
was a clue and key to much of Cyril’s success and also to a reputation
he gained at the Museum of being difficult to get along with, namely,
the trait which psychologists term “overcompensation’’. He always was
aggressive when challenged.

In retrospect I marvel that we remained friendly, since I ventured
to argue with him rather hotly on various issues. Predictably, he was
laissez-faire capitalist in philosophy, often in a rage against the socialist
trends of the day. In view of our present national debts and deficits I
am becoming convinced that his opinions made far more sense than
mine.

Certainly our entomological relations always were very cordial. A
mountain of correspondence passed between us as we worked out details
of a major joint project, a study of nearctic “Argynnis’’. This was for
several years a shared labor, with results which at that time proved to
be somewhat controversial. As Scott has pointed out, in the latest But-
terflies of North America, we made a break in tradition toward syn-
thesis, away from the (European-fueled) tendency to finer and finer
splitting. Cyril’s role in all of this sometimes has been underestimated.
He was in every respect the senior author. He did a lion’s share of the
work and definitely was the ““maker-possible” for my contributions. A
bit of review may be of historical interest:

166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

I can pinpoint the exact moment when this project was born. It was
sparked by our shared exasperation that nobody seemed able to identify
our western material, and ignited one day when Cyril was showing me
some California specimens which he had purchased with the under-
standing that they were to have been identified by the collector. But
they arrived minus names, aside from one specimen which bore a label
stating “this looks like an oddball’’. So it did, and to our eyes so did all
the rest.

From that time onward we each began a serious study of those
enigmas, Cyril working with the literature and with the problems of
nomenclature hinging on locating and identifying type materials, while
I accumulated specimens and solicited all my collector friends for lo-
cality data. Apparently we each had intuitively analyzed our respective
strengths and adapted to integrate them. He trusted and never ques-
tioned my developing ideas concerning the speciations, while I certainly
was in no position to question his grasp of the literature or his plans
for organizing our subjects. It made for a smooth-working partnership.

Wilkinson has noted some of the instances of Cyril’s quiet philan-
thropies. I suspect Cyril derived considerable enjoyment in introducing
me to the world beyond my native turf. He paid my way for trips to
Boston, New York, Philadelphia and Pittsburgh, where I expanded our
argynnid data and was able to see numerous types. An incidental but
priceless feature of those journeys was the opportunity to meet and talk
with people such as Andrei Avinoff at Pittsburgh, Vladimir Nabokov
and Nathan Banks at the Museum of Comparative Zoology, and of
course the entomologists then at the American Museum, including Lutz,
Mitchener, Klots, and that very gracious gentleman, W. P. Comstock.
Also, Cyril took me to meetings where I heard lectures by such legendary
figures as “William T. Davis of Staten Island” (the two are inseparable)
and Robert Cushman Murphy, the great authority on oceanic birds.
All in all, quite an education for a youth from the backwoods! These
debts are gratefully acknowledged.

Also, I should express my appreciation for his cautious guidance as
we came nearer to our goal of revising the North American argynnids.
It had become apparent fairly early that the records seldom indicated
more than 6 or 8 distinct kinds of populations of those butterflies in
any single general area, a fact of significance when compared against
the listings then current, which ran to 100 “species” or more. Collations
of local data sets also indicated numerous instances of intergrading. I
felt that we were on the verge of solving the puzzle, but Cyril then
urged that we had a duty to enlarge the perspective to include whatever
might appear when native argynnids were compared with those on the
other continents. A fairly comprehensive genitalic survey of the Nearc-

VOLUME 42, NUMBER 8 167

tic species had been completed, but the task of studying the world
argynnines was slowed by difficulties in procuring the needed material.

Thus, it was 1947 before our Systematic Catalogue of Speyeria finally
was published. A few corrections have been required, both to Cyril’s
nomenclatorial and other data and to my concepts of the speciations,
but these have been gratifyingly few considering the tangle we were
dealing with. It was, indeed, as Scott has noted, a turning point in the
philosophy of butterfly “‘species’’, but unfortunately we were too much
captives of the times to have broken loose from the addiction to “‘sub-
species’. In fact, back at that time it would have been unwise to have
reduced the number of such taxa, even though many are hardly more
than unstable local color forms. As Cyril had warned me, we encoun-
tered considerable resistance to our radically altered classification, the
big difficulty lying with our downgrading of many taxa traditionally
hallowed as “species”. To help soften those outrages to custom we
thought it essential to retain numerous subspecific names and even to
propose some new ones for the same purpose of indicating some of the
connecting links and the widespread continuity of intergrading within
the series discriminated as constituting polytypic species.

After those prolonged associations it seemed only natural that we
should remain close and affectionate friends. When failing health forced
him to curtail his entomological activities he presented me with the
entire contents of his laboratory. I remain surrounded by reminders of
his kindness. :

In closing, it seems fitting to recall one of Cyril’s most striking man-
nerisms. He never lingered when the time came to part—a wave of
the hand, a “bye-bye’’ and he would turn abruptly and walk away.
Fond recollections remain.

Journal of the Lepidopterists’ Society
42(3), 1988, 168-183

ANNOTATED BIBLIOGRAPHY OF THE
ENTOMOLOGICAL PUBLICATIONS OF
CYRIL F. DOS PASSOS (1887-1986)

RONALD S. WILKINSON
228 Ninth Street, N.E., Washington, D.C. 20002

This bibliography includes all of Cyril dos Passos’ entomological
publications known to me except his abstracts of current literature
published in early issues of The Lepidopterists News. All publications
have been personally examined; in most cases separates or reprints had
been furnished to me by the author during his lifetime with a bibli-
ography in mind, and a search of the literature has revealed additional
publications.

Arrangement of entries is chronological according to actual dates of
publication, which were determined by examination of journal issues,
and through correspondence and other methods. Items identified by
asterisk (*) have not been dated precisely, but these do not affect the
chronology. Titles of publications are exact, although I have uniformly
italicized generic and specific names. Citations of place of publication
are followed by stated date and, in parentheses, actual date if it differs
or is more precise. The summaries of content are, of necessity, somewhat
uneven; as might be expected, those of notes or brief papers may include
details that would not have been mentioned had the work been of
greater length. All new names proposed by dos Passos have been in-
cluded. When new species, subspecies, or forms were named, I have
provided the type locality, either the name of the collector of the
holotype or the collection from which it was selected (the latter if the
collector is not specifically named in the paper), and the holotype
repository, all as indicated by dos Passos. All fixations of type localities
and designations of lectotypes and neotypes have been documented. In
the summaries, names are given the standing accorded to them by dos
Passos, and are printed as they appeared in print (forms are italicized,
for example).

I am grateful for the extensive assistance of Maria Amalia dos Passos
and F. Martin Brown, and for the kind cooperation of L. Paul Grey,
who has informed me that he holds, entrusted to him by the author,
an incomplete dos Passos manuscript not yet prepared for publication.

The following are some abbreviations used: AMNH: American Mu-
seum of Natural History, New York City; CM: Carnegie Museum of
Natural History, Pittsburgh; USNM: U.S. National Museum, Smithson-
ian Institution, Washington, D.C.; j.a.: junior author; s.a.: senior author;
t.l.: type locality. Postal abbreviations are used for States.

VOLUME 42, NUMBER 3 169

10.

1934

With L. P. Grey, j.a. A list of the butterflies of Maine with notes concerning some
of them. Can. Entomol. 66:188-192, Aug. 1934 (2 Sep. 1934). 110 taxa reported
including subspecies, forms.

With L. P. Grey, j.a. Additions and corrections to “‘A list of the butterflies of Maine.”
Can. Entomol. 66:278, Dec. 1934 (81 Dec. 1934). 7 taxa added; 2 deleted.

1935

. Some butterflies of southern Newfoundland with descriptions of new subspecies

(Lepid. Rhopal.). Can. Entomol. 67:82-88, Apr. 1935 (4 May 1935). Discussion of
collection made in 1934 by H. Mclsaac; Coenonympha inornata mcisaaci, n. ssp.
(t.l. Doyles Station, Newfoundland, H. MclIsaac); Oeneis jutta terrae-novae, n. ssp.
(t.l. Doyles Station, Newfoundland, H. MclIsaac); Argynnis atlantis canadensis, n.
ssp. (t.l. Doyles Station, Newfoundland, H. Mclsaac); Phyciodes tharos arctica, n.
ssp. (t.l. Table Mountain, Port au Port, Newfoundland, G. C. Hall); all holotypes in
AMNH;; Mclsaac’s collection included 21 taxa; 12 additional taxa listed as occurring
in Newfoundland.

1936

. Further notes on the butterflies of southern Newfoundland. Can. Entomol. 68:98,

May 1936 (6 Jun. 1936). In 1936 H. Mclsaac collected 2 species not previously
reported from Newfoundland.

. The life history of Calephelis borealis (Lepidoptera). Can. Entomol. 68:167-170, 1

pl. incl. 6 figs., Aug. 1936 (29 Aug. 1936). C. borealis appears to be single-brooded
in NJ; females observed ovipositing on Senecio obovatus Muhlenberg; insect reared;
egg, instars of larva, pupa discussed, figured.

. Some early stages of Brenthis montinus Scudder (Lepidoptera—Nymphalidae).

Can. Entomol. 68:239-241, 1 pl. incl. 4 figs., Nov. 1936 (5 Dec. 1936). Specimens
confined over various plants; all eggs on Solidago cutleri Fernald except some on
sides of breeding cages; perhaps eggs are dropped on ground, fall into detritus in
which larvae, upon emerging, hibernate; egg, first instar larva discussed, figured.

1938

Some new subspecies of North American Lycaenidae (Lepid.). Can. Entomol. 70:
45-48, 1 pl. incl. 16 figs., Mar. 1938 (2 Apr. 1938). Material from various collections
described as Lycaena nivalis browni, n. ssp. (t.1. Snowslide Canyon, 8 mi [13 km]
from Montpelier, ID, W. J. Gertsch); Plebeius saepiolus gertschi, n. ssp. (t.l. Cedar
Breaks, nr. Cedar City, UT, W. J. Gertsch); Plebeius icarioides buchholzi, n. ssp.
(t.l. White Mts., AZ, 8500 ft [2591 ml], E. Y. Dawson); Plebeius acmon lutzi, n. ssp.
(t.l. Snowslide Canyon, 8 mi [13 km] from Montpelier, ID, W. J. Gertsch); all
holotypes in AMNH; holotypes, allotypes figured.

. Synonymic notes on Aglais milberti (Godart) with the description of a new subspecies

(Lepidoptera—Nymphalidae). Can. Entomol. 70:72-78, 1 pl. incl. 6 figs., Apr. 1938
(14 May 1938). Type locality of A. milberti fixed; Godart’s type figured; A. m.
rothkei Gunder jr. synonym of milberti; Vanessa furcillata Say distinct form of
milberti; V. m. var. subpallida Cockerell distinct form of milberti; Cockerell’s type
figured; Aglais m. viola, n. ssp. (t.1. Doyles Station, Newfoundland, H. Mclsaac);
holotype in AMNH; holotype, allotype figured.

. The types of Lepidoptera described by J. D. Gunder. Am. Mus. Novit. No. 999, 16

pp., 26 Jul. 1938. Gunder’s collection of North American Lepidoptera (chiefly west-
ern Rhopalocera), recently acquired by AMNH, contains type material for 171 of
212 taxa described by him; all types listed; references given to original descriptions,
type localities, collectors’ names, disposition of types when not at AMNH.

A new race of Euphydryas chalcedona Dbldy. & Hew. from Arizona (Rhopalocera—
Nymphalidae). Can. Entomol. 70:199-200, 1 pl. incl. 4 figs., Oct. 1938 (5 Nov. 1938).
Material received for several years as E. hermosa (Wright) described as Euphydryas

170

ie

12.

13.

14.

15.

16.

17.

18.

ug;

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

chalcedona klotsi, n. ssp. (t.1. Roosevelt Lake, AZ, D. K. Duncan); holotype in AMNH;
holotype, allotype figured.

A new race of Basilarchia archippus Cramer from Louisiana (Rhopalocera—Nym-
phalidae). Can. Entomol. 70:248, 1 pl. incl. 4 figs, Dec. 1938 (31 Dec. 1938). A
previously undescribed “race” in Gulf States, Basilarchia archippus watsoni, n. ssp.
(t.l. Alexandria, LA, J. Woodgate); holotype in AMNH; holotype, allotype figured.

1939

A catalogue of the original descriptions of the Rhopalocera found north of the
Mexican border. Part two: the Satyridae. Bull. Cheyenne Mountain Mus. 1, part 2,
13 pp., 20 Apr. 1939. Synonymic catalogue with type localities and full citations to
descriptions.

1940

A new subspecies of Erora laeta Edwards from Arizona and New Mexico (Rho-
palocera: Lycaenidae). Am. Mus. Novit. No. 1052, 2 pp., 15 Mar. 1940. Erora laeta
sanfordi, n. ssp. (t.l. White Mts., AZ, 8000 ft [2438 m], D. K. Duncan); holotype in
AMNH.

A new subspecies of Erebia discoidalis Kirby (Rhopalocera: Satyridae). Am. Mus.
Novit. No. 1053, 2 pp., 22 Mar. 1940. Material from Alberta to AK described as
Erebia discoidalis mcdunnoughi, n. ssp. (t.l. White Horse, AK, J. A. Kusche); ho-
lotype in AMNH.

A new species of Incisalia from southern California (Rhopalocera, Lycaenidae).
Can. Entomol. 72:167-168, Aug. 1940 (31 Aug. 1940). Incisalia doudoroffi, n. sp.
(t.l. Big Sur, Monterey Co., CA, M. Doudoroff); holotype in AMNH.

On the occurrence of Papilio polydamas Linnaeus within the United States. Can.
Entomol. 72:188, Sep. 1940 (30 Sep. 1940). P. p. lucayus not only subspecies in U.S.
as P. p. polydamas also occurs here; TX specimens in collections acquired by dos
Passos.

1942

With C. D. Michener, s.a. Taxonomic observations on some North American Strymon
with descriptions of new subspecies (Lepidoptera: Lycaenidae). Am. Mus. Novit.
No. 1210, 7 pp., 5 figs., 13 Nov. 1942. Strymon of calanus group discussed; S.
liparops (Boisduval & LeConte) synonym of S. favonius (J. E. Smith), so species
usually called liparops becomes strigosus Harris; Strymon strigosus aliparops, n.
ssp. (t.l. Glenwood Springs, CO, Oslar); holotype in AMNH,; S. liparops (Fletcher)
homonym of liparops (Boisduval & LeConte), renamed Strymon strigosus fletcheri,
n. name; lectotype designated (cotype of Thecla strigosa liparops Fletcher, USNM);
neotype designated for S. edwardsii (Grote & Robinson); genitalia figured.

With L. P. Grey, j.a. Two new North American subspecies of Argynnis, with some
revisional notes (Lepidoptera: Nymphalidae). Am. Mus. Novit. No. 1214, 6 pp., 1
fig., 8 Dec. 1942. Material from Gunder collection and others described as Argynnis
utahensis linda, n. ssp. (t.1. Heyburn Peak, Sawtooth-Boise, ID, 9500-10,000 ft [2896—
3048 m], C. W. Herr); Argynnis coronis carolae, n. ssp. (t.l. Charleston Park, Clark
Co., NE, E. Schiffel); both holotypes in AMNH; both holotypes figured; A. pfoutsi
Gunder a synonym of A. platina Skinner; A. albrighti Gunder appears to be form
of A. mcdunnoughi Gunder, a subspecies of A. utahensis Skinner; A. semivirida
McDunnough correctly placed with A. nevadensis W. H. Edwards, a species distinct
from A. utahensis; A. chitone W. H. Edwards a subspecies of A. hesperis W. H.
Edwards; A. snyderi Skinner not a subspecies of A. coronis Behr; A. monticola Behr
a synonym of A. zerene Boisduval; A. malcolmi Comstock a race of A. zerene; A.
conchyliatus Comstock might be subspecies rather than form of A. zerene.

1943

Some new subspecies of Incisalia from North America (Lepidoptera, Lycaenidae).
Am. Mus. Novit. No. 1230, 5 pp., 1 Jun. 1943. Scudder, not Minot, is author of

VOLUME 42, NUMBER 3 L7G

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Incisalia; type localities fixed for I. augustus (Kirby), I. augustus croesioides Scud-
der, I. iroides (Boisduval), I. henrici (Grote & Robinson), lectotypes designated for
these taxa; Incisalia iroides annetteae, n. ssp. (t.l. New Mexico, ex J. D. Gunder
collection); Incisalia augustus helenae, n. ssp. (t.1. Doyles Station, Newfoundland,
H. Mclsaac); Incisalia henrici margaretae, n. ssp. (t.l. 8 mi [13 km] E Deland, FL,
B. Heineman); all holotypes in AMNH.

A new Riodinid record. Can. Entomol. 75:108, Jun. 19438 (23 Jul. 1943). H. A.
Freeman furnished 4 specimens of Apodemia walkeri Godman & Salvin, Browns-
ville, TX, 2-9 Jun. 1940, a new U.S. record.

A correction. Can. Entomol. 75:178, Sep. 1943 (21 Oct. 1943). In 3 above Polygonia
comma (Harris) reported from Newfoundland; specimens were P. marsyas (W. H.
Edwards).

1945

Some collections of Lepidoptera. J. New York Entomol. Soc. 53:62, Mar. 1945 (4
May 1945). Since 1985 dos Passos acquired collections of E. H. Blackmore, Victoria,
British Columbia; T. E. Bean (IL, Alberta material); M. Rothke, Scranton, PA; R.
F. Sternitzky, San Francisco, CA; O. Bryant (including his Arctic material); L.
Doerfel, Newark, NJ; most paratypes retained; other types now in AMNH.

With L. P. Grey, j.a. A new species and some new subspecies of Speyeria (Lepi-
doptera, Nymphalidae). Am. Mus. Novit. No. 1297, 17 pp., 30 figs., 10 May 1945.
Material from various collections described as Speyeria wenona, n. sp. (t.l. Cerro
Potosi, Municipio de Galeana, Nuevo Leon, Mexico, 12,000 ft [8658 mJ], R. A.
Schneider); Speyeria cybele letona, n. ssp. (t.l. City Creek Canyon, Salt Lake City,
UT, 4500 ft [1872 m], W. L. Phillips); Speyeria coronis simaetha, n. ssp. (t.l. Black
Canyon, Cascade Mts., nr. Brewster, WA, J. C. Hopfinger); Speyeria zerene myr-
tleae, n. ssp. (t.l. San Mateo, CA, W. F. Breeze); Speyeria z. sinope, n. ssp. (t.l. Estes
Park area, Rocky Mt. National Park, CO, 8000 ft [2488 m], R. Weist); Speyeria z.
cynna, n. ssp. (t.1. Humboldt National Forest, Ruby Valley, Elko Co., NV, E. Schiffel);
Speyeria callippe elaine, n. ssp. (t.l. Butte Falls, OR, ex J. D. Gunder collection);
Speyeria c. sierra, -n. ssp. (t.l. Gold Lake, Sierra Co., CA, C. Hill); Speyeria c.
harmonia, n. ssp. (t.1. Mt. Wheeler, Snake Range, nr. UT border, NV, 8000 ft [2438
mj], ex J. D. Gunder collection); Speyeria montiviga [sic.] [montivaga] secreta, n.
ssp. (t.l. Estes Park area, Rocky Mt. National Park, CO, 8000 ft [2438 m], R. Weist);
Speyeria hydaspe conquista, nu. ssp. (t.l. Little Tesuque Canyon, nr. Sante Fe, NM,
8000 ft [2438 ml], A. B. Klots); Speyeria atlantis lurana, n. ssp. (t.l. Harney Peak,
Black Hills, SD, A. C. Frederick); Speyeria a. wasatchia, n. ssp. (t.l. Payson Canyon,
Payson, UT, L. D. Pfouts); Speyeria a. tetonia, n. ssp. (t.l. Teton Mts., WY, ex J. D.
Gunder collection); Speyeria a. viola, n. ssp. (t.l. Trail Creek, Sawtooth Mts., ID,
7400 ft [2256 m], C. W. Herr); all holotypes in AMNH,; all holotypes figured.
With L. P. Grey, j.a. A genitalic survey of Argynninae (Lepidoptera, Nymphalidae).
Am. Mus. Novit. No. 1296, 29 pp., 54 figs., 14 Sep. 1945. Genitalia generally
discussed, distinctive characteristics given for genera Boloria (21 species), Brenthis
(3 species), Argynnis (18 species), Speyeria (7 species), Euptoieta (2 species); 3
independent genitalic studies of Argynninae conducted (present and those of B. C.
S. Warren and F. A. T. Reuss); agreements and disagreements reviewed; Brenthis
and Argynnis should be restricted to Palearctic species; Palearctic Brenthis should
be set apart from Holarctic Boloria, which may require several genera or subgenera;
Nearctic Speyeria distinct from Argynnis; genitalia figured.

1946

“1945.”" The photography of types of Lepidoptera. Bull. Brooklyn Entomol. Soc.
n.s. 40:166-169, 4 figs., Dec. 1945 (15 Mar. 1946). Improvement on apparatus of J.
D. Gunder described, illustrated; type specimens may be photographed with all
their labels, without reflections, shadows; cooperative effort proposed to photograph
all types of Lepidoptera.

26. With B. C. S. Warren, s.a., and L. P. Grey. Supplementary notes on the classification

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of Argynninae (Lepidoptera, Nymphalidae). Proc. Roy. Entomol. Soc. London ser.
B, 15:71-73, 15 Jun. 1946. New tribal division, Boloriidi, proposed to include Boloria,
Proclossiana, Clossiana; the other tribe, Argynnidi, includes Brenthis, Yramea,
Issoria, Speyeria, Fabriciana, Mesoacidalia, Damora, Pandoriana, Childrena, Ar-
gyreus, Argyronome; Reuss’s Neoacidalia synonym of Speyeria.

1947

Notes on Grinnell’s types of Erynnis Schrank (Lepidoptera, Hesperiidae). Am. Mus.
Novit. No. 1337, 3 pp., 24 Feb. 1947. E. callidus (Grinnell) should include E. c.
callidus (Grinnell), E. c. pernigra (Grinnell), E. c. lilius (Dyar); lectotype designated
for Thanaos callidus Grinnell.

Erebia youngi Holland, its subspecies and distribution (Lepidoptera, Satyridae). Am.
Mus. Novit. No. 1348, 4 pp., 14 Jul. 1947. E. herscheli Leussler a subspecies of E.
youngi Holland; Erebia y. rileyi, n. ssp. (t.l. Mt. McKinley National Park, AK, ex J.
D. Gunder collection); holotype in AMNH.

With L. P. Grey, j.a. Systematic catalogue of Speyeria (Lepidoptera, Nymphalidae)
with designations of types and fixations of type localities. Am. Mus. Novit. No. 1370,
30 pp., 12 Dec. 1947. 109 names published prior to end of 1946 found attributable
to Speyeria; these represent 13 valid species; remaining 96 names considered sub-
species; 58 synonyms; valid species are S. diana (Cramer), S. cybele (Fabricius), S.
aphrodite (Fabricius), S. idalia (Drury), S. nokomis (W. H. Edwards), S. edwardsii
(Reakirt), S. coronis (Behr), S. zerene (Boisduval), S. callippe (Boisduval), S. egleis
(Behr), S. atlantis (W. H. Edwards), S. hydaspe (Boisduval), S. mormonia (Bois-
duval); checklist followed by catalogue, in which names and citations supplemented
by type localities, deposition of types, distribution, references to figures, descriptions
of preparatory stages when applicable; Speyeria mormonia eurynome ab. igeli, n.
name (type is type of Argynnis eurynome ab. eris Igel); Speyeria mormonia eu-
rynome ab. fieldi, n. name (type is type of Argynnis eurynome clio trans. form
gunderi Field); type localities fixed for S. diana (Cramer), S. c. cybele (Fabricius),
S. a. aphrodite (Fabricius), S. a. alcestis (W. H. Edwards), S. idalia (Drury), S. n.
nokomis (W. H. Edwards), S. edwardsii (Reakirt), S. c. coronis (Behr), S. c. snyderi
(Skinner), S. c. halcyone (W. H. Edwards), S. z. zerene (Boisduval), S. z. hippolyta
(W. H. Edwards), S. z. platina (Skinner), S. c. callippe (Boisduval), S. c. rupestris
(Behr), S. c. juba (Boisduval), S. c. laurina (Wright); S. e. egleis (Behr), S. e. adiaste
(W. H. Edwards), S. a. atlantis (W. H. Edwards), S. a. hesperis (W. H. Edwards),
S. a. irene (Boisduval), S. a. electa (W. H. Edwards), S. a. lais (W. H. Edwards), S.
h. hydaspe (Boisduval), S. h. rhodope (W. H. Edwards), S. m. mormonia (Boisduval),
S. m. erinna (W. H. Edwards), S. m. arge (Strecker), S. m. artonis (W. H. Edwards),
S.m. eurynome (W. H. Edwards), and in synonymies for Papilio daphnis Cramer,
P. daphnis Martyn, Argynnis cypris W. H. Edwards, A. monticola Behr, A. liliana
var. baroni W. H. Edwards, A. wrighti Wright, A. nevadensis r. meadii trans. form
gerhardi Gunder, A. adiante Boisduval, A. montivaga Behr 1863, A. montivaga
Behr 1864, A. astarte W. H. Edwards 1862, A. astarte W. H. Edwards 1864, A.
cornelia W. H. Edwards, A. clio W. H. Edwards, and A. eurynome trans. form
brucei Gunder; types designated in synonymies for A. astarte W. H. Edwards 1862,
A. astarte W. H. Edwards 1864, A. montivaga Behr 1864; lectotypes designated
for S. cybele carpenterii (W. H. Edwards), S. c. charlottii (Barnes), S. aphrodite
alcestis (W. H. Edwards), S. a. columbia (Hy. Edwards), S. nokomis nitocris (W.
H. Edwards), S. n. coerulescens (Holland), S. edwardsii (Reakirt), S. c. coronis
(Behr), S. c. semiramis (W. H. Edwards), S. c. snyderi (Skinner), S. z. zerene
(Boisduval), S. z. hippolyta (W. H. Edwards), S. z. behrensi (W. H. Edwards), S. z.
bremnerii (W. H. Edwards), S. z. platina (Skinner), S. c. callippe (Boisduval), S. c.
liliana (Hy. Edwards); S. c. rupestris (Behr), S. c. juba (Boisduval), S. c. laura (W.
H. Edwards), S. c. nevadensis (W. H. Edwards), S. c. macaria (W. H. Edwards), S.
c. meadii (W. H. Edwards), S. e. egleis (Behr), S. e. adiaste (W. H. Edwards), S. e.
atossa (W. H. Edwards), S. e. oweni (W. H. Edwards), S. a. atlantis (W. H. Edwards),
S. a. hesperis (W. H. Edwards), S. a. nikias (Ehrmann), S. a. nausicaa (W. H.

VOLUME 42, NUMBER 3 bs

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Edwards), S. a. chitone (W. H. Edwards), S. a. irene (Boisduval), S. a. lais (W. H.
Edwards), S. h. hydaspe (Boisduval), S. h. rhodope (W. H. Edwards), S. h. sakuntala
(Skinner), S. m. mormonia (Boisduval), S. m. bischoffui (W. H. Edwards); S. m. opis
(W. H. Edwards), S. m. washingtonia (Barnes & McDunnough), S. m. erinna (W.
H. Edwards), S. m. arge (Strecker), S. m. artonis (W. H. Edwards), S. m. eurynome
(W. H. Edwards), and S. m. luski (Barnes & McDunnough), and in synonymies for
A. cypris W. H. Edwards, A. monticola Behr, A. inornata W. H. Edwards, A.
wrighti Wright, A. adiante Boisduval, A. montivaga Behr 1868, and A. atlantis ab.
chemo Scudder; neotypes designated for S. cybele leto (Behr), S. a. aphrodite (Fa-
bricius), S. idalia (Drury), S. n. nokomis (W. H. Edwards), S. coronis halcyone (W.
H. Edwards), and in synonymies for P. daphnis Cramer, P. daphnis Martyn, A.
aphrodite f. arizonensis Elwes, A. clio W. H. Edwards.

1948

The eye colors of some Colias collected in New Jersey (Lepidoptera, Pieridae). Proc.
Entomol. Soc. Washington 50:35-88, Feb. 1948 (27 Feb. 1948). 53% of individuals
of C. philodice-eurytheme complex collected near Mendham had black eyes, 47%
yellow-green when alive; percentages given by sex; no references found to black
eyes in complex; yellow-green is normal color.

The care of a collection and library. Lepid. News 2:27, Mar. 1948 (6 Apr. 1948).
Hints for protection, maintenance of entomological collections, leather bindings.
Critics and criticisms. Lepid. News 2:41, Apr. 1948 (7 Jun. 1948). Ethics of scientific
criticism, prompted by short critical reviews included in notices of current ento-
mological literature in News.

Notes on the disappearance of Polygonia gracilis at Rangeley, Maine, in 1947.
Lepid. News 2:59, May 1948 (30 Jun. 1948). Very wet spring had sc.ious effect on
all 3 Polygonia species at Rangeley (faunus, progne, gracilis); no adults seen during
summer; gracilis, a rare and local insect, may not reappear.

The occurrence of anthoxanthins in the wing pigments of some Nearctic Oeneis
(Rhopalocera: Satyridae). Entomol. News 59:92-96, Apr. 1948 (2 Jul. 1948). Chem-
istry of pigments in wings of Nearctic Oeneis assists greatly in their systematic
arrangement without conflicting with result obtained by genitalic examination;
presence of anthoxanthins in scales of O. uhleri (Reakirt) and O. taygete Geyer
groups suggests need for rearrangements; O. nahanni Dyar a subspecies of uhleri
or should be placed next to it; O. chryxus ivallda (Mead) should have specific
standing; describes test for anthoxanthins not injurious to specimens.

1949

New butterflies from Mount McKinley National Park, Alaska, with a review of
Erebia rossii (Rhopalocera, Satyridae). Am. Mus. Novit. No. 1389, 17 pp., 28 figs.,
6 Jan. 1949. Oeneis mckinleyensis, n. sp. (t.l. McKinley Park, AK, ex C. F. dos
Passos collection); holotype in AMNH; holotype, allotype, 3 paratypes figured; sub-
species of E. rossii (Curtis) reviewed, lectotype designated for E. r. kuskoquima
Holland; Erebia r. gabrieli, n. ssp. (t.l. Mount McKinley Park, AK, 3500 ft [1067
m], ex G. P. Engelhardt and C. F. dos Passos collections); holotype in AMNH;
holotype, allotype, 2 paratypes figured.

The distribution of Oeneis taygete Geyer in North America with descriptions of
new subspecies (Lepidoptera, Satyridae). Am. Mus. Novit. No. 1399, 21 pp., 16 figs.,
26 Jan. 1949. Type locality fixed for O. taygete, neotype designated; Oeneis t.
gaspeensis, n. ssp. (t.l. Mt. Albert, Quebec, A. E. Brower); holotype in AMNH;
Oeneis t. fordi, n. ssp. (t.l. Kuskokwim River, AK, A. Stecker); holotype in CM;
Oeneis t. edwardsi, n. ssp. (t.1. San Juan Mts., Hinsdale Co., CO, B. Rotger); holotype
in AMNH,; holotypes, allotypes figured.

[Letter to editor.] Lepid. News 3:19-20, Feb. 1949 (7 Apr. 1949). Actions of Section
on Nomenclature and F. Hemming in amending Régles at 1948 International Con-
gress of Zoology defended against criticisms of C. W. Sabrosky.

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JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The photography of types of Lepidoptera. Lepid. News 3:41-42, 1 fig., Apr.-May
1949 (11 Jul. 1949). Revision and condensation of 25 above.

A visit to the home of the late William Henry Edwards at Coalburg, West Virginia.
Lepid. News 3:61-62, 1 fig., Jun. 1949 (23 Sep. 1949). Account of visit to State Dept.
of Archives and History, Charleston, WV, to consult Edwards’ journals, notebooks,

_other papers; and to house the entomologist built in 1869, where other Edwards

manuscripts examined; figure depicts house.

Notes on two Incisalia types (Lepidoptera, Lycaenidae). Can. Entomol. 81:180-181,
Jul. 1949 (25 Oct. 1949). Neotypes designated for I. hadros Cook & Watson, I.
henrici var. solatus Cook & Watson.

1950

A correction. Lepid. News 4:15, 1950 (20 May 1950). dos Passos erred in note to
literature abstract in News 3:109; combination Malacosoma fragile correct as to
ender.

i epaeeeanee Butterflies and Moths: Trans. Lepid. Soc. Japan 1:40-42, Aug.
1950.* Summary of current American activity in letter invited by journal editor;
translated by him into Japanese.

With D. B. Stallings, s.a. The Lepidopterists’ Society: Report of the Organization
Committee. Lepid. News 4:38, 1950 (16 Nov. 1950). Committee formed to consider
proposed constitution and by-laws drafted by dos Passos completed work, submits
finished texts for Society ratification; temporary Society officers appointed.

1951

The entomological reminiscences of William Henry Edwards with an introduction
and annotations. J. New York Entomol. Soc. 59:129-186, Sep. 1951 (23 Aug. 1951).
Previously unpublished autobiographical MS written by Edwards in old age, edited
and with introduction by dos Passos.

On the proposal that the trivial name “ajax’’ Linnaeus, 1758 (as published in the
binomial combination “Papilio ajax’’) should be suppressed by the International
Commission on Zoological Nomenclature under its plenary powers. [Reference Z.
N. (S.) 192.] Bull. Zool. Nomen. 2:349-350, 28 Sep. 1951. In recent years ajax used
for 2 different Nearctic butterflies which have valid names, Papilio polyxenes as-
terius Stoll and P. marcellus Cramer; ajax not properly applicable to either; it is
desirable to suppress name.

1952

Application to the International Commission on Zoological Nomenclature to recon-
sider and rephrase in part their decision suspending the “Régles” concerning “Papilio
plexippus” Linnaeus, 1758, insofar as that decision refers to a figure in Holland’s
“Butterfly book.” [Reference Z. N. (S.) 323.] Bull. Zool. Nomen. 6:278-283, 23 Jul.
1952. Original description of P. plexippus applies to 2 species, 1 American, 1 Oriental;
Commission decided to apply name to the American species, as figured by W. J.
Holland in Butterfly book; Holland’s figure of Danaus plexippus menippe (Hiibner),
so when Opinion is rendered reference should be made to an accurate figure of D.
p. plexippus (Linnaeus).

With L. P. Grey, s.a., and A. B. Klots. The “niobe /cydippe /adippe” problem (Class
Insecta, Order Lepidoptera, Family Nymphalidae) with suggestions for its solution.
[Reference Z. N. (S.) 79.] Bull. Zool. Nomen. 6:323-325, 29 Aug. 1952. Papilio niobe
Linnaeus 1758 presents no nomenclatorial problem; P. cydippe Linnaeus 1761, a
synonym of niobe, long misdetermined as a different butterfly, the “High Brown
Fritillary”; P. adippe Linnaeus 1767, a new name for cydippe and synonym of
niobe, also misdetermined as “High Brown Fritillary’’; to settle scientific name of
latter, Commission should suppress certain usages, validate name adippe for insect
as from 1775 when used by Denis & Schiffermiiller.
[Book review.] Die Schmetterlinge Mitteleuropas. By Walter Forster & Theodor

VOLUME 42, NUMBER 3 175

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A. Wohlfahrt. Lepid. News 6:79-80, 1952 (17 Nov. 1952). First installments of vols.
1 and 2 reviewed.

In support of the application to suspend the rules to (a) validate seven generic names
of Linnaeus as of 1758, and designate their type species (b) suppress the generic
name Phalaena” Linnaeus, 1758, give preference to its typical subgenus “Noctua,”
declare “Noctuidae” the correct name for the family, and (c) validate one generic
name of Linnaeus as of 1767 and designate its type species (Class Insecta, Order
Lepidoptera). [Reference Z. N. (S.) 462.] Bull. Zool. Nomen. 9:153-154, 30 Dec.
1952. Generic names in question except Phalaena (Bombyx, Noctua, Geometra,
Pyralis, Tortrix, Tinea, Alucita) in constant use for very long time; to upset their
usage would cause greater confusion than uniformity; unfortunate to suppress Pha-
laena but not to do so will result in suppressing almost equally well-known Noctua;
advisable to settle Bombyx and Pyralis as generic names as of 1758.

1953

[Book review.] Die Schmetterlinge Mitteleuropas. By Walter Forster & Theodor
A. Wohlfahrt. Lepid. News 7:26, 1953 (20 Apr. 1953). Second installments of vols.
1 and 2 reviewed.

Shall the “Régles” be amended so as to regulate the fixation of type localities and
if so upon what terms and conditions? [Document 1/58.] Bull. Zool. Nomen. 8:102-
108, 25 Jun. 1953. F. Hemming suggested that to reduce instability provisions should
be added concerning fixation of type localities; fixation of localities a well-established,
desirable practice; rules proposed; some Hemming ideas questioned as basis for
discussion of article to amend Régles.

On the question whether and subject to what conditions the concept of a “neotype”’
should be officially recognized by an appropriate amendment to the “Régles.”’
[Document 2/13.] Bull. Zool. Nomen. 8:121-127, 30 Jun. 1953. In recent years it
has been practice among some zoologists to designate neotypes when types lost or
destroyed; arguments presented for recognition of neotypes; rules proposed, com-
ments made on F. Hemming’s suggestions regarding neotypes as basis for discussion
of article to amend Régles.

1954

With F. Hemming, s.a. Proposed limitation to the purposes of the law of priority
of the suppression of the name “Argus” Bohadsch, 1761 (Class Gastropoda) effected
in “Opinion” 185, in order to prevent the confusion which would otherwise arise
in the Class Insecta, Order Lepidoptera. [Reference Z. N. (S.) 714.] Bull. Zool. Nomen.
9:281-283, 22 Oct. 1954. An Opinion of Commission suppressed for all nomencla-
torial purposes generic name Argus Bohadsch 1761, and unless action taken, Argus
Scopoli 1763 becomes available for a genus of Lepidoptera, replacing either Ly-
sandra Hemming 1933 or Polyommatus Latreille 1804; either result would cause
serious confusion; proposals submitted to restrict previous decision to prevent emer-
gence of Argus Scopoli.

With E. L. Bell, s.a. The lectotype of Megathymus aryxna Dyar (Lepidoptera,
Megathymidae). Am. Mus. Novit. No. 1700, 5 pp., 20 Dec. 1954. Opinions differ as
to what constitutes type series of M. aryxna and which specimen is lectotype because
Dyar did not designate holotype in description; history reviewed and it is concluded
lectotype is specimen figured by H. Druce in Lepidoptera-Heterocera section of
Biologia Centrali-Americana, ed. F. D. Godman & O. Salvin; this permits recog-
nition of M. evansi Freeman as valid name.

1955

“1954.” [Book review.] Die Schmetterlinge Mitteleuropas. By Walter Forster &
Theodor A. Wohlfahrt. Lepid. News 8:170-171, 1954 (7 Jan. 1955). Third, fourth,
fifth installments of vols. 1 and 2 reviewed.

With E. L. Bell, j.a. Request for a ruling as to the specimen to be accepted as the
lectotype of “Megathymus aryxna”™ Dyar, 1905 (Class Insecta, Order Lepidoptera).

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[Reference Z. N. (S.) 889.] Bull. Zool. Nomen. 11:289-294, 30 Dec. 1955. Bell and
dos Passos (54 above) identified lectotype as specimen figured by H. Druce; in same
year D. B. Stallings and J. R. Turner identified lectotype as specimen in USNM to
which Dyar attached label stating the name aryxna was restricted to that specimen;
arguments presented against latter conclusion and for former; suggested ruling

- provided to Commission.

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1956

A bibliography of general catalogues and check lists of Nearctic Rhopalocera. Lepid.
News 10:29-34, 1956 (10 Aug. 1956). “Catalogue” and “check list” defined and
discussed in introduction followed by chronologically arranged bibliography of 78
items including “‘not only the strictly Nearctic works but also all general Palaearctic
check lists and catalogues that refer to... circumpolar insects.”

1957

“1955.” With L. P. Grey, j.a. A new name for Argynnis lais Edwards (Lepidoptera,
Rhopalocera). J. New York Entomol. Soc. 63:95-96, 1955 (8 Mar. 1957). A. lais W.
H. Edwards 1883 a primary homonym of A. lais Scudder 1875; insect renamed
Speyeria atlantis helena, n. name (type is lectotype of A. lais W. H. Edwards in
CM); question arises whether new name for a homonym should be proposed in
original genus in which homonym described or in genus to which homonym trans-
ferred, and it would be well to amend Régles to cover the problem.

A newly discovered announcement of the proposed publication of the Sammlung
exotischer Schmetterlinge by Jacob Hubner. J. Soc. Bibliog. Nat. Hist. 3:206, 2 pls.,
Jan. 1957* (date stamps suggest U.S. receipt mid-May 1957). dos Passos’ incomplete
copy of Hiibner’s Ziefer volume of text (1805-[1823]) to the Sammlung europdischer
Schmetterlinge (1796-[1838]) contained 2-page letter press announcement dated 21
Sep. 1806 of proposed publication of the work on exotics; announcement donated
to AMNH,; plates reproduce the pages.

“1956.” Additions and corrections to “A bibliography of general catalogues and
check lists of Nearctic Rnopalocera.”’ Lepid. News 10:213-214, 1956 (21 Jun. 1957).
14 entries added, typos corrected.

“1956.” William Phillips Comstock, 1880-1956. J. New York Entomol. Soc. 64:1-
5, 1 pl. (portrait), 1956 (23 Dec. 1957). Obituary, bibliography.

1958

With A. B. Klots, j.a. Proposal for the amendment of Article 28 of the existing
“Régles” as amended at Copenhagen (1953) so as to give preference to the principle
of page priority in the selection of generic and specific names and for other purposes.
[Reference Z. N. (S.) 1291; Document 15/1.] Bull. Zool. Nomen. 15:285-292, 11
Feb. 1958. Argument in favor of reinstating “page precedence principle’ in place
of “first reviser principle’; page and line priority objective, while first reviser prin-
ciple highly subjective; when 2 or more names proposed at same time in same
publication for same genus or other taxon, first name published should prevail; text
proposed for draft Régles which provides page, line, word precedence.

In W. I. Follett. Views of the committees on nomenclature: (a) of the American
Society of Ichthyologists and Herpetologists; and (b) of the Society of Systematic
Zoology on the relative status of specific names based on modern patronymics having
the terminations “-i” and “-ii” respectively. [Document 32/4.] Bull. Zool. Nomen.
15:677-685, 18 Apr. 1958. Follett publishes statements by 12 taxonomists; dos Passos’
opinion (p. 681) is that original spellings whether ending in -i or -ii should be
retained without emendation; -i ending should be recommended to authors but if
they do not use it their spellings should be valid and not subject to emendation;
second similar name in a genus whether ending in -ii or -i or vice versa should be
considered junior homonym.

With A. B. Klots, j.a. Proposal for the amendment of Article 21 of the “Régles”’ (i.e.
Draft Article 22) so as to make its operation entirely objective in cases where a

VOLUME 42, NUMBER 3 Laz.

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73.

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person other than the nominal author of the book or paper concerned is responsible
for a name and its indication, definition or description. [Reference Z. N. (S.) 1326;
Document 34/1.] Bull. Zool. Nomen. 15:695-702, 25 Apr. 1958. Article should state
that author of scientific name is person who publishes it in connection with indication,
definition, description, unless express statement in same publication that some other
person responsible; if 1 person responsible for name and another for rest, that shall
constitute joint authorship; text proposed.

“1957.” [Book review.] Die Schmetterlinge Mitteleuropas. By Walter Forster &
Theodor A. Wohlfahrt. Lepid. News 11:176, 1957 (13 May 1958). Sixth installment
of vol. 2 reviewed.

Proposals for the amendment and correction of the draft “Régles” concerning the
establishment of neotypes (Article 20). [Document 41/3.] Bull. Zool. Nomen. 15:
816-821, 23 May 1958. Provisions regarding neotypes reviewed, thought too strict,
impracticable; language proposed in Bradley draft should be adopted with exception
of several provisions, cited but not quoted.

Article 22, Section 5(c)(1) and Section 6(b). [Reference Z. N. (S.) 1344; Document
42/1.] Bull. Zool. Nomen. 15:824, 23 May 1958. Regarding draft Régles, citation of
dates when generic combination changed, dos Passos gives examples of what he
considers proper citations; in new combinations author’s name only should appear
in parentheses, not date.

Proposed relaxation of the ban on intemperate language and proposed relaxation
of the ban on names calculated to give personal and other types of offence. [Reference
Z. N. (S.) 1296; Documents 19/3 and 23/3.] Bull. Zool. Nomen. 15:857, 28 May
1958. Elimination of these provisions may lead some to assume falsely that zoologists
have come to feel differently about such matters; matters could be treated as effec-
tively by omitting them from present position in Régles, incorporating them in
Code of Ethics.

Support for the proposal included by Professor Chester Bradley in the suggested
annexe to Subsection (6) of Section 4 of Article 7 of the draft “Régles.” [Reference
Z. N. (S.) 1848; Document 44/1.] Bull. Zool. Nomen. 15:935, 13 Jun. 1958. Regarding
status of names in preprints when paper concerned not published later in regular
manner, dos Passos agrees with Bradley's addition, suggests it be made to apply
after a certain date.

Citation of corrected and emended names. [Z. N. (S.) 1269; Document 9/3.] Bull.
Zool. Nomen. 15:974, 13 Jun. 1958. Regarding draft Régles, when scientific name
misspelled or otherwise written incorrectly, all emendations should be noted as such;
in appropriate cases incorrect spelling should be placed in synonymy followed by
lapsus calami.

Citation of dates in round brackets for bibliographical references. [Reference Z. N.
(S.) 1294; Document 17/4.] Bull. Zool. Nomen. 15:975, 13 Jun. 1958. Regarding
draft Régles, dos Passos objects to proposed deletion of Article 22, Recommendation
10(B) relating to citation of dates; Paris decisions concise, logical, not restrictive,
pedantic; provide instant knowledge where to find a citation.

In R. V. Melville. [Draft “Régles,” Article 28, Section 4(a): The diaeresis symbol,
Reference Z. N. (S.) 1018; Document 72/1.] Bull. Zool. Nomen. 15:1158-1162, 2
Jul. 1958. In draft, diaeresis symbol excluded from category of diacritic marks;
Melville publishes statements by 3 taxonomists; dos Passos’ opinion (p. 1161) is that
diaeresis symbol be retained.

The Satyrid butterflies of northwestern North America (Lepidoptera: Satyridae).
Proc. Tenth Intern. Congr. Entomol., Montreal, August 17-25, 1956 1:673-681, Dec.
1958. Survey of species of Coenonympha, Cercyonis, Oeneis, Erebia of AK, Yukon,
British Columbia, western Alberta; brief history of collecting in area followed by
discussion of each biotic province, catalogue with references.

1959

“1958.” Frank Edward Watson, 1877-1947. J. New York Entomol. Soc. 66:1-6,
Mar.—Jun. 1958 (20 Jan. 1959). Biographical sketch, bibliography.

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“1958.” The dates and authorships of the names proposed in volume 9 of Encyclo-
pédie méthodique by Godart and Latreille, 1819-1824]. Lepid. News 12:119-120,
1958 (26 Jan. 1959). Although title of work dated 1819, only first part published
that year; all names in it should be ascribed to Godart; second part published in
1824; some of its names should be ascribed to Godart, others to Latreille; explanatory

~ tables of generic and specific names provided.

76.

ie

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0:

80.

81.

82.

“1958.” The dates and authorships to be ascribed to the generic and specific names
proposed by Boisduval and LeConte and by LeConte alone in the Lépidoptéres de
l’ Amerique septentrionale, 1829-1833-[1834]. Lepid. News 12:121-122, 1958 (26
Jan. 1959). Discovery of a set of the work in original wrappers and other research
resulted in new evidence; tables provide authorship, dates of names.

“1958.” The authorship of the names proposed in the Natural history of the rarer
lepidopterous insects of Georgia (1797). Lepid. News 12:191-192, 1958 (30 Apr.
1959). Internal evidence reveals relative roles of J. E. Smith and J. Abbot in producing
work; Smith explains he alone responsible for systematic names, definitions, so all
names proposed in book can be attributed to Smith alone.

“1958.” The authorship and dates of publication of the names of some Rhopalocera
proposed in the Reise der oesterreichischen Fregatte Novara um die Erde in den
Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B. von Willerstorf-
Urbair, 1864-1867-[1875]. Lepid. News 12:193-194, 1958 (30 Apr. 1959). Summary
of pertinent bibliographical data followed by tables of generic, specific names, dated
and ascribed jointly to C. Felder and son R. Felder.

“1958.” The dates and authorships of some names proposed by Cramer and Stoll
in De uitlandsche kapellen voorkomende in de drie waereld-deelen Asia, Africa
en America, and by Stoll alone in Aanhangsel van het werk, De uitlandsche kapellen
voorkomende in de drie waereld-deelen Asia, Africa en America, door den heere
Pieter Cramer [1775} 1791. Lepid. News 12:195-198, 1958 (30 Apr. 1959). Names
cannot be dated from text because many not binomials; specific name often appears
alone; names must be dated from indexes (often published later than text) in which
generic names appear in conjunction with specific names and references to text
figures; table provides pertinent data.

1960

“1959.”’ Further notes on the dates of publication of some generic and specific names
proposed by Boisduval and LeConte in the Lépidoptéres de [ Amerique septen-
trionale, 1829-1833-[1834]. J. Lepid. Soc. 13:212, 1959 (1 Aug. 1960). Information
from correspondent about another copy in wrappers (76 above) led to redating a
number of names.

Taxonomic notes on some Nearctic Rhopalocera. 1. Hesperioidea. J. Lepid. Soc. 14:
24-36, 1960 (15 Dec. 1960). Systematic changes incorporated in forthcoming check-
list of Nearctic Rhopalocera explained; contrary to present practice, list will proceed
from lower butterflies to higher; result in Hesperioidea is complete reversal of order
used by W. H. Evans (1951-55); genera in his work used but there will be changes
in systematic arrangement of species; treatment of Papilionoidea will accord more
or less with plan of B. C. S. Warren (1947) from lowest to highest; within genera
listing of species by J. H. McDunnough (1938) followed except when improvement
desirable; most names in recent literature resulting from splitting of genera given
subgeneric standing; effort will be made to comply with Régles but nomenclature
code of N. Banks and A. N. Caudell (1912) preferable to Régles in present state,
will be followed except when modified by Régles; taxonomic notes follow on species
within Hesperioidea, in which no new names proposed but some changed in rank
or relegated to synonymy, many other corrections made.

1961

“1960.” [Book review.] Butterflies of Formosa in colour. By Takashi Shir6zu. J.
Lepid. Soc. 14:243, 1960 (& Sep. 1961).

VOLUME 42, NUMBER 3 179

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1962

The dates of publication of the Histoire générale et iconographie des lépidoptéres
et des chenilles de l Amerique septentrionale, by Boisduval and LeConte 1829-
1833[-1834]. J. Soc. Bibliog. Nat. Hist. 4:48-56, Jan. 1962* (date stamps suggest U.S.
receipt mid-Feb. 1962). Detailed bibliographical summary of work, including in-
formation from 3 copies in original wrappers; review of previous relevant biblio-
graphical contributions followed by tables that apply publication dates of livraisons
to scientific names.

“1961.” Taxonomic notes on some Nearctic Rhopalocera. 2. Papilionoidea. J. Lepid.
Soc. 15:209-225, 1961 (19 Jun. 1962). Continuation of 81 abcve; no new names
proposed but some changed in rank or relegated to synonymy, many other correc-
tions made; brief supplemental note to part 1 (Hesperioidea) on p. 225.

The authorship of three scientific names of Nearctic Rhopalocera variously credited
to Boisduval or Lucas. J. Lepid. Soc. 16:45-46, 1962 (80 Aug. 1962). Authorship of
Papilio eurymedon, P. rutulus, P. zelicaon ascribed to P. H. Lucas, who published
names before J. Boisduval.

1963

The status of infrasubspecific names. [Reference Z. N. (S.) 1569.] Bull. Zool. Nomen.
20:67-70, 18 Mar. 1963. New article should be added to Code to deal with these
names; text proposed, practically same as that in Bradley Draft but not adopted in
1958; if proposal adopted, emendations to Articles 1, 15, 17(9), 45c will be necessary.
A name first published as a synonym is not thereby made available. Article 11(d).
[Reference Z. N. (S.) 1570.] Bull. Zool. Nomen. 20:70, 18 Mar. 1963. Code should
be amended to state that a name first published as synonym not thereby made
available unless prior to 1958 it has been recognized, removed from synonymy, and
used as name of a taxon.

Neotypes—Article 75. [Reference Z. N. (S.) 1571; Document 17/1.] Bull. Zool.
Nomen. 20:71-72, 18 Mar. 1963. Recognition of neotypes by Code was step in right
direction but some provisions respecting their designation so strict and unnecessary
that article will likely be ignored or workers discouraged from designating neotypes;
additions, deletions proposed.

Form of citation—Article 51b(1). Date in a changed combination—Article 22, Rec-
ommendation 22B. [Reference Z. N. (S.) 1576; Document 22/2.] Bull. Zool. Nomen.
20:77-78, 18 Mar. 1963. Article 51b(1) of Code should be amended to state that
name of subsequent user of a scientific name, if cited, to be separated by comma;
Article 22, Recommendation 22B should be repealed because placing date in pa-
rentheses when combination is changed can affect and make improper the date
citation which in that particular case should be outside parentheses.

Calephelis Grote and Robinson, 1869, (Insecta, Lepidoptera): Proposed use of the
plenary powers to designate a type-species in conformity with current usage [Ref-
erence Z. N. (S.) 1563.] Bull. Zool. Nomen. 20:313-320, 12 Jul. 1963. History of uses
of generic names Nymphidia Boisduval & LeConte, Calephelis Grote & Robinson,
Lephelisca Barnes & Lindsey for North American riodinids reviewed; it is proposed
to retain Calephelis with type species Erycina virginiensis Guérin-Méneville and
invalidate others.

Supplemental notes to previous taxonomic notes on some Nearctic Rhopalocera. J.
Lepid. Soc. 17:103-104, 1963 (8 Nov. 1963). Since publication of 2 papers (81 and
84 above) designed to explain systematic changes incorporated in forthcoming check-
list of Nearctic Rhopalocera, communications received from other workers; these,
other supplemental matters discussed.

1964

A synonymic list of the Nearctic Rhopalocera. N. p. [New Haven, CT], 1964. vi,
145 pp. Lep. Soc. Mem. No. 1 (Feb. 1964, in litt.). List “almost a catalogue’; effort
made to give generic synonymies in addition to specific, and to cite type species of
each generic name used; subjective, objective generic synonyms differentiated; taxa

180

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JOURNAL OF THE LEPIDOPTERISTS SOCIETY

provided with authors’ names, publication dates; 687 species numbered; 1002 sub-
species recognized, 1 questioned; 96 species asterisked as “of doubtful North Amer-
ican occurrence’; of these, 89 are species in which nominate subspecies not Nearctic
but Nearctic subspecies exist; infrasubspecific names included.

1965

Peale’s Lepidoptera Americana (1833). J. New York Entomol. Soc. 73:18-26, 5 figs.,
Mar. 1965 (19 Apr. 1965). T. R. Peale’s work, “the first book begun by an American
author on American lepidoptera that was published in this country,’ discontinued
after one number; that part discussed and bibliographically described; 8 copies
located; wrappers, subscribers’ list illustrated.

With H. Ruckes, s.a. In memoriam: Ernest Layton Bell, 1876-1964. J. New York
Entomol. Soc. 73:49-56, 1 fig. (portrait), Jun. 1965 (16 Jun. 1965). Obituary, bib-
liography.

Review of the Nearctic species of Pieris ““napi’’ as classified by androconial scales
and description of a new seasonal form (Lepidoptera: Pieridae). J. New York Ento-
mol. Soc. 73:135-137, Sep. 1965 (17 Sep. 1965). As result of B. C. S. Warren’s papers
on androconial scales and their bearing on speciation in Pieris, classification of P.
bryoniae Ochsenheimer, P. napi (Linnaeus), P. narina Verity, and subspecies re-
viewed; Pieris narina mogollon gen. aest. warreni n. form (t.l. White Mts., AZ, ex
F. R. Sternitzky collection); holotype in AMNH.

With H. Ruckes, s.a. Ernest Layton Bell (1876-1964). J. Lepid. Soc. 19:190-191,
1965 (24 Sep. 1965). Obituary differing from 94 above.

Addenda et corrigenda to the “Synonymic list of Nearctic Rhopalocera.”’ J. Lepid.
Soc. 19:192, 1965 (24 Sep. 1965). Page laid into later copies of Synonymic list sold
by Society.

With L. P. Grey, j.a. Notes on certain lectotypes designated by the authors in their
Systematic catalogue of Speyeria (Lepidoptera: Nymphalidae). Trans. Am. Entomol.
Soc. 91:351-360, Sep. 1965 (30 Sep. 1965). Continuation of catalogue, prompted by
F. M. Brown’s study of argynnid names proposed by W. H. Edwards; lectotypes of
S. coronis coronis (Behr), S. callippe inornata (W. H. Edwards), and S. atlantis
hesperis (W. H. Edwards) redesignated as neotypes; neotype designated for S.
callippe nevadensis (W. H. Edwards); lectotype redesignated for S. mormonia
artonis (W. H. Edwards); neotype redesignated for S. mormonia clio (W. H. Ed-
wards).

1966

The discovery of additional journals of Frank E. Watson. J. New York Entomol.
Soc. 74:188, Dec. 1966 (29 Dec. 1966). Entomological journals for 1896-1905, 1914—
22, 1926-31, 1934-47 located, donated to AMNH, which now has all Watson journals
except those for 1932-33, presumed lost.

Pieris narina oleracera (Harris) in New Jersey (Lepidoptera: Pieridae). J. New York
Entomol. Soc. 74:222-223, Dec. 1966 (29 Dec. 1966). NJ records cited by earlier
workers but discounted by later ones as misdeterminations verified by capture of a
male by M. A. dos Passos near Springdale, Sussex Co., 8 Jul. 1966.

1968

With B. C. S. Warren, j.a. The homonymy of Papilio aglaja Linnaeus 1758 (Insecta,
Lepidoptera, Pieridae and Nymphalidae): Request for validation. Z. N. (S.) 1791.
Bull. Zool. Nomen. 25:68-71, 27 Sep. 1968. Linnaeus named 2 insects P. aglaja in
1758 ed. of Systema naturae, then in 1767 ed. renamed pierid, retained aglaja for
nymphalid; pierid usage has 1758 page priority over nymphalid usage which is
therefore homonym; this long recognized but recent application seeks to resurrect
nymphalid name, recognize 2 uses of aglaja in different families; situation brought
about by adoption of first reviser rule, repeal of priority rule; Linnaeus not reviser
in 1767, primary junior homonym not an available name; consequences of decision
recognizing 1767 ed. as revision would be chaotic; Commission asked to deny

VOLUME 42, NUMBER 3 181

102.

103.

104.

105.

106.

107.

108.

application insofar as it seeks to suspend rule concerning homonyms and permit 2
uses of name, asked to recognize specific name charlotta Haworth for nymphalid,
aglaja Linnaeus for pierid, and to take other appropriate actions.

1969

A revised synonymic list of the Nearctic Melitaeinae with taxonomic notes (Nym-
phalidae). J. Lepid. Soc. 23:115-125, 1969 (29 May 1969). 2 revisions, Ist by H. L.
Higgins, 2nd by D. L. Bauer, published before Synonymic list (92 above) rendered
its arrangement of Melitaeinae genera, species somewhat obsolete, but checklist
already in press; revised synonymic list of subfamily presented in format of 92; 8
fewer species-level taxa, owing primarily to relegation to subspecies; taxonomic
notes follow to explain changes, placement of names.

A name for Polygonia satyrus marsyas auctorum (Lepidoptera: Nymphalidae).
Trans. Am. Entomol. Soc. 95:153-159, 2 figs., Mar. 1969 (6 Jun. 1969). Misled by
false locality labels, W. H. Edwards described European P. c-album as American
species marsyas in 1870; marsyas usually considered U.S. West Coast population of
P. satyrus (W. H. Edwards), so desirable to propose new name for that population,
Polygonia s. neomarsyas, n. ssp. (t.l. Salmon Meadows, Brewster, WA, J. C. Hop-
finger); holotype in dos Passos collection but will be deposited in CM; holotype,
allotype figured.

With B. C. S. Warren, s.a. The homonymy of Papilio aglaja Linnaeus 1758 (Insecta,
Lepidoptera, Pieridae and Nymphalidae): Request for validation Z. N. (S.) 1791.
A further note in opposition to this application. Bull. Zool. Nomen. 26:67-68, 8 Aug.
1969. Further evidence provided to support application has not established that
Linnaeus a first reviser in 1767; not conducive to stability of nomenclature to alter
long-accepted usages; application and another to same end should be denied.
Lethe eurydice (Johansson) and L. fumosus (Leussler), sibling species (Lepidoptera:
Satyridae). J. New York Entomol. Soc. 77:117-122, Jun. 1969 (24 Oct. 1969). L.
eurydice has been considered single species with 4 subspecies; rather, 2 sibling species
involved which occur in different environments, have constant superficial differ-
ences, probably different foodplants; bibliographical synonymies provided for L.
eurydice, L. fumosus n. comb.; species discussed; arrangement of names proposed
in checklist form.

With A. B. Klots, j.a. The systematics of Anthocharis midea Htibner (Lepidoptera:
Pieridae). Entomol. Am. 45:1-34, 11 figs., 1969 (29 Dec. 1969). Species placed in
subgenus A. (Falcapica) Klots; neotypes designated for 3 species-group names avail-
able for species: genutia Fabricius, midea Hubner, lherminieri Godart; systematics,
geographic variation discussed; bibliographical synonymies provided for species,
nominate subspecies; Anthocharis midea annickae, n. ssp. (t.l. West Rock, New
Haven, CT, C. L. Remington); holotype in AMNH,; life history, foodplants, parasites
discussed; relevant types figured including holotype, allotype of annickae.

1970

A revised synonymic catalogue with taxonomic notes on some Nearctic Lycaenidae.
J. Lepid. Soc. 24:26-38, 1970 (26 Mar. 1970). Revision by H. K. Clench appeared
when Synonymic list (92 above) in press, rendered its arrangement of Theclinae
obsolete; revised synonymic list of subfamily presented in format of 92; Harken-
clenus, n. g. proposed; taxonomic notes follow to explain changes, placement of
names.

1972

Designation of a lectotype for Erebia youngi Holland. Entomol. Rec. J. Var. 84:
238-241, 1 pl. incl. 4 figs., Oct. 1972 (15 Oct. 1972*). Since Holland’s description a
very similar Asiatic species, E. dabanensis Erschoff, discovered in AK, E. kozhan-
tshikovi Sheljuzhko may occur there also; necessary to determine genitalically wheth-
er these species confused in Holland’s type series; on dissection of 2 male syntypes
1 found to be dabanensis; 2nd youngi, latter designated lectotype; E. herscheli

182

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Leussler a local race of youngi; E. youngi rileyi dos Passos similarly proven to be
dabanensis so rileyi falls as synonym.

1973

The great advantages of zoological nomenclature as contrasted with the many
disadvantages of popular names!?! News. Lepid. Soc. 15 May 1973*:2-3. In light
vein, suggested that many common names more stable than scientific names.

The correct name for the subspecies of Limenitis weidemeyerii occurring in Arizona
(Nymphalidae). J. Res. Lepid. 12:21-24, Mar. 1973 (18 Dec. 1973, in litt.) Confusion
in literature reviewed; synonymy provided; name angustifascia Barnes & Mc-
Dunnough a jr. synonym of sinefascia Dyar et al., the correct name.

1974

With W. D. Field, s.a., and J. H. Masters. A bibliography of the catalogs, lists,
faunal and other papers on the butterflies of North America north of Mexico
arranged by state and province (Lepidoptera: Rhopalocera). Washington, DC:
Smithsonian Institution Press, 1974. [ii], 104 pp. Smiths. Contrib. Zool. No. 157 (20
Feb. 1974). 2987 selected publications listed in geographical units (Greenland in-
cluded) and in supplemental bibliography of items that cover more than | state or
province.

With J. R. Heitzman, s.a. Lethe portlandia (Fabricius) and L. anthedon (Clark),
sibling species, with descriptions of new subspecies of the former (Lepidoptera:
Satyridae). Trans. Am. Entomol. Soc. 100:52-99, frontis., 20 figs., Mar. 1974 (16
May 1974). L. portlandia has been considered as having 4 subspecies and 1 synonym;
rather, 2 sibling species involved: portlandia, having 3 subspecies (2 named here),
L. anthedon; the 2 species occur in different environments, have different foodplants;
bibliographical synonymies provided for portlandia, its nominate subspecies, for
Lethe portlandia floralae, n. ssp. (t.l. Rock Springs, Orange Co., FL, S. Roman);
holotype in AMNH,; and Lethe portlandia missarkae, n. ssp. (t.l. 5 mi [8 km] S of
Fayetteville, Washington Co., AR, 1300 ft [396 ml], J. R. Heitzman); holotype in
AMNH,; the 2 subspecies discussed; most specimens referred to in literature as
portlandia are anthedon, so bibliography provided; arrangement of names proposed
in checklist form; holotypes, allotypes of new subspecies figured, as are other relevant
types, some genitalia.

1977

“1976.” A note on Oeneis jutta harperi, its author and date of publication (Satyr-
idae). J. Res. Lepid. 15:211-213, Dec. 1976* (date stamps suggest receipt late Apr.
1977). Previous publications of name harperi as subspecies of O. jutta (Hiibner)
invalid according to Code; name validly published here as Oeneis j. harperi, n. ssp.;
t.l. fixed; types mentioned in literature presumably in P. W. Chermock collection.
A taxonomic note on Polygonia faunus arcticus Leussler (Lepidoptera: Nymphal-
idae). Pan-Pac. Entomol. 53:179-180, Jul. 1977 (28 Nov. 1977). Leussler’s arcticus
is subspecies of P. hylas, not P. faunus, as type specimens indicate; name should be
written as Polygonia hylas arcticus Leussler, n. comb.

1978

Correction—Note on Polygonia faunus arcticus. Pan-Pac. Entomol. 54:42, Jan. 1978
(26 Apr. 1978). Phrase concerning type locations added to 114 above.

1981

A little-known, anonymous work on American and European butterflies and moths
(1906), which should be attributed to William Beutenmiiller (Lepidoptera: Nym-
phalidae). J. New York Entomol. Soc. 89:143-145, Jun. 1981 (24 Sep. 1981). Dis-
cussion and description of A manual of American and European butterflies and
moths reproduced in natural colors with their common and scientific names; Mrs.
Beutenmiller probably executed plates.

VOLUME 42, NUMBER 3 183

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118.

119.

120.

1982

“1981.”" With A. B. Klots, s.a. Studies of North American Erora (Scudder) (Lepi-
doptera, Lycaenidae). J. New York Entomol. Soc. 89:295-331, 34 figs., Dec. 1981
(19 Feb. 1982). Genus Erora characterized, discussed, as are E. laeta (W. H. Ed-
wards), E. g. quaderna (Hewitson), E. q. sanfordi dos Passos; early stages, ethology,
ecology, geographic distribution of laeta, quaderna sanfordi discussed; bibliograph-
ical synonymies, lists of distributional records included; early stages, types figured.
“1981.” Some little-known U.S. publications on Lepidoptera I. [Edited and with
abstract and foreword by L. P. Grey.] J. Res. Lepid. 20:111-115, Summer 1981 (20
Sep. 1982). Periodicals The Lepidopterist (1916-17), Lepidoptera (1918-21), The
Lepidopterist (1918-31) discussed, known numbers listed with dates of publication,
pagination, inclusions.

“1981.” Some little-known U.S. publications on Lepidoptera II. [Edited and with
abstract by L. P. Grey.] J. Res. Lepid. 20:115-122, Summer 1981 (20 Sep. 1982).
Information similar to 117 above provided for The Butterfly Farmer (1913-14),
Lorquinia (1916-19), Southwest Science Bulletin (1920), Butterfly Park Nature
Club News (1929-31), The Lepidopterists' News (1933), The Entomologists’ Ex-
change Association (1936), The Entomologists Exchange News (1937-42), The
Butterfly Club (1946-47), Club Notes, Moth and Butterfly Club (?1947-53), Notes
on Moths and Butterflies (1958-55), for relevant material in Sierra Club Bulletin
(1913), Hobbies (1936).

1983
With C. D. Ferris, s.a., J. A. Ebner, and J. D. Lafontaine. An annotated list of the
butterflies (Lepidoptera) of the Yukon Territory, Canada. Can. Entomol. 115:823-
840, 6 figs., Jul. 1983 (22 Jun. 1983, in litt.) 95 taxa reported including subspecies;
some listed as questionable; various species figured.

Journal of the Lepidopterists’ Society
42(3), 1988, 184-195

BIOLOGY OF SPEYERIA ZERENE HIPPOLYTA
(NYMPHALIDAE) IN A MARINE-MODIFIED
ENVIRONMENT

DaAvID V. MCCORKLE

Biology Department, Western Oregon State College,
Monmouth, Oregon 97361

AND

PAUL C. HAMMOND
2435 E. Applegate, Philomath, Oregon 97370

ABSTRACT. This paper examines life history and adaptations of Speyeria zerene
hippolyta (Edwards) along the Oregon and Washington coasts where cold wind, rain,
and fog persist during much of the year. The butterfly uses an open grassland habitat on
salt-spray meadows and higher headlands adjacent to the ocean, where the larvae feed
on the common Viola adunca J. E. Smith. Four unusual adaptations to this environment
are seen in S. zerene hippolyta that are absent from the closely related S. z. bremnerii
(Edwards) of the inland Willamette Valley: small body size and extensive dark basal
suffusion which enhance body heating from solar radiation; normal flight activity under
cool, cloudy or foggy conditions; prolonged larval development which coordinates adult
emergence with the most favorable weather conditions in late summer and fall; and much
individual variation in larval development rate and adult emergence which compensates
for variable and unpredictable weather from year to year.

Additional key words: Speyeria zerene bremnerii, Viola adunca, adaptation, grass-
land, coastal habitat.

Speyeria zerene (Boisduval) is a complex polytypic species with 14
recognized subspecies (Grey & Moeck 1962). The subspecies occupy a
wide diversity of habitats ranging from coastal rainforests in the Pacific
Northwest to arid sagebrush plains in the Great Basin. Speyeria z.
hippolyta (Edwards), informally known as the ““Hippolyta Silverspot’’
or “Oregon Silverspot’’, is restricted to a cool, wet, marine-modified
environment adjacent to the Pacific Ocean in western Washington and
Oregon. This subspecies is of special concern because of its decline
toward extinction and its official classification as a threatened species
(Hammond & McCorkle 1983).

The closely related S. z. bremnerii (Edwards) occupies inland areas
of the Pacific Northwest from Vancouver Island S through the Puget
Sound trough and Willamette Valley of western Oregon. The primary
difference in adult phenotype between these subspecies is the small
wing of S. z. hippolyta although extinct Oregon populations of S. z.
bremnerii also differed in having reduced basal suffusion on the dorsal
wing surfaces (Fig. 3). In addition, S. z. hippolyta differs in several
aspects of life history and developmental physiology which appear to
be specific adaptations to the coastal environment. An investigation of

VOLUME 42, NUMBER 3 185

these characteristics is the subject of the present paper. It should be
noted that both Moeck (1957) and Howe (1975) confused this coastal
subspecies with a population of dwarfed S. z. conchyliatus (Comstock)
endemic to the volcanic ash and pumice fields along the E slope of the
Oregon Cascade Range.

MATERIALS AND METHODS

Field and museum studies were conducted from 1960 to 1986 to-
gether with laboratory rearing of larvae. Most public and many private
collections in Washington and Oregon were examined. In 1963 and
1964, one of us (McCorkle) developed a technique for rearing Speyeria
larvae using a modification of a procedure (Magnus 1958) for the Eu-
ropean fritillary Argynnis paphia L. A variant of the former technique
was described by Mattoon et al. (1971), and was used in the present
study, except that larvae were kept over winter in hollow wooden blocks
and reared in small jars instead of nylon sleeves.

Capture-recapture studies were done at the Rock Creek study site
in Lane Co., Oregon, during 1980 using the 1-2-4-7 marking system
described by Ehrlich and Davidson (1961). Sex, wing length, general
condition, time, place, and type of activity at time of capture and
recapture were recorded.

Voucher specimens are deposited in the Systematic Entomology Lab-
oratory at Oregon State University, Corvallis.

BIOLOGY OF STAGES
Oviposition

Speyeria zerene hippolyta is usually a grassland butterfly that lives
on open salt-spray meadows and grassy headlands adjacent to the Pacific
Ocean, where the larvae feed on the common blue violet, Viola adunca
J. E. Smith. Based on more than 100 observations, females oviposit
singly among vegetation near host plants. Females are apparently stim-
ulated to oviposit by some volatile compound emanating from violets.
We found that females oviposit only in the presence of violets, but that
direct physical contact with the host is not necessary. We observed
oviposition up to 20 cm downwind of even dried violet leaves.

During oviposition behavior, the butterflies flew near the ground,
working their way upwind. When violets were near, they paused to
climb in meadow vegetation, probing with curved abdomen until a
suitable site was contacted, and an egg deposited. We even observed
females crawling into knee-deep layers of grass that overgrew violets
by late summer. Oviposition observations and location of larvae indicate
that females favor sunny sites, and usually avoid N slopes of steep
meadow rises.

186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Eggs are cream colored when first laid, but if fertile, darken to pinkish
tan by the second day. Eggs began to hatch 16 days after oviposition
with ambient room temperature varying from 21 to 24°C (N = ca.
1000). A large quantity of lipid is stored in Speyeria eggs which appears
to serve as a food reserve during larval diapause. This lipid, in the form
of a light oil, is readily observed in yolk by dissecting eggs.

Larva

Newly hatched larvae (N = ca. 1000) usually wander short distances.
As in other Speyeria, first instar S. z. hippolyta soon enter a diapause
that lasts over winter. After selecting a suitable site, they spin a thin
silk mat on which they rest until spring. Larvae exhibit considerable
resistance to desiccation during diapause. In the laboratory, they sur-
vived a month or more without moisture, but this stress may diminish
survival by spring. Thus, in years with delayed fall rains, the earliest
hatched larvae may be at a disadvantage. When moisture is available,
laboratory larvae touched their mouths to a wet surface, and some
nearly doubled in size within a short time.

The following description of the last (6th) instar is based on larvae
from about 10 family lines. The spiny larva (Fig. 1) is dark brown with
a pair of pale lines running down the back, each of which has a row
of black patches running parallel to it on the outside. These black patches
are located on both sides of each body segment adjacent to the pale
dorsal lines. Lateral parts of the body are finely and irregularly mottled
with pale yellow. The head is mostly black, but the occipital area is
yellow. Spine bases, especially those of the lateral rows, are straw colored
similarly to the lines on the back. This color pattern resembles that of
the inland S. zerene bremnerii (Hardy 1958), and blends with dried
leaves where larvae take refuge when not feeding.

The first instar possesses unbranched setae or hairs. Later instars bear
branched spines in the pattern illustrated by Scott and Mattoon (1982)
for S. nokomis (Edwards). We reared all 18 species of Speyeria, and
this setal pattern is consistent throughout the genus. However, it should
be noted that a lateral spine is present on the 10th abdominal segment
that was omitted from the Scott-Mattoon setal map. Superficially, this
spine appears to be on the 9th segment. Spine branches or spinules are
attached to the primary shaft in such a way that they can swing in
toward the shaft apex when the larva withdraws from an entanglement.
When pushed outward, the spinules lock into the outstretched position.
In larger larvae, these sharp spines may provide protection from pred-
ators such as mice and, perhaps, shrews (unpubl. obs.).

In addition to spines, Speyeria larvae possess what may also serve as
a defense against predators in the form of a fleshy, eversible osmmeterium.

VOLUME 42, NUMBER 8 187

Fics. 1-3. S. zerene. 1, Mature 6th instar of S. z. hippolyta; 2, Pupa of S. z. hippolyta;
3, Reared males of S. z. hippolyta (left) and Willamette Valley S. z. bremnerii (right).

Whenever this structure is extruded, a disagreeable musky smell be-
comes apparent. This odor is faint in the small S. zerene hippolyta, but
is much stronger in larger-bodied species such as S. coronis (Behr) and
S. edwardsii (Reakirt). Unlike the long dorsal osmeteria of papilionid
larvae, Speyeria osmeteria are short, wedge-shaped organs located ven-
trally just behind the head and before the first pair of thoracic legs. In
addition, the musky smell of Speyeria osmeteria is distinctly different
from the more pungent, aromatic smell of papilionid osmeteria.

As with most Speyeria, older larvae of S. z. hippolyta retreat to shelter
sites sometimes several centimeters from host violets. These sites may

188 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

hippolyta

10 11 12 13 14 15

ph
ro)
o
~
fo)
o

Development Time (Weeks)

Fic. 4. Laboratory development time from first instar to adult in S. zerene hippolyta
and S. z. bremnerii reared at 22° + 1°C.

provide thermal advantages as well as cover from predators (McCorkle
1980). Some species may be mostly nocturnal (Dornfeld 1980:75). How-
ever, one field larva of S. z. hippolyta found during daytime in the
present study was feeding, and laboratory larvae of many Speyeria feed
both night and day. Their feeding is typically rapid so that they are
exposed from cover only briefly. In the constantly heated laboratory,
larvae probably grow faster than in nature. Their growth rate in the
field may be delayed especially by cooler spring weather. Nevertheless,
in the laboratory, they still required from one to three months to grow
through six instars (Fig. 4, Table 1), and they spent about two weeks
in the pupal stage.

Larval feeding signs were evident on violets in the Rock Creek mead-
ow on 15 April 1980, indicating that at least some larvae were well into
feeding activity. On 8 May 1986, one early 4th instar was observed.
On 5 July, we found one nearly mature 6th instar (not parasitized) and
several violets with fresh feeding signs, indicating that at least some
larvae were still active. On the same date, however, the first adult male
was taken, although normally the males do not begin to eclose until
after 10 July with peak emergence in early August. Thus, the minimum
natural larval feeding span is from mid-April to mid-June, some two
months. It may be that a few larvae continue to feed well into August,
producing the fresh adults present in early September.

Pupa and Adult

The pupa (Fig. 2) is smooth, rounded, and mostly dark brown with
variable paler areas on abdomen and wing covers, and a dark, transverse
band on the anterior edge of each abdominal segment. As in most
Speyeria, the pupal chamber consists of several leaves drawn together
with silk, and the pupa is usually attached in a hanging position to the
top of the chamber.

VOLUME 42, NUMBER 3 189

nt50 VU 0

10 20 10 20 10 20 10 20
June July Aug Sept

Fic. 5. Time range in field presence of fresh adults of S. zerene hippolyta (central
Oregon coast) and S. z. bremnerii (Willamette Valley).

As noted above, the first adult male was taken on 5 July, and a nearly
mature larva was also found on that date. By early August, males were
common (Fig. 5) (22 marked 1-8 August), and females had begun to
appear (2 marked). Between 20 and 22 August, 22 new males and 7
new females were marked, including several fresh females and a few
fresh males. On 4 September, 15 new males and 16 new females were
marked. Of these, five females were fresh, three males were fairly fresh,
and one very fresh. It is thus apparent that in this colony, males eclosed
from mid-July until at least the end of August. The first females eclosed
by the end of July, and eclosion continued through August to mid-
September, with aging specimens surviving into October. Recapture
results (McCorkle 1980) indicate that some adults live for at least three
weeks, and disperse widely up to 2-3 km (1-2 miles). Butterflies fly
inland and seek shelter along forest margins when strong winds are
blowing off the ocean (McCorkle 1980).

ADAPTATIONS TO COASTAL ENVIRONMENT

Both coloration and wing size appear to have a strong genetic de-
termination, since S. z. hippolyta differs consistently from the larger
and paler Willamette Valley S. z. bremnerii both in the field and in
laboratory rearings (Figs. 3, 6). The difference between forewing lengths
(Fig. 6) is highly significant (P, < 0.0001). Data from Willamette Valley
was obtained before the apparent extinction of these populations around
1977 (Hammond & McCorkle 1988).

There is evidence that small size and extensive dark basal suffusion
as shown in S. z. hippolyta are adaptations to enhance solar heating,
as would be needed in a marine-modified environment with persistent
cold wind and frequent fog (McCorkle 1980). Butterflies, being het-
erothermic, usually depend on solar radiation to elevate body temper-
ature sufficiently to allow flight necessary for foraging, mate seeking,
escape from predators, and oviposition (Watt 1968, Douglas 1978,

190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Time required for larval and pupal development in Speyeria zerene pop-
ulations reared at 21-23°C. Each entry represents progeny of one female, with number
of individuals in parentheses.

No. weeks
' Subspecies Locality Males Females
bremnerii Benton Co., Oregon 6 (2) 7 (3)
5-6 (5) 6 (5)
Thurston Co., Washington 6-7 (3) 8-9 (3)
6 32) 7-8 (3)
hippolyta Pacific Co., Washington 8-9 (3) 9-10 (5)
8-9 (7) ee)
Clatsop Co., Oregon 7-9 (11) 8-10 (12)
Lincoln Co., Oregon 8-9 (9) 9-10 (10)
Lane Co., Oregon 7-9 (7) 8-10 (9)
hippolyta-like Del Norte Co., California 9-13 (8) 12-14 (7)
8-9 (5) 9-10 (7)
10-13 (6) 11-18 (8)
behrensii Curry Co., Oregon 7-8 (5) 8-9 (9)
7-8 (2) 9 (5)
gloriosa Josephine Co., Oregon 7-8 (11) 8-10 (12)
7-8 (7) 9-10 (6)
myrtleae Marin Co., California 7-8 (11) 8-10 (14)
7-9 (11) 8-10 (10)
8-9 (12) 8-10 (12)

McCorkle 1980). Rapid ovarian development is also thermodependent
(Watt 1968).

Speyeria use a dorsal basking position in which the wings are extended
in a horizontal plane from the body, thus exposing the dark basal
suffusion to solar radiation. In dorsal basking butterflies, heat first ab-
sorbed by the basal part of the wings is then absorbed by the thorax,
and usually a thick coat of long hairs serves as insulation for retention
of thoracic heat (Douglas 1978, McCorkle 1980). Douglas (1978:43)
suggests that large butterflies are at a disadvantage under cool conditions
because they heat up more slowly, while smaller butterflies warm quick-
ly to an adequate thoracic temperature. In a cold, windy environment,
small butterflies also have a second advantage because the smaller
wingspan allows flight closer to the ground where wind velocity is lower.
Wind is a problem in maintaining body temperatures because heat is
lost from the body surface by forced convection (Douglas 1978:69).

Since most Speyeria require high body temperatures for normal
activity, they usually fly only in full sunshine, or under cloudy conditions
when the air temperature is higher than 21°C (70°F). However, field
observations of S. z. hippolyta revealed that it engages in normal activity
under cloudy or foggy conditions with air temperatures as low as 16°C

VOLUME 42, NUMBER 3 191

No. individuals

24

S. zerene’ bremnerii

Ss wild

20

16 n=48
X=31
12 oo:
LZ reared
n=7

M32

24

S. zerene hippolyta

20
LSS wild
16 MGS
X=27

a 2 =
AZ reared

n=36

Se

SSeletselets

Forewing length (mm)

Fic. 6. Frequency distribution of male forewing length in wild and reared S. zerene
hippolyta (central Oregon coast) and S. z. bremnerii (Willamette Valley).

(60°F), although the butterflies retreat to shelter under windy conditions.
For example, during early September 1982, we observed approximately
a dozen males in mate-searching flight, six males and females nectaring
on flowers, and three females engaged in oviposition under windless,
cloudy-foggy conditions with air temperature only 16°C (60°F). Similar
observations have been made consistently during subsequent years. In
sharp contrast, observations of the similarly colored S. a. atlantis (Ed-
wards) in the Appalachians of West Virginia during 1977 revealed

192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

virtually no evidence of similar activity under cool, cloudy conditions.
These adults (N = 100+) engaged in nectaring and flight only in full
sunshine, and retreated to shelter in trees when clouds covered the sun
at a temperature of approximately 21°C (70°F). In more dramatic con-
trast, the large S. idalia (Drury) favors minimum air temperatures of
24-27°C (75-80°F) for normal flight activities based on observations
(N = 100+) in Nebraska during 1983, and continues to fly at temper-
atures of 41°C (105°F) or more. This species exhibits little basking
behavior in the field, and the basal wing areas completely lack dark
suffusion. Even the black hindwings of S. idalia are mostly pale orange
at the wing base.

Because S. a. atlantis and S. zerene hippolyta are nearly identical
in their dark basal suffusion, and are of similar small size, the expected
thermal regulation and behavioral activity of these species should be
similar. Therefore, the above observations suggest that S. z. hippolyta
employs an additional mechanism that allows activity under low tem-
peratures and cloudy conditions. This mechanism might include changes
in enzyme systems that would allow normal physiological function at
low body temperatures, but this possibility has not yet been studied.

Finally, S. z. hippolyta along the central Oregon coast is also highly
unusual in that adult emergence in the field extends over a long period
of some 8 weeks from early July to early September, and it requires
7-14 weeks to complete larval and pupal development in the laboratory
(Figs. 4, 5). These developmental patterns appear to be an additional
adaptation to the coastal environment. By contrast, most western Spey-
eria emerge over a relatively short period of about three to four weeks
in June and July. For example, the Willamette Valley form of S. z.
bremnerii usually emerged during late June and early July in the field,
and required only 5-9 weeks to complete larval and pupal development
in the laboratory. As shown in Figs. 4 and 5, the sharp differences in
the developmental patterns between S. z. hippolyta and S. z. bremnerii
are maintained consistently both in field and laboratory, suggesting that
a strong genetic component is involved.

There are two apparent reasons for these developmental patterns in
S. z. hippolyta. In most years, adverse conditions of cold wind, fog,
and rain persist along the ocean through May, June, and July, and
weather more favorable for adult flight usually does not develop until
August and September. The retarded larval development of S. z. hip-
polyta coordinates adult emergence with the best weather conditions
in the coastal environment. In addition, adverse stormy weather may
develop in some years along the coast during either August or Septem-
ber. The great range of individual variation in developmental rate and
adult emergence suggests an additional adaptation to variable and un-

VOLUME 42, NUMBER 3 193

predictable weather conditions. Thus, early emerging butterflies may
be more successful in reproduction one year, and late butterflies the
next year, depending on each year’s storm patterns. Since coastal weath-
er is so variable from year to year, genes for both early and late emer-
gence (fast and slow development) would tend to be maintained.

During the past 20 years, we have reared most of the geographic
subspecies recognized in Speyeria under similar laboratory conditions
at temperatures of 21-23°C (70-72°F). The fastest rate of larval and
pupal development was observed in certain forms of S. atlantis, S.
egleis (Behr), S. callippe (Boisduval), and Oregon S. zerene bremnerii,
all of which required only five to six weeks for males and six to seven
weeks for females. Most subspecies of S. zerene and S. coronis required
six to seven weeks for males and seven to eight weeks for females. These
include Sierran-type S. z. zerene, Rocky Mountain S. z. sinope dos
Passos & Grey, and Great Basin S. z. gunderi (Comstock). Even very
large-bodied species such as S. c. cybele (Fabricius), S. idalia, and S.
nokomis caerulescens (Holland) required only a similar amount of time,
while S. nokomis apacheana (Skinner), S. diana (Cramer), and S. ed-
wardsii required seven to eight weeks for males and eight weeks for
females.

However, except for typical S. z. bremnerii itself, all populations of
S. zerene within the bremnerii subspecies group as defined by Grey
and Moeck (1962) exhibit a relatively long developmental time of 7-9
weeks for males and 8-10 weeks for females (Table 1). Our field emer-
gence data (Fig. 5), indicate especially long and variable development
times for S. z. hippolyta populations along the central Oregon coast
from Lane Co. N to Tillamook Co.

Hammond & McCorkle (1983) noted hippolyta-like populations of
S. zerene along the coast of Del Norte Co., California, N of Crescent
City. These are disjunct from Oregon hippolyta, and are separated by
intervening populations of an S. z. behrensii-gloriosa intergrade in
Curry Co., Oregon. Two of three family lines reared from the Del
Norte populations exhibited an extended development time of 10-13
weeks for males and 11-14 weeks for females (Table 1). Thus, the
extremely variable developmental rates observed in field emergence of
S.z. hippolyta are also seen in some family lines reared in the laboratory.
In sharp contrast, two family lines of S. z. behrensii from Curry Co. to
the north, and three family lines of S. z. myrtleae from Marin Co.,
California to the south did not exhibit this extended development.
Moreover, wild populations of S. z. myrtleae usually emerge in the
field during early to mid-July, a full month earlier than the hippolyta-
like populations. The manner in which these emergence patterns relate
to respective local weather is undocumented as yet.

194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

There are two close parallels to S$. zerene hippolyta within the Ar-
gynninae that also exhibit adaptations to cold, wet marine-modified
grasslands. One is S. mormonia bischoffi (Edwards) in coastal S Alaska
from the Kodiak region N to Anchorage. The second is Fabriciana
(Mesoacidalia) aglaja scotica Watkins in the Hebrides and Orkney
Islands N of Scotland. Both species display extensive dark suffusion over
much of the dorsal wing surfaces. Ford (1945) indicated that F. a.
scotica often exhibits reduced wing size on small, wind-swept islands.

In the San Francisco Bay area of California, both Speyeria c. callippe
and S. coronis coronis also have heavy melanic basal suffusion in a cool,
foggy coastal environment (Hovanitz 1941), although the California
habitat is less severe compared to the northern coastal grasslands oc-
cupied by the previous three species.

ACKNOWLEDGMENTS

Part of this research was funded by the U.S. Forest Service. We particularly thank M.
G. Parsons and M. D. Clady of this agency for assistance and encouragement.

LITERATURE CITED

DORNFELD, E. J. 1980. The butterflies of Oregon. Timber Press, Forest Grove, Oregon.
276 pp.

Douc as, M. M. 1978. The behavior and biophysical strategies of thermoregulation in
temperate butterflies. Ph.D. Dissertation, University of Kansas, Lawrence. 231 pp.
Diss. Abs. Int. 39 (7) 31832B-3133B. Order No. 78-24790.

EHRLICH, P. R. & S. E. Davipson. 1961. Techniques for capture-recapture studies of
Lepidoptera populations. J. Lepid. Soc. 14:227-229.

ForD, E. B. 1945. Butterflies. Collins, London. 368 pp.

Grey, L. P. & A. H. Mogeck. 1962. Notes on overlapping subspecies. I. An example in
Speyeria zerene. J. Lepid. Soc. 16:81-97.

HAMMOND, P. C. & D. V. MCCoRKLE. 1983. The decline and extinction of Speyeria
populations resulting from human environmental disturbances (Nymphalidae: Ar-
gynninae). J. Res. Lepid. 22:217-224.

Harpy, G. A. 1958. Notes on the life histories of three species of Lepidoptera from
southern Vancouver Island, British Columbia. Proc. Entomol. Soc. British Columbia
50:27-28.

HovANITz, W. 1941. Parallel ecogenotypical color variation in butterflies. Ecology 22:
259-284.

Howe, W.H. 1975. The butterflies of North America. Doubleday & Co., Garden City,
New York. 633 pp.

MAGNus, D. B. E. 1958. Experimental analysis of some “overoptimal”’ sign-stimuli in
the mating behaviour of the fritillary butterfly Argynnis paphia L. Proc. 10th Intern.
Congr. Entomol. 2:405-418.

MATTOON, S. O., R. D. Davis & O. D. SPENCER. 1971. Rearing techniques for species
of Speyeria. J. Lepid. Soc. 25:247-256.

McCorkLe, D. V. 1980. Ecological investigation report: Oregon silverspot butterfly
(Speyeria zerene hippolyta). USDA Forest Service, Siuslaw National Forest, Corvallis,
Oregon. 117 pp.

MoEcK, A. H. 1957. Geographic variability in Speyeria. Milwaukee Entomol. Soc. Spec.
Pap. 48 pp.

VOLUME 42, NUMBER 3 195

ScoTT, J. A. & S.O. MATTOON. 1982. Early stages of Speyeria nokomis (Nymphalidae).
J. Res. Lepid. 20:12-15.

Watt, W. B. 1968. Adaptive significance of pigment polymorphisms in Colias butter-

flies. I. Variation of melanin pigment in relation to thermoregulation. Evolution 22:
437-458.

Received for publication 16 September 1986; accepted 7 April 1988.

Journal of the Lepidopterists’ Society
42(3), 1988, 196-203

MARKING LEPIDOPTERA AND THEIR OFFSPRING:
TRACE ELEMENT LABELLING OF COLIAS EURYTHEME
(PIERIDAE) WITH RUBIDIUM

JANE LESLIE HAYES AND CATHRYN L. CLAUSSEN

Southern Field Crop Insect Management Laboratory,
Agricultural Research Service, USDA,
Stoneville, Mississippi 38776

ABSTRACT. Trace elements can be used to label lepidopteran eggs via treated adults,
but sources and importance of sample variability are relatively unexplored. Female Colias
eurytheme (Boisduval) reared on rubidium (Rb)-treated foodplants and their eggs were
analyzed for Rb by atomic absorption spectrophotometry. Parent (3 untreated and 4
treated females) and mean egg (25/female) element content were significantly correlated.
Compared with untreated adults and eggs, treated samples were reliably marked, although
significant egg-to-egg variability in Rb concentration was found within and between sib-
groups. Analysis-day differences were not significant. Adult sizes and element exposures
may have contributed to between-group variance. Maternal and genetic influences were
potential sources of within sib-group variance.

Additional key words: alfalfa butterfly, internal marker, adults, eggs, atomic absorp-
tion spectrophotometry.

Many methods for marking and monitoring lepidopteran populations
have been proposed, tested, and employed with various degrees of
success (Southwood 1978). An internal marking method that shows great
potential for use in dispersal studies of phytophagous insects is that of
trace element labelling (Berry et al. 1972). These labels can be used in
concentrations low enough to permanently mark but not adversely
affect the insect or its host. The mark can be acquired without handling
the insect because it is obtained as the animal feeds on treated plant
tissue (Stimmann et al. 1973) or nectaries (Culin & Alverson 1986).
Insects in 7 orders, including 10 species of Lepidoptera in 5 families,
have been successfully labelled (Hayes & Hopper 1987:table 1). Dis-
persal tests employing trace elements as adult markers have been con-
ducted with three lepidopteran species: Heliothis zea (Boddie) and
Spodoptera frugiperda (J. E. Smith) (both Noctuidae) (Graham et al.
1978); and Pectinophora gossypiella (Saunders) (Gelechiidae) (Van
Steenwyk et al. 1978).

One recently revealed advantage of trace element use over external
markers of Lepidoptera is that the mark is passed along to reproductive
products, including eggs (Legg & Chiang 1984, Hayes & Hopper 1987)
and spermatophores (Graham & Wolfenbarger 1977). Detection of a
parental mark in the egg or spermatophore provides, much like a genetic
marker, a potential means of assessing gene flow in the field. In mea-
suring dispersal, recovery of eggs may prove superior to recovery of
adults because the marked adult transmits multiple signals through the

VOLUME 42, NUMBER 3 197

distribution of marked eggs (Jones et al. 1980). Also, the possibility of
using elemental marking of reproductive products in behavioral, phys-
iological, or developmental studies is suggested, but has not been much
explored (Engebretson & Mason 1981).

It remains to be seen whether recovery of marked eggs will prove
feasible in large-scale field studies. However, the method has been used
successfully to monitor small-scale egg dispersal by Trichoplusia ni
(Htibner) (Geometridae) to adjacent crops (G. Ballmer pers. comm.).
Among the many questions that need to be addressed is that of vari-
ability among eggs. Individual variability among adults can arise from
exposure differences and differences in size or weight, and can occur
through time. It has yet to be determined whether variability is passed
along to offspring and whether there are intrinsic differences among
eggs from the same female.

To investigate these areas and develop methods for more efficient
and possibly expanded utilization of trace-element labelling, we ana-
lyzed marked adult Colias eurytheme (Boisduval) (Pieridae) and their
eggs by atomic absorption spectrophotometry (AAS) for trace-element
content. In addition to the parent-offspring relation, we examined egg-
to-egg variability within sib-groups and among offspring of different
parents. Differences between preparation dates were also considered
possible sources of variability. Adult samples were prepared from bodies
and head capsules and compared with egg samples to examine cost-
effective adult sample preparation procedures.

MATERIALS AND METHODS

Insects. Colias eurytheme, the alfalfa butterfly, was used because of
its tractability in the laboratory and greenhouse (Taylor et al. 1981),
and because it is considered a model system for other Lepidoptera
(Watt et al. 1974). Like Heliothis spp. and a number of other econom-
ically important Lepidoptera, Colias spp. are highly mobile, po-
lyphagous, and distribute their eggs individually over a potentially large
area (Tabashnik 1980). Experimental insects were obtained from a
colony originating from eggs collected in November 1986 on commer-
cial vetch, Vicia villosa Roth (Leguminosae), on the grounds of the
Jamie Whitten Delta States Research Center, Stoneville, Mississippi.

Treatments. Adults were reared in the greenhouse from eggs or
neonate larvae on vetch plants treated with rubidium (in chloride form).
Host plants were grown from seed in vermiculite, and treated weekly
with 1 g RbCIl/1 water (1000 ppm), initially by foliar application, then
by watering the potting soil after insects were placed on the plants.
Freshly treated host plants were provided as needed until pupation.
Control insects were reared in a similar manner on untreated plants.

198 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Treated and untreated pupae were placed in separate 2-] cardboard
cartons with organdy top cloths and moistened paper towel liners.
Eclosing adults were marked on the left hindwing with a felt tip pen
(/ = treated; //= untreated), and placed in a mating cage. The cage,
a 0.6 X 0.6 X 0.38-m wood frame covered with transparent plastic, was
provisioned daily with honey-water (1:3) soaked cotton balls, and held
at 12°C without light. To stimulate mating, temperatures were elevated
to 30°C and light was provided by two banks of fluorescent lights for
2-4 h/day. The cage was checked at 15-30-min intervals for the oc-
currence of mating. Pairs were removed and held at room temperature
(ca. 25°C) until spermatophore transfer was completed. The males were
uniquely marked and returned to the cage. For oviposition, females
were placed individually on host plants covered by plastic bags and
maintained at room temperature (ca. 25°C) and LD 12:12. Ovipositing
females were fed daily with honey-water and transferred to fresh host
plants as needed.

Twenty-five eggs from each treated female (N = 4) and each un-
treated female (N = 3) were collected separately by sib-group, and
frozen. Eggs were obtained from day 1-2 of oviposition. When a female
died, wings were removed and the body frozen.

Sample digestion. Individual eggs were digested, following the meth-
od of Hayes and Hopper (1987), by placing an egg directly into the
sample cuvette along with 0.025 ml ultrapure nitric acid (HNO, ), heat-
ing by microwave for 8-12 min at a low setting, and then diluting with
0.5 ml deionized distilled water (DDI).

The head and body of each female were placed in separate 7-ml
scintillation vials with 0.2 ml and 0.4 ml ultrapure nitric acid, respec-
tively. Digestion was allowed to occur at room temperature for 24 h,
then samples were microwaved for 8-12 min. Digested material was
then diluted with DDI, 4.0 and 5.0 ml, respectively.

AAS analysis. Samples were analyzed using a Perkin-Elmer 3030
with an HGA 400 graphite furnace and AS-40 autosampler. An elec-
trodeless discharge lamp for Rb was used. Wavelength was set at 780
nm. Char and atomization temperatures were 800° and 1900°C (Slavin
1984). Elements were atomized off the wall of pyrolitically-coated
graphite tubes.

Data. Initially, 10 eggs from each female (treated and untreated)
were individually digested and analyzed by AAS for presence or absence
of a detectable Rb signal (=day 1). To increase the sample size and
examine between-analysis-day variability, an additional 15 eggs from
each female were prepared and analyzed 14 days later (=day 2). To
examine within-preparation variability, two separate aliquots of a single
preparation from each head and body were decanted and analyzed on

VOLUME 42, NUMBER 3 199

different days. Mean values for head and body samples were used in
subsequent analyses since no significant differences were found between
dates (Mann-Whitney U-test).

Data were analyzed to determine reliability of mark detection for
each female and her eggs. Mark thresholds for both eggs and adults
(heads and bodies) were determined and compared by two methods:
(1) using the high-range value of untreated controls, and (2) using the
conservative method of Stimmann (1974), which assumes a normal
distribution, three standard deviation units above the mean of untreated
control samples. All Rb concentration values are given in units per egg
or body part. Variation in egg weight within and between sib-groups
was considered negligible for our purposes (mean dry wt = 0.111 mg,
SE = 0.0017, N = 10/female). Variation in head and body weight was
more extreme (mean dry wt of heads = 0.897 mg, SE = 0.1536; body
wt = 16.54 mg, SE = 3.374). Within and between sib-group differences
were examined by analysis of variance. Parent-offspring relation was
evaluated by correlation of Rb content of a female body or head with
the mean Rb content of her eggs. Data analyses were performed using
SAS software.

RESULTS AND DISCUSSION

The mean quantity of Rb (in ppm) found in the body, head capsule,
and eggs of treated and untreated females is given in Table 1. Samples
prepared from the bodies of treated females were found to be 100%
reliably marked when compared to thresholds derived from samples
prepared from untreated adults. However, only 10-20% of the samples
prepared from heads alone produced detectable signals, and head results
were not significantly correlated with body results (r = 0.80). It is
apparent that detectable quantities of Rb were not evenly distributed
throughout the insect’s tissues. The time and expense of digesting whole
insects makes it advantageous to use the smallest sample that provides
consistent results. For Heliothis spp. it has been found that a single
wing is an adequate substitute for a whole moth (Hayes in press). For
butterflies, the wing is not as practical because of large size and the
frequent need to retain wings for morph determinations. Thus, Colias
samples prepared from wingless and headless bodies were used, and
they produced reliably detectable signals. If spermatophores are rou-
tinely dissected from females, or abdomens are removed for electro-
phoretic analysis, it would be ideal to be able to rely on a preparation
from the thorax alone. However, a feasibility test for use of the thorax
has not yet been conducted.

More than 90% of eggs (N = 100; 4 females) were determined to be
detectably marked regardless of method used (92% exceeded range of

200

TABLE 1. Quantity of Rb (mean & range in ppm) in untreated and treated female
Colias eurytheme (body and head) and their eggs (for 2 analysis days). Mark thresholds,
both high range of untreated controls (Mark 1) and Stimmann value calculated from
mean of controls (Mark 2), are provided along with mark determination (yes/no) or

percentage of marks.

Female no. N

Threshold values:

Body 10
Head 10
Eggs (day 1) 30
(day 2) 45
1
Body
Head
Eggs (day 1) 10
(day 2) 15)
2
Body
Head
Eggs (day 1) 10
(day 2) U5)
3
Body
Head
Eggs (day 1) 10
(day 2) 15
4
Body
Head
Eggs (day 1) 10
(day 2) 15
+)
Body
Head
Eggs (day 1) 10
(day 2) 15
6
Body 2
Head 2
Eggs (day 1) 10
(day 2) 15
Ff
Body
Head
Eggs (day 1) 10
(day 2) it)

Mean

0.0039*
0.0021:
0.0004
0.0011

0.0182
0.0046
0.0006
0.0013

0.0047
0.0125
6.0002
0.0011

0.0739
0.0208
0.0035
0.0038

0.0999
0.0058
0.0035
0.0038

0.0378
0.0106
0.0022
0.0045

0.0995
0.0101
0.0063
0.0070

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Rb concentration

Range

Untreated

0.0000-0.0012
0.0003-—0.0014

0.0002-—0.0014
0.0008—0.0022

0.0000-0.0006
0.0004—0.0028

Treated

0.0011-—0.0030
0.0031-—0.0048

0.0016-—0.0160
0.0030-0.0051

0.0011-0.0032
0.0019-0.0077

0.0033-—0.0169
0.0049-0.0086

Mark 1

0.0185
0.0140
0.0014
0.0028

no

no
0%
0%

no

no
0%
0%

no

no
0%
0%

yes
yes
90%

100%

yes
no

100%

100%

yes
no

70%
73%

yes

no
100%
100%

*Mean of two aliquots per sample; for further explanation see Materials and Methods.

Mark 2

VOLUME 42, NUMBER 3 201

nonmarks; 91% exceeded Stimmann’s value). The proportion of de-
tectably marked eggs per analysis day for each sib-group is given in
Table 1. Significant correlation was found on both analysis days between
maternal Rb content of the body and mean quantity of Rb in offspring
(Table 1; day 1 r = 0.88, P < 0.01; day 2 r = 0.84, P < 0.01).

Analysis of variance revealed significant differences between treated
and untreated samples (P < 0.0001; Table 1). More than 60% of the
variance was due to within-sib-group variance, more than 35% to be-
tween-group variance, and less than 1% was attributable to analysis-
day variance. Low day-to-day analysis difference is reassuring because
in field tests large numbers of samples must be processed over several
days.

Between-group variance dropped to less than 20% for the untreated
sib-groups when treated and untreated groups were analyzed sepa-
rately. However, within-sib-group variance remained above 60%. Be-
tween-group differences could be attributed to differing insect sizes
and element exposures. Eggs from a large female or one that has fed
consistently on well-treated foliage may show higher levels of Rb than
those from a smaller female or one that has fed inconsistently on a
treated host plant.

Within-group differences cannot be understood as easily. Each fe-
male was mated only once, and use of the first eggs oviposited should
lessen age effects. Since Rb is a potassium mimic, results suggest that
the female does not supply her eggs with consistently similar quantities
of necessary metabolites. Alternatively, inherent differences from egg
to egg (genomic differences) may result in the observed Rb content
variance. The parent-offspring correlation analysis reveals significant
associations which may indicate some degree of inherent relation. Fur-
ther investigation of these hypotheses could provide valuable insights
into development, and might dictate an expanded role for the use of
trace elements as an experimental tool.

In the final analysis, specimen-to-specimen, in particular egg-to-egg,
variability does not present difficulties for use of this marking technique
in field operations. Despite high individual variability, our findings
indicate that labelled parents and offspring are readily distinguishable
from unlabelled specimens.

Trace element marking has been reported previously with only one
other butterfly species, Pieris rapae (L.), and then only in the adult
stage (Stimmann 1974). The potential to exploit this marking method
among all Lepidoptera is great. It seems well justified since mark-
release-recapture studies using external markers are commonly used to
study pest and nonpest lepidopteran population attributes (Ehrlich &
Davidson 1960), but have received considerable criticism (Morton 1984).

202 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Along with problems arising from handling insects, insufficient recap-
ture numbers are a persistent problem. By labelling the egg, the adult
signal is amplified, and the concomitant ability to directly assess gene
flow is a definite advantage. Increasing the number of unique marks
will also improve recapture efficiency per unit area. Along with Rb,
other elements such as cesium and strontium are promising adult and
possible egg markers.

ACKNOWLEDGMENTS

We thank D. W. Hubbard and K. Reed for their assistance. Support was provided, in
part, by a grant to JLH from the Whitehall Foundation.

LITERATURE CITED

BERRY, W. L., M. W. STIMMANN & W. W. WoLF. 1972. Marking native phytophagous
insects with rubidium: A proposed technique. Ann. Entomol. Soc. Amer. 65:236-238.

CULIN, J. D. & D. R. ALVERSON. 1986. A technique to mark adult Heliothis zea using
rubidium chloride-spiked artificial nectar sources. J. Agric. Entomol. 3:56—60.

EHRLICH, P. R. & S. E. DAvIDSON. 1960. Techniques for capture-recapture studies of
Lepidoptera populations. J. Lepid. Soc. 14:227-229.

ENGEBRETSON, J. A. & W. H. Mason. 1981. Depletion of trace elements in mated male
Heliothis virescens and Drosophila melanogaster. Comp. Biochem. Physiol. 68A:
023-525.

GRAHAM, H. M. & D. A. WOLFENBARGER. 1977. Tobacco budworm: Labelling with
rubidium in the laboratory. J. Econ. Entomol. 70:800-892.

GRAHAM, H. M., D. A. WOLFENBARGER, J. R. Nosky, N. S. HERNANDEZ, J. R. LLANES
& J. A. TAMAyo. 1978. Use of rubidium to label corn earworm and fall armyworm
for dispersal studies. Environ. Entomol. 7:485-438.

HAYES, J. L. Detection of single and multiple trace element labels in the individual eggs
of diet-reared Heliothis virescens (F.) (Lepidoptera: Noctuidae). Ann. Entomol. Soc.
Amer. In press.

HAYES, J. L. & K. R. Hopper. 1987. Trace element labelling of Heliothis spp.: Labelling
of individual eggs from moths reared on treated host plants, pp. 311-315. In Cotton,
T. N. (ed.), Proc. Beltwide Cotton Production Research Conf., Dallas, Texas.

JONES, R. E., N. GILBERT, M. Guppy & V. NEALIS. 1980. Long-distance movement of
Pieris rapae. J. Anim. Ecol. 49:629-642.

Lecc, D. E. & H. C. CHIANG. 1984. Rubidium marking technique for the European
corn borer (Lepidoptera: Pyralidae) in corn. Environ. Entomol. 13:579-583.

Morton, A. C. 1984. The effects of marking and handling on recapture frequencies
of butterflies. Symp. R. Entomol. Soc. 11:55-58.

SLAVIN, W. 1984. Graphite furnace AAS, a source book. Perkin Elmer Corp., Ridgefield,
Connecticut. 230 pp.

SOUTHWOOD, T. R. E. 1978. Ecological methods. Chapman & Hall, London. 524 pp.

STIMMANN, M. W. 1974. Marking insects with rubidium: Imported cabbage worm
marked in the field. Environ. Entomol. 3:327-328.

STIMMANN, M. W., W. W. WoLF & W. L. BERRY. 1973. Cabbage loopers: Biological
effects of rubidium in the larval diet. J. Econ. Entomol. 66:324-326.

TABASHNIK, B. E. 1980. Population structure of pierid butterflies. III. Pest populations
of Colias philodice eriphyle. Oecologia 47:175-183.

TAYLOR, O. R., J. W. GRULA & J. L. HAYES. 1981. Artificial diets and methods for
continuous rearing of sulfur butterflies, Colias eurytheme and C. philodice (Pieridae).
J. Lepid. Soc. 35:281-289.

VOLUME 42, NUMBER 3 203

VAN STEENWYK, R. A., G. R. BALLMER, A. Z. PAGE, T. J. GANJE & H. T. REYNOLDS.
1978. Dispersal of rubidium-marked pink bollworm. Environ. Entomol. 7:608—-613.

Watt, W. B., P. C. Hocu & S. G. MILLS. 1974. Nectar resource use by Colias butterflies:
Chemical and visual aspects. Oecologia 14:353-374.

Received for publication 22 September 1987; accepted 8 April 1988.

Journal of the Lepidopterists’ Society
42(3), 1988, 203

ANNOUNCEMENT

JOURNAL COVER ILLUSTRATIONS AND FEATURE PHOTOGRAPHS

Journal submission categories are being broadened to include front cover illustrations
and feature photographs. Submissions in these new categories, like all submissions to the
Journal, may deal with any aspect of Lepidoptera study. Submissions in both categories
must be accompanied by brief captions that include scientific names.

Cover illustration subjects are well depicted by past covers created before submissions
were invited. Submissions should be no larger than letter size, with the caption for the
inside front cover on a separate sheet. Drawings may be more suitable than photographs
because drawings can usually better withstand the coarse reproduction necessitated by
present cover stock texture. Submissions will be selected for artistry, novelty, and general
appeal. There are no author page charges for cover illustrations.

Feature photographs might show unusual behaviors, unusual habitats, type localities,
specimens in nature illustrating identifying marks, or other subjects. Photographic sub-
missions should be mounted on white cardboard no larger than letter size, with a brief
caption on a separate sheet. Feature photographs must be suitable for reduction to either
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caption included). Submissions will be selected for photographic quality, scientific merit,
and general appeal. Regular page charges will apply to accepted feature photographs.

Submissions in both new categories should be sent to the Journal editor.

WILLIAM E. MILLER, Editor

Journal of the Lepidopterists’ Society
42(3), 1988, 204-212

AN APPRAISAL OF GAZORYCTRA HUBNER
(HEPIALIDAE) AND DESCRIPTION OF A
NEW SPECIES FROM ARIZONA AND NEW MEXICO

DAVID L. WAGNER!

Department of Entomology, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118

AND

NORMAN B. TINDALE

Honorary Research Associate, South Australian Museum,
North Terrace, Adelaide, S.A., Australia, 5000

ABSTRACT. Gazoryctra includes 10 species from North America, and 8 from N
Europe and Asia. Seven Nearctic species of Hepialus are transferred to Gazoryctra:
hyperboreus Méschler, lembertii Dyar, mathewi H. Edwards, novigannus Barnes &
Benjamin, pulcher Grote, roseicaput Neumoegen & Dyar, and sciophanes Ferguson.
Gazoryctra wielgusi is described based on 94 males from the White Mountains of Arizona
and adjacent New Mexico. A checklist of Gazoryctra is included.

Additional key words: Gazoryctra wielgusi, Hepialus, systematics, Holarctic.

Gazoryctra Hiibner are medium-sized swift moths found in high
latitudes or alpine habitats of the Holarctic Region. They are handsome
moths, with brown, orange, or pinkish forewings and silvery white
maculations. Adults of most species fly in late summer or fall. They are
exceptionally strong fliers, particularly the diurnal arctic-alpine species.
Many have very brief periods of diurnal activity, flying for only 20 or
30 min during evening twilight.

All previously known Gazoryctra were described in the nominotyp-
ical genus Hepialus Fabricius (or Epialus Lederer). North American
Gazoryctra have been referred to as the “hyperboreus group” by Barnes
and Benjamin (1926) and Ferguson (1979). Members of this group were
incorrectly placed in Phymatopus Wallengren by Pfitzner (1912, 1987—
38). Viette (1949) alone recognized that some Nearctic hepialids should
be classified in Gazoryctra.

The purpose of this paper is to provide adult, pupal, and larval
characters for the recognition of the genus, to clarify which elements
of the Holarctic hepialid fauna belong to Gazoryctra, to validate no-
menclature changes for the North American species, and to describe a
new species from the southwestern United States.

' Present address: Department of Ecology and Evolutionary Biology, 75 North Eagleville Rd., Room 312, U-43, Storrs,
Connecticut 06268.

VOLUME 42, NUMBER 3 205

Background

Hubner ({1820]:198) established three genera for the heterogeneous
Palearctic swift moth fauna occurring in N Europe: Gazoryctra, Phar-
macis, and Triodia. In his synopsis of the generic classification of Eu-
ropean Hepialidae, Viette (1949) designated Bombyx ganna as type
species of Gazoryctra. He characterized Gazoryctra as having a lobed
valva and a toothlike trulleum in male genitalia, and further noted that
small apical tibial spurs were present on middle and hind legs. He
transferred to Gazoryctra the Palearctic Hepialus macilentus Evers-
mann and the Nearctic H. confusus H. Edwards and H. mcglashani
H. Edwards. Later, Viette (1953) noted that Gazoryctra species also
had prominent earlike lobes in the intersegmental membrane between
abdominal segments 8 and 9 (socii of Robinson 1977, tergal lobes of
Nielsen and Kristensen in press), and added a fifth species to the genus,
H. fuscoargenteus Bang-Haas.

Many lepidopterists have overlooked or ignored the generic concepts
of Hubner [1820], Wallengren (1869), Borner (1925), and Viette (1948,
1949), and have continued to treat most Holarctic swift moths as mem-
bers of the nominotypical genus Hepialus (McDunnough 1939, Heath
1976, Ferguson 1979, Davis 1983). In North America, Hepialidae have
been classified into two genera: Hepialus (type species: humuli Lin-
naeus) and Sthenopis Packard (type species: qguadriguttatus Packard).
Most lepidopterists have recognized Sthenopis from the time of its
proposal in 1865 (Packard 1865, Kirby 1892, Neumoegen & Dyar 1894,
Wagner & Pfitzner 1911, Forbes 1923, McDunnough 1939, Davis 19838).
However, our studies indicate that Hepialus humuli and members of
Sthenopis (with Zenophassus Tindale, Aenetus Walker, and perhaps
others) have a common ancestor not shared by most other Holarctic
Hepialidae. Synapomorphies for these taxa include (1) a metatibial
hairpencil in males; (2) swollen metatibiae in males (members of the
Sthenopis regius group and some Aenetus lack hairpencils and swollen
metatibiae, but these appear to be reversals within the clade); (8) tri-
angular forewings with falcate apices; (4) forewing scales with rounded
apices; and (5) absence (loss) of the epiphysis in all but Aenetus and
Sthenopis argenteomaculatus (Harris). In addition, all members of this
group are larger than most other Hepialidae and exoporian Lepidoptera,
with forewing lengths typically exceeding 4 cm. Consequently, if Sthen-
opis is to be retained as a distinct genus, then names proposed for more
distantly related taxa like Gazoryctra warrant at least generic status.

Gazoryctra in North America

So that we could reliably assign the hepialid described here to a
genus, we prepared dissections of all the Holarctic hepialid generotypes.

206 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

We also examined all North American hepialid extant primary types,
and prepared genitalia and wing slides for all named Nearctic and
many European species, including all examined by Viette (1949).

In addition to the characters given by Viette (1949), eight others
were found to be shared by Gazoryctra species: (1) forewing subcosta
forked (vein Sc, well developed); (2) halves of tegumen meeting dor-
soanteriorad, but free over much of midline; (3) dorsal margin of teg-
umen bearing dense oval patch of spinules; (4) caudal portion of trul-
leum not fused to tegumen; and (5) pulvilli large and setose. In the
larva, (6) claw elongate, with basal tooth ending before 4; (7) D2 and
SD setae on prothorax not grouped. In the pupa, (8) caudal band of
teeth encircling abdominal segment 7 broken ventrolaterad.

Our studies indicate that Gazoryctra is the largest genus of North
American Hepialidae. In addition to the two Nearctic species identified
by Viette (1949), seven other described hepialids were found to share
this list of characters and are transferred here to the genus Gazoryctra.

Checklist

In what follows, subspecific names are indicated by a), and are but
tentatively recognized. Author names followed by year do not neces-
sarily refer to literature cited in this paper.

Gazoryctra Hibner [1820]
Gazoryctes Kirby 1892, missp.

confusa (H. Edwards 1884) (Hepialus)
fuscoargentea (Bang-Haas 1927) (Hepialus)
sordida (Nordstrom 1929) (Hepialus), infrasubsp.
a) postmaculata (Landin 1943) (Hepialus)
ganna (Hibner [1810]) (Bombyx)
arctica (Boheman 1848) (Hepialus)
reducta (Deutsch 1925) (Hepialus), infrasubsp.
confluens (Hellweger 1914) (Hepialus), infrasubsp.
chishimana (Matsumura 1931) (Hepialus), infrasubsp.
hyperborea (Méschler 1862) (Epialus), new combination
lembertii (Dyar 1894) (Hepialus), new combination
macilenta (Eversmann 1851) (Hepialus)
gerda (Staudinger 1897) (Hepialus)
a) nesiotes (Bryk 1942) (Hepialus)
mathewi (H. Edwards 1875) (Epialus), new combination
matthewi (H. Edwards 1884), missp.
mceglashani (H. Edwards 1887) (Hepialus)
mceglachanii (Pfitzner 1912), missp.
noviganna (Barnes & Benjamin 1926) (Hepialus), new combination
novigana (Pfitzner 1937-38) (Hepialus), missp.
a) mackiei (Barnes & Benjamin 1926) (Hepialus)
pulchra (Grote [1865]) (Hepialus), new combination
roseicaput (Neumoegen & Dyar 1893) (Hepialus), new combination
mutata (Barnes & Benjamin 1926) (Hepialus), infrasubsp.
demutata (Barnes & Benjamin 1926) (Hepialus), infrasubsp.

VOLUME 42, NUMBER 3 207

Fics. 1, 2. Gazoryctra wielgusi. 1, Holotype male; 2, Paratype males from Ditch
Camp. White scales replaced by silvery gray scales in lower right-hand specimen. Label
data in text.

sciophanes (Ferguson 1979) (Hepialus), new combination
wielgusi Wagner & Tindale, new species

Gazoryctra wielgusi, new species

This pink and silvery moth is so far known only from a restricted area in the mountains
of E Arizona and W New Mexico. Terminology for genital structures follows Birket-
Smith (1974), Ueda (1978), and Nielsen and Robinson (1983); for wing veins, Nielsen and
Robinson (1983); and for scale ultrastructure, Downey and Allyn (1975) and Kristensen
(1978b).

Male (Figs. 1-11). Forewing length 15-18 mm (N = 94). Head. Antenna with 29-32
segments (N = 10), flagellomeres slightly compressed with abundant short setose sensory
setae (Figs. 6, 7), yellow to orange-brown. Head vestiture dense admixture of buff and
darker piliform scales; dark or dark-tipped scales prominent over frons, labial palpus,
and ventral region. Labial palpus with 2 subequal segments (Fig. 3), vom Rath’s organ
dorsosubapical. Thorax. Pro- and mesothoracic dorsum with brown-tipped and buff
piliform scales intermixed; metathorax buff. Procoxae and pro- and mesofemora dark-
scaled. Tibiae and tarsi with elongate salmon-colored lamellar scales and contrasting dark
fusiform scales (Fig. 8). Venation (Fig. 4). R,,, branched at mid-length; hindwing vein
CuP obscure in some specimens, and 2A differentiated from wing cuticle. Seales (dorsal
surface over median region) (Figs. 10, 11). Broadest beyond middle, apices 3- or 4-
toothed; secondary ridges prominent; windows small, circular to elliptical, diameter less
than ¥4 interridge distance, surrounded by ring of unmodified cuticle, separated by 1 to
3 transverse flutes; window membrane occasionally present; flutes prominent with per-
pendicular secondary ribbing between adjacent flutes. Forewing tan or brown to peach
or salmon with peppering of darker scales; heavily maculated with silvery white (rarely
silvery gray) markings, these outwardly edged with dark scales; submarginal spots nearly
always present, occasionally fused with oblique submarginal band; white spots or streaks
also along subcosta and base of inner margin. Hindwing uniformly brown with orange
or salmon-colored scales along margin, at apex, and extending basally along veins; apex
faintly patterned. Fringe of both wings orange or salmon. Abdomen. Dorsum of segments
1 and 2 uniformly covered with long pale piliform scales; segments 3 to 8 with both long
buff scales and lamellar salmon-colored scales. Genitalia (Fig. 5). Tergal lobes prominent,
densely setose dorsad and laterad, hemispherical, with ventrolateral digitate lobe extend-
ing below margin of tegumen. Caudal margin of tegumen with 2 sets of ventrally pro-
jecting, strongly melanized processes, upper pair digitate and angled ventrad, apices with
single prominent tooth and several smaller distal teeth; lower pair gradually tapering to

208 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fics. 8-5. Gazoryctra wielgusi. 3, Labium: scale = 0.2 mm (DLW Slide 86-63); 4,
Wing venation: scale = 0.5 cm (DLW Slide 86-62); 5, Male genitalia: scale = 0.5 mm.

points, apices approximate over midline. Trulleum long, narrow, tapering to strongly
melanized spine. Valva densely setose, boot-shaped; lower lobe rounded, curving mesad;
inner margin notched above basal articulation. Juxta elongate, constricted in middle;
roughly pentangulate. Vinculum often emarginate ventrolaterad.

Female. Unknown.

Diagnosis. A heavily maculated species with silvery white streaks and spots along base
of inner margin, subcosta, and termen between medial veins. It can be readily separated
from other strongly marked Gazoryctra—confusa, hyperborea, pulchra, and roseicaput —
by its dark brown hind wings with contrasting orange or salmon-colored fringe. The
oblique submarginal band is continuous, never broken into separate spots as is often the
case in other members of the genus. It is the only salmon-colored Gazoryctra in the S
Rocky Mountains. Male genitalic characters distinguishing wielgusi from at least some
other members of the genus follow: tergal lobes nearly as high as broad, valva boot-
shaped, vinculum ventrolaterally emarginate, trulleum very elongate.

Distribution. White Mountains of E Arizona and adjacent ranges in New Mexico
between 2400 and 2800 m elev.

Material examined. 94 males. Holotype: Male, Arizona, Apache Co., 14.4 km E McNary,
Ditch Camp, 2400 m, 25-VII-1974, R. S. Wielgus, at ultraviolet and white light, DLW
Slide No. 86-66. Deposited in Los Angeles County Museum. Paratypes: Arizona: Apache

VOLUME 42, NUMBER 3 209

Fics. 6-9. Gazoryctra wielgusi male. 6, Antenna showing abundance of sensory setae
and cuticular projections. Scale = 43 um; 7, Distal antennal segments. Scale = 43 um; 8,
Protibia. Fusiform scales (arrows) appear dark and spinelike against salmon-colored squa-
mose scales. Scale = 60 um; 9, Metathoracic tibial spurs. Scale = 100 um.

Co., same data as above, 25-VII/14-VIII-1974/80, R. S. Wielgus, 58 males (Fig. 2), and
26/29-VII-1978, N. B. & M. Tindale, 2 males; White Mtns., Greer, 2600 m, 4-VIII-1962,
E. & I. Munroe, 4 males; Greer, 0.8 km S, Government Springs, 6- VIJI-1977, R. S. Wielgus,
2 males, and 28/30-VII-1978, N. B. & M. Tindale, 10 males; Greer, 19.2 km SW, Winn
Cmpgd., 2800 m, 26-VII-1986, R. Robertson, 12 males; Greenlee Co., Hannagan Meadow,
“12-IV-1975”, A. Menke, 3 males. New Mexico: Catron Co., Gila Wilderness, along route
78, Willow Creek Cmpgd., ca. 2400 m, 28-VII-1978, 1947 h MST, R. S. Wielgus, 1 male;
Indian Creek nr. Gilita Cmpgd., 2400 m, 29-VII-1978, 1948 h MST, R. S. Wielgus, 1
male.

The three specimens of G. wielgusi in the USNM bearing the data “CALIFORNIA:

210 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 10, 11. Gazoryctra wielgusi scales from medial region of male forewing. 10,
Scale showing typical exoporian arrangement of primary and secondary ridges. Scale =
30 wm; 11, Scale ultrastructure: scutes and flutes well developed. Scale = 1.15 wm.

Greenlee Co., Hannagan MDW., April 12, 1975, A. Menke” are mislabeled as there is
no Greenlee Co., California. Moreover, all Gazoryctra species are summer- or fall-flyers.
Presumably, Menke captured the moths in Hannagan Meadow during a collecting trip
to the Southwest in August 1975 (A. Menke pers. comm. ).

Paratypes are deposited at Arizona State University, Tempe; Australian National Insect
Collection, Canberra; British Museum (Natural History), London; California Academy
of Sciences, San Francisco; Canadian National Collection, Ottawa; Los Angeles County
Museum, California; South Australian Museum, Adelaide; United States National Museum,
Washington, D.C.; University of California, Berkeley and Davis; Zoologische Staats-
sammlung, Munich.

Biology. Gazoryctra wielgusi has a very brief period of adult activity. All specimens
have been captured in early evening just after onset of darkness. In late July, the main
flight occurs between 1945 and 2000 h MST (N = 17); by mid-August flight starts as early
as 1930 h MST (N = 7). All specimens (males) were collected at light; typically they are
the first moths to arrive at sheets. Adults are most numerous after afternoon rains and
may even fly during strong rains (R. S. Wielgus pers. comm. ).

Our records are from mesic areas in conifer forests. The locality at Ditch Camp is an
open ponderosa pine forest with abundant grasses in open areas and nearby alders. Higher-
elevation localities have more understory shrubbery. Spruce is dominant at the two sites
in New Mexico. Nothing is known of the early stages. The larvae presumably are po-
lyphagous, subterranean feeders, as are other Holarctic hepialids (Heath 1976, Wagner
1985, 1987). Recorded hosts for other Gazoryctra include Betula, Phlox, Picea, and grasses
(Wagner 1985, Tham et al. 1985).

Etymology. We name this moth after Ronald S. Wielgus whose seemingly inexhaustible
collecting efforts produced most of the known specimens and biological data.

Discussion

Gazoryctra is confined to the Holarctic Region; 10 Nearctic and 3
Palearctic species are recognized. No member is recognized from both

VOLUME 42, NUMBER 3 DANS |

faunas, although the markings of both ganna and macilenta approach
those of hyperborea from North America.

Gazoryctra appears to represent one of the most primitive genera of
Hepialidae. No synapomorphies have been identified that link Gazo-
ryctra to other hepialids. In the past, the absence of tibial spurs has
been used to define Hepialidae (Borror & White 1970, Kristensen 1978a,
Nielsen & Robinson, 1983). Yet members of Gazoryctra possess a pair
of small tibial spurs (Fig. 9, Viette 1949, Wagner 1985). In addition,
the trulleum is free from the tegumen caudad in Gazoryctra, but fused
in more derived Hepialidae (Nielsen & Scoble 1987).

Both adults and immature stages of Gazoryctra are rare in collections.
Only three new hepialids have been described from North America
since the turn of the century, and all belong to Gazoryctra. Moreover
no specimens of sciophanes and wielgusi were known before 25 years
ago. The biology is not known in detail for any species.

ACKNOWLEDGMENTS

M. A. Tenorio assisted with scanning electron microscopy and darkroom work. E. S.
Nielsen, G. S. Robinson, M. J. Scoble, W. J. Pulawski, V. Lee, and T. E. Miller offered
many helpful suggestions on the manuscript. We thank P. T. Dang, Biosystematics Re-
search Institute, Ottawa, Canada; D. R. Davis, Smithsonian Institution, Washington, D.C.;
J. P. Donahue, Los Angeles County Museum, California; and R. G. Robertson for loans
of material. This work was supported in part by a Tilton Fellowship to DLW from the
California Academy of Sciences, San Francisco, California.

LITERATURE CITED

Items marked by * were not seen by present authors.

BARNES, W. & F. H. BENJAMIN. 1926 (1925). The hyperboreus group of Hepialus
(Lepidoptera: Hepialidae). Pan-Pac. Entomol. 2:81-84.

BIRKET-SMITH, S. J. R. 1974. Morphology of the male genitalia of Lepidoptera. II.
Monotrysia, Zeugloptera, and discussion. Entomol. Scand. 5:161-183.

*BORNER, C. 1925. Lepidoptera, pp. 358-387. In Brohmer, P. (ed.), Fauna von Deutsch-
land. Quelle & Meyer, Leipzig.

Borror, D. J. & R. E. WHITE. 1970. A field guide to the insects of America north of
Mexico. Houghton Mifflin, Boston. 404 pp.

BROWN, F. M. 1964 (1963). Dates of publication of the various parts of the Proceedings
of the Entomological Society of Philadelphia. Trans. Am. Entomol. Soc. 89:305-308.

Davis, D. R. 1983. Hepialidae, p. 2. In Hodges, R. W., et al. (eds.), Check list of the
Lepidoptera of America north of Mexico. E. W. Classey Ltd., London.

Downey, J. C. & A. C. ALLYN. 1975. Wing-scale morphology and nomenclature. Bull.
Allyn Mus. 31:1-32.

FERGUSON, D. C. 1979. A new ghost moth from the southern Appalachian Mountains
(Hepialidae). J. Lepid. Soc. 33:192-196.

FORBES, W. T. M. 1923. The Lepidoptera of New York and neighboring states. Cornell
Univ. Agr. Exp. Stat. Mem. 68:1-—729.

HEATH, J. 1976. Hepialidae, pp. 166-170. In Heath, J. (ed.), The moths and butterflies
of Great Britain and Ireland. Curwell Science Press, London.

HEMMING, F. 1937. Hiibner: A bibliographical and systematic account of the ento-
mological works of Jacob Hiibner. Roy. Entomol. Soc. Lond. Vol. 1 605 pp; Vol. 2
271 pp.

212 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

HUBNER, J. [1820]. Verzeichniss bekannter Schmettlinge. Privately printed, Augsburg.
Pp. 198-199. (Dating after Hemming 1937.)

KirBy, W. F. 1892. A synonymic catalogue of Lepidoptera Heterocera. (Moths). Vol.
1. Sphinges and Bombyces. Gurney & Jackson, London. Pp. 879-894.

KRISTENSEN, N. P. 1978a. A new familia of Hepialoidea from South America, with
remarks on the phylogeny of the subordo Exoporia (Lepidoptera). Entomol. Germ.
4:272-294.

1978b. Ridge dimorphism and second-order ridges in wing scales in Lepidoptera:
Exoporia. Int. J. Insect Morphol. Embryol. 7:297-299.

McDuNNOUGH, J. H. 1939. Check list of the Lepidoptera of Canada and the United
States of America. Part IJ. Microlepidoptera. Mem. So. California Acad. Sci. 2:1-171.

NEUMOEGEN, B. & H. G. Dyar. 1894. Preliminary revision of the Bombyces of America
north of Mexico. J. New York Entomol. Soc. 2:147-174.

NIELSEN, E. S. & N. P. KRISTENSEN. In press. Primitive ghost moths: Morphology and
taxonomy of the Australian genus Fraus Walker (Lepidoptera: Hepialidae s. lat.). E.
J. Brill, Leiden.

NIELSEN, E. S. & G. S. ROBINSON. 1983. Ghost moths of southern South America
(Lepidoptera: Hepialidae). Entomonograph Vol. 4. Scandinavian Science Press, Co-
penhagen. 192 pp.

NIELSEN, E. S. & M. J. SCOBLE. 1987. Afrotheora, a new genus of primitive Hepialidae
from Africa (Lepidoptera: Hepialoidea). Entomol. Scand. 17:29-54.

PACKARD, A. S. 1865 (1864). Synopsis of the Bombycidae of the United States. Proc.
Entomol. Soc. Philadelphia 3:331-396. (Dating after Brown 1964.)

PFITZNER, R. 1912. Familie: Hepialidae, pp. 433-439, pls. 58, 54. In Seitz, A. (ed.), Die
Gross-Schmetterlinge der Erde. 1 (8). Kernan, Stuttgart.

1937-38. Familie: Hepialidae, pp. 1289-1302, pls. 99, 185. In Seitz, A. (ed.),
Die Gross-Schmetterlinge der Erde. 6 (4). Kernan, Stuttgart. (Often cited as R. Pfitzner
& M. Gaede.)

ROBINSON, G.S. 1977. A taxonomic revision of the genus Callipielus Butler (Lepidoptera:
Hepialidae). Bull. Brit. Mus. (Nat. Hist.) (Entomol.) 35:101-121.

THAM, E., H. ELMQuisT & H. TREI. 1985. Larval biology of H. fuscoargenteus Bang-
Haas (Lep.: Hepialidae). Entomol. Tidskr. 106:43—44.

UepDA, K. 1978. The male genital structure of some hepialid moths with a historical
review of their terminology. [Japanese, English summary.] Ty6 to Ga 29:191-206.

VIETTE, P. E. L. 1948. Lépidoptéres Homoneures. Faune de France. Vol. 49. Lech-
evalier, Paris. 83 pp.

1949. Contribution a l’etude des Hepialidae (12° note). Genres et synonymie.

Lambillionea 49:101-104.

1953. Contribution a |’étude des Hepialidae (30° note). Lambillionea 53:32-35.

WacGNneER, D. L. 1985. The biosystematics of Hepialus F. s. lato, with special emphasis
on the californicus-hectoides species group. Ph.D. Dissertation, University of Cali-
fornia, Berkeley. 391 pp. Diss. Abstr. Order No. DA8610260.

1987. Hepialidae, pp. 347-349. In Stehr, F. (ed.), Immature insects. Kendall-
Hunt, Dubuque, Iowa.

WAGNER, H. & R. PFITZNER. 1911. Hepialidae. Lepidopterorum Catalogus 4:1—26.

*WALLENGREN, H.D.J. 1869. Skandinaviens Heterocer-fjarilar. 2 vols. C. Bulow, Lund.
256 pp.

Received for publication 20 February 1987; accepted 25 April 1988.

Journal of the Lepidopterists’ Society
42(3), 1988, 213-217

TEMPORAL TRENDS IN FREQUENCIES OF
MELANIC MORPHS IN CRYPTIC MOTHS
OF RURAL PENNSYLVANIA

THOMAS R. MANLEY

Route #1, Box 269, Port Trevorton, Pennsylvania 17864
Curatorial Affiliate in Entomology, Peabody Museum of Natural History,
Yale University, New Haven, Connecticut 06511

ABSTRACT. Five species of moths with recorded melanic forms were light-trapped
for 10-16 years during 1971-86 at a remote mountain valley in E-central Pennsylvania.
Observed melanic frequencies were: Biston betularia cognataria 0.52 (1971-78) and 0.38
(1979-86); Epimecis hortoria 0.34; Phigalia titea 0.14; Charadra deridens 0.64; Catocala
ultronia 0.001. All but Biston showed constant melanic frequencies during their sampling
periods.

Additional key words: Geometridae, Noctuidae, Biston betularia cognataria, Ly-
mantria dispar.

Many eastern North American nocturnal moths that escape daytime
predation by cryptic resting behavior have a moderate frequency of
heritable dark forms. A simple working hypothesis to explain the adap-
tive advantage of this phenomenon is that these are “industrial melan-
ics’, favored by industrial and automotive pollutants in the atmosphere
(Kettlewell 1973). Such an hypothesis is only vaguely supported in North
America because of inadequate records. If trends showing systematic
increases or decreases in frequency of melanics can be documented,
the causes of such shifts might be found.

A second major body of relevant evidence results when enough lo-
calities are sampled for melanic frequencies. With these possibilities in
view, I have been obtaining melanism data from daily light trap samples
in a rural wooded valley for 16 years. The first eight years (1971-78)
of records for Biston betularia cognataria (Guenée) (Geometridae) were
presented and discussed previously (Manley 1981). In the present paper,
data are given for B. b. cognataria for the next eight years (1979-86)
as well as the first eight, and comparative records are included for
Epimecis hortoria (F.), Phigalia titea (Cramer) (both Geometridae),
Charadra deridens (Guenée), and Catocala ultronia (Hiibner) (both
Noctuidae). The observed melanic frequencies are compared with each
other and with published records from other areas.

MATERIALS AND METHODS

Sampling (dusk to dawn) was conducted nightly from April through
early September each year using a fluorescent 15-watt blacklight and
a mercury-vapor light trap. The trap was located in an isolated mountain
valley 12 km NE of Klingerstown, Schuykill Co., Pennsylvania. Except

214 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

TABLE 1. Melanic frequencies in four species of moths near Klingerstown, Pennsyl-
vania. The two annual broods of Biston betularia cognitaria are combined. Dash signifies
no observation.

Biston b. cognitaria Charadra deridens Epimecis hortoria Phigalia titea
Fre- Fre- Fre- Fre-

Year quency N quency N quency N quency N
1ST 0.52 588 — —_ _ — — —
72 0.51 669 = = a == = —
73 0.56 828 0.68 35 — — — —
74 0.52 272 0.58 52 0.47 15 — —
75 0.52 102 0.62 50 0.35 15 — —
76 0.53 219 0.80 5 0.38 21 — —
ehh 0.51 244 0.67 15 0.33 55 0.14 176
78 0.46 226 0.71 7 0.23 13 On 189
79 0.48 452 0.73 pl 0.29 68 0.15 148
80 0.52 68 0.86 a 0.25 24 0.17 219
81 0.43 30 0 0 0.14 7 0.20 257
82 0.36 100 0 0) 0.22 40 0.15 31
83 0.34 466 0) 0) 0.42 52 0.15 180
84 0.33 196 0.67 12 0.31 74 0.18 211
85 0.34 239 0.75 4 0.37 87 0.15 93
86 0.38 301 0.75 4 0.38 60 0.10 33

Total 0.47 5000 0.65 202 0.33 531 0.15 1537

for a few open fields, the area is tree-covered. Details of site, sampling
methods, and regional sources of air pollution potentially affecting air
quality and melanic frequency are given in Manley (1981). Sampling
extended over a period of 10 or more years for each species. Specimens
were pinned with full data and are part of the Manley Collection,
Peabody Museum of Natural History, Yale University.

The data format in Table 1 allows quick comparisons with Owen
(1961, 1962), Sargent (1971, 1974), and Klots (1964, 1968). G-tests using
Williams’s correction (Sokal & Rohlf 1981) were employed to test whether
melanic frequencies differed from year to year.

RESULTS

Biston betularia cognataria appears to undergo large fluctuations in
population density every 4—5 years (Table 1). In 1975, the first brood
consisted of 3 trapped specimens followed by a second brood of 99,
with the melanic frequency of 0.52 equal to the first 8-year average
(Manley 1981). The melanic frequency declined from 0.52 in 1980 to
0.33 in 1984, while numbers trapped rebounded to levels before the
population crash of 1980-81. Populations after 1981 show a six-year
(1981-86) melanic frequency of 0.34, compared to 0.48 for the 6-year
period (1975-80) following the 1975 crash. This contrasts with a fre-
quency of 0.53 for the first 4 years (1971-74).

The 1971-78 trends in Biston b. cognitaria were earlier interpreted

VOLUME 42, NUMBER 3 215

partly as a gradual decline in melanic frequency (Manley 1981). The
new data here strengthen this suggestion (linear regression of melanic
frequency against year: slope = —0.016 + 0.002, N = 16, t = 7.27, P <
0.001). However, an equally plausible explanation is the population
crash of 1981 (1971-80 data versus 1981-86 data: G = 182.47, df = 1,
P < 0.001; samples in 1971-80 set are homogeneous, as are samples in
1981-86 set, P > 0.25 by G-tests).

Epimecis hortoria (Table 1) exhibits a stable frequency of 0.33 for
its melanic form “carbonaria”’ (G = 9.98, df = 12, P > 0.50). This
moth has become increasingly abundant at the light trap since its pop-
ulation crash in 1981. Owen (1961, 1962) reported a 1957 sample of 8
specimens from Westmoreland Co., just E of Pittsburgh, as 100% me-
lanic, and a 1959 sample at Lebanon, Hunterdon Co., New Jersey, as
0.90.

The melanic form “deplorens” of Phigalia titea is distinct, with no
apparent intermediates; it (Table 1) maintained a stable frequency of
0.15 during the sampling period (G = 8.66, df = 9, P > 0.50). Owen
(1961) reported 1960 melanic frequencies in Michigan ranging from
0.11 to 0.14; Sargent (1971) reported 1968-70 melanic frequencies at
Leverett, Massachusetts, at 0.20.

My samples of Phigalia titea were taken 25 March through 5 May.
Since this moth begins flying on warm March nights, a portion of the
total possible sample of it was probably not taken. My samples reflect
only warm nights in late March with no continuous sampling until
April. Nevertheless, there does not appear to be substantive change in
melanic frequency of this species.

Melanic frequency for Charadra deridens during 1973-86 (Table 1)
was stable at 0.65 (G = 3.89, df = 9, P > 0.50). Klots (1968) at Putman,
Windham Co., Connecticut, reported limited 1961-66 samples (N =
28) to be 0.89 melanic, and a laboratory reared sample (N = 39) to be
2:1 melanic.

Charadra deridens samples have been small since 1981-83, when no
moths were taken (Table 1). The 10-year melanic frequency of 0.65 is
the highest of any melanic moth sampled to date at this locality.

Catocala ultronia, the most abundant Catocala in central Pennsyl-
vania, was sampled during 1968-78. In my sampling, only 2 of 1520
specimens were the melanic form “nigrescens’’, all others being color
variants of form “‘celia’’. Sargent (1974) reported the 1968-74 melanic
frequency in Leverett, Massachusetts, to be 0.17 (N = 586).

DISCUSSION

Except for Catocala ultronia, the species discussed in this paper show
strong melanic tendencies. Melanic frequencies differ from those ob-
served at Leverett, Massachusetts (Sargent 1974), even for the same

216 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

years. Melanic frequency appears to fluctuate independently at least
in part among species. Some factors affecting melanic frequencies are
presented in Manley (1981).

The 1980-81 season heralded severe reductions in the populations of
Biston, Epimecis and Charadra, whereas Phigalia produced the largest
sample of its 10-year period, only to be reduced to 31 individuals in
1982. An explanation for the sudden reductions in population densities
could in part be local infestation of the deciduous woods by the gypsy
moth, Lymantria dispar (L.). A partial defoliation in 1980 was followed
by severe defoliation in 1981 and aerial spraying with Dylox or Dimilin
by the Pennsylvania Department of Forestry. Spraying was discontin-
ued in 1982, and no noticeable defoliation has occurred since.

Four species in this study are polyphagous. More than 50 species of
trees and shrubs are recorded for Biston betularia cognitaria, with
Salix, Populus, Betula, and Alnus preferred (Rindge 1975, McGuffin
1977). Prentice (1963) lists 25 species of hard and softwoods for Phigalia
titea, with Tilia, Ulmus, Betula, Populus, Acer, and Quercus preferred.
Epimecis hortoria prefers Liriodendron, Sassafras, and Prunus, and is
rarely found on other deciduous trees (Forbes 1948). Charadra deridens
prefers Ulmus, Betula, and Quercus (Forbes 1954). Only Quercus among
the preferred food plants is normally eaten by the gypsy moth. Aerial
spraying to control gypsy moth may have been a critical factor in
reducing populations of the polyphagous species. The rapid recovery
of Biston and Phigalia following reductions in 1981-82 (Table 1) could
be attributed to the wide range of food plants acceptable to these species.
Similarly, the preferred food plants of Epimecis are not those eaten by
larvae of the other species in this study, thus perhaps accounting for a
rapid recovery in 1982. Charadra deridens, a Quercus feeder (Forbes
1954), appears to have been severely reduced in the defoliated area, as
none were trapped from 1981 to 1983, and it remains rare in the area
(Table 1).

The population reduction of Phigalia in 1982 does not coincide with
the 1981 reductions of the other species (Table 1) in that the annual
sample was taken before the aerial spraying in 1981.

Defoliation by gypsy moth opens the forest canopy, allowing light
penetration which could aid predators in finding active adults, espe-
cially ovipositing females. The tendency of birds to seek the safety of
trees escaping defoliation could increase the density of predators on
larvae feeding on those trees, as well as on ovipositing females.

Air pollution over Pennsylvania is often from industrial areas in the
Ohio Valley and Gulf Coast (Brown 1987). Local sources of pollution
do not greatly increase the quantity of oxides of sulfur and nitrogen in
the sampling area, since the area is not industrial. Despite reports of a

VOLUME 42, NUMBER 3 ag lab

two- to three-fold increase in ozone and other oxidants in the Appa-
lachians between 1962 and 1976 (West 1977), air pollution may not be
a critical factor in fluctuations in melanic frequencies at this site. Despite
high levels of polluted air over Pennsylvania, four of the five species
sampled in this report have stable melanic frequencies. The determining
factors appear to be localized biological factors, like gypsy moth.

The data here suggest that the widespread aerial spraying for gypsy
moth control may be having a catastrophic effect on many species of
lepidopterous insects in the eastern United States.

ACKNOWLEDGMENTS

I thank L. F. Gall, C. L. Remington (Yale University) and D. F. Schweitzer (The Nature
Conservancy) for reviewing and criticizing the manuscript, and L. J. Kopp (Klingerstown,
Pennsylvania) for operating the light traps on his property and caring for the samples.

LITERATURE CITED

Brown, M. H. 1987. Toxic wind. Discover. Nov. 1987:42—49.

ForBES, W. T. M. 1948. Lepidoptera of New York and neighboring states. Part II.
Geometridae. Mem. Cornell Univ. Agr. Exp. Sta. 274, 263 pp.

1954. Lepidoptera of New York and neighboring states. Part III. Noctuidae.
Mem. Cornell Univ. Agr. Exp. Sta. 329, 433 pp.

KETTLEWELL, B. 1973. The evolution of melanism. Clarendon Press, Oxford. 423 pp.

Kiots, A. B. 1964. Notes on melanism in some Connecticut moths. J. New York Entomol.
Soc. 72:142-144.

1968. Melanism in Connecticut Charadra deridens (Guenée) (Lepidoptera:
Noctuidae). J. New York Entomol. Soc. 76:58-59.

MANLEY, T. R. 1981. Frequencies of the melanic morph of Biston cognataria (Geo-
metridae) in a low-pollution area in Pennsylvania from 1971 to 1978. J. Lepid. Soc.
39:257-265.

McGuFFIN, W.C. 1977. Guide to the Geometridae of Canada (Lepidoptera) II. Subfam-
ily Ennominae. 2. Mem. Entomol. Soc. Canada 101, 191 pp.

OwEN, D. F. 1961. Industrial melanism in North American moths. Amer. Nat. 95:227-
233.

1962. The evolution of melanism in six species of North American geometrid
moths. Ann. Entomol. Soc. Amer. 55:695-703.

PRENTICE, R. M. 1963. Forest Lepidoptera of Canada. Canada Dept. Forestry, Forest
Entomol. Pathol. Br. Publ. No. 1013, 3:283-543.

RINDGE, F.H. 1975. A revision of the New World Bistonini (Lepidoptera: Geometridae).
Bull. Amer. Mus. Nat. Hist. 156 (2):69-156.

SARGENT, T. D. 1971. Melanism in Phigalia titea (Cramer) (Lepidoptera: Geometridae).
J. New York Entomol. Soc. 79:122-129.

1974. Melanism in moths of central Massachusetts (Noctuidae, Geometridae).
J. Lepid. Soc. 28:145-152.

SOKAL, R. R. & F. J. ROHLF. 1981. Biometry. Freeman, San Francisco. 859 pp.

WEsT, D. A. 1977. Melanism in Biston (Lepidoptera, Geometridae) in the rural central
Appalachians. Heredity 39:75-81.

Received for publication 1 June 1987; accepted 4 April 1988.

Journal of the Lepidopterists’ Society
42(3), 1988, 218-230

A NEW SPECIES OF OCALARIA
(NOCTUIDAE: CATOCALINAE) AND ANALYSIS OF
SOME MORPHOLOGICAL CHARACTERS USEFUL

FOR ELUCIDATING NOCTUID PHYLOGENY

IAN J. KITCHING

Department of Entomology, British Museum (Natural History),
Cromwell Road, London SW7 5BD, Great Britain

ABSTRACT. Ocalaria cohabita is described from 24 specimens captured on Barro
Colorado Island, Panama. A key to adults of Ocalaria is provided, followed by a mor-
phological comparison of O. cohabita, O. oculata (Druce) and O. quadriocellata (Walker).
Particular emphasis is placed on characters that may prove useful in elucidating noctuid
phylogeny, including features of head, legs, wings, abdominal segments 2 and 8, and
genitalia.

Additional key words: morphology, Panama, Ocalaria oculata, O. quadriocellata,
systematics.

Greig and DeVries (1986) described the gregarious diurnal roosting
behavior of a small noctuid moth from Parque Nacional Corcovado in
Costa Rica. This moth was an undescribed species of Ocalaria Schaus.
Adults of this species were subsequently captured near the Smithsonian
Tropical Research Institute on Barro Colorado Island, Panama, and sent
to me for identification. Comparison with illustrations of Corcovado
specimens showed that the Panamanian material was conspecific. The
species is described below.

Ocalaria cohabita, new species

(Figs. 1-5, 10, 14, 16, 17, 20, 21, 24, 26)

Diagnosis. Forewing underside discal eyespot with double pupil, smaller pupil sur-
rounded by iridescent deep-blue scales, lacking marginal blue-green scales; hindwing
underside with solid brown band between postmedian and subterminal lines; male antenna
serrate.

Male. Head. Haustellum unscaled; labial palp mainly dark gray-brown, first segment
with some off-white scales dorsally and lateroventrally, smooth-scaled laterally but scales
on dorsal and ventral edges projecting as a loose fringe, 2nd and 3rd segments uniformly
dark gray-brown, smooth-scaled on all surfaces, subequal in length, elongate, ascending
in life, at rest recurved dorsally over head (Greig & DeVries 1986:fig. 4), 2nd segment
gibbous; antenna long, almost equal to forewing length, dorsally uniformly scaled dark
gray-brown, ventrally serrate, sensory setae distinct, white; eyes large, bulbous, bare,
unlashed; frons narrow, unscaled on lower half except for extreme edges, scales on dorsal
half projecting anteriorly forming median ridge; frons, vertex, occiput dark gray-brown
except for band of off-white scales bordering eye posteriorly. Thorax. Patagia, thoracic
scaling concolorous with head, tegulae similar but with weak, transverse, median white
band. Legs. All uniformly dark gray-brown dorsally, off-white ventrally; apexes of tibial
and tarsal segments off-white, weakly so on forelegs, more strongly marked on hindlegs;
tibiae unspined. Forewing length 13.4-16.0 mm (holotype 16.0 mm). Wings (Figs. 1, 2).
Forewing ground color dark gray-brown, transverse bands pale gray, with weak purple

VOLUME 42, NUMBER 3 219

iridescence in oblique lighting; space on forewing enclosed by costa, discal cell, antemedian
line pale orange; eyespot black with off-center white pupil, surrounded by ring of pale
orange; apical spot black with basal small white spot; ventral pattern similar to upper but
more colorful; forewing pale gray, ventral of discal cell as far as postmedian line, eyespot
as upper surface but with additional smaller white pupil posterodistal to main pupil,
smaller pupil surrounded by deep blue iridescent scales visible only in oblique lighting,
area between postmedian and subterminal lines violet-gray with central brown area,
anterior part suffused with orange scales; pupil of apical eyespot much larger than on
upper surface, rounded or square, subequal to main pupil of discal eyespot; distally veins
R, and M,, together with costal area to apical eyespot, orange; hindwing ground color
pale gray, suffused costally with brown scales, discal lunule, antemedian, postmedian,
and subterminal lines dark gray-brown, distinct; brown band present between postmedian
and subterminal lines, bounded basally and distally by pale gray. Abdomen. Dorsally
and ventrally transversely striped, anterior half of each segment pale gray, posterior half
dark gray-brown. Genitalia (Fig. 3). Uncus with small apical hook; saccus ovoid; juxta
ill-defined; valve simple, lacking appendages, with a strong constriction on the costa
basally; aedeagus cylindrical, with bluntly-pointed apical process; vesica without cornuti.

Female. As described for male except antenna filiform, wings broader, more rounded,
not as acutely pointed (Figs. 1, 2). Ratio of forewing length to maximum width perpen-
dicular to costa averaging 2.28 in males (n = 12) and 2.11 in females (n = 11), the 0.17
difference being significant (P = 0.001, 1-tailed Mann-Whitney test because larger ratio
expected in males). Genitalia (Fig. 4). Segment 8 annular; antrum membranous, undif-
ferentiated from ductus bursae; corpus bursae membranous, ovoid, lacking signa; ductus
seminalis broader than ductus bursae, arising from posterior end of corpus bursae.

Types. Holotype male: Panama, Barro Colorado Island, 20 May 1986, N. Greig. Para-
types: 1 female: Panama, Barro Colorado Island, 28 May 1986, P. J. DeVries (BMNH
noctuid slide #12816); 11 males, 11 females: Panama, Barro Colorado Island, 18 August
1986, P. J. DeVries (BMNH noctuid slides #12803-12815). In British Museum (Natural
History).

Life history. Nothing is known about the immature stages or larval hosts of any species
of Ocalaria.

Variation. There is no major variation in wing pattern either between or within sexes.
The pale orange forewing costal band may be weakly subdivided medially by ground
color scales. However, older moths become worn and faded, eventually presenting a
“washed-out” pale coloration. This phenomenon affects wings and body scales equally
and appears due to a combination of scale loss and fading of brown pigments.

Key to Adults of Ocalaria

Ocalaria currently contains 7 described species (provenances and
numbers examined are those of specimens held in BMNH): dioptica
(Walker) (=macrops (Felder & Rogenhofer)) (Brazil: Amazonas, n =
2; Peru: Amazonas, n = 1; Bolivia, n = 2; French Guiana, n = 1),
guarana Schaus (Brazil: Rio de Janeiro, n = 2; Sao Paulo, n = 1), oculata
(Druce) (Guatemala: San Isidro, n = 56), pavina Schaus (Brazil: Rio de
Janeiro, n = 9), pavo Schaus (Costa Rica: Sixola), quadriocellata (Walk-
er) (Brazil: Cuiaba, n = 11) and cohabita,; new species.

The couplet for O. pavo is based on a small water-color painting in
BMNH, which lacks specimens of this species. This type of painting,
probably commissioned by G. F. Hampson, has been found to be re-
alistic and reliable as an identification guide (A. Watson pers. comm.).

220 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 1, 2. Ocalaria cohabita. 1, Holotype male; 2, Paratype female.

1. Discal eyespot on forewing underside with single white pupil (occasionally double
in O. quadriocellata, not surrounded by deep blue iridescence), marginal iri-
descent blue-green scales; hindwing underside without solid brown band be-
tween postmedian and subterminal lines; male antenna bipectinate 2
Discal eyespot on forewing underside with double pupil, smaller pupil surrounded
by iridescent deep blue scales, lacking marginal blue-green iridescence; hind-
wing underside with solid brown band between postmedian and subterminal

lines: amalevantennarsenrate sass see eee ane ne cohabita, new species
2. Area between postmedian and subterminal lines on upperside of both wings solid
dark brown, edged with continuous pale cream lines pavo Schaus

Area between postmedian, subterminal lines on upperside of both wings not solid
dark brown, pale line bordering inner edge of subterminal line interrupted

a mnecmecmccrmcccenrancnin ts, 3

3. Discal eyespot on forewing upperside subequal to apical eyespot; brown streak
present in discal fold between median and postmedian lines ............. pavina Schaus

Discal eyespot on forewing upperside much larger than apical eyespot; brown
streak in discal fold between median, postmedian lines absent cece ee 4

4. Forewing upperside, distal to discal eyespot, with conspicuous white spot often
divided in half by brown scaling along vein Mg .......-.- quadriocellata Walker
Forewing upperside without such spot 0 eee 5

5. Forewing underside with white spot or streak below apical eyespot; pupil of apical
eyespot small, no more than half width of eyespot in diameter, surrounding
black-seales conspicuous: uso) 0. ee 6

Forewing underside without white spot or streak below apical eyespot; pupil of
apical eyespot large, subequal in diameter to width of eyespot, almost obliterating
surroanding ring) of ‘black scales. en) mann oculata Druce

6. Ground color dark brown; forewing upperside postmedian line cream, continuous
across wing; discal eyespot on forewing underside without diagonal cream line
below 2h eg dy den doe a ee guarana Schaus

Ground color off-white suffused with pale brown; forewing upperside postmedian
line indistinct, not continuous across wing; discal eyespot on forewing underside
with diagonal cream line below, originating from posterobasal corner of the
eyespot i.) a ce ae ee eee et dioptica Walker

Comparative Morphology of O. cohabita,
O. oculata, and O. quadriocellata

In a previous cladistic analysis of Plusiinae (Kitching 1987), I de-
scribed several characters useful in elucidating generic and higher level

VOLUME 42, NUMBER 3 Pig |

Fic. 8. Ocalaria cohabita, paratype male genitalia, BMNH noctuid slide #12804
(genitalia) and #12805 (aedeagus). Scale line = 1 mm.

interrelations on the proboscis, female frenulum, basal abdominal seg-
ments and abdominal segment 8. I have examined these structures in
three Ocalaria (O. cohabita, O. oculata and O. quadriocellata) and
here describe the results to facilitate future analyses of higher classi-
fication of Catocalinae. Ocalaria oculata and O. quadriocellata were
chosen for detailed examination because they are well represented in
the BMNH collection; the remaining species (several undescribed) were
not because it was not my intention to revise the genus. Comparisons
are also drawn, where appropriate, with equivalent conditions in Plu-
siinae and the taxa employed as outgroups in that study.

Labial palps. In all three Ocalaria species, the dorsal margin of
segment 2 of the labial palp is markedly convex. This feature is more
conspicuous in O. quadriocellata (Fig. 6) and O. oculata than in O.
cohabita (Fig. 5) but there is no sexual dimorphism. The function of
the bulge is unknown; examination at 400 x revealed no obvious sensory
structures or differential scale arrangements.

Antennae. Female O. cohabita have filiform antennae (Fig. 16) with
relatively short subventral setae; males have longer, more conspicuous
subventral setae, while the ventral surface of each segment is produced
into a rounded triangular flange (Fig. 14), giving a serrate appearance

222 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fic. 4. Ocalaria cohabita, paratype female genitalia, BMNH noctuid slide #12816.
Scale line = 1 mm.

to the antenna. Apically, this flange bears two setae and a sensillum
styloconicum.

Antennae of female O. quadriocellata are also filiform, although the
segments are longer than in O. cohabita. In contrast, male antennae of
O. quadriocellata are strongly bipectinate (Fig. 12). Each pair of pec-
tinations, which arise at the base of a segment, are long, slender, parallel-
sided and bear a strong apical seta. Male O. oculata are similar but the
pectinations are even longer and thinner (Fig. 15). Female O. oculata
are unique among the three taxa studied in also having bipectinate

VOLUME 42, NUMBER 3 Peper:

Fics. 5-11. Ocalaria structures. 5, 6. Labial palps. 5, O. cohabita male; 6, O. oculata
female. 7-9. Foretibiae. 7, O. oculata female; 8, O. quadriocellata female; 9, O. oculata
male. 10, 11. Male sternite 8. 10, O. cohabita; 11, O. quadriocellata. Stippling indicates
extent of hairpencil scale insertions. Scale lines = 1 mm.

224 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

antennae (Fig. 13), although the branches are shorter than in either
male O. oculata or O. quadriocellata.

Proboscis. Proboscides in all three species are short, stout, and similar
in general appearance to those of such genera as Oncocnemis Lederer
(Cuculliinae), Magusa Walker (Amphipyrinae), Stictoptera Guenée
(Stictopterinae) and Paectes Hubner (Euteliinae). Ocalaria also shares
with these and other genera a nodular apex to the proboscis and ridged
styloconic sensilla (which appear stellate in apical view), all of which
suggests that this form of proboscis is plesiomorphic within Noctuidae.
Concomitantly, the form of proboscis found in Plusiinae is apomorphic
for that subfamily, although certain features appear to have been con-
vergently derived in Cucullia Schrank and Calophasia Stephens (both
Cuculliinae).

Epiphysis. The epiphysis in Ocalaria exhibits considerable variation
in length. In both sexes of O. cohabita and female O. quadriocellata,
it is small, being only about a quarter the length of the fore-tibia (Fig.
8). However, in male O. quadriocellata and O. oculata, the epiphysis
is highly elongate and often exceeds the fore-tibial apex (Fig. 9). Female
O. oculata, which have smaller antennal pectinations, also have a shorter
epiphysis, but one that is still elongate compared to female O. cohabita
(Fig. 7). This close correlation between epiphysis length and degree of
development of antennal pectinations is strong circumstantial evidence
to support the hypothesis that the primary role of the epiphysis is
keeping the antenna clean. A long epiphysis is necessary in male O.
quadriocellata and O. oculata to clean the long pectinations efficiently.

Wings. Venation of Ocalaria is typical quadrifine noctuid, although
hindwing vein M, is somewhat weaker than either M, or M, and does
not arise close to the base of M;. A potential Ocalaria apomorphy in
hindwing shape is the shallow concavity just beyond the midpoint of
the costal margin, although this is weakly expressed in O. oculata. All
Ocalaria examined have a trisetose female frenulum in which the setae
are subequal in length, further corroborating the hypothesis that this
state is plesiomorphic within Noctuidae (Kitching 1987).

Abdominal segment 2. In all noctuid genera examined so far, the
anterior edge of tergite 2 (T2) bears an inflected flange. At its simplest,
the flange is concave, uniformly narrow and difficult to discern in slide-
mounted material, as in Stiriinae, Cuculliinae, Heliothinae, and basal
plusiine tribe Omorphinini. The more derived Abrostolini display two
subdorsal lobes directed medially. These fuse in Argyrogrammatini and
Plusiini but leave a central rounded emargination. The ventral edge of
the flange in Stictopterinae and Euteliinae is straight, with a median
dorsal triangular inflection of T2 itself.

In general, the form of the T2 flange is highly invariant in large

VOLUME 42, NUMBER 3 225

5 | 16

Fics. 12-16. Ocalaria antennae. 12, O. quadriocellata male; 13, O. oculata female;
14, O. cohabita male, lateral view; 15, O. oculata male; 16, O. cohabita female, lateral
view. Scale line = 1 mm.

226 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

_——_—_—_— SS eee
a a
17 18 19

Fics. 17-19. Male tergite 8. 17, O. cohabita; 18, O. oculata; 19, O. quadriocellata.
Posterior edge uppermost, with transverse line marking posterior margin of abdominal
segment 8. Scale line = 1 mm.

taxonomic groups, such as tribes or subfamilies. However, in Ocalaria,
the T2 flange is remarkably variable between species. The putatively
basal O. cohabita shows a sclerotized dorsal inflection of T2 similar to
that in Stictopterinae and Euteliinae, but with the median emargination
typical of “higher” plusiines. In addition, the ventral edge of the flange
is somewhat cut back laterally. This flange is sexually dimorphic in O.
cohabita, a feature almost certainly correlated with the dimorphism
observed in sternite 2 (St2). The female (Fig. 21) differs from the male
(Fig. 20) in that the median emargination is not parallel-sided, while
the dorsal inflection is much broader and has a straight ventral edge.

Ocalaria oculata and O. quadriocellata differ markedly in lacking
the sclerotized dorsal inflection and in the extreme specialization of the
flange. Ocalaria quadriocellata has a T2 flange formed of two broad,
ventrally-directed, well-separated rounded lobes (Fig. 22). This trend
is more noticeable in O. oculata, in which the lobes are long, narrow,
and closer to the lateral edges of T2 than to the center (Fig. 28).

St2 in O. oculata and O. quadriocellata is typically noctuid in form,
with no marked diagnostic features. It is a roughly square sclerite (Fig.
25), with convex lateral and posterior edges. The anterolateral corners
are produced into a pair of apodemes, from which a sclerotized bar
arises laterally. This passes anteriorly to the counter-tympanal hood.
The anterior margin of St2 is broadly U-shaped.

Females of O. cohabita have a broadly similar St2, but it is slightly
squarer, with marginally concave lateral edges, and the anterior margin
is more V-shaped (Fig. 24). St2 of male O. cohabita, however, is mark-
edly different (Fig. 26), which may be the first record of sexual di-
morphism in this structure in quadrifine noctuids. The whole sclerite

VOLUME 42, NUMBER 3 Hie

20

21

22

23

Fics. 20-23. Tergite 2 flange. 20, O. cohabita male; 21, O. cohabita female; 22, O.
quadriocellata male; 23, O. oculata male. Scale line = 1 mm.

228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

is elongate, with distinctly concave lateral margins. The anterior edge
is V-shaped, while the posterior margin is produced as a broad, trian-
gular point. Posterolaterally are two secondary sclerites in the inter-
segmental membrane between St2 and St8. In addition, the anterior
corners of St3 (Fig. 26) are produced around the posterolateral edges
of these secondary sclerites, a feature similar to that seen in numerous
trifine species, where it is associated with the lever of male basal ab-
dominal hair pencils (Birch 1970). The posterior margin of St3 is also
produced to a median point, but is more obtuse than that of St2. Finally,
on the anterior edge of St4 (Fig. 26) are two concavities that appear to
house shallow glandular pockets. The function of this complex in male
O. cohabita is a matter for conjecture but suggests that adult biology,
particularly courtship, of this species might be usefully studied.

Male abdominal segment 8. The form of the male eighth tergite
(T8) and sternite (St8) displays species-level diagnostic features in a
number of noctuid subfamilies; for example, Plusiinae (Dufay 1970),
Stiriinae (Hogue 1963), Heliothinae (M. J. Matthews pers. comm.). In
addition, these sclerites have yielded characters useful in recognizing
suprageneric taxa in Plusiinae (Kitching 1987).

The form of T8 in Ocalaria is characteristic, consisting of a median
sclerotized longitudinal bar tapered posteriorly and expanded ante-
riorly. The three species differ in minor details: in O. cohabita, the bar
does not reach the posterior margin of the segment and the anterior
expansion has concave margins and a truncate apex (Fig. 17); in O.
oculata, the anterior expansion is drawn out laterally into narrow points
(Fig. 18); while in O. quadriocellata, the anterior expansion has a
rounded apex (Fig. 19). In all three taxa, there appear to be two shallow
pockets associated with the lateral edges of T8 anteriorly.

A8 bears a median weak tuft of hairs, barely differentiated into a
pair of hair pencils, arising from a shallow, membranous, ventral pocket.
St8 forms a thin sclerotized bar anterior to this pocket, with two concave
bars running longitudinally on either side of the hair tuft. The antero-
lateral corners of St8 are produced as blunt triangular lobes. The form
of St8 is similar in all three species, but whereas the anterior bar forms
a blunt median point in O. oculata and O. quadriocellata (Fig. 11), in
O. cohabita, it is broadly rounded with a somewhat straight central
section (Fig. 10).

Male genitalia. Ocalaria oculata is very similar to O. cohabita (Fig.
3) except that the valve is of more uniform width basally, the saccus
is acutely pointed and the aedeagal process is absent. In addition, the
spines on the vesica basally are larger and more strongly sclerotized.
Genitalia of O. quadriocellata are similar to O. oculata but the sacculus

VOLUME 42, NUMBER 3 229

Fics. 24-26. Ocalaria sternites. 24, 25. Sternite 2. 24, O. cohabita female; 25, O.
quadriocellata male; 26, Sternites 2-4 O. cohabita male. Scale line = 1 mm.

bears a small rounded lobe basally and the saccus forms a less acute
point.

Female genitalia. The posterior margin of St8 bears a fringe of
persistent, brown, setose scales that may be apomorphic for Ocalaria.
Ocalaria oculata is essentially similar to O. cohabita (Fig. 4) but differs
in that the corpus bursae is not differentiated from the ductus bursae,
being merely a long, slightly broader membranous sac; the ductus bursae
is broader with a median annulus of longitudinal ridges and sclerotized
granulations; and the ostium bursae is adorned with sclerotized gran-
ulations. The anal papillae are diagonally cut back dorsally to the origin
of the posterior apodemes. Ocalaria quadriocellata is broadly similar

230 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

to O. cohabita, except that the dorsoposterior parts of the anal papillae
are drawn out into blunt, slightly downcurved points.

ACKNOWLEDGMENTS

This study was carried out partly during the tenure of a Junior Research Fellowship
awarded by the Trustees of the British Museum (Natural History), which is gratefully
acknowledged. I thank my colleagues at BMNH for support and comments, and the staff
of the BMNH Photographic Unit for the photographs. Special thanks go to Nancy Greig
and Philip DeVries for the opportunity to describe this species.

LITERATURE CITED

BircH, M. C. 1970. Structure and function of the pheromone-producing brush-organs
in males of Phlogophora meticulosa (L.) (Lepidoptera: Noctuidae). Trans. Roy. Ento-
mol. Soc. Lond. 122:277-292.

DuFay, C. 1970. Insectes Lépidoptéres Noctuidae Plusiinae. Faune Madagascar 31:1-
198.

GrEIG, N. & P. J. DEVRIES. 1986. Observations on the diurnal gregarious roosting of
Ocalaria sp. (Noctuidae) in Costa Rica. J. Lepid. Soc. 40:124—-126.

Hocug, C. L. 1968. A definition and classification of the tribe Stiriini (Lepidoptera:
Noctuidae). Contr. Sci. Los Angeles 64:1—129.

KITCHING, I. J. 1987. Spectacles and Silver Ys: A synthesis of the systematics, cladistics
and biology of the Plusiinae (Lepidoptera: Noctuidae). Bull. Brit. Mus. Nat. Hist.
(Entomol.) 54:75-261.

Received for publication 4 November 1987; accepted 6 April 1988.

Journal of the Lepidopterists’ Society
42(3), 1988, 230

ADDITIONAL MANUSCRIPT REVIEWERS, 1987

. The following persons from whom the editor received manuscript reviews in 1987 were
inadvertently omitted from the manuscript reviewer list published in the May issue (42:

152-153). The Journal acknowledges with gratitude the contributions of all manuscript
reviewers.

Paul H. Arnaud, San Francisco, CA

Richard A. Arnold, Pleasant Hill, CA

Howard D. Baggett, Tampa, FL

G. G. Grant, Sault Ste. Marie, ON, Canada
Daniel H. Janzen, Philadelphia, PA

Ian J. Kitching, London, UK

Rudolf E. J. Lampe, Niirnberg, West Germany
Dorothy Pashley, Baton Rouge, LA

Austin B. Platt, Catonsville, MD

J. R. G. Turner, Leeds, UK

William E. Miller, Editor

Journal of the Lepidopterists’ Society
42(3), 1988, 231-235

A NEW SESIA CLEARWING MOTH FROM MICHIGAN
(SESIIDAE)

THOMAS D. EICHLIN

Insect Taxonomy Laboratory, A&I, Division of Plant Industry,
California Department of Food and Agriculture,
Sacramento, California 95814

AND

WILLIAM H. TAFT

Adjunct Assistant Curator, Department of Entomology,
Michigan State University, East Lansing, Michigan 48824

ABSTRACT. Sesia spartani, new species was discovered in Michigan (30 males) in
traps baited with a sex attractant consisting of 50:50 mixture of (3,13) Z,Z-ODDOH/
(3,13) E,Z-ODDOH. The new species is described, illustrated, and compared with S.
tibialis (Harris). The two differ in structure of male antennae, genital morphology, re-
sponse to sex attractants, and seasonal occurrence.

Additional key words: Sesia spartani, S. tibialis, attractants.

Use of synthetic sex attractants has resulted in the discovery of several
new species of sesiids in the Western Hemisphere (Duckworth & Eichlin
1977a, 1977b, Greenfield & Karandinos 1979, Brown et al. 1985, Eichlin
1986, 1987).

During studies employing sex attractant baits to survey the Sesiidae
of Michigan, a new species closely related to Sesia tibialis (Harris) was
discovered and is described below. Males of S. tibialis (Fig. 2) are known
to be very responsive to the Z,Z isomer of 3,13-octadecadien-1-ol acetate
(Z,Z-ODDA) (Duckworth & Ejichlin 1978:28), a major component of
many clearwing moth pheromone systems (elucidated by Tumlinson
et al. 1974). In Saskatchewan, Underhill et al. (1978) found the best
attractant was probably a 80:20 blend of Z,Z-ODDA/Z,Z-ODDOH.
The new species was discovered in the Lower Peninsula of Michigan
when males were captured in traps baited with sex attractants consisting
of a 50:50 mixture of the Z,Z and E,Z alcohols (Z,Z-ODDOH/E,Z-
ODDORH). Traps baited with other sex attractants including that known
to be effective for S. tibialis were deployed in the same areas throughout
the collecting season but failed to capture any S. spartani.

Sesia Fabricius

Genus Sesia is characterized by the following: Head with haustellum reduced, about
% length of labial palpus; antenna strongly clavate, ventrally ciliate-unipectinate on male;
horizontal flat plate of scales projecting somewhat over middle of eye. Forewing vein R,
terminating at apex, R, below. Hindwing veins M, and Cu, joined at corner of cell or
very short-stalked. Genitalia unique, generally as shown in Figs. 5 and 6.

932 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fics. 1, 2. Adult males of Sesia spp. 1, S. spartani, Shiawassee Co., Michigan (ho-
lotype); 2, S. tibialis, Isabella Co., Michigan.

Sesia spartani, new species

Male (Fig. 1). Head with vertex brown-black, some white posteriorly; front brown-
black; occipital fringe white dorsally, yellow laterally; labial palpus roughened with long
hairlike scales ventrally on basal segment, yellow with brown toward base; haustellum
short, less than % length of labial palpus; antenna brown-black with yellow-orange at tip,
unipectinate, individual middle segments with ramus about 8 times as long as the distance
between 2 adjacent rami (dorsoapical view of antenna), produced from dorsoapical portion
of segment (Fig. 3). Thorax brown-black with yellow behind collar, around wing bases,
subdorsally on posterior half of mesothorax, and variously on metathorax. Abdomen
brown-black but with broad yellow bands on segments 5, 6, and 7, less so on 4 dorsally;
tip of abdomen mostly yellow with anal tuft poorly defined. Wings hyaline; narrow
margins and discal spot brown. Wing length 8-9 mm (30 n). Legs with coxae brown-
black edged with yellow; femora yellow dorsally, brown-black ventrally; tibiae yellow,
hind tibia roughened dorsally by semierect, thin, elongate scales; tarsi yellow-orange.
Genitalia as in Fig. 6.

Female. Unknown.

Host. Probably species of Salicaceae, particularly Populus tremuloides Micheaux (quak-
ing aspen), based on observed larval damage to trees at the type locality and at other
sites where S. spartani was collected.

Distribution. Sesia spartani has been collected in the Lower Peninsula of Michigan in
Clinton, Shiawassee, and Lake counties.

Types. Holotype: Male, MICHIGAN: Shiawassee Co., Bath, 13 June 1987, Coll. William
H. Taft; Rose Lake Wildlife Research Area, T5N RIE Sec 20; ZZOH/EZOH, 50:50;
deposited in Entomology Museum, Michigan State University, East Lansing (MSU). Para-
types (29 males): 6, Shiawassee Co.: V-29-1987 (1); VI-9-1987 (2); T5N RIE Sec 20, 13
June 1987 (2); same as last except 14 June 1987 (1). 22, Clinton Co.: T5N R2W Sec 31,
13 June 1987 (3); same as last except 14 June 1987 (2); 16 June 1987 (4); 17 June 1987
(7); 19 June 1987 (2); 20 June 1987 (4). 1, Lake Co.: T17N R14W Sec 12, 14 June 1987
(all collected by William H. Taft using traps baited with ZZOH/EZOH 50:50).

Paratypes are deposited in MSU; U.S. National Museum of Natural History, Washington,
D.C.; California Department of Food and Agriculture, Sacramento; Canadian National
Collection, Ottawa, Ontario; and Field Museum, Chicago, Illinois.

Discussion. Sesia spartani is superficially similar to S. tibialis. However, male genitalia

VOLUME 42, NUMBER 3 233

Fics. 3, 4. Segments from near middle of left antennae of Sesia spp. as viewed
ventrally on proximal surface by scanning electron microscope. 3, S. spartani; 4, S.
tibialis.

differ considerably (Figs. 5, 6): S. spartani uncus deeply and acutely divided, cleft of S.
tibialis uncus less deep, broadly rounded; saccus about % length of valve, only % on S.
tibialis; valve more produced ventroposteriorly and with more thick, dark spines near
center than on S. tibialis; gnathos narrowing apically and of different form than for S.
tibialis; and aedeagus with jagged plate posteriorly, S. tibialis lacking jagged plate. Some
specimens of S. tibialis from Michigan (Fig. 2) are nearly lacking yellow dorsally on
abdominal segments 4 and 5, while on S. spartani segment 5 is mostly yellow, and 4 has
some yellow powdering. Sesia tibialis from elsewhere usually has yellow banding on all
segments.

The collecting sites were low muck soil depressions in scattered locations. These habitats
are characterized by large stands of regrowth quaking aspen mixed with willows (Salix
spp.), elm (Ulmus sp.), red maple (Acer rubrum L.), and black cherry (Prunus serotina
Ehrh.). The undergrowth is dogwood (Cornus sp.), viburnum (Viburnum sp.), and blue-
berries (Vaccinium sp.).

234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

6

Fics. 5, 6. Male genitalia of Sesia spp. viewed ventrally, left valve removed. 5, S.
tibialis (from Duckworth & Eichlin 1978); 6, S. spartani.

The S. spartani males were captured in Multi-pher #1® plastic pheromone traps.
Collecting dates for S. spartani were 29 May-20 June 1987. Paranthrene dollii (Neu-
moegen) was collected with S. spartani during late May and early June. At the time of
capture, the growing season was 250-300 degree-days (base 50) above normal; conse-
quently, in normal years S. spartani may fly later in June or in early July. It appears to
fly two weeks to a month earlier than does S. tibialis in Michigan. Sesia tibialis has not
been found in counties where S. spartani originated, but has been collected as far south
as Newaygo, Isabella, and Midland counties, and is known from Nova Scotia and New
England to British Columbia, and from the Rocky Mountains to the Pacific Coast. The
Michigan habitats for both species appear to be similar.

This species is named for the Spartans, a nickname applied to Michigan State University
athletic teams.

ACKNOWLEDGMENTS

We thank J. W. Snow and K. Scarborough, Southeastern Fruit and Tree Nut Research
laboratory, USDA, Byron, Georgia, for generously providing pheromones and technical
assistance; C. S. Papp, Sierra Graphics and Typography, Sacramento, California, for artistic
help; and T. Montenegro, Insect Taxonomy Laboratory, State Department of Food and
Agriculture, Sacramento, California, for technical assistance. We especially thank the
Michigan State University Entomology Department and Museum, East Lansing, for fiscal
assistance and encouragement, also J. M. Scriber, T. Ellis, D. Smitley, and S. Dutcher;

VOLUME 42, NUMBER 3 Ze

and for reviewing the manuscript, R. Fischer, F. Stehr, Michigan State University, M. D.
Greenfield, Department of Biology, University of California, Los Angeles, and J. B.
Heppner, Florida State Collection of Arthropods, Gainesville.

This paper is Michigan Agricultural Experiment Station Journal Article No. 12496.

LITERATURE CITED

Brown, L. N., T. D. EICHLIN & J. W. SNow. 1985. A new species of clearwing moth,
Carmenta laurelae (Sesiidae), from Florida. J. Lepid. Soc. 39:262-265.

DuckworTH, W. D. & T. D. EICHLIN. 1977a. Two new species of clearwing moths
(Sesiidae) from eastern North America clarified by sex pheromones. J. Lepid. Soc.
31:191-196.

1977b. A new species of clearwing moth from south-central Texas (Lepidoptera:

Sesiidae). Pan-Pac. Entomol. 53:175-178.

1978. The clearwing moths of California (Lepidoptera: Sesiidae). Calif. Dept.
Food & Agric., Occas. Pap. Entomol. 27:1-80.

EICHLIN, T. D. 1986. Western Hemisphere clearwing moths of the subfamily Tinthiinae
(Lepidoptera: Sesiidae). Entomography 4:315-378.

1987. Three new Western Hemisphere clearwing moths (Lepidoptera: Sesiidae:
Sesiinae). Entomography 5:531-540.

GREENFIELD, M. D. & M. G. KARANDINOS. 1979. A new species of Paranthrene (Lep-
idoptera: Sesiidae). Proc. Entomol. Soc. Wash. 81:499-504.

TUMLINSON, J. H., C. E. YONCE, R. E. DOOLITTLE, R. R. HEATH, C. R. GENTRY & E.
R. MITCHELL. 1974. Sex pheromones and reproductive isolation of the lesser peach
tree and the peach tree borer. Science 185:614-616.

E. W. UNDERHILL, W. STECK, M. D. CHISHOLM, H. A. WORDEN & J. A. G. HOWE. 1978.
A sex attractant for the cottonwood crown borer, Aegeria tibialis (Lepidoptera:
Sesiidae). Can. Entomol. 110:495-—498.

Received for publication 23 December 1987; accepted 14 April 1988.

Journal of the Lepidopterists’ Society
42(3), 1988, 236-239

TWO NEW SPECIES OF RHYACIONIA PINE MOTHS
FROM MEXICO (TORTRICIDAE: OLETHREUTINAE)

WILLIAM E. MILLER

Department of Entomology, University of Minnesota,
St. Paul, Minnesota 55108

ABSTRACT. Rhyacionia cibriani is described from five males and five females, and
R. rubigifasciola from one male and one female. The former is differentiated from R.
jenningsi Powell by genital and other characters including longer antennal pecten. The
latter is differentiated from all congeners, none of which it resembles closely, by genital
characters including a ridge separating sacculus and valval neck in the male, and an
anally emarginate sterigma in the female. Pinus hartwegii Lindl. and P. oocarpa Schiede
are the respective hosts, the larvae boring in branchlets. The new species bring the number
of Rhyacionia known in Mexico and the Neotropics to nine species.

Additional key words: taxonomy, Eucosmini, Rhyacionia cibriani, R. rubigifasciola,
Neotropics.

Pines, the larval hosts of Rhyacionia, are numerous in Mexico, 21 of
the 30 pine species occurring there being absent in the U.S. (Critchfield
& Little 1966). The 2 new species described here bring the number of
Rhyacionia known in Mexico and the Neotropics to 9 species (Powell
& Razowski in press), and the number described worldwide to 34 (Miller
1985, Obraztsov 1964, Powell & Miller 1978). This is the fourth paper
in a series in which I describe new Neotropical olethreutines in various
genera whose hosts and modes of feeding make them of economic
interest or importance (Miller 1966, 1986, 1987).

In both species described here, hindwing vein M, is bent at its base,
and hindwing veins M, and CuA are either stalked or connate and
approximate toward their bases. These character states place the species
in Eucosmini (Obraztsov 1958). Features that place them in Rhyacionia
are italicized in descriptions. Venation was ascertained under a stereo-
microscope from temporary preparations made by touching xylol to
wings while light passed through them (Zimmerman 1978).

Rhyacionia cibriani, new species

(Figs. 1-3)

Male. Forewing length 11.0 to 12.0 mm (holotype 11.0 mm) (5n). Head. Labial palpus
clothed with brown-banded white scales, length of 2nd segment 2x eye diam., length of
3rd segment % that of 2nd; vestiture of vertex similar to that of labial palpus; antennal
pecten length 1 to 1% flagellar length, 1% to 2x flagellar diam. Thorax. Dorsal vestiture
beige, ventral paler; front and middle leg scaling similar to that of labial palpus, hind
leg paler, tarsi indistinctly white-banded; forewing with veins M,and M, connate, termen
straight or convex, costal fold absent, upper side yellowish to coppery red, tinged with
lavender in cell area, crossed by irregular striae near middle (Fig. 1), underside pale
grayish yellow; hindwing upper side gray, underside paler than forewing underside.
Genitalia (Fig. 2) (3n). Valva lacking costal hook, a ridge from sacculus to neck terminating
in a nipplelike process at mid-neck, neck constricted to nearly % maximum sacculus

VOLUME 42, NUMBER 3 237

Fics. 1-6. 1-8. Rhyacionia cibriani from type locality. 1, Wings of paratype; 2, Male
genitalia of holotype; 3, Female genitalia of paratype. 4-6. R. rubigifasciola from type
locality. 4, Wings of holotype; 5, Male genitalia of holotype; 6, Female genitalia of
paratype. Additional information keyed to figure numbers appears in Type Data section.
Some negatives reversed.

238 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

width, pollex present and its length about 4% maximum cucullus width; uncus and socii
rudimentary or absent; aedeagus curved and tapering toward apex, vesica with 3 to 4
cornuti. ;

Female. Forewing length 9.5 to 11.0 mm (5n). Similar exteriorly to male except for
shorter antennal pecten. Genitalia (Fig. 3) (3n). Sternum 7 emarginate; sterigma nearly
square in outline, laterally inflected, with a broad and evenly rounded longitudinal ridge;
ductus bursae sclerotized only near ostium bursae; corpus bursae with 1 thornlike signum,
sometimes a tiny 2nd one.

Type data. Holotype male, Paso de Cortez, Méx., Mexico, 12 March 1984, No. 1133,
Pinus hartwegii Lindl., D. Cibrian, genit. prep. WEM 1910844 (Fig. 2), in U.S. National
Museum of Natural History, Washington, D.C. Four paratype males, same data as holotype
except 5-9 April 1984, 2 genit. preps. WEM 910842 and WEM 84885; 5 paratype females,
same data as holotype except 5-16 April 1984 (Fig. 1), 3 genit. preps. WEM 910843 (Fig.
3), WEM 2210841, and WEM 53882, in U.S. National Museum of Natural History; Essig
Museum, University of California, Berkeley; University of Minnesota, St. Paul; and Lab.
de Entomologia Forestal, Universidad Autonoma Chapingo, Chapingo, Mexico.

Discussion. Rhyacionia cibriani most resembles R. jenningsi Powell, but differs in size,
structure, and forewing pattern as follows. Rhyacionia cibriani has a 40% greater average
forewing length, 100% greater relative length of 2nd palpus segment, and 350 to 400%
longer relative antennal pecten length than R. jenningsi; the lavender hue of the R.
cibriani forewing cell is lacking in R. jenningsi; the nipplelike process on the male valva
in R. cibriani is lacking in R. jenningsi; the 7th female sternum is more deeply emarginate
in R. cibriani and the sterigma more square than in R. jenningsi. The foregoing character
states for R. jenningsi are documented in Powell and Miller (1978).

Pinus hartwegii is classified in Ponderosae (Critchfield & Little 1966), a Pinus sub-
section whose members are hosts to several Rhyacionia species (Powell & Miller 1978).

The species is named for David Cibridn-Tovar, who reared adults from larvae boring
in Pinus hartwegii branchlets.

Rhyacionia rubigifasciola, new species

(Figs. 4-6)

Male. Forewing length 8.5 mm (ln). Head. Labial palpus clothed with silvery white
scales sometimes tinged with orange or gray, length of 2nd segment 14x eye diam.,
length of 3rd segment % that of 2nd; vestiture of vertex silvery white except for orange
near antennal bases; antennal pecten length 0.8% flagellar length, 0.8x flagellar diam.
Thorax. Dorsal vestiture similar to vertex; front and middle legs orange, banded with
white, hind leg paler except for tarsi; forewing with veins M, and M, connate, termen
convex, costal fold absent, upper side with 4 orange spindle-shaped fasciae extending
from costa to dorsum, 2 less tapered ones from costa to termen, all separated by silvery
white (Fig. 4), underside pale gray; hindwing upper side gray, underside paler than
forewing underside. Genitalia (Fig. 5) (In). Valva lacking costal hook, sacculus separated
from neck by a ridge, neck scarcely constricted dorsoventrally, concave anally, pollex
present but not well defined in outline; uncus absent; socii tiny, inflected, nearly obscured
by tergum, aedeagus apically expanded, forked, with several tiny apical spurs; vesica
with 6 cornuti.

Female. Forewing length 8.5 mm (ln). Similar exteriorly to male. Genitalia (Fig. 6)
(In). Sternum 7 not markedly emarginate; sterigma rounded in outline, emarginate on
anal margin, lamella antevaginalis scoop-shaped; ductus bursae sclerotized in an incom-
plete ring for a short distance at % its length from ostium bursae; corpus bursae with 2
thornlike signa.

Type data. Holotype male, Sta. Lucia, Sinaloa, Mexico, 1 July 1981 (Fig. 4), No. 802,
Pinus oocarpa Schiede, D. Cibrian & T. Méndez, genit. prep. WEM 108842 (Fig. 5), in
Essig Museum, University of California, Berkeley. One paratype female, same data and
depository as holotype except genit. prep. WEM 234851 (Fig. 6).

Discussion. Rhyacionia ribigifasciola does not clearly resemble any congener (Miller
1985, Obraztsov 1964, Powell & Miller 1978). It differs from all in male valval outline

VOLUME 42, NUMBER 3 239

and in the ridge separating sacculus and neck; also in shape of the female sterigma with
its anal emargination. The larvae bore in Pinus oocarpa branchlets.

Pinus oocarpa is classified in Oocarpae (Critchfield & Little 1966), a Pinus subsection
whose members are hosts to only one other Rhyacionia species, R. pasadenana (Kearfott)
(Powell & Miller 1978). The new species does not appear closer morphologically to R.
pasadenana than to other Rhyacionia species, however.

ACKNOWLEDGMENTS

I thank J. A. Powell and H. O. Yates III for specimen loans and other assistance; and
D. C. Ferguson for serving as special Journal editor for this paper.

LITERATURE CITED

CRITCHFIELD, W. B. & E. L. LITTLE. 1966. Geographic distribution of the pines of the
world. U.S. Dept. Agr. Misc. Publ. 991, 97 pp.

MILLER, W. E. 1966. A new species of moth destructive to pine cones in Mexico
(Tortricoidea). J. Lepid. Soc. 20:251-258.

1985. Nearctic Rhyacionia pine tip moths: A revised identity and a new species

(Lepidoptera: Tortricidae). Great Lakes Entomol. 18:119-122.

1986. New species of the genus Cydia that attack seeds of Mexican conifers

(Lepidoptera: Tortricidae), pp. 5-7. In Cibrian-Tovar, D., B. H. Ebel, H. O. Yates,

and J. T. Méndez-Montiel (eds.), Insectos de conos y semillas de las coniferas de

Mexico/Cone and seed insects of the Mexican conifers. U.S. Dept. Agr. Gen. Tech.

Rep. SE-40, 110 pp.

1987. A new species of Gretchena (Tortricidae) injurious to planted Neotropical
walnut. J. Lepid. Soc. 41:151-158.

OprRAZTSOV, N.S. 1958. Die Gattungen der palaearktischen Tortricidae II. Die Unter-
familie Olethreutinae. Tijdschr. Entomol. 101:229-261.

1964. Die Gattungen der palaearktischen Tortricidae II. Die Unterfamilie Ole-
threutinae 5. Tijdschr. Entomol. 107:1-48.

POWELL, J. A. & W. E. MILLER. 1978. Nearctic pine tip moths of the genus Rhyacionia:
Biosystematic review (Lepidoptera: Tortricidae, Olethreutinae). U.S. Dep. Agr., Agr.
Handb. 514, 51 pp.

POWELL, J. A. & J. RAZOwsKI. In press. Tortricidae: Olethreutinae. In Heppner, J. B.
(ed.), Atlas of Neotropical Lepidoptera. Checklist: Part 2. Pyraloidea-Tortricoidea.
E. J. Brill, Leiden.

ZIMMERMAN, E. L. 1978. Insects of Hawaii 9, Microlepidoptera I. University Press of
Hawaii. 882 pp.

Received for publication 1 March 1988; accepted 29 March 1988.

Journal of the Lepidopterists’ Society
42(3), 1988, 240-244

BOOK REVIEWS

THE BUTTERFLIES OF COSTA RICA AND THEIR NATURAL HISTORY: PAPILIONIDAE, PIERI-
DAE, NYMPHALIDAE, by Philip J. DeVries (illus. by P. J. DeVries, Jennifer Clark and R.
Cubero). 1987. Princeton Univ. Press. xxii + 327 pp. Hardback, ISBN 0-691-08420-3,
$60.00; paperback, ISBN 0-691-02403-0, $22.50.

Costa Rica is an exceptionally delightful Central American country with a happy and
friendly people (they abolished their military some years ago and used the money re-
covered for education), gorgeous landscape, and a surprisingly complete complement of
beautiful Neotropical butterflies which are ably described, discussed, and made known
in intimate detail in this book.

For the three families covered, this is a very good book on Neotropical Lepidoptera,
and will be most useful (especially at the low price for the paperback edition) to anyone
interested in these families from central México southwards. It could be a better book
still; it contains a rather high density of errors of nomenclature, fact, writing, and inter-
pretation which should be corrected in the second edition (which surely will be necessary
as several thousand copies have already been sold). A list of these has been given to the
author; only some more important facets can be commented on here, probably most
usefully to those who have already bought the book. If you haven't, you should as soon
as you plan to work on the Neotropical fauna in the three families.

DeVries has lived for almost 10 years in Costa Rica, working at the National Museum
of Natural History with top Costa Rican scientists, and maintaining ties with many foreign
scientists studying natural history in this Country (mostly through the Organization for
Tropical Studies; a general volume entitled Costa Rican Natural History, edited by Daniel
Janzen, recently appeared, and covers many aspects of this work), and also in other
Central American countries (especially the Muyshondts, active in El Salvador). He is
responsible for the discovery of many species in Costa Rica, verification of their habitats
and foodplants, and description of their early stages, and is surely well qualified to produce
this book. The text was written mostly during periods in Austin, Texas, and the British
Museum; the color plates, of high quality but reduced a little too much for optimal
usefulness, were prepared in the latter institution, at times using non-Costa-Rican or older
specimens, with faded colors er tattered wings.

Strong points of the book include maps of the Country with roads, cities, topography,
and parks (but with a wrong scale; 50 km on the scale given are actually 82 km on the
map), a refreshing emphasis on natural history and juvenile biology with much valuable
new data, a useful list of butterfly-enemy groups (p. 18), a sensible and eclectic position
on systematics (p. 32), a recommendation for taking notes rather than specimens in the
field (p. 33) and use of binoculars and simple camera equipment to increase the value
and number of observations (p. 37), a good grasp of the Costa Rican faunal regions
showing much first-hand experience (though perhaps not enough yet in cloud-infested
Atlantic coastal forests), recognition of certain behavioral traits such as “only new males
visit wet sand’, an excellent job of correlating illustrations with types in difficult groups
like Memphis, Adelpha, Phyciodes (s.l.), and Euptychiini which will be useful to many
(if all are correct), and palatability data on many species. Line drawings of juveniles are
strikingly clear and correct.

DeVries’ youthful enthusiasm, which made this book possible and contributed greatly
to its notable authority, style, grace, and the breadth and interest of its natural history
accounts, gets the best of him in diverse parts of the book. His statement in the Preface
that, when he began to work in Costa Rica in the 1970's, “nobody was willing to do a
detailed study of the Neotropical butterfly fauna” is bound to earn some grumbles from
the many “nobodies” who have willingly and tirelessly labored in this task since Miller
(1870's) and Moss (1900's) started doing broad work on natural history of Brazilian
Lepidoptera; several dozen are now active, many in fact quoted in the large Bibliography
(382 references). An unexpected unfamiliarity with recent work in Mexico, such as that
on Parides (mostly published in the Rev. Soc. Mex. Lepid. mentioned on p. 53 but cited
only 3 times in the Bibliography), and in South America since Miiller and Moss, sometimes

VOLUME 42, NUMBER 3 241

leads to unnecessary affirmations or contradictions; in species ranges, 10 genera and 58
species common in southeastern Brazil are indicated as reaching only the Amazon Basin,
10 others have their southern limits shrunken appreciably (sometimes all the way to
Central America) and many groups are indicated as “reaching their maximum diversity
in the Amazon Basin” when in fact this occurs on the lower Andean slopes (as indicated
correctly in Prepona)—Amazonian to be sure but only a small, special part of the Basin,
over almost all of which butterfly diversity is quite low. Peterson’s classic 1948 work on
larvae is not mentioned (p. 6), and the list of parasites omits mites (p. 17). The affirmation
that “the effects of parasitoids on populations of tropical butterflies are unstudied” (p.
17) makes one wonder what the author accepts as a study (there are dozens of purported
scientific papers published in this area). Likewise, the generalization that “toxins in adult
butterflies are probably entirely directed at vertebrate predators’ (p. 23) is indefensible.
That Neotropical diversity “remains one of the great challenges in evolutionary biology”
(p. 57) again ignores much serious work done recently by many Neotropical and other
scientists. In a perhaps Freudian slip, several medicinal (officinal) plant species are spelled
“officionale”, and misspelled plant families (new synonyms?) include Vochysiaceae
(“Vouchysiaceae’’, p. 66, 68), Canellaceae (“Cannelaceae’’, p. 61), Quiinaceae (“Quiini-
aceae’, p. 109, but correct on p. 112), and Verbenaceae (“Verbenaeceae’’, p. 76). Ith-
omiinae venations on p. 215 are so poorly drawn that they will confuse, not help the
reader. The late Walter Forster would surely be disappointed at DeVries’ summary
dismissal of almost all his Euptychiine genera, many of which are sound natural groupings.
Nomenclatural corrections and changes necessary (*) or suggested include at least the
following (which should be noted also on the check-list, pp. 291 ff):

Page Name given Should be Comments

18 *Tyranidae Tyrannidae —

Al *Hamadryas iphtheme __H. iphthime —

65 *Parides alopius (delete) Nicaraguan record an
error.

*P. dares (delete) A unique hybrid (photin-

us X montezuma).

67 *P. arcas P. eurimedes Arcas a homonym.

69 *Battus belus varus B. latinus B. varus is Amazonian,

female latinus mono-
morphic throughout

range.
70 B. crassus B. crassus lepidus Transandean subspecies.
is Papilio victorinus, P. (probably one species) Would be useful to hy-
cleotas bridize these in Costa
Rica.
ek Eurytides ilus, E. bran- (probably one species) —
chus
78 *E. protesilaus dariensis (includes E. macrosilaus) Should separate.
89 *Archonias eurytele Charonias eurytele Well differentiated gen-
era; also Fig. 7.
97 *Perrhybris pyrrha P. pamela Pyrrha a homonym; also
Fig. 7:
99 Ascia limona A. buniae limona Probably conspecific.
Tis *Prepona omphale (includes P. laertes) Should separate.
1S *P. meander amphima- (two different species) Separate.
chus
116 Zaretis ellops, Z. itys (probably one species) Widespread polymor-
phism all over Neo-
tropics; seasonal.
144, Dynamine hecuba, D. (probably subspecies of —
145 sosthenes South American spe-

cies)

242

142
153

154
161

162
163
182
183
185

186
188

191
193
192

207
221
224
226
230
240
241

249

257 ff

262

275

276

D. glauce

*M yscelia orisis

Eunica venusia, E. au-
gusta

*Cantonephele

*Haematera pyramus

*Pseudonica

Diaethria marchallii
*Turnera ulmnifolia
*Actinote leucomelas

*A. melampeplos, A.

guatemalensis
*Heliconius sappho
Dione juno

*Eueides lybia lybioides
(first mention, (Fabr.))
Heliconius doris
*Fueides isabella zora-
con
*Napeogenes peredia
*Callithomia hexia
*Ithomia diasa
*Prestonia portabellensis
*Codyris zavaleta sorites
* Hyaliris
Antirrhaea miltiades, A.
tomasia
Brassolis isthmia

Euptychiini (esp. Eup-

tychia mollina, Cissia
terrestris )

*Callitaera (for polita)

*Cissia libye

*Cissia hesione

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

D. artemisia glauce

M. orsis

E. volumna venusia, E.
caelina augusta

Catonephele

Callidula pyrame

Nica

D. clymena marchallii
T. ulmifolia
Altinote ozomene nox

A. pellenea subspecies
(both)

H. sapho

D. juno huascuma

E. lybia olympia
Laparus doris
E. isabella zorcaon

N. peridia

C. hezia

I. diasia

P. portobellensis

G. zavaleta caesiopicta
Hyalyris

(should be one species)

B. sophorae isthmia

(many Amazonian names
pulled in incorrectly for
Central American spe-
cies. Nomenclature
here a big step forward
but still lacks refine-
ment)

Cithaerias

C. libyoidea (Butler) (or
Magneuptychia)

Pareuptychia ocirrhoe

Senior synonym, same
species.

Senior synonym, Nica
used on p. 156.

Well separated genus,
older name.

Central American sub-
species.

Well differentiated ge-
nus.

See also p. 46, fig. 7.

Senior synonym.

Popular misspelling.

Reason for separation not
convincing.

Probably monotypic ge-
nus.

Older synonym, correct
on p. 260.

Transandean species has
different chromosomes
from Guianan type.

Hesione a homonym.

Plate errors include illustration of Papilio paeon as P. cresphontes (Plate 4; P. paeon
not mentioned in text); inversion of names for Danaus gilippus and D. eresimus (Plate
33); and illustration of a probable Caligo oileus (indicated as “not illustrated” in text) as
C. illioneus on Plate 46, no. 2. Figure 7 (a “tipped in” plate following p. 26) was missing

from the paperback copy purchased for my University library.

A selection of additional irksome errors (unfortunately, there are many more, some
quite misleading) includes the following. There is a general lack of detailed information
on variation in juveniles (very frequent in my experience), accentuated by sketchy, often
inscrutable or generalized, or even wrong (as in Papilio, Hypothyris) descriptions, and
few mentions of the number of replicates of rearing (thereby many opportunities to help

VOLUME 42, NUMBER 3 243

in juvenile field identification lost). There occurs an implicit advocation of collection in
nature reserves (p. 33); recent work on endangered species in Brazil suggests that even
limited collecting of adults or juveniles of rare or local species in some kinds of habitats
can seriously depress subsequent generations. Species lists for the Carrillo Belt are repeated
(pp. 47, 49). Many place names mentioned in text are missing from maps (but usually
are present in the gazetteer, pp. 285-287, where only one locality—Rincon, Osa—was
found misplaced, 160 km to the NW). Data on canopy faunas are anecdotal, and seem
overemphasized, or else are much more important in Costa Rican topography than in
the flatter Amazon Basin. Diversity comparisons (p. 52) use only two sites in each of the
six regions; only near-asymptotic lists, which these are probably not, can give a reliable
picture. A number of generalizations presented in family, tribe or generic accounts are
not obeyed by many included taxa, such as “tailless and sexually dimorphic troidines”
(not Parides photinus or P. montezuma) with “white woolly scent scales” (not P. eu-
rimedes ), and “mimetic Dismorphiines’’ (not the majority listed). Inference of larval host
plant from pupal placement for Papilio cleotas (fide W. Haber) would permit P. an-
chisiades to eat my back door, over 80 m from the nearest Citrus they actually fed upon;
some go much farther. The book’s author doubts that male Battus visit sand (normal all
over South America), that male Eutresis visit pyrrolizidine alkaloid sources (frequent in
Venezuela and Colombia), that pterin pigments occur outside Pieridae (present in most
butterflies, at times in large quantities), or that mimetic charaxines occur outside Consul
(he illustrates several without commenting on their mimicry). He mixes up the characters
in describing seasonal forms of Eurema proterpia (p. 105, compare Plate 10), and uses
an-idi ending for “subfamily or tribal status” (p. 127). He indicates 10 species in Liby-
theana (there are 3), 2 in Baeotus (there are 3), “a few’ in Eutresis (probably only 2),
5 in Brassolis (probably all 1), 10 in Cithaerias (probably 3 or less), 1 in Dulcedo (few
know about the 2nd high-elevation west Colombian species, D. mimica). He restricts
Nessaea to swamp forests, though it occurs on mesic hillsides in many parts of South
America. Female behavior of Callidula pyrame (“Haematera’’) is generalized to both
sexes. The red basal dots on the ventral hindwing of Héliconius are transferred to the
forewing. Microtia is placed in the Melitaeini on p. 198, in the Phyciodini on p. 205. A
dropped line on p. 213 grows wing-pads on Lycorea larvae. The interesting mimicry
situations involving Caligo atreus-Antirrhaea pterocarpha-female Catoblepia orgetorix
and Drucina leodonta-Tithorea tarricina need comment; the second is simply denied (it
occurs in Panamanian Chiriqui), the first involves birds’ wariness at the large size and
eyespots of Caligo who can even keep birds away from their feeders.

These diverse “gripes” could go on, but will tire the reader. They are not intended to
subtract from the value of the book, but to add to that of future editions, avoiding a wide
circulation of inconsistencies and misinformation. In relation to most current books cov-
ering parts of the Neotropical butterfly fauna, this one stands out in general as carefully
written, taxonomically accurate, biologically important, ecologically interesting, and sane.
This is, in part, a negative reflection on the others.

It seems interesting to compare this book with the only other well illustrated modern
single-country butterfly guide for the Neotropics, Barcant’s 1970 Butterflies of Trinidad
and Tobago. Prepared by an amateur and lifetime Trinidad resident, and aimed at nature
lovers, children, and collectors, this book received such a negative and unfair review in
this journal that its author remained deeply embittered until his death last year. It covered
the complete lot of families (with rather inscrutable plates for Lycaenidae and an outdated
list for Hesperiidae); its color illustrations were generally of high quality and esthetic
balance, showing recently captured specimens with no damage. Emphasis was likewise
on natural history, though with less information on juveniles and foodplants, to the point
of using an arrangement based on habitat rather than taxonomy; coverage of historical,
traditional and folklore aspects was strong, with common names in Trinidad given for
many species. Barcant was not familiar with modern ecological patter and theory as is
DeVries, which sometimes detracted and other times helped his book (many such aspects
have notoriously short half-lives). Both books give many specific details on collecting
localities, adult habitats, mimicry, seasonality, physical geography and general methods;
Barcant is more “folksy” while DeVries is more “objective ’.

244 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

It is to be hoped that DeVries’ book reaches a wider readership today than did Barcant’s
18 years ago, and will continue to stimulate and aid those who study the fullness of
biological aspects of Neotropical butterflies, and who wish to publish such well illustrated
regional accounts in the future. All of these, like the Barcant and DeVries books, should
be of immeasurable assistance in biological and ecological studies in the Neotropics, as
well as useful and enjoyable to amateurs.

KEITH S. BROWN Jr., Departamento de Zoologia, Instituto de Biologia, Universidade
Estadual de Campinas, Campinas, Séo Paulo 13.081 Brazil.

Journal of the Lepidopterists’ Society
42(3), 1988, 244-245

BUTTERFLIES OF NORTH DAKOTA: AN ATLAS AND GUIDE, by Ronald Alan Royer. 1988.
Science Monograph No. 1, Division of Science, Minot State University, Minot, North
Dakota 58701. Format 14 x 20 cm, 192 pp., 12 pp. color plates, 1 b/w plate, 30 pp.
maps. Plastic spiral binding. Soft cover. $14.95.

The author has brought together in a single volume a guide and atlas which describes
with text, color plates, and distribution maps all 142 species of butterflies known to occur
in North Dakota. The book succeeds in fulfilling the author’s goal of providing access to
the Lepidoptera literature for the state.

The book begins with an Introduction which includes an explanation of the binomial
system, North Dakota environments, terminology (wing surface and venation), and scope
and use of the book. This is followed by the Guide, which includes a narrative for each
species. Each species is introduced with the common and binomial name (including full
author name), and corresponding plate number. The Atlas includes a State map with
counties named followed by a State map for each species with the counties of record spot
marked. There are five maps on the left side of each page with space on the right for
notes. Plates are 60% natural size photographs of actual specimens. Opposite each plate,
the binomial name, sex, view (dorsal or ventral), collection locality, and date collected
are arranged according to how specimens appear in the plate. Next is a Hypothetical and
Erroneous Records List followed by a list of names and addresses of lepidopterist orga-
nizations and suppliers. The Bibliography follows, then a 121 word Glossary, and finally
the Index of Butterflies listed by binominal and common names with page numbers for
the Guide, Atlas, and Plate sections.

Some of the nice things about this volume have already been mentioned, but still others
are obvious when you pick it up—the sturdy binding and quality paper are suited for
years of use. The cover is dominated by a photograph of Hesperia dacotae (Skinner), a
nice touch. Coverage is complete and you could not ask for more information in the
species descriptions. The author follows the 1981 Miller & Brown generic naming system,
and there are no taxonomic surprises.

The faults with the book are few considering the wealth of information presented. The
map in the Introduction shows only the major life zones. A more detailed map should
have named the major rivers, drainage systems, and geographical features. Repeating the
named counties map would have been helpful, too. Reading this section makes one feel
the book was written for North Dakota collectors already acquainted with the State rather
than for collectors who find themselves in North Dakota. The terminology section would
have benefited with an explanation of how to distinguish the sexes, and with a generalized
diagram of external morphological characters. The chapter might also have included a
brief discussion of butterfly evolution, clarifying the heirarchy used in the book. There
are no keys except one to the Papilionoidea.

Most of my comments concern the Guide and Plates chapters. The desired information
is there, but would have been easier to locate if headings such as Description, Similar

VOLUME 42, NUMBER 3 245

Species, Life Cycle, Flight, Habitat, and Range were inserted in bold type in the text.
The text is not cross-indexed except for a Plate number with each species description.
The Guide, Atlas, and Plates sections should have included page numbers for the cor-
responding sections which would have helped tie the chapters together. The color plates
are of good quality, and my only change would have been to adjust background colors
of Plates II and IV to increase contrast. Illustrated specimens should have been numbered,
with those numbers repeated in the Plate legend. This would have helped Plate II, as the
extreme example, where 56 specimens are pictured, and searching the legend for the
binomial name is tedious. A simple checklist at the end would have been useful to some
collectors, or perhaps a box to check by each distribution map.

The faults with the book are few, and my criticisms also apply to a number of other
popular books and field guides. This book is a valuable source of information. Whether
or not you are ever fortunate enough to collect in North Dakota, this handsome book is
a must for the naturalist.

R. D. PETERSON II, Biosciences Research Laboratory, ARS-USDA, P.O. Box 5674,
Fargo, North Dakota 58105.

Journal of the Lepidopterists’ Society
42(3), 1988, 245

THE MOTHS OF BORNEO: SUPERFAMILY BOMBYCOIDEA: FAMILIES LASIOCAMPIDAE,
EUPTEROTIDAE, BOMBYCIDAE, BRAHMAEIDAE, SATURNIIDAE, SPHINGIDAE, by Jeremy D.
Holloway. 1987. Southdene Sdn. Bhd., P.O. Box 10139, Kuala Lumpur 50704, Malaysia.
199 pages, 20 color plates. Paperback. About $35.00.

This book deviates from other faunistic treatments by including more detail on phy-
logeny and ecology, particularly hostplants. The color plates were produced by Bernard
D Abrera, so are predictably of high quality. All known species in these families from
Borneo are treated in the text and depicted in color, thus including a considerable portion
of the Indo-Australian moth fauna. The text draws from observations and works published
in Asia by resident entomologists, and manifests Holloway’s own extensive field experience
in the region; the result is far beyond what could be produced from study of museum
specimens alone. Where new or controversial taxonomic decisions are enacted, the author
faithfully provides justification or explanation.

Inclusion of Sphingidae within Bombycoidea is unexpected. Upon reading the discussion
of characters to justify this, I was a little disappointed, but apparently seven synapomophies
do link sphingids to other bombycoids. Such a large superfamily, now comprising 18 or
14 families worldwide, makes it difficult to designate nomenclaturally the closer rela-
tionships within the group; one wishes for a category between superfamily and family
levels (or between suborder and superfamily levels) to remedy the situation. Holloway’s
discussion of the phylogeny of those families makes the book useful to Lepidoptera
taxonomists around the world, even to those who profess no interest in moths of south-
eastern Asia. The book is well done. I found no typographical errors. I believe those who
acquire it will wish to purchase other volumes in the series, most of which are as yet
unpublished.

RICHARD S. PEIGLER, 323 Van Gordon Street, #19-523, Lakewood, Colorado 80228.

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EDITORIAL STAFF OF THE JOURNAL
WILLIAM E. MILLER, Editor

Dept. of Entomology
University of Minnesota
St. Paul, Minnesota 55108 U.S.A.

Associate Editors and Editorial Committee:
M. DEANE BOWERS, BOYCE A. DRUMMOND III, DouGLas C. FERGUSON,
LAWRENCE F. GALL, ROBERT C. LEDERHOUSE, THOMAS A. MILLER,
THEODORE D. SARGENT, ROBERT K. ROBBINS

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories
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Literature Cited: References in the text of Articles should be given as Sheppard (1959)
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SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London.
209 pp.

196la. Some contributions to population genetics resulting from the study of

the Lepidoptera. Adv. Genet. 10:165-216.

In General Notes and Technical Comments, references should be shortened and given
entirely in the text as P. M. Sheppard (1961, Adv. Genet. 10:165-216) or (Sheppard, P.
M., 1961, Sym. R. Entomol. Soc. London 1:23-30) without underlining.

Illustrations: Only half of symmetrical objects such as adults with wings spread should
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CONTENTS

CYRIL FRANKLIN DoS Passos (1887-1986). Ronald S. Wilkin-

SOM oie ee et kate ee ek ee 3S
MEMORIES OF CyRIL F. pos Passos (1887-1986). Lionel Paul
Grey 6 ee ee A

ANNOTATED BIBLIOGRAPHY OF THE ENTOMOLOGICAL PUBLICA-
TIONS OF CyRIL F. pos Passos (1887-1986). Ronald S. Wil-
kinson 62

BIOLOGY OF SPEYERIA ZERENE HIPPOLYTA (NYMPHALIDAE) IN A
MARINE-MODIFIED ENVIRONMENT. David V. McCorkle &
Paul C. Hammond oo

MARKING LEPIDOPTERA AND THEIR OFFSPRING: TRACE ELEMENT
LABELLING OF COLIAS EURYTHEME (PIERIDAE) WITH RU-
BIDIUM. Jane Leslie Hayes & Cathryn L. Claussen

AN APPRAISAL OF GAZORYCTRA HUBNER (HEPIALIDAE) AND DE-
SCRIPTION OF A NEW SPECIES FROM ARIZONA AND NEW
Mexico. David L. Wagner &© Norman B. Tindale _.

TEMPORAL TRENDS IN FREQUENCIES OF MELANIC MORPHS IN
CRYPTIC MOTHS OF RURAL PENNSYLVANIA. Thomas R.
Manley oo

A NEW SPECIES OF OCALARIA (NOCTUIDAE: CATOCALINAE) AND
ANALYSIS OF SOME MORPHOLOGICAL CHARACTERS USEFUL
FOR ELUCIDATING NOCTUID PHYLOGENY. Ian J. Kitch-

{) ORION REM ae EL
A NEW SESIA CLEARWING MOTH FROM MICHIGAN (SESI-
IDAE). Thomas D, Eichlin & William H. Taft ——e

TWO NEW SPECIES OF RHYACIONIA PINE MOTHS FROM MEXICO
(TORTRICIDAE: OLETHREUTINAE). William E. Miller

ee eew en neenene

BooK REVIEWS

The Butterflies of Costa Rica and their Natural History: Papilionidae, Pier-

idae, Nymphalidae. | Keith S. Brown Jr. i.e
Butterflies of North Dakota: An Atlas and Guide. R. D. Peterson II
The Moths of Borneo: Superfamily Bombycoidea: Families Lasiocampidae,

Eupterotidae, Bombycidae, Brahmaeidae, Saturniidae, Sphingi-
dae. Richard S. Peigler

ANNOUNCEMENT
Journal Cover Illustrations and Feature Photographs.

ADDITIONAL MANUSCRIPT REVIEWERS, 1987

155

164

168

184

196

204

213

ee O

: a .
—— So ee eS OO

Volume 42 1988 Number 4

ISSN 0024-0966

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

olf! ‘95°

6 December 1988

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

JULIAN P. DONAHUE, President GEORGE W. GIBBS, Vice
JERRY A. POWELL, Immediate Past President

President RONALD LEUSCHNER, Vice
JOHN N. ELIOT, Vice President President
RICHARD A. ARNOLD, Secretary JAMES P. TUTTLE, Treasurer

Members at large:

JOHN W. BROWN M. DEANE BOWERS FREDERICK W. STEHR
MOGENS C. NIELSEN RICHARD L. BROWN JOHN E. RAWLINS
FLOYD W. PRESTON PAUL A. OPLER Jo BREWER

The object of the Lepidopterists’ Society, which was formed in May 1947 and for-
mally constituted in December 1950, is “to promote the science of lepidopterology in all
its branches, .... to issue a periodical and other publications on Lepidoptera, to facilitate
the exchange of specimens and ideas by both the professional worker and the amateur
in the field; to secure cooperation in all measures’ directed towards these aims.

Membership in the Society is open to all persons interested in the study of Lepi-
doptera. All members receive the Journal and the News of the Lepidopterists’ Society.
Institutions may subscribe to the Journal but may not become members. Prospective
members should send to the Treasurer full dues for the current year, together with their
full name, address, and special lepidopterological interests. In alternate years a list of
members of the Society is issued, with addresses and special interests. There are four
numbers in each volume of the Journal, scheduled for February, May, August and
November, and six numbers of the News each year.

Active members—annual dues $25.00
Student members—annual dues $15.00
Sustaining members—annual dues $35.00
Life members—single sum $500.00
Institutional subscriptions—annual $40.00

Send remittances, payable to The Lepidopterists’ Society, to: James P. Tuttle, Treasurer,
3838 Fernleigh Ave., Troy, Michigan 48083-5715, U.S.A.; and address changes to: Julian
P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, California
90007-4057 U.S.A. For information about the Society, contact: Richard A. Arnold, Sec-
retary, 50 Cleaveland Rd., #3, Pleasant Hill, California 94523-3765, U.S.A.

For back issues of the Journal and News, write the Publications Coordinator at the
address below about availability and prices. Prices of The Lepidopterists’ Society com-
memorative volume 1945-1978 are $12.00 ($8.00 to members and subscribers); of A cat-
alogue/checklist of the butterflies of America north of Mexico, clothbound, $19.00 ($12.00
to members and subscribers), paperbound, $10.50 ($7.00 to members and subscribers). Order
from the Publications Coordinator, Ronald Leuschner, 1900 John St., Manhattan Beach,
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Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for $40.00
(institutional subscription) and $25.00 (active member rate) by the Lepidopterists’ Society,
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Cover illustration: Captive female cecropia moth, Hyalophora cecropia (L.) (Saturni-
idae), released to wild and resting on staghorn sumac, Rhus typhina L. (Anacardiaceae).
Submitted by Monica Miller, 5560 Library Road #201, Bethel Park, Pennsylvania 15102.

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

JOURNAL OF

Tue LEPIDOPTERISTS’ SOCIETY

Volume 42 1988 Number 4

Journal of the Lepidopterists’ Society
42(4), 1988, 247-262

SYSTEMATIC POSITIONS OF ACENTRIA EPHEMERELLA
(DENIS & SCHIFFERMULLER), NYMPHULINAE, AND
SCHOENOBIINAE BASED ON MORPHOLOGY OF
IMMATURE STAGES (PYRALIDAE)

STEVEN PASSOA

Department of Entomology, University of Illinois, 320 Morrill Hall,
Urbana, Illinois 61801

ABSTRACT. Acentria ephemerella (Denis & Schiffermiller), the one known species
of its genus, is sometimes placed in Schoenobiinae, but it lacks three important autapo-
morphies of that subfamily: larval prothoracic sac, exposed pupal mesothoracic coxae,
and deep pitlike pupal mesothoracic spiracle. Apomorphies such as spinelike pupal frontal
setae, lack of pupal mesothoracic spiracle, and reduced posterior pupal abdominal spiracles
confirm that Acentria belongs in Nymphulinae. No larval or pupal characters were found
to support Acentria as a separate family or subfamily (Acentropidae or Acentropinae).
Several synapomorphies suggest Nymphulinae and Schoenobiinae are sister groups. They
share long exarate pupal appendages and reduction of larval L2 seta on abdominal
segments 1-8. The unisetose L group on abdominal segment 9 in other subfamilies of
Crambiformes may be used as a synapomorphy to define a clade separate from Nym-
phulinae and Schoenobiinae in which the L group is bisetose on segment 9.

Additional key words: larva, pupa, cladogram, systematics.

Acentria ephemerella (Denis & Schiffermiller), formerly Acentria
(=Acentropus) nivea (Olivier), has a long and varied systematic history
(Speidel 1981, 1984). It was placed in Schoenobiinae because of a
reduced proboscis, tubular CuP (1A) forewing vein, and lack of hind-
wing Cu pecten (hair fringe) (Hampson 1895, Forbes 1926, 1938). Other
workers (Marion 1954, Roesler 1973, Leraut 1980, Goater 1986) thought
Acentria should be in Acentropinae or Acentropidae largely because
the adult lacks a praecinctorium. Nigmann (1908) and Speidel (1981)
cited enlarged anterior abdominal pupal spiracles as an autapomorphy
of Nymphulinae and thus considered Acentria to be in this subfamily
because of its similar pupa. Larval chaetotaxy confirmed this view.
Hasenfuss (1960) placed Acentria in Nymphulinae based on a bisetose
L group on abdominal segment 9, and unusual arrangement of larval

248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

stemmatal (ocular) setae. Speidel (1984) suggested Acentria ephemer-
ella is the correct name for A. nivea and recommended acceptance of
Acentropinae instead of Nymphulinae as the valid subfamily name.
The latter change, in agreement with Minet (1982), is not followed here
because Nymphulinae has been stable and unambiguous in most check-
lists. Fletcher and Nye (1984) placed Acentria with Nymphulinae in
their Pyraloidea catalogue. However, Yoshiyasu (1985) doubted the
placement of Acentria with Nymphulinae because enlarged abdominal
spiracles are also found in some aquatic Crambinae. The possibility
remained that enlarged spiracles had arisen in certain species because
of aquatic habits instead of common ancestry. Minet (1982, 1985) also
considered Acentria to be a nymphuline, based on several apomorphies
of the tympanum. As was traditional in the U.S., Munroe (19838) listed
Acentria with Schoenobiinae. Batra (1977), Berg (1942), Buckingham
and Ross (1981), Speidel (1981), and Yoshiyasu (1985) either illustrated
stages of A. ephemerella or discussed its biology.

Only three workers have published Pyralidae cladograms (Fig. 1).
Roesler (1973), relying mostly on adult morphology, recognized an
Acentropidae-Crambidae complex. Yoshiyasu (1985) doubted the va-
lidity of Roesler’s characters. He called attention to variability in the
Nymphulinae radial vein and maxillary palpi, as well as to the presence
of aquatic species in other subfamilies. More importantly, some key
portions of Roesler’s (1973) cladogram are defined by plesiomorphic
features (lack of specialized scales in the male genitalia, for example).
Kuznetzov and Stekolnikov (1979) considered Schoenobiinae and Nym-
phulinae to be unrelated, based almost exclusively on genital muscu-
lature. However, they studied very few species and paid only superficial
attention to immature stages. Yoshiyasu (1985), considering characters
of all stages, linked Schoenobiinae, Nymphulinae, and Acentria as sister
groups but was unable to place this clade in an overall scheme. Instead,
three clades were extended to a single point with dotted lines and a
question mark at their bases (Fig. 1A). Thus, convincing evidence from
adult (Minet 1982), larval (Hasenfuss 1960) and pupal (Nigmann 1908)
morphology suggests Acentria belongs with Nymphulinae in spite of
recent doubts (Yoshiyasu 1985, Goater 1986).

This paper examines apomorphic larval and pupal characters of
Acentria ephemerella to provide additional evidence on the systematic
position of Acentria. The relation of Nymphulinae to Schoenobiinae,
and their taxonomic position within Crambiformes are also discussed.

METHODS

Morphological information on pyralid immature stages came from
Passoa (1985), literature illustrations, and borrowed material. Unpub-

VOLUME 42, NUMBER 4 249

cAey
SO SAV“s Cy Poe
OS SQ LF CS
SSW Were acEN CRAMB

AS

F MLE PL SF
PM OWSQEY MD? oS
LEESON ONS

CRAMBIF
C D

Fic. 1. Systematic position of Acentria, Nymphulinae, and Schoenobiinae after var-
ious authors. A, Yoshiyasu (1985). B, Roesler (1973). C, Kuznetsov & Stekolnikov (1979).
D, Present study, with major apomorphies numbered as follows: 1—larva with prothoracic
sac; 2—pupal mesothoracic spiracle pitlike; 3—pupal mesothoracic coxae exposed; 4—
stemmatal setae in line with each other; 5—pupal frontal setae enlarged and spinelike;
6—pupal anterior abdominal spiracles enlarged and on conelike projections, posterior
abdominal spiracles reduced; 7—pupal mesothoracic spiracle lost; 8—V1 lost on larval
thorax; 9—L2 on larval abdominal segments reduced; 10—tegumen-vinculum plate de-
veloped, transtilla lost; 1l1—pupal appendages exarate with metathoracic legs exposed;
12—praecinctorium present; 13—larva with unisetose L group on AQ. Abbreviations:
ACEN—Acentropidae; Acen—Acentropinae; Anchy—Ancylolomiinae (Anchylolomiinae
of Yoshiyasu 1985); CRAMB—Crambidae; Cramb—Crambinae; CRAMBIF—Crambi-
formes; Cybal—Cybalomiinae; Ever—Evergestinae; Glap—Glaphyriinae; Muso—Mu-
sotiminae; Nymph—Nymphulinae; Odon—Odontiinae; PYRAL—Pyraliformes; Pyr-
aus—Pyraustinae; Schoen—Schoenobiinae; Scop—Scopariinae.

lished keys and a data matrix of larval characters by workers at the
U.S. National Museum (C. Heinrich, H. Capps, and D. Weisman)
(““USNM Tables’) provided information on pyralid genera in that col-
lection. Literature on Crambiformes immature stages included general
works such as Fracker (1915), Mosher (1916), Peterson (1962), and
Neunzig (1987) for the U.S., Hasenfuss (1960) for Europe, Nakamura
(1981) for Asia, and Gerasimov (1947, 1949) for the U.S.S.R. Important
articles on New World Nymphulinae immatures were selected from
Munroe (1981, 1982). Yoshiyasu (1985) published a review on Japanese
Nymphulinae and their systematic position. Schoenobiinae immatures
were discussed by Passoa and Habeck (1987). Crawford (1961), Mauston
(1970), and Tan (1984) provided descriptions of Crambini larvae and
pupae. Agarwal and Chaudhry (1966), Passoa (1985), and Rothschild
(1967) described Chilini immatures. Works on New World Pyraustinae

250 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

included Allyson (1981, 1984), and Passoa (1985). Khot’ko and Mol-
chanova (1975) studied Old World species. Some African pyralids were
illustrated by Breniere (1979). Indian pyralids were described by Ma-
thur and Singh (1963) and Mathur (1954, 1959).

Preserved larvae, and usually pupae, of the following species were
examined:

Nymphulinae

Acentria ephemerella (Denis & Schif- | Neargyractis slossonalis (Dyar)
fermiller) Petrophila longipennis (Hampson)

Nymphula depunctalis Guenée P. bifascialis (Robinson)
N. fluctuosalis (Zeller) P. avernalis (Grote)
Munroessa sp. P. jaliscalis (Schaus)
Synclita sp. Usingeriessa onyxalis (Hampson)
Parapoynx diminutalis Snellen Eoparargyractis sp.
P. obscuralis (Grote)

Schoenobiinae
Rupela albinella (Cramer) Scirpophaga (=Schoenobius, Tryporyza)
R. horridula Heinrich incertulas (Walker)
R. leucatea (Zeller) S. excerptalis (Walker) (=S. intacta
R. sp. Snellen)

Donacaula sp.
D. maximella (Fernald)

This list represents 9 of 16 Nymphulinae genera and 3 of 5 Schoe-
nobiinae genera in the U.S. (Munroe 1983). Acentria contains only one
species, A. ephemerella (Speidel 1984). Larva and pupa terminology
follows Stehr (1987) and Mosher (1916). Munroe (1972) and Minet
(1982, 1983, 1985) were used to characterize adult subfamilies.

CHARACTER POLARITY

Certain assumptions are necessary before a cladistic study of Acen-
tria, Nymphulinae, and Schoenobiinae can proceed. Pyralidae is as-
sumed monophyletic because of apomorphies in the tympanum (Minet
1982, 1983, 1985) and venation (Munroe 1972). All Pyralidae clado-
grams (Fig. 1) agree there are two lineages, Crambiformes (sometimes
called Crambidae) and Pyraliformes (sometimes called Pyralidae in a
restricted sense). Crambiformes, which include Nymphulinae, Schoe-
nobiinae, and Acentria, are apomorphically defined, in part, by a prae-
cinctorium in the tympanum (Minet 1982). Although tympanic mor-
phology of Midiliformes and other pyralids differ, larval characters, as
discussed further on, support Minet’s (1982) placement of this taxon
within Crambiformes. Pyraliformes, which include all remaining py-
ralid subfamilies, are the sister group to Crambiformes, and thus com-
prise the outgroup. Minet (1985) apomorphically defined Pyraliformes
by a tympanic “paraspina” and sclerotized pinaculum rings around

VOLUME 42, NUMBER 4 5a)

larval seta SD1. Speidel (1984) mentioned scale morphology, dimorphic
labial palps, and reduction of proboscis, ocelli, and leg spurs as apo-
morphies of Acentria. A shortened, stout gnathos, and broad basal
portion of the apophysis united Acentria and Kasania on a single clade.

Unless stated otherwise, Watrous and Wheeler’s (1981) method of
outgroup comparison was used to polarize characters. This method is
especially appropriate when most characters have two states, and rel-
atives are easily defined. In spite of criticisms (Farris 1982, Clark &
Curran 1986), outgroup comparison appears to be tke most reliable
way to determine polarity (Donoghue & Cantino 1984). All morpho-
logical features relevant to the systematic positions of Acentria, Nym-
phulinae, and Schoenobiinae are mentioned below even if their polarity
is somewhat uncertain. References under each morphological feature
usually provide illustrations.

Larval Characters

Stemmatal (ocular) setae. Hinton (1946) considered $1 close to stemmata 3 and 4, S2
level with stemma 5, and S3 below all stemmata as the usual arrangement in Lepidoptera.
This trend is also true in Pyralidae where all Pyraliformes and Crambiformes except
Nymphulinae show this arrangement (Hasenfuss 1960, Yoshiyasu 1985). Two states occur
in Crambiformes: setae in nonlinear arrangement or in line with each other. Since all
Pyraliformes (the outgroup) have a nonlinear arrangement, this is considered plesio-
morphic. The alternative state in Crambiformes, stemmatal setae in a line with each
other, is apomorphic.

Mandible. Based on study of Pyralidae mandibles (Passoa 1985, Neunzig 1987, Peterson
1962), presence of a dentate ridge under the first scissorial tooth is an unusual modification.
Inner teeth are sometimes present on the first molar ridge, especially in Pyraustinae
(Peterson 1962, Passoa 1985), but in the latter case they do not form a ridge. Two character
states occur in Crambiformes: ridge absent or present. Since all Pyraliformes lack a ridge
(Passoa 1985), this is plesiomorphic. A dentate ridge, the alternative state, is apomorphic.

Thoracic V1 seta. Hinton (1946) stated V1 was present on all first and last instar
Lepidoptera he examined. In Crambiformes, two character states occur: V1 absent (Yo-
shiyasu 1985) or present (Passoa 1985). Since V1 is present in Pyraliformes (Passoa 1985),
this is plesiomorphic. Therefore, loss of this seta is considered apomorphic.

Rothschild (1967) speculated V1 may not be lost in Tryporyza (Schoenobiinae) but
instead could have migrated to the coxae as in some Tineidae and Psychidae (Hinton
1946). The extreme reduction in body setal length of Nymphulinae and Schoenobiinae
(setae may be difficult to see even under a compound microscope), coupled with lack of
knowledge about coxal setae and their homologies, makes evaluation of Rothschild’s
hypothesis impossible at present. In any event, either case would be apomorphic as V1
is not found on the coxa in the outgroup (Pyraliformes).

Prothoracic membranous sac. The Schoenobiinae membranous sac is apparently a
unique structure not homologous to other lepidopteran cervical glands (Passoa & Habeck
1987). In Crambiformes, two character states occur: prothoracic sac present or absent.
All Pyraliformes lack a prothoracic sac (Passoa 1985). Therefore, presence of a membra-
nous prothoracic sac is apomorphic.

L2 seta on abdominal segments. Hinton (1946) remarked that Ll and L2 are mac-
roscopic and frequently subequal in length throughout Lepidoptera. This is true for all
Pyraliformes and Crambiformes except Schoenobiinae (Hasenfuss 1960) and Nymphul-
inae (Neunzig 1987, Yoshiyasu 1985). Therefore, when L1 and L2 are subequal in length,
this is plesiomorphic. A very short, almost microscopic, abdominal L2 seta is apomorphic.

252 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Thoracic L seta. Neunzig (1987) noted that all Pyraliformes have three setae in the L
group of mesothorax and metathorax. In Crambiformes, two character states occur: L
group bisetose (some Nymphulinae and Schoenobiinae) or L group trisetose (most Cram-
biformes). Therefore, loss of the thoracic L seta is considered apomorphic.

L2 on AQ. All Pyraliformes have L1, L2, and L3 present on A9, whereas L2 is either
present (Schoenobiinae and Nymphulinae) or absent (most Crambiformes) in other pyralid
larvae (Neunzig 1987, Hasenfuss 1960). L1 is always present in Pyralidae while L3 is
always absent in Crambiformes. Because outgroup comparison demands a character
distribution in which a feature is present or absent in the group being studied, L2 is the
only seta that can be polarized at present. Since L2 is present in the outgroup, the
plesiomorphic state within Crambiformes occurs when L2 is present (bisetose condition).
In contrast, the apomorphic state occurs when L2 is lost (unisetose condition).

Yoshiyasu (1985) also considered loss of L setae in Crambiformes to be apomorphic
but he polarized both bisetose and unisetose conditions as apomorphies. Unfortunately,
this idea cannot be confirmed by outgroup comparison until more information is available
on the sister group of Pyralidae. If the unisetose condition is apomorphic, the bisetose
condition may be part of a trend from trisetose (plesiomorphic state) to a unisetose L
group on AQ.

Extra pinacula. When present, pinacula are located only around setal bases in Pyral-
iformes and most other Lepidoptera (Hinton 1946, Passoa 1985). In Crambiformes, two
character states occur. There may be extra pinacula (apparently lacking setae) on the
thorax and abdomen of Crambinae, a few Pyraustinae, and Scopariinae (MacKay 1972,
Passoa 1985) while extra pinacula are absent in Nymphulinae and Schoenobiinae. There-
fore, development of secondary pinacula is considered apomorphic. When extra pinacula
are lacking, this is plesiomorphic.

Pupal Characters

Frontal setae. Frontal setae are about as thick as other body setae in Pyraliformes
(Passoa 1985). In Crambiformes, they are thin in all subfamilies except Nymphulinae
(Passoa 1985) and several described Musotiminae (Nakamura 1977, for example). There-
fore, thin setae are plesiomorphic while thick spinelike frontal setae are apomorphic.

Mesothoracic spiracle. Outgroup comparison is of limited value here since both clades
have equal character distributions. In Pyraliformes, all subfamilies except Galleriinae and
some Phycitinae have a mesothoracic spiracle (Passoa 1985). Among Crambiformes, all
subfamilies except Nymphulinae have a mesothoracic spiracle. Loss of the mesothoracic
spiracle is considered apomorphic by parsimony since three independent losses (Nym-
phulinae, Galleriinae, and some Phycitinae) is a more likely evolutionary scenario than
independent gain of this spiracle many times in other pyralid subfamilies. Moreover,
Mosher (1916) found a mesothoracic spiracle on nearly all other Lepidoptera studied.
This supports the contention that a mesothoracic spiracle was probably present in ancestors
of Pyralidae.

No Pyraliformes examined during this study have a deep pitlike mesothoracic spiracle.
In Crambiformes, all subfamilies except Schoenobiinae lack a deep pit. Therefore, a
pitlike mesothoracic spiracle is considered apomorphic while absence of a pitlike meso-
thoracic spiracle is plesiomorphic.

It should be noted that some Pyraustinae (for example, Spoladea and Asciodes) have
pits adjacent to the mesothoracic spiracle while a few Epipaschiinae have the spiracle set
in a shallow concavity. This should not be confused with the situation in Schoenobiinae
where only a deep pit can be found and no trace of the spiracle is visible inside the pit.

Anterior abdominal spiracles on Al-3. All Pyraliformes lack enlarged anterior ab-
dominal spiracles set on conelike projections (Passoa 1985). In Crambiformes, two char-
acter states exist. Nearly all species of Crambiformes except Nymphulinae (Speidel 1984)
and Thopeutis forbesellus (Fernald) (Crambinae) lack enlarged anterior abdominal spi-
racles set on conelike projections. Therefore, lack of enlarged anterior abdominal spiracles
is plesiomorphic while their presence on conelike projections is apomorphic. Speidel (1981)

VOLUME 42, NUMBER 4 253

also considered enlarged anterior abdominal spiracles of Nymphulinae pupae to be apo-
morphic.

Posterior spiracles. All Pyraliformes examined during this study have anterior and
posterior spiracles subequal in diameter. In Crambiformes, two character states exist.
Most species, except Nymphulinae and a few Pyraustinae, have spiracles subequal in
diameter throughout the abdomen. This is considered plesiomorphic. Reduced posterior
spiracles are considered apomorphic.

Mesothoracic and metathoracic coxae. All Pyraliformes and all Crambiformes except
Schoenobiinae have hidden mesothoracic and metathoracic coxae. Thus, exposed meso-
thoracic and metathoracic coxae are apomorphic while hidden coxae are plesiomorphic.
Davis (1986) noted that only the forecoxa is exposed in higher Lepidoptera, and thus he
considered exposed mesothoracic coxae to be apomorphic.

Metathoracic legs. All Pyraliformes have obtect appendages; the metathoracic legs, if
not hidden, have only their tips exposed. This is also true of most Crambiformes, except
Nymphulinae and Schoenobiinae which have exarate appendages with metathoracic legs
clearly exposed. Fully exposed metathoracic legs and exarate appendages are considered
apomorphic while partially hidden metathoracic legs are plesiomorphic.

Adult Characters

Proboscis. Most pyralids have the proboscis well developed and scaled but some Cram-
biformes (Schoenobiinae) and Pyraliformes (Peoriinae) lack a proboscis (Munroe 1972).
This character distribution (present or absent in each clade) limits the usefulness of
outgroup comparison. Instead, a reduced proboscis is considered apomorphic by parsi-
mony since two independent reductions are more likely than many acquisitions.

Forewing CuP. Forewing CuP is another difficult character to polarize by outgroup
comparison since it may be either a fold or a tubular remnant in each clade of Pyralidae
(E. G. Munroe pers. comm.). Perhaps a fully developed vein was gradually lost until only
a tubular remnant remained at the distal end of the forewing. This reduction of CuP
continued so only a fold now marks its former position. Since Common (1970) noted a
trend in higher Lepidoptera where anal and radial veins are gradually lost in advanced
forms, reduction of CuP to a fold is tentatively called apomorphic. Further studies on
Pyraloidea ancestors would help polarize this character, but dugeoneids, which Minet
(1982) believed could be the sister group of the Pyralidae, have CuP developed.

Another possibility, independent reacquisition of CuP in Schoenobiinae, some Nym-
phulinae and some Pyraliformes, is equally parsimonious with the reduction of CuP in
most Crambiformes, most Pyraliformes, and some Nymphulinae. CuP reduced to a fold
would be plesiomorphic while gain of a tubular remnant would be apomorphic. This
polarization of CuP is especially attractive if morphological studies show the sister group
of Pyralidae is not Dugeoneidae (dugeoneids have CuP developed).

Praecinctorium. The praecinctorium is either present (Crambiformes) or absent (Py-
raliformes) in Pyralidae. Dugeoneidae, a tentative sister group of Pyralidae, lacks a prae-
cinctorium. Thus, presence of a praecinctorium is apomorphic whereas its absence is
plesiomorphic.

Acentria probably lost the praecinctorium secondarily because it may be vestigially
present in the tympanum (Minet 1985). Given presence of a praecinctorium as a ground-
plan apomorphy of Crambiformes, absence or extreme reduction of praecinctorium must
be an apomorphic reversal.

Tegumen-vinculum plate. All Pyraliformes lack the t-v plate (Yoshiyasu 1985). This
is also true for all Crambiformes except Nymphulinae and Schoenobiinae (Yoshiyasu
1985). Therefore, presence of the t-v plate is considered apomorphic.

Cu hindwing pecten. Munroe (1972) noted that cubital pecten occurs in both Cram-
biformes and Pyraliformes, and this limits outgroup comparison as a method of analysis.
However, parsimony would indicate that several independent gains of cubital pecten are
more likely than numerous losses. This suggests that presence of pecten is apomorphic
while its absence is plesiomorphic. Roesler (1973) also considered presence of pecten to
be apomorphic.

254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

BIOLOGY

Aquatic habitat. Nearly all Pyraliformes are terrestrial, and do not form cases entirely
of leaf fragments. This is true of all Crambiformes except Nymphulinae. When restricted
to exclude Musotiminae, Nymphulinae include species which are always aquatic and
frequently form cases. Thus, aquatic habit is apomorphic while terrestrial living is ple-
siomorphic. Speidel (1981) also considered aquatic living to be apomorphic.

TAXONOMIC AFFINITIES OF ACENTRIA

The above characters and their polarities provide additional infor-
mation on the systematic position of Acentria ephemerella. Schoeno-
biinae larvae are apomorphically defined by a membranous prothoracic
sac (Hasenfuss 1960, Passoa & Habeck 1987), which is absent from
Acentria (Yoshiyasu 1985). Acentria also lacks the pitlike mesothoracic
spiracle and exposed mesothoracic coxae (Figs. 2, 3) that apomorphi-
cally define Schoenobiinae pupae (Passoa & Habeck 1987). Therefore,
immature stages of Acentria demonstrate the genus is misplaced in
Schoenobiinae. Hampson (1895) and Forbes (1938) claimed affinity
between A. ephemerella and Schoenobiinae because of a reduced pro-
boscis, tubular forewing CuP, and absence of hindwing cubital pecten.
Lack of cubital pecten is plesiomorphic; thus absence of this feature
does not indicate relation (individuals sharing symplesiomorphies may
not be relatives). The tubular forewing CuP may be apomorphic, but
this character is found in both Nymphulinae and Schoenobiinae (Mun-
roe 1972), and thus does not clarify the systematic position of Acentria.
The single apomorphic adult character that Acentria and Schoenobiinae
have in common, a reduced proboscis, perhaps arose through conver-
gence since both taxa are associated with a similar (moisture-rich) aquat-
ic environment. Although a reduced proboscis is usually considered
characteristic of Schoenobiinae (Forbes 1938), some Nymphulinae also
have the proboscis reduced (Yoshiyasu 1985), so a species with reduced
mouthparts could be a member of either subfamily. Adult morphology,
like that of immatures, provides little evidence that Acentria belongs
in Schoenobiinae.

As mentioned earlier, there may be strong selection for enlargement
of anterior abdominal spiracles in pupae of aquatic pyralids. These
spiracles were considered autapomorphic for Nymphulinae (Speidel
1984), but they also occur in some aquatic Crambinae of Asia (Yoshiyasu
1985), Thopeutis forbesellus (Fernald) of the United States, and a few
terrestrial Pyraustinae genera such as Lygropia, Microthyris, Spoladea,
and Marasmia (Passoa 1985). Nevertheless, other pupal apomorphies
indicate Acentria is related to Nymphulinae. Enlarged spinelike frontal
setae are found on most Nymphulinae pupae (Yoshiyasu 1985), and are
apomorphic for this subfamily. Acentria has these enlarged setae (Figs.

VOLUME 42, NUMBER 4 290

Fics. 2, 3. 2, Ventral view of Acentria ephemerella pupa. Scale line = 0.8 mm. 3,
Dorsal view of Acentria ephemerella pupal antenna and thorax. Scale line = 0.25 mm.

4, 5) which indicates a close relation to Nymphulinae. In addition, very
few pyralid subfamilies (Galleriinae, Nymphulinae, and some Phyci-
tinae) lack a mesothoracic spiracle (Passoa 1985). Among Crambi-
formes, only Nymphulinae show this loss. Acentria has no mesothoracic
spiracle (Fig. 3) and, as is typical for Nymphulinae, has enlarged an-
terior abdominal spiracles set on conelike projections (Fig. 2). This
spiracular arrangement, when combined with much reduced posterior
spiracles, is autapomorphic for Nymphulinae. Thopeutis forbesellus
(Crambinae) has anterior abdominal spiracles set on weak conelike
projections, but the abdominal spiracles are all equal in diameter. Some
Pyraustinae have enlarged anterior abdominal spiracles (Passoa 1985),
but unlike Nymphulinae, lack conelike projections and have posterior
spiracles at least half the diameter of anterior ones. These examples
show, as Yoshiyasu (1985) suspected, that convergence has produced
enlarged spiracles and conelike projections in other Crambiformes.

256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 4,5. 4, Micrograph of Acentria ephemerella labrum, pillifers, frons, and vertex.
Arrow points to spinelike frontal seta. Scale line = 100 microns. 5, Micrograph of Acentria
ephemerella spinelike frontal seta. Scale line = 10 microns.

However, it still remains possible to define Nymphulinae pupae easily
by their frontal setae and other spiracular modifications.

Besides the aquatic habit (Nigmann 1908) and stemmatal setal ar-
rangement (Hasenfuss 1960), another larval apomorphy may help re-
solve the systematic position of Acentria ephemerella. Many Nymphul-
ini have a dentate ridge on the mandible (Yoshiyasu 1985, “semicircular
arrangement” of Lange 1956) which also occurs in Acentria (Yoshiyasu
1985). This contrasts with Argyractini larvae which have the mandible
more elongated, flattened, and usually without the dentate ridge (Lange
1956). Other characters (Lange 1956, Speidel 1984) such as diet of
submerged plants, prothoracic shield chaetotaxy, ability to make cases
of leaf fragments, lack of gills on body, lack of palmate setae on labrum,
and three enlarged pupal spiracles would indicate A. ephemerella lacks
apomorphies of Argyractini and belongs in Nymphulini as defined by
Lange (1956). Speidel (1984) did not use mandibles, pupal spiracles, or
labral setae in his Nymphulinae cladogram. Since the tribal classification
proposed by North American workers can be difficult to apply to certain
Asiatic genera, for example Nymphicula (Yoshiyasu 1980), these fea-
tures merit further attention.

In summary, Acentria is misplaced in Schoenobiinae because im-
mature stages radically differ. In spite of some morphological special-
izations, there seems little reason to consider this genus separate from
Nymphulinae. Placement of Acentria in its own family or subfamily
was based, in part, on lack of a praecinctorium which is unusual among
Crambiformes. Minet (1985), while studying the tympanum, found a
possible praecinctorium vestige, and thus placed Acentria in Nymphul-
inae. No characters in immature stages were found to exclude Acentria
from Nymphulinae as a separate taxon, although crochet arrangement

VOLUME 42, NUMBER 4 257

is somewhat unusual. Since differences between the tympanum of Acen-
tria and other nymphulines may not be as great as previously thought,
and several additional larval and pupal apomorphies confirm its relation
to Nymphulinae and exclude it from known Schoenobiinae, there seems
little doubt that transfer of Acentria to Nymphulinae by Hasenfuss
(1960) was correct.

It is worth noting that Neoschoenobia decoloralis Hampson, another
disputed taxon placed in Nymphulinae (Inoue 1982, cited by Yoshiyasu
1985) and Schoenobiinae (Lewvanich 1981), might be a member of
Schoenobiinae because it has exposed pupal coxae and lacks enlarged
pupal spiracles and stemmatal setae in a straight line. Since illustrations
by Yoshiyasu (1985) do not show a mesothoracic spiracle or a larval
prothoracic sac, it seems wise to retain this species in Nymphulinae,
although preserved specimens should be examined for these features.

RELATION BETWEEN NYMPHULINAE
AND SCHOENOBIINAE

Historically, the systematic position of Schoenobiinae has been de-
bated. Borner (cited by Munroe 1958) thought Crambinae and Schoe-
nobiinae were close relatives. Roesler (1973) considered them unrelated
based on maxillary palpi and cubital pecten. Kuznetsov and Stekolnikov
(1979) included Crambinae, Schoenobiinae, and Nymphulinae as the
most primitive members of their Crambidae.

Larval and pupal features indicate Crambinae and Schoenobiinae
are not closely related phenetically or cladistically. Crambinae larvae
have a unisetose L group on AQ, and well developed extra pinacula on
both thorax and abdomen (Passoa 1985, Tan 1984). Schoenobiinae lar-
vae, in contrast, frequently have a bisetose L group on AQ and no
pinacula (Passoa & Habeck 1987). Pupal structure is also radically
different. Crambinae pupae either have a well developed cremaster
(Crambini) or processes on the head or body (Chilini). Metathoracic
legs are not exposed or are barely visible. Schoenobiinae pupae, in
contrast, always have exposed metathoracic legs, and never have a
cremaster or appendages on the head or body. In fact, it is difficult to
find any synapomorphies in immature stages to link these two groups.

Immature stages do support the hypothesis of Passoa (1985) and
Yoshiyasu (1985) that Schoenobiinae and Nymphulinae are related.
Bollman (1955) and Allyson (1976) distinguished Schoenobiinae by their
reduced L2 seta, but minute L setae are common on many Nymphulinae
(Yoshiyasu 1985, Neunzig 1987). Additional apomorphies to unite
Schoenobiinae and Nymphulinae include fully exposed metathoracic
legs and exarate appendages. Other synapomorphies listed by Yoshiyasu
(1985) are V1 lost on the larval thorax, and absence of transtilla with

258 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

development of the t-v plate in male genitalia. One exception is Rupela
albinella (Passoa & Habeck 1987) which has V1 present, but this may
merely represent a reversion to the primitive state. All other known
species in both subfamilies lack V1, so loss of this seta is probably a
groundplan apomorphy. Finally, several characters merit further in-
vestigation as synapomorphies of the Nymphulinae-Schoenobiinae clade.
These are mesothoracic pupal spiracle (does the pit in Schoenobiinae
contain a spiracle, or is the pit the spiracle itself); absence of pupal
cremaster (unknown polarity); L setae of thorax bisetose (apomorphic
but its distribution within the clade needs study); loss of pinacula (un-
known polarity); and CuP tubular at margin (unknown polarity). In
addition, a bisetose L group on A9 was thought characteristic of only
Nymphulinae (Yoshiyasu 1985, Hasenfuss 1960, Bollman 1955) but this
condition is also found in several Schoenobiinae genera (Passoa & Ha-
beck 1987). Some illustrations show a unisetose L group on AQ in
Schoenobiinae (Hasenfuss 1960) but these probably represent cases where
L2 was overlooked. Chaetotaxy of Schoenobiinae larvae is difficult to
study without slide mounts of larval skin. Further study may also show
the bisetose L group on AQ is a synapomorphy of the two subfamilies.

RELATIONS OF NYMPHULINAE—SCHOENOBIINAE
CLADE IN CRAMBIFORMES

Relation of the Nymphulinae-Schoenobiinae clade to other subfam-
ilies has been unclear. Yoshiyasu (1985) defined a clade uniting all
Crambiformes, except Pyraustinae and its relatives, by a reduced tran-
stilla. However, certain exceptions to this generalization limit its use as
a synapomorphy. Yoshiyashu (1985) characterized Pyraustinae and Gla-
phyriinae by their well developed transtilla, but E. G. Munroe (pers.
comm.) stated that the transtilla varies widely in these groups. One
alternative to a clade defined by transtilla morphology, with far fewer
exceptions, involves L setae on A9. The Nymphulinae-Schoenobiinae
clade is separated from remaining Crambiformes by the number of L
setae on AY. Other Crambiformes subfamilies, without exception, have
a unisetose L group on AQ (loss of seta L2 is an apomorphy), which
defines them as a monophyletic group. This seta is present (bisetose
condition) in nearly all Nymphulinae (restricted sense) and Schoeno-
biinae larvae. Although the above phylogeny accepts some parallel
evolution with the presence of a unisetose L group in a single Nym-
phulinae species (Yoshiyasu 1985) and in published figures of some
Schoenobiinae (if these figures are correct), this represents only a very
small number of species. Parallel evolution appears to be normal in the
evolution of both Macrolepidoptera (Michener 1949) and Microlepi-

VOLUME 42, NUMBER 4 259

doptera (Kristensen 1984), so perhaps pyralids have also followed this
trend. It seems unrealistic to expect a group with thousands of species
to be defined by a single trait without parallelisms, so choice of a clade
based on the L setae may represent the case with minimum homoplasy.
Use of the unisetose L group on AQ as a synapomorphy supports Minet’s
(1982) contention that Midiliformes belong in Crambiformes since a
Midila larva in the U.S. National Museum has a unisetose L group on
A9. Moreover, separation of Musotiminae from Nymphulinae is sup-
ported by the fact that Musotima has a unisetose L group on AQ
(Nakamura 1977) unlike the bisetose L group of other Nymphulinae
(Hasenfuss 1960).

In conclusion, this study calls attention to the role of immature insects
on Pyralidae classification and phylogeny. Modifications of pupae are
especially diverse and in need of study. Future studies will probably
use more larval and pupal characters, especially if the sister group of
Pyralidae can be confirmed.

ACKNOWLEDGMENTS

Comments of G. Godfrey and M. Berenbaum on a draft of this manuscript were most
helpful. This study could not have been completed without a gift of Acentria immatures
from G. Buckingham, or German translations by S. Davis. Gerasminov’s papers were
translated by L. Khotko. E. MacLeod, W. LaBerge, S. Heydon, and A. Solis advised me
on cladistic methodology often. Scanning electron micrographs were provided by S.
Heydon. R. Woodruff, T. Harrison, and J. Apperson helped with figures. I especially
thank E. Munroe who provided direction and ideas. The following individuals and curators
loaned specimens: S. Allyson (Canadian National Collection); R. Hodges (United States
National Museum); D. Habeck (University of Florida Collection, Florida State Collection
of Arthropods); G. Godfrey and A. Brigham (Illinois Natural History Survey); and S.
Frommer (University of California, Riverside).

REFERENCES CITED

AGARWAL, R. A. & J. P. CHAUDHRY. 1966. Larval pinaculi as an identifying character
of crambids with special reference to sugarcane borers. Punjab Agr. Univ. J. Res. 3:
301-312.

ALLYSON, S. 1976. North American larvae of the genus Loxostege Hubner (Lepidoptera:
Pyralidae: Pyraustinae). Can. Entomol. 108:89-104.

1981. Last instar larvae of Pyraustini of America north of Mexico. Can. Entomol.

113:463-518.

1984. Description of last-instar larvae of 22 species of North American Spilo-
melini (Lepidoptera: Pyralidae: Pyraustinae) with a key to species. Can. Entomol.
116:13801-1334.

BaTRA, S. W. T. 1977. Bionomics of the aquatic moth, Acentropus niveus (Olivier), a
potential biological control agent for Eurasian watermilfoil and hydrillia. J. N.Y.
Entomol. Soc. 85:1438-152.

BERG, K. 1942. Contributions to the biology of the aquatic moth Acentropus niveus
(Oliv.). Vidensk. Medd. Dansk. Naturh. Foren. 105:59-189.

BOLLMANN, H. G. 1955. Die Raupen mitteleuropdischer Pyraustinae (Leidoptera—
Pyralidae). Beitr. Entomol. 5:521-639.

260 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

BRENIERE, J. 1979. Reconnaissance des principaux lépidoptéres du riz de L’Afrique de
L’Ouest. Agron. Tropicale 31:213-231.

BUCKINGHAM, G. R. & B. M. Ross. 1981. Notes on the biology and host specificity of
Acentria nivea (= Acentropus niveus). J. Aquat. Plant Manage. 19:32-36.

CLARK, C. & D. J. CURRAN. 1986. Outgroup analysis, homoplasy, and global parsimony:
A response to Maddison, Donoghue, and Maddison. Syst. Zool. 35:422—426.

Common, I. B. F. 1970. Lepidoptera. In Waterhouse, D. F. (ed.), The insects of Australia.
Melbourne Univ. Press. 1029 pp.

CRAWFORD, C. S. 1961. The bionomics of destructive Microlepidoptera of grass fields.
Ph.D. Diss., Washington State Univ., Pullman. 140 pp. (Diss. Abs. 22(11):4120, Order
No. 62-926.)

Davis, D. R. 1986. A new family of monotrysian moths from austral South America
(Lepidoptera: Palaephatidae), with a phylogenetic review of the Monotrysia. Smiths.
Contrib. Zool. 434, 202 pp.

DONOGHUE, M. J. & P. D. CANTINO. 1984. The logic and limitations of the outgroup
substitution approach to cladistic analysis. Syst. Bot. 9:192-202.

Farris, J. S. 1982. Outgroups and parsimony. Syst. Zool. 31:328-334.

FLETCHER, D. S. & I. W. B. Nye. 1984. The generic names of moths of the world.
Vol. 5. Pyraloidea. British Museum (Nat. Hist.), London. 185 pp.

ForBEs, W. T. M. 1926. The relationships of some aberrant pyralids (Lep.). J. N.Y.
Entomol. Soc. 34:331-336.

1938. Acentropus in America (Lepidoptera: Pyralididae). J. N.Y. Entomol. Soc.
46:338.

FRACKER, S. B. 1915. The classification of lepidopterous larvae. Ill. Biol. Monogr. 2:1-
161.

GERASIMOV, A. M. 1947. Larvae and pupae of pyralids (Pyralididae: Lepidoptera). I
(Russian). Entomol. Obozr. 29:165-181.

1949. Larvae and pupae of pyralids (Lepidoptera: Pyralididae). II (Russian).
Entomol. Obozr. 30:365-369.

GoaTER, B. 1986. British pyralid moths. A guide to their identification. Harley Books,
Great Horklesley, England. 175 pp.

Hampson, G. F. 1895. On the classification of the Schoenobiinae and Crambinae, two
subfamilies of moths, of the family Pyralidae. Proc. Zool. Soc. Lond. 1895:897-974.

HASENFuss, I. 1960. Die Larvalsystematik der Zunsler (Pyralidae). Abh. Larvalsyst.
Insekten 5. Akademie-Verlag, Berlin. 263 pp.

HINTON, H. E. 1946. On the homology and nomenclature of the setae of lepidopterous
larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Roy. Entomol.
Soc. Lond. 97:1-87.

KHoT ko, E. I. & R. V. MOLCHANOVA. 1975. On the morphology of the immature
stages of some species of the subfamily Pyraustinae (Lepidoptera: Pyralidae). Ento-
mol. Rev. 53:105-112.

KRISTENSEN, N. P. 1984. Studies-on the morphology and systematics of primitive Lep-
idoptera (Insecta). Steenstrupia 10:141-191.

KUZNETZOV, V. I. & A. A. STEKOLNIKOV. 1979. Classification and phylogenetic rela-
tionships of the families and subfamilies of the Pyraloidea (Lepidoptera) of the
Palearctic fauna with regard of functional morphology of the male genitalia (Russian).
Trudy Zool. Inst. Leningr. 82:43-74.

LANGE, W. H. 1956. A generic revision of the aquatic moths of North America: (Lep-
idoptera: Pyralidae: Nymphulinae). Wasmann J. Biol. 14:59-144.

LERAUT, P. 1980. Liste systematique et synonymique des Lépidoptéres des France,
Belgique, et. Corse. Suppl. Alexanor Bull. Soc. Entomol. France. 334 pp.

LEWVANICH, A. 1981. A revision of the Old World species of Scirpophaga (Lepidoptera:
Pyralidae). Bull. Brit. Mus. (Nat. Hist.) (Entomol.) 42:185-298.

MacKay, M. R. 1972. Larval sketches of some Microlepidoptera, chiefly North Amer-
ican. Mem. Can. Entomol. 88, 83 pp.

MARION, H. 1954. Revision des Pyraustidae de la faune francaise (suite). Rev. Franc.
Lepid. 14:181-188, 221-227.

VOLUME 42, NUMBER 4 261

MaTHurR, R. N. 1954. Immature stages of Indian Lepidoptera No. 9—Pyralidae, sub-
family Pyraustinae. Indian For. Rec. (N.S.) 8(11):241-265.

1959. Immature stages of Indian Lepidoptera No. 12—Pyralidae, sub-family
Pyraustinae. Indian For. Rec. (N.S.) 9(10):183-210.

MATHUuR, R. N. & P. SINGH. 1963. Immature stages of Indian Lepidoptera No. 13—
Pyralidae, sub-family Pyraustinae. Indian For. Rec. (N.S.) 10(6):117-148.

Mauston, G. W. 1970. The Crambinae of North Dakota with descriptions of some
previously undescribed immature forms. Ph.D. Diss., North Dakota State Univ.,
Fargo. 107 pp. (Diss. Abs. Int. 31(6):3460 B, Order No. 70-25,501.)

MICHENER, C. D. 1949. Parallelisms in the evolution of the saturniid moths. Evolution
3:129-141.

MINET, J. 1982 (1981). Les Pyraloidea et leurs principales divisions systematiques (Lep:
Ditrysia). Bull. Soc. Entomol. France 86:262-280.

1983. Etude morphologique et phylogénétique des organes tympaniques des

Pyraloidea. I. Généralités et homologies (Lep. Glossata). Ann. Soc. Entomol. France

19:175-207._

1985. Etude morphologique et phylogénétique des organes tympaniques des
Pyraloidea. 2—Pyralidae; Crambidae, primiére partie. (Lepidoptera Glossata). Ann.
Soc. Entomol. France 21:69-86.

MosHER, E. 1916. A classification of the Lepidoptera based on characters of the pupa.
Bull. Ill. State Lab. Nat. Hist. 12:1-159.

MUNROE, E. 1958. Hampson’s Schoenobiinae (Lepidoptera: Pyralidae). Proc. Tenth
Intern. Cong. Entomol. (1956) 1:301-302.

1972. Pyraloidea. Pyralidae (part). In Dominick, R. B., et al. (eds.), The moths

of America north of Mexico. Fasc. 18.1A. E. W. Classey, London. 134 pp.

1981. Lepidoptera, pp. 254-257. In Hurlbert, S., G. Rodriguez & N. D. Dos

Santos (eds.), Aquatic biota of tropical South America. Part I. Arthropoda. San Diego

State Univ., San Diego, California. 342 pp.

1982. Lepidoptera, pp. 401-405. In Hurlbert, S. & A. Villalobos-Figueroa (eds.),

Aquatic biota of Mexico, Central America and the West Indies. San Diego State

Univ., San Diego, California. 529 pp.

1983. Pyralidae. In Hodges, R. W., et al. (eds.), Check list of the Lepidoptera
of America north of Mexico. E. W. Classey Ltd. & Wedge Entomological Research
Foundation, London. 284 pp.

NAKAMURA, M. 1977. Notes on the immature stage of Musotina acclaralis Walker (Lep:
Pyralidae). Kita-Kyushu no Konchu 23:33-36.

1981. Key to the classification of the Japanese lepidopterous pupae. Tyo to Ga
32:1-12.

NEUNZzIG, H. H. 1987. Pyralidae (Pyraloidea), pp. 462-494. In Stehr, F. W. (ed.),
Immature insects. Kendall/Hunt, Dubuque, Iowa. 754 pp.

NIGMANN, M. 1908. Anatomie und Biologie von Acentropus niveus Oliv. Zool. Jahrb.
Abt. Syst. 26:489-560.

Passoa, S. 1985. Taxonomy of the larvae and pupae of economically important Pyralidae
in Honduras. M.S. Thesis, Univ. Florida, Gainesville. 486 pp.

Passoa, S. & D. H. HaBeEcK. 1987. A description of the larva and pupa of Rupela
albinella (Cramer), a pest of rice in Latin America (Pyralidae: Schoenobiinae). Fla.
Entomol. 70:368-375.

PETERSON, A. 1962. Larvae of insects. An introduction to the Nearctic species. Part I.
Lepidoptera and plant infesting Hymenoptera. Edwards Bros., Ann Arbor, Michigan.
315 pp.

ROESLER, R. U. 1973. Phycitinae. In Amsel, H. G., F. Gregor & H. Reisser (eds.),
Microlepidoptera Palaearctica. Vol. 4. G. Fromme, Vienna. 752 pp.

ROTHSCHILD, G. H. L. 1967. Descriptions of larval and pupal stages of four lepidopterous
rice borers in Malaysian Borneo (Sarawak). Bull. Entomol. Res. 57:343-352.

SPEIDEL, W. 1981. Die Abgrenzung der Unterfamilie Acentropinae (Lepidoptera, Pyr-
aloidea). Atalanta 12:117-128.

262 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1984. Revision der Acentropinae des palaearktischen Faunengebietetes (Lepi-
doptera, Crambidae). Neue Entomol. Nachr. 12:1-157.

STEHR, F. W. 1987. Immature insects. Kendall/Hunt, Dubuque, Iowa. 754 pp.

TAN, C. L. 1984. Biology and taxonomy of sod webworms (Lepidoptera: Pyralidae)
destructive to turfgrasses in Florida. M.S. Thesis, Univ. Florida, Gainesville. 121 pp.

Watrous, L. E. & Q. D. WHEELER. 1981. The outgroup comparison method of char-
acter analysis. Syst. Zool. 30:1-11.

YoOsHIyasu, Y. 1980. A systematic study of the genus Nymphicula Snellen from Japan
(Lepidoptera: Pyralidae). Tyo to Ga 31:1-28.

1985. A systematic study of the Nymphulinae and Musotiminae of Japan (Lep-

idoptera: Pyralidae). Sci. Reports Kyoto Pref. Univ., Agr. 37:1-162.

Received for publication 4 December 1987; accepted 28 April 1988.

Journal of the Lepidopterists’ Society
42(4), 1988, 263-268

EFFECT OF LARVAL PHOTOPERIOD ON MATING AND
REPRODUCTIVE DIAPAUSE IN SEASONAL FORMS OF
ANAEA ANDRIA (NYMPHALIDAE)

THOMAS J. RILEY

Department of Entomology, Louisiana Agricultural Experiment Station,
Louisiana State University Agricultural Center, Baton Rouge, Louisiana 70803

ABSTRACT. In experiments conducted under simulated field conditions in Baton
Rouge, Louisiana, laboratory-reared summer-form female Anaea andria Scudder from
13- and 14-h larval photoperiods underwent oocyte maturation and mated within two
days of eclosion. Laboratory-reared winter-form females from 13-h larval photoperiods
did not mate, and showed little oogenesis 20 days after eclosion in simulated field con-
ditions. Data from field-collected specimens numbering 55 winter forms and 26 summer
forms support the experimental results, and indicate that female winter forms remain
unmated and in reproductive diapause in the fall. Results suggest that larval daylength,
by determining adult seasonal form, also regulates reproductive diapause and mating in
A. andria.

Additional key words: Charaxinae, Croton capitatus, seasonal dimorphism.

Anaea andria Scudder (Nymphalidae: Charaxinae) is distributed from
Texas to Nebraska, E to West Virginia, Georgia, and the Florida pan-
handle (Opler & Kriezek 1984). It is common in the southern Mississippi
basin and Gulf Coast where it occurs with its primary host plant, Croton
capitatus Michx. (Euphorbiaceae), an annual herb.

Adult Anaea andria are characterized by distinct seasonal wing di-
morphism induced by larval photoperiod (Riley 1980, 1988). Winter-
form butterflies emerging in fall and surviving until the following spring
are characterized by apically acute and falcate forewings, well devel-
oped hindwing tails and anal angle projections, and brighter and more
contrasting coloration than summer-form butterflies. Summer forms
have non-falcate forewing apices, reduced tails and anal-angle projec-
tions on hindwings, and lighter overall coloration. Photoperiods of 14
h or more result primarily in summer-form adults. Decreasing photo-
periods result in a greater percentage of winter-form individuals (Riley
1988).

In Louisiana, summer-form A. andria occurs from May to September
when actively growing host plants are available. The winter form begins
to appear in late August, and survives until June of the following year.
Its appearance in the fall is followed shortly by the beginning of Croton
capitatus senescence.

The occurrence of two distinct seasonal forms, one when food plants
are abundant, and another when they are in decline, suggests that a
corresponding difference in female reproductive status may also occur.
In this paper, effects of larval daylength on reproductive diapause and

264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

mating in laboratory-reared seasonal forms of A. andria, and the mating
and reproductive status of field-collected seasonal forms are reported.

METHODS

Experimental insects were collected in East Baton Rouge and East
Feliciana parishes, Louisiana. Larvae were collected from host plants
in the field, or reared from eggs deposited on container-grown host
plants by reared and field-collected butterflies confined in 2-m* Saran®
screen outdoor cages (Chicopee Manufacturing Co., Cornelia, Georgia
30531). Adults were collected using traps similar to those described by
Rydon (1964) baited with bananas.

Larvae were reared in clear plastic 26 x 20 x 10-cm boxes containing
a raised 3-mm mesh metal grid to prevent undue larval contact with
feces. Larvae were fed fresh Croton capitatus leaves, and the boxes
cleaned daily or as needed. Pupation occurred on tops and sides of the
boxes and on plant material.

Seasonal forms of A. andria were produced by rearing the third instar
in environmental chambers with controllable photoperiods. To obtain
summer-form butterflies, 14 h of light was used. Winter and summer
forms were obtained using a 13-h photoperiod. Temperature in the
chambers was 27°C during the experiments.

Laboratory-reared males and females of each seasonal form were
maintained in separate outdoor cages. Eight hours after eclosion, fe-
males were numbered with permanent ink on the underside of a hind-
wing and released into the cages. Males were also caged eight hours
after eclosion, and remained in the cages until death. Virgin females
were always caged with males of equal or greater age.

Well-ripened bananas were provided for adult food. Cage location
provided exposure to full sun 6 h/day. One cage corner was covered
with plywood to provide a shaded resting area for the butterflies. The
experiment was conducted from 1 June to 81 October 1986; conse-
quently, insects were exposed to a changing natural photoperiod. Each
seasonal form was caged during the time of year when each can be
found in the field, summer forms from 1 June to 23 September, winter
forms from 6 August to 31 October.

Stage of oogenesis, number of mature eggs, and mating status were
determined by dissection and examination of the female reproductive
system. Summer forms were dissected 2 and 3 days after eclosion, and
winter forms 10 and 20 days after eclosion. Stage of oogenesis is de-
scribed using a scale of 0-5, 0 denoting no evident oocyte development,
and 5 the presence of mature eggs (further explained in Table 1). Insects
were judged to be in reproductive diapause if oocyte development 10

VOLUME 42, NUMBER 4 265

TABLE 1. Percentage mated, stage of oogenesis, and number of mature eggs/female
in 2- and 3-day-old mated and unmated laboratory-reared summer-form A. andria under
simulated field conditions.

No. mature eggs/female

Percent Mean stage of
Age (days) N mated! oogenesis”? Mean* Range
2 18 83.3a 4.la 0.6a 0-4
3 44 90.9a 4.9b 44.2b 0-82

1 Not significant according to Fisher’s Exact Test.

20-5 scale. 0 = no visible oocyte formation; 1 = beginnings of oogenesis; 2 = slight enlargement of oocytes; 3 = some
oocytes 50% mature; 4 = greater oocyte enlargement, no oocytes at median oviduct; 5 = mature oocytes at median
oviduct.

3 Means in columns followed by the same letter do not differ significantly according to F-test (P < 0.01).

days after eclosion was rated <2.0. Mated status of females was deter-
mined by spermatophore presence in the bursa copulatrix.
Percentage mating was analyzed using Fisher’s Exact Test; all other
variables were subjected to analysis of variance (SAS Institute 1985).
Voucher specimens are in the Louisiana State University Entomology
Museum.

RESULTS

Laboratory-reared butterflies. Winter-form butterflies resulted only
from the 18-h photoperiod. Twenty-seven females and 30 males were
reared. Twelve females were dissected after 10 days, and 15 dissected
after 20 days of caging with winter-form males. None of the female
winter forms had mated, and none of their ovarioles showed any sign
of oogenesis. No courtship behavior or mating attempts were seen. It
was therefore concluded that winter-form females remain in repro-
ductive diapause for at least 20 days after eclosion. Male mating be-
havior and female attractiveness may also be inhibited in winter forms.

Summer-form butterflies resulted from both photoperiods, 40 females
and 41 males from the 14-h, and 22 females and 25 males from the
13-h. Comparison of summer-form data from both photoperiods in-
dicates that oogenesis, number of mature eggs/female, and mating were
not significantly affected by these larval photoperiods.

Age was the most important factor affecting stage of oogenesis and
number of mature eggs per female in recently eclosed summer forms.
Two-day-old females showed significantly less oogenesis (F = 31.51; df
= 1,58; P < 0.01), and carried fewer mature eggs (F = 59.18; df = 1,
58; P < 0.01) than 3-day-olds (Table 1). The age x photoperiod in-
teraction was not significant.

Percentage of mated 2- and 3-day-olds did not differ (Table 1). In
several instances, courtship of virgin females was observed within hours
of their release into the outdoor cage. Although age of the males involved

266 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

was not known, it was concluded that mating behavior of male and
female summer forms is not suppressed in recently eclosed butterflies.
These observations suggest that most female summer forms are mated
within a few days of emergence.

Among unmated summer forms, four were three days old, and three
were two days old. Mean number of mature eggs/female for the 3-day-
olds was 23.5 (range 0-48) while none of the 2-day-olds contained
mature eggs, evidence that oogenesis in summer forms progresses with
age, independent of mating.

Field-collected butterflies. Forty-two winter-form females were
trapped during September-November 1984 and September 1985. They
showed little evidence of oogenesis, and all were unmated (Table 2).
Thirteen overwintered winter forms trapped in April and May 1985
were mostly mated and contained fully mature eggs (Table 2).

Summer-form females, collected in September 1984 and June to mid-
September 1985, all carried fully mature eggs and all were mated
(Table 2).

DISCUSSION

Laboratory results and field observations indicate that newly eclosed
summer-form females undergo rapid oogenesis and are mated within
two days of emergence. This enables summer forms to immediately
begin ovipositing and larvae to complete development before onset of
unsuitable environmental conditions. Conversely, winter-form females
remain unmated and in reproductive diapause for a considerable time
after adult eclosion. They are thus relieved of the physiological burden
of producing and carrying mature or maturing eggs when environ-
mental conditions are not favorable for oviposition and larval devel-
opment. Reproductive status appears to be linked to adult seasonal form
since summer forms from both 18- and 14-h photoperiods underwent
rapid oogenesis while winter forms from the 13-h photoperiod remained
in diapause.

Field observations of courtship, mating, and feeding behavior in A.
andria support these conclusions (unpubl. data). No courtship activity
has been seen in winter forms during fall. However, newly eclosed male
and female winter forms are readily attracted to fermented fruit baits,
indicating a possible feeding response linked to preparation for over-
wintering. In spring, winter-form males have been observed exhibiting
strong territorial behavior, chasing other males, patrolling along forest
edges and then returning to the same perch, engaging in courtship
behavior, and attempting to mate with females. Baits placed near male
territories and perches in spring have proven relatively unattractive
and trapping ineffective. These observations lend support to the ex-

VOLUME 42, NUMBER 4 267

TABLE 2. Collection month, percentage mated, and stage of oogenesis in female A.
andria collected 1984-85 in East Baton Rouge and East Feliciana parishes, Louisiana.

Percent Mean stage of
Month collected Seasonal form N mated oogenesis!
September Winter 18 0.0 0.0
October Winter 23 0.0 0.5
November Winter 1 0.0 Ie,
April Winter? 5 80.0 5.0
May Winter? 8 100.0 5.0
June though September Summer 26 109.0 5.0

1Same scale as in Table 1.
2 Overwintered butterflies.

perimental results and suggest different behavioral priorities in winter
forms before and after overwintering. Factors initiating oogenesis and
mating in winter forms that have overwintered are unknown.

My experience with bait traps during summer in Louisiana aud
Missouri, and that of Vernon Brou, Abita Springs, Louisiana, who op-
erates bait traps year-round, indicate that summer forms are both trapped
and collected less frequently than winter forms. They are also less
common in collections. The total number of field-collected summer
forms in collections of the author, V. Brou, and the Louisiana State
University Entomology Museum is 77 compared to 241 winter forms
(Riley 1988). This discrepancy may be due to collecting bias, but may
also indicate greater behavioral priority for reproduction vs. feeding in
summer forms, similar to winter-form behavior in the spring. These
observations along with the experimental results suggest that larval
daylength, by determining adult seasonal form, is also a major factor
regulating mating and reproduction in recently emerged A. andria.

Photoperiod is well documented as a diapause inducing and regu-
lating stimulus for insects (Beck 1980, Danilevsky et al. 1970, Tauber
et al. 1986). It is an ideal environmental cue for A. andria. The nym-
phalid Polygonia c-aureum L. is very similar in its response to daylength
(Hidaka & Aida 1968, Fukuda & Endo 1966, Endo 1970, 1972). In P.
c-aureum, reproductive diapause and seasonal wing dimorphism are
determined by photoperiod but are under independent neuroendocrine
control. Pheromone production and mating receptivity of female P.
c-aureum are also hormonally regulated and under photoperiodic con-
trol (Endo 1978). The results presented here suggest that a similar
interaction between daylength and neuroendocrine system could be
controlling wing dimorphism, diapause, and mating in A. andria.

ACKNOWLEDGMENT

Approved for publication by the Director of the Louisiana Agricultural Experiment
Station as manuscript No. 87-17-1490.

268 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

LITERATURE CITED

BECK, S. D. 1980. Insect photoperiodism. Academic Press, New York. 387 pp.

DANILEvsky, A. S., N. I. GORYSHIN & U. P. TYSHCHENKO. 1970. Biological rhythms in
arthropods. Ann. Rev. Entomol. 15:201-244.

ENbo, K. 1970. Relation between ovarian maturation and activity of the corpora allata
in seasonal forms of the butterfly, Polygonia c-aureum L. Devel. Growth Differ. 11:
297-304.

1972. Activation of corpora allata in relation to ovarian maturation in the seasonal

forms of the butterfly, Polygonia c-aureum L. Devel. Growth Differ. 14:263-274.

1973. Hormonal regulation of mating in the butterfly, Polygonia c-aureum L.
Devel. Growth Differ. 15:1-10.

FUKUDA, S. & K. ENDO. 1966. Hormonal control of the development of seasonal forms
of the butterfly, Polygonia c-aureum L. Proc. Japan Acad. 42:1082-1087.

Hipaka, T. & S. AIDA. 1963. Daylength as the main factor of seasonal form determi-
nation in Polygonia c-aureum (Lepidoptera, Nymphalidae). Zool. Mag. 72:77-83
(Japanese, English summary).

OPLER, P. A. & G. O. KrIZEK. 1984. Butterflies east of the Great Plains. Johns Hopkins
Univ. Press, Baltimore. 294 pp.

RILEY, T. J. 1980. Effects of long and short day photoperiods on the seasonal dimorphism
of A. andria (Nymphalidae) from central Missouri. J. Lepid. Soc. 34:330-337.

1988. Effect of larval photoperiod on incidence of adult seasonal forms in Anaea
andria (Lepidoptera: Nymphalidae). J. Kansas Entomol. Soc. 61:224—227.

RyDON, A. 1964. Notes on the use of insect traps in East Africa. J. Lepid. Soc. 18:
o1-58.

SAS INSTITUTE INC. 1985. SAS/STAT® guide for personal computers, version 6 edition.
SAS Institute Inc., Cary, North Carolina. 378 pp.

TAUBER, M. J., C. A. TAUBER & S. MASAKI. 1986. Seasonal adaptations of insects. Oxford
Univ. Press, New York. 411 pp.

Received for publication 28 August 1987; accepted 3 August 1988.

Journal of the Lepidopterists’ Society
42(4), 1988, 269-275

SYSTEMATIC STATUS AND DISTRIBUTION OF THE
LITTLE-KNOWN CHARAXINE PREPONA WERNERI
HERING & HOPP

KURT JOHNSON

Department of Entomology, American Museum of Natural History,
Central Park West at 79th Street, New York, New York 10024

AND

HENRI DESCIMON

Laboratoire de Systématique évolutive, Université de Provence,
3 Place Victor Hugo, F-13331, Marseille, Cedex 3, France

ABSTRACT. Prepona werneri, hitherto of uncertain systematic status, and since 1925
recorded from only the holotype male, is authenticated from eight additional specimens.
Genitalia dissection and review of characters defining Archaeoprepona Fruhstorfer and
Prepona Boisduval indicates werneri belongs in Prepona sensu stricto. Most specimens
are from hydric forest habitat in the Choco and Cauca areas of endemism, Colombia,
but one has data indicating occurrence southward in the upper Rio Putumayo region.
The disjunct distribution is probably relict, reflecting former wider occurrence of per-
humid biomes.

Additional key words: Apaturidae, Archaeoprepona, Neotropics, biogeography.

Of all “Prepona’”’ butterflies, P. werneri Hering & Hopp (1925) has
been the most problematical. Previously recorded only from the ho-
lotype male (Hering & Hopp 1925, Le Moult 1932-33), its melanic
appearance, unusual under-surface wing pattern, and hitherto unex-
amined genitalia have made it a taxon of uncertain status. The most
recent treatment of Neotropical Nymphalidae (D’Abrera 1987) does
not mention the species. From fieldwork and survey of public and
private collections, we recently located eight additional specimens of
P. werneri. Only two of these were collected since 1929, and it appears
unlikely that more specimens will soon be available for study. We
therefore summarize below our current determinations of the taxonomic
status, biology, and biogeography of this seldom-collected charaxine
butterfly.

Taxonomy of “Prepona’’ Butterflies

Despite accumulation of specimens in private and public collections,
there has not been wide agreement on the systematics of “Prepona”’
butterflies. Because of overall similarity in the striking blue and black
markings of the wing upper surfaces, many authors have treated ‘‘Pre-
pona”’ as a monophyletic group (Comstock 1944, Barcant 1970, Brown
& Heinemann 1972, Riley 1975). However, as early as 1915, Fruhstorfer
defined two subgroups of ““Prepona’’. One he described as genus Ar-

270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

chaeoprepona (type species demophon Linnaeus) (Fig. 2C), which he
regarded as “primitive” (Fruhstorfer 1915). The other, including taxa
placed with Boisduval’s (1836) Prepona (type species demodice Bois-
duval) (Fig. 2D), he noted as sharing all morphological characters with
Agrias Doubleday, from which it differed only in wing pattern. Le
Moult (1932-33) also noted the heterogeneity of the group and proposed
Pseudoprepona (type species demophon L., a junior objective synonym
of Archaeoprepona). The above distinctions were followed by a number
of authors (Orfila 1950, Rydon 1971, Descimon et al. 1973-74, Johnson
& Quinter 1982).

As defined hitherto in the literature, some obvious phenetic differ-
ences separate Archaeoprepona from Prepona (Table 1). Authors rec-
ognizing these differences have considered both groups as genera (Sti-
chel 1939, Papworth 1982) and even tribes (Rydon 1971) (Table 1).
Our concern when considering the taxonomic status of werneri has
been whether Prepona and Archaeoprepona are monophyletic groups.
Our unpublished numerical cladistic analyses on taxa placed in these
groups (Table 1), including outgroups Charaxes, Polyura, Palla, Eu-
xanthe and Comstock’s (1961) Anaea sensu lato, do not conflict with
generic usage of Prepona and Archaeoprepona as reviewed in Table
1. Therefore, based on male genitalia (Fig. 2A, B) and historical usage,
P. werneri can be reliably placed in Prepona sensu stricto.

Prepona werneri Hering & Hopp
(Figs. 1A, B, C, 2A)

Diagnosis. Male. Upper surface of wings: ground darker black-brown than congeners,
with blue stripes of deeper azure color (not silverish or blue-green) restricted thinly caudad
the forewing discal cell and in a median arc across hindwing. Under surface of wings:
hindwing with variably suffused median band, area basad discal band variously marked
with dark blotches, two large eyespots each submarginal in cells RS and CuAI (or a third
in cell M3), forewing with disjunct or continuous apical and postmedian lines. Genitalia
(Fig. 2A). Typical of general configuration of Prepona sensu stricto (Fig. 2D).

Female. Unknown.

Distribution. Principally Chocé and Cauca regions (region names follow areas of ende-
mism postulated by Brown 1976, 1982), Colombia, with a single specimen having data
indicating upper Putumayo region.

Known specimens. In addition to the type male (Zoologisches Museum der Humbolt
Universitat zu Berlin, ZMH), labelled “Origin, Prepona werneri Hering & Hopp, Rio
Micay, Columbien, Februar 1925, 1000m” (Fig. 1A, B, C), eight male specimens are
reported here for the first time: (1) Rio Guayabal, Colombia, February 1929, anonymous
private collection (examined by first author); (2) Rio Bravo, Prov. Valle, Colombia, March
1985, anonymous private collection (noted by collector as only specimen taken at locality
in many years of collecting, examined by David Matusik, Field Museum of Natural
History, FMNH, photograph examined by us); (3) Frontino, Colombia, no other data,
anonymous private collection (photograph furnished to first author), one male; (4) Cis-
neros, Colombia, 6 May 1928 (purchased from Le Moult collection February 1968 for
Niedhoffer collection), Milwaukee Public Museum (MPM) (photograph examined; gen-
italia dissected, illustrated in Fig. 2D); (5) Rio Micay, Cordillera Occidentale, Colombia,

VOLUME 42, NUMBER 4 DAG |

TABLE 1. Main characters in literature differentiating Archaeoprepona and Prepona.

Character
location
(authors) Prepona Archaeoprepona
Wing upper Androconia well-defined, brush- Androconia diffuse, softly
surface (1-6) like, with rigid setae hairy
Hindwing un- Eyespots large, usually two, post- Eyespots small, undifferentiat-
der surface median to marginal, cells RS ed, marginal, cells RS to
(1-6) and CuA1 CuA2
Male genitalia Slender in all parts (especially un- _ Stout in all parts; gnathos flat,
(1, 3-5) cus and valvae); gnathos rod- without spines
like, with prominent radial
spines
Female genita- Sterigma Y-shaped Sterigma circular
lia (8, 7)

Taxa included: Prepona amesia Fruhstorfer, brooksiana Godman & Salvin, deiphile Godart, demodice Godart, dex-
amenes Herbst, eugenes Bates, eal epeane Staudinger, gnorima Bates, laertes Hiibner, omphale Hiibner, pheridamas
Cramer, praeneste Hewitson, pylene Hewitson, neoterpe Hewitson, xenagoras Hewitson, Archaeoprepona amphimachus
Fabricius, camilla Godman & Salvin, chalciope Hiibner, demophon Linnaeus, demophoon Hiibner, licomedes Cramer,
phaedra Godman & Salvin, meander Cramer (Rydon 1971 included chromus Guérin-Méneville and priene Hewitson
in his genus Noreppa and treated genera as tribes).

Authors: (1) Fruhstorfer (1915, 1916)***; (2) Stichel (1939)**; (3) Orfila (1950)***; (4) Rydon (1971)***; (5) Descimon
et al. (1973-74)***; (6) Papworth (1982)**; (7) Johnson and Quinter (1982)*. * Emphasized certain characters, ** grouped
taxa based on these characters.

February—April 1928, collector Kruger, sold by Niepelt 31 May 1928, in Biedermann
Collection, Zurich, Switzerland (examined by second author); (6) Cisneros, Rio Dagua
valley, 1000 m, 28 February 1928, collector Hopp, sold by Staudinger 15 May 1928, in
Biedermann Collection (examined by second author); (7) Queremal, Colombia, November
1986, collector Julian Salazar, Manizales Museum (K. S. Brown Jr. pers. comm., sole South
American deposition known to him); (8) Upper Rio Putumayo valley, 1981, local collectors,
obtained by David Matusik (FMNH), deposited in American Museum of Natural History
(AMNH) (Fig. 1D). !

No specimens were located at Allyn Museum of Entomology (AME), British Museum
(Natural History) (BMNH), Carnegie Museum of Natural History (CMNH), Field Mu-
seum of Natural History, Rijkmuseum van Natuurlijke Historie (Leiden, Netherlands)
(RMNH), Museum National d’Histoire Naturelle (Paris), National Museum of Natural
History (Smithsonian Institution).

Variation. Variation in the Chocé and Cauca samples appears slight (Fig. 1C), but the
single Putumayo specimen (Fig. 1D) is distinctive, as follows: hindwing with emphatic
medial band, area basad discal band with heavy blotched markings, three large submar-
ginal eyespots (cells RS, M3, CuAl1), and forewing with subapical stripe connected to
postmedian stripe across vein M3.

Biology. The few acquirers of P. werneri provide the only sources of information about
the butterfly’s biology. Most specimens now in public (6) or private (3) collections derive
from the pre-World War II era of highly financed butterfly sampling in the Neotropics.
Initially, commercial interest prompted collection of P. werneri at several localities on
the Pacific slopes of the Colombian Cordillera (mostly Chocé region). These sites proved
extremely inhospitable (Hering & Hopp 1925), being rain forest with extraordinarily high
precipitation; Gentry (1982) cites Chocé as the rainiest tropical forest in the world.
Consequently, commercial interest in the insect waned, and only one specimen has since
been recorded from the region (specimen 6 above). Specimens are so few that most private
owners, to avoid deluges of buy offers, request anonymity.

Biogeography. Most specimens of P. werneri are from the Choco region, though one
(Queremal, Colombia) is near its eastern margin with the Cauca region. Very likely the
extremely hydric Choco region was a “forest refugium” during Pleistocene glaciations

272 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 1. Prepona werneri. A, B, Le Moult’s (1932-33) figure of the holotype of P.
werneri. A, Under surface. B, Upper surface. C, Reproduction of Hering and Hopp’s
(1925) original figure of holotype (showing distinctive markings characterizing Choco
and Cauca specimens). D, Drawing indicating distinctive markings on divergent specimen
from upper Rio Putumayo region.

(Brown 1976, 1982), explaining the occurrence of a number of highly insular and seldom
collected butterflies in it and immediately adjacent areas. Brown emphasizes the close
geographic proximity of the Chocé and Cauca regions, and includes them in his “North-
western Region” cluster (Brown 1976). He notes zones of hybridization between their
taxa. If represented only by Choco and Cauca specimens, P. werneri might be charac-
terized as a seldom collected, insular cis-Andean species, typifying limited hydric habitat.
However, a larger view of its taxonomy and biogeography is necessitated by specimen 8
above from the upper Putumayo region of south-central Colombia. This collection is

VOLUME 42, NUMBER 4 273

Fic. 2. Male genitalia of Archaeoprepona and Prepona, and male genitalia and
abdominal androconia of P. werneri. A, Topotypical P. werneri, lateral view of genitalia
with aedeagus removed (aedeagus, lateral view, beneath) and (x) ventral view, juxta, (y)
lateral view, abdominal androconia at first and second abdominal spiracles. B, P. werneri
specimen from upper Rio Putumayo region (dashed lines indicating areas of genitalia
not available for study because of prior damage to abdomen). C, Archaeoprepona, type
species demophon, Rio de Janeiro, Brazil, same format except for x and y. D, Prepona
type species demodice, Rio de Janeiro, Brazil, same format except for x and y. Females
of Archaeoprepona and Prepona are illustrated in Orfila (1950).

particularly striking, since the Andes are usually considered as a very efficient barrier
against faunal exchange. The Putumayo region is located disjunctly southwest of the
Chocé and Cauca regions and included in Brown’s (1976) “Andean Foothills” cluster.
Brown notes very little hybridization between taxa of the Putumayo and Choco-Cauca
regions. Faunal elements of the Putumayo region are mostly trans-Andean. Thus, occur-
rence of P. werneri in the Putumayo region appears biogeographically significant. It
seems likely that disjunct distribution in P. werneri is relict, reflecting former more
widespread occurrence of perhumid biomes. Compared to the rest of the Andes, uplift
of its northern elements was relatively recent (Gansser 1973). Consequent separation of
P. werneri into cis-Andean and trans-Andean nuclei associated with general climatic
drying appears more likely than dispersal across theeAndes in present or recent times. If
further documented, the Putumayo P. werneri could be construed as a subspecies.

274 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

ACKNOWLEDGMENTS

Dale Jenkins (AME), K. S. Brown Jr. (Universidade Estadual de Campinas, Brazil) and
two anonymous reviewers made helpful comments on the manuscript. We particularly
thank David Matusik (FMNH) for obtaining the Putumayo specimen of P. werneri for
AMNH, and A. M. Young (MPM) for providing a Choco specimen for dissection. For
location of additional material we thank David Matusik, H. J. Hannemann (ZMH), and
three anonymous private collectors. The following also assisted in efforts to locate spec-
imens: Philip Ackery (BMNH); L. D. Miller (AME); R. deJong (RMNH); J. E. Rawlins
(CMNH); and K. S. Brown Jr (various South American inquiries). F. H. Rindge (AMNH)
kindly facilitated assistance at AMNH including obtaining archival material. Robert
Aronheim (Oakton, Virginia) generously provided an AMNH patron grant in 1981 for
initial study of Agrias and Prepona butterflies which was subsequently followed by a
similar grant from Joel B. Grae (Harrison, New York).

LITERATURE CITED

BARCANT, M. 1970. Butterflies of Trinidad and Tobago. Collins, London. 314 pp.

BOISDUVAL, J. B. A. D. 1836. Histoire naturelle des insectes. Species general de Leépi-
doptéres. Paris, Libr. Encycl. de Roret. 690 pp.

Brown, F. M. & B. HEINEMANN. 1972. Jamaica and its butterflies. F. W. Classey,
London. 478 pp.

BRowN, K.S. 1976. Geographical patterns of evolution in Neotropical forest Lepidoptera
(Nymphalidae: Ithomiinae and Nymphalinae—Heliconiini), pp. 118-160. In Des-
cimon, H. (ed.), Biogeographie et Evolution en Amerique Tropicale, Laboratoire de
zoologie de l’ecole normale superieure, No. 9, 344 pp.

1982. Paleoecology and regional patterns of evolution in Neotropical forest
butterflies, pp. 255-308. In Prance, G. T. (ed.), Biological diversification in the tropics.
Columbia Univ. Press, New York. 714 pp.

Comstock, W. P. 1944. Butterflies of Porto Rico. New York Acad. Sci. Scientific Survey
of Porto Rico and the Virgin Islands, Insects of Porto Rico and the Virgin Islands
12(4):421-622.

1961. Butterflies of the American tropics: the genus Anaea, Lepidoptera, Nym-
phalidae. American Museum of Natural History, New York. 214 pp.

D’ABRERA, B. 1987. Butterflies of the Neotropical region. Part 4. Nymphalidae (Partim).
Hill House, Victoria, Australia. Pp. 528-678.

DESCIMON, H., J. MAST DE MAEGHT & J. R. STOFFEL. 1973-74. Contribution 4 l’etude
des Nymphalides néotropicales. Description de trois nouveaux Prepona mexicains.
Alexanor 8:101-105, 155-159, 235-240.

FRUHSTORFER, H. 1915. Beitrag zur Morphologie der Prepona und Agrias. Entomol.
Rundsch. 32:45-47.

1916. Prepona Bsd., pp. 550-566. In Seitz, A. (ed.), 1910-1919. The Macrolep-
idoptera of the world. 5. The American Rhopalocera. Alfred Kernen, Stuttgart. Pp.
993-1139.

GANSSER, A. 1973. Facts and theories of the Andes. J. Geol. Soc. London 129:93-131.

GeNnTRY, A. H. 1982. Phytogeographic patterns as evidence for a Choco refuge, pp.
112-136. In Prance, G. T. (ed.), Biological diversification in the tropics. Colombia
Univ. Press, New York. 714 pp.

HERING, M. & H. W. Hopp. 1925. Sammelausbeute des H. Werner Hopp aus der Choco
Kolumbiens. Deut. Entomol. Z. (Iris) 38:194—195.

JOHNSON, K. & E. L. QUINTER. 1982. Commentary on Miller and Brown vs. Ehrlich
and Murphy et al.: Pluralism in systematics and the worldwide nature of kinship
groups. J. Res. Lepid. 21:255-269.

Le MouLtT, E. 1932-33. Etudes sur les Prepona. Nov. Entomol. II, 1, Suppl. 1: 16 pp.;
III, Suppl.

OrFILA, R. N. 1950. Las especies Argentinas de “Prepona’’ Boisd. (Lep. Nymph.).
Cienc. Zool. 1(7):273-321.

VOLUME 42, NUMBER 4 275

PAPWORTH, H. 1981-82 [1982]. A review of the Trinidad butterflies hitherto placed in
the genus Prepona (Lepidoptera: Nymphalidae). Living World, J. Trinidad and
Tobago Field Naturalist’s Club, unnumbered, pp. 4-8.

RiLey, N. D. 1975. Field guide to the butterflies of the West Indies. Collins, London.

244 pp.
RyYDON, A. H.B. 1971. The systematics of the Charaxidae (Lepidoptera: Nymphaloidea).

Entomol. Rec. J. Var. 83:219-233, 283-287, 310-316, 336-341, 384-388.
STICHEL, H. 1939. Pp. 628-664. In Bryk, F., Lepidopterorum Catalogus. 93: Nym-
phalidae III. Subfamily Charaxinae II. 794 pp.

Received for publication 1 April 1987; accepted 27 May 1988.

Journal of the Lepidopterists’ Society
42(4), 1988, 276-280

A NEW EUPTYCHIA SPECIES FROM
NORTHWESTERN MEXICO (SATYRIDAE)

LEE D. MILLER AND JACQUELINE Y. MILLER

Allyn Museum of Entomology of the Florida Museum of Natural History,
3621 Bay Shore Road, Sarasota, Florida 34234

ABSTRACT. A new euptychiine satyr, Euptychia rubrofasciata, is described based
on 15 males and 4 females from NW Mexico, and compared with other similarly red-
suffused species. A possible Selaginella foodplant association is discussed, and a mimetic
assemblage involving satyrids is suggested.

Additional key words: Euptychiini, Euptychia rubrofasciata, mimicry, Selaginella.

Mexican and northern Central American euptychiine Satyridae are
unusual in that several species are strongly laved with red on the upper
surface. This condition is shown in such diverse species as Euptychia
fetna Butler, Megisto rubricata (W. H. Edwards), a few species of
Cyllopsis (L. Miller 1974) and Paramacera (L. Miller 1972), Cissia
pellonia (Godman & Salvin), and C. cleophes (Godman & Salvin). These
red-patterned elements are rare in Euptychiini, and they are almost
unknown in members of the tribe outside Mexico and northern Central
America. Recently, Douglas Mullins showed us a series of a red-pat-
terned species from Sonora, Mexico, that is totally unlike any other in
this complex of “look alikes’”. Later, James Brock and Jerry Powell sent
additional specimens. This insect is the most ornate of the Mexican red-
laved euptychiines, and is undescribed. A name for it is required for
Brock and Mullins’s forthcoming book on the butterflies of Sonora.

Euptychia rubrofasciata L. & J. Miller, new species
(Figs. 1-9)

Male (Figs. 1, 2). Head clothed with fuscous dorsal setae and somewhat paler hairs
ventrad; area immediately behind eye narrowly white. Eyes rich brown, only slightly
hirsute. Antennae plain brown above, light brown and narrowly ringed on shaft, dark
brown beneath; tip black. Palpi clothed with long fuscous ventral setae and short lateral
white hairs. Thorax and abdomen clothed with short fuscous dorsal and gray-brown
ventral hairs. Legs clad with short gray-brown hairs.

Upper surface of forewing fuscous, grayer and paler distad of cell, with a darker fuscous
submarginal shade and a single smooth, dark fuscous marginal line; wing laved with
brick-red in posterior part of cell and just posteriad of cell, and with a darker red fascia
from end of cell to middle of Cu,-2A, a blackish fuscous subapical black ocellus in M,-
M,, and a smaller one in M,—M,, each with a single silver pupil and narrow, dull ocherous
ring. Upper surface of hindwing also fuscous, slightly paler subapically, with submarginal
darker fuscous shade and a double dark fuscous marginal line. Wing laved with brick-
red just outside and posteriad of cell, a red fascia outside cell from apex to near inner
angle, blackish fuscous ocelli in Rs—-M, (large and diffuse), M,-M, (very small, almost a
point and occasionally absent), and a well-defined, quite large ocellus in Cu,—Cu,, all
ocelli consisting of a white pupil and a narrow, dull ocherous iris.

Under surface of forewing light gray-brown slightly shaded with red in and just
posteriad of cell, with three brick-red fascia from near costa to inner margin, one across

VOLUME 42, NUMBER 4 reg

Fics. 1-4. Euptychia rubrofasciata. 1,2, Holotype é, upper (1) and under (2) surfaces.
3, 4, Paratype 2, upper (3) and under (4) surfaces. Scale line represents 10 mm.

cell, one just outside cell, and one beyond ocelli, the last two connected by brick-red
streaks between veins from M, to 2A; ocelli as on upper surface, but black with silver
pupils and ocherous then fuscous rings surrounding both (not individual rings). Under
surface of hindwing likewise gray-brown with three reddish fascia as described for fore-
wing, and dark brown double marginal lines; six black ocelli with silver pupils and ocherous
and fuscous rings from Sc+R, to Cu,-2A, the ones in Rs-M, and Cu,—Cu, large and
prominent, the one in Cu,—2A of moderate size, the others quite small; ocelli in anterior
three cells with rings coalesced.

Forewing length of holotype 6 17.6 mm, of the 14 4 paratypes 17.3 to 19.2 mm, averaging
18.0 mm.

Male genitalia (Figs. 5-8) simple and lightly sclerotized; no superuncus as in most
Euptychia (comparative illustrations in Forster 1964:81); uncus only slightly curved ven-
trad; brachia represented by only a very narrow sclerotized ring completely surrounding
anus; valvae relatively unadorned, curved dorsad; penis short and straight with no obvious
adornment.

Female (Figs. 3, 4). Head, thorax, abdomen, and appendages as in 6, except thorax
and abdomen below somewhat tanner.

Upper surface of forewing somewhat lighter than that of 6 and more extensively laved
with reddish fulvous, rusty fascia across cell and just beyond it, reddish streaks between
veins from M, to 2A, a fuscous submarginal fascia and double marginal fuscous lines;
blackish-brown coalesced ocelli with silver pupils in M,—-M, (large and prominent) and
M,—-M, (very small) with coalesced narrow ocherous and fuscous rings. Hindwing above
with similar ground color, red shading slightly more extensive than in 6, and white-

278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

i
| i" Nil
} Vi

Fics. 5-9. Genitalia of Euptychia rubrofasciata. 5-8, Holotype 6. 5, Uncus, tegumen,
saccus, and associated structures, left lateral view. 6, Right valva, internal view. 7, Penis,
dorsal view. 8, Left lateral view. 9, Paratype 2, ventral view, genit. prep. M-7336-6 (J.
Y. Miller). Scale line represents 0.5 mm.

VOLUME 42, NUMBER 4 279

pupilled fuscous ocelli in Rs—M, (large), M;—Cu, (very small, not present in all specimens),
Cu,-Cu, (large), and Cu,—2A (small and absent in one specimen), each with ocherous
and fuscous rings; red-brown submarginal fascia, two fuscous marginal lines.

Under surface of forewing somewhat less gray than in é with similar markings except
ocelli. Under surface of hindwing also less gray than in 6, but marked similarly with
larger ocelli and more prominent ocherous rings.

Forewing length of the 4 2 paratypes 17.6 to 20.0 mm, averaging 19.2 mm.

Female genitalia (Fig. 9) very lightly sclerotized with 8th segment heavily clothed in
scales; papillae anales densely setose with 6-10 elongated setae posteriad; sterigma simple,
lamella postvaginalis membranous with numerous folds, and lamella antevaginalis indi-
cated by a lightly sclerotized plate; ductus bursae and corpus bursae membranous and
strongly folded; attachment of ductus seminalis near atrium.

Described from 15 males and 4 females from the Sierra Madre Occidental of Sonora
and Chihuahua, Mexico.

Holotype ¢ (Figs. 1, 2). MEXICO: Sonora, 13 mi (21 km) E El Novillo, 12 August
[19]85 (J. P. Brock); é genitalia preparation M-7341-v (Lee D. Miller).

Paratypes. All MEXICO: Sonora, 8 é, 1 9, same data as holotype, 1 2 (Figs. 3, 4), Rte.
16, 10 mi (16.1 km) E Trinidad, “Cypress” Canyon, 7 August [19]86 (D. D. M[ullins]); 2
6, 2 2, San Nicholas-Yecora Rd., 4.1-10.3 mi (5.6-16.5 km) E Santa Rosa, 7.viii.1986 (J.
P. Brock) (1 with 2 genitalia preparation M-7346-v (J. Y. Miller); 3 6, creek at 3000 ft
(909 m), 6 mi (9.6 km) W Yecora, 31.vii.1984 (J. P. Brock); Chihuahua, S[ierra] Madre
Occid[ental], Yepachic Rd., Canyon Rio Tomochic (oak/grass hillside), 31 July [19]84 (D.
D. Mfullins]); Sinaloa, 1 4, 2 mi (3.2 km) SW Potrerillos, 4200’ (1280 m) viii.7/8.[19]86
(J. Brown & J. Powell).

Disposition of type-series. Holotype 6, 2 6 and 1 2 paratypes in Allyn Museum of
Entomology; 1 ¢ paratype in collection of California Insect Survey; remaining 11 6 and
3 2 paratypes to be returned to J. P. Brock and D. D. Mullins for eventual distribution
to other collections.

Etymology. The name refers to the unique brick-red fascia on both surfaces of all
wings.

Discussion. That this insect proved to be a member of Euptychia came as a surprise.
It is the largest known Euptychia, and superficially more closely resembles Cissia. How-
ever, the 6 genitalia are unmistakably Euptychia, the abbreviated brachia fused with the
tegumen. The 2 genitalia are simple and very lightly sclerotized, this also in keeping with
the apomorphic condition for Euptychia.

The only published life history information about Euptychia sensu lato is that by Singer
et al. (1971) who found the white congener, E. westwoodi, feeding as a larva on the
lycopsid Selaginella. Those authors suggested that Selaginella might have “rather potent
biochemical defenses,” since few herbivores attack them, and that these defenses might
convey some protection to Euptychia. These toxic chemical defenses have yet to be proven
(J. Beckner pers. comm.), but seem reasonable. The Mexican E. fetna feeds also on
Selaginella (J. Llorente and others pers. comm.). Euptychia westwoodi appears to be in
a mimetic complex involving lycaenids and riodinids (Singer et al. 1971:1342).

We suggest that E. rubrofasciata also feeds as a larva on Selaginella. This is supported
by Brock (pers. comm.), who writes “. . . nearly all the Euptychia were found on a shady
[canyon] wall loaded with a Selaginella species.’’ He further mentioned that he identified
the Selaginella because it was so abundant and conspicuous at the spot where the new
species was most abundant. Mullins (pers. comm.) independently confirms this habitat
preference.

Assuming the above foodplant and its toxicity to predators, the present new species
and E. fetna may be Muellerian mimics, and the other red-laved euptychiines (and
perhaps other butterflies) could be Batesian mimics of them.

ACKNOWLEDGMENTS

We thank D. D. Mullins for sending the first specimens of this insect for description,
and J. P. Brock and J. A. Powell for additional ones. Thanks are due J. Llorente for

280 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

discussions concerning E. fetna and to J. Beckner for discussions about the properties of
Selaginella.

LITERATURE CITED

FORSTER, W. 1964. Beitrage zur Kenntnis der Insecktenfauna Boliviens XIX. Lepidop-
tera III Satyridae. Veroff. Zool. Staatssamml. Mutinchen 8:51-188.

MILLER, L. D. 1972. Revision of the Euptychiini. 1. Introduction and Paramacera
Butler. Bull. Allyn Mus. 8, 18 pp.

1974. Revision of the Euptychiini. 2. Cyllopsis R. Felder. Bull. Allyn Mus. 20,
98 pp.

SINGER, M. C., P. R. EHRLICH & L. E. GILBERT. 1971. Butterfly feeding on lycopsid.
Science 172:1341-1342.

Received for publication 19 May 1988; accepted 30 June 1988.

Journal of the Lepidopterists’ Society
42(4), 1988, 281-284

A NEW SPECIES OF ETHMIA FROM THE FLORIDA KEYS
(OECOPHORIDAE: ETHMIINAE)

J. B. HEPPNER

Florida State Collection of Arthropods, Center for Arthropod Systematics,
Bureau of Entomology, Florida Department of Agriculture & Consumer Services,
P.O. Box 1269, Gainesville, Florida 32602

ABSTRACT. Ethmia powelli is described from Upper Matecumbe Key based on 123
specimens. It is related to E. humilis Powell and E. julia Powell, in the confusella species-
group, by genitalic characters, and is distinguished from E. farrella Powell by the small
wingspan and fewer forewing black spots.

Additional key words: Ethmia powelli, E. farrella.

The genus Ethmia was monographed for the known New World
fauna by Powell (1973). In Florida seven species are now recorded
(Florida Lepidoptera Survey), mostly being Caribbean elements present
in southern Florida. The new species was collected after the publication
of Powell’s (1973) monograph and is described here to make the name
available for a revision of Kimball (1965). Description of the species
has awaited collection of more specimens, but only recently has one
additional individual been collected. Capuse (1981) reviewed the Cuban
ethmiines but did not include the species described from Florida. Spec-
imens are deposited with the Florida State Collection of Arthropods
(FSCA) and my own collection (JBH), with paratypes distributed to the
University of California, Berkeley (UCB), and the National Museum
of Natural History, Washington, D.C. (USNM).

Ethmia powelli Heppner, new species

(Figs. 1-4)

Forewing length 4.0—-4.7 mm (N = 100) (male); 4.1-4.9 mm (N = 23) (female).

Male (Fig. 1). Head. Silvery gray-white with black central mark on vertex; labial palpus
silvery gray-white with lateral black mark on each segment and black laterally near base.
Thorax. Silvery white; legs white, with fore- and mid-tibiae and tarsi marked with black;
hind legs white. Forewing. Ground color silvery white with numerous black spots (costal
spots at base and % from base; cubital area with elongated spots near base and at hindwing,
with a small round spot near dorsal margin; a large elongate spot mid-wing and another
along tornus); terminal black spots extending along costa on apical 4; fringe silvery; venter
silvery gray. Hindwing. Unicolorous pale gray with dark gray at margin; fringe gray;
venter similar. Abdomen. Silvery white with darker gray dorsum. Genitalia (Fig. 3).
Tegumen with slightly bulbous terminal points; vinculum rounded, without saccus; valva
subquadrate with prolonged distal end having 3 large spines and 2 smaller truncated
central spines, with a large curved hooklike process on dorsal margin near apex; anellus
an elongated tube (troughlike), dorsally open; aedeagus similar to that of E. humilis, with
bulbous phallobase; cornutus indistinct.

Female (Fig. 2). Similar to male; forewing terminal black spots slightly larger on average
than in male. Genitalia (Fig. 4). Setose ovipositor; posterior apophyses 3x length of
anterior apophyses; sterigma composed of fused anterior apophyses extensions; ductus

282 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 1, 2. Ethmia powelli paratypes, Islamorada, Monroe Co., Florida. 1, Male, 2,
Female.

bursae with sclerotized collar at ostium, then spiralled to ovate corpus bursae; signum a
small sclerotized depression.

Type material. Holotype: male, 1 mi [1.6 km] SW Islamorada, Upper Matecumbe Key,
23-VI-1974, J. B. Heppner (slide JBH 1773) (FSCA). Paratypes: 99 males, 23 females,
same data as holotype. Paratypes distributed to FSCA, UCB, USNM, and author’s personal
collection. Additional specimen: Key Largo, Monroe Co., 30-VIII-1986 (1 male), L. C.
Dow (FSCA).

Hosts. Unknown. One species in the confusella species-group feeds on Bourreria ovata
(Boraginaceae).

Remarks. Thus far, Ethmia powelli has been collected only twice in the Florida Keys.
There are no records of it from any Neotropical locality; thus, the species may be native
to Florida. Relations of E. powelli by some genital characters appear nearest to E. humilis

Fic. 3. Ethmia powelli male genitalia, aedeagus omitted (JBH 1773).

VOLUME 42, NUMBER 4 283

Fic. 4. Ethmia powelli female genitalia (JBH 1774).

284 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Powell and E. julia Powell in the confusella species-group. This is a primarily tropical
group with several species also occurring in S Florida (includes records of West Indian
species recently found in the Florida Keys). Forewing maculation, however, is more
similar to E. farrella Powell. In the key to species in Powell (1973), E. powelli keys to
couplet 118, differing from E. farrella in having fewer black spots on the forewings, and
in being significantly smaller, 4.0-4.9 mm versus 6.5-7.0 mm for E. farrella. The male
genitalia are particularly diagnostic, having 5 spines on the distal end of the valva, and
the central 3 of these being truncated; E. humilis has only 3 curved spines on the valva,
likewise for E. julia. Female genitalia are not very similar to the other species; the sterigma
is most similar only to the Central and South American Ethmia catapeltica Meyrick. The
female ductus bursae in E. humilis is not coiled as in most Ethmia species and the sterigma
is very different in E. farrella.

Ethmia powelli appears to be one of the smallest species in Ethmia. The species is
named in honor of Professor J. A. Powell, University of California, Berkeley.

ACKNOWLEDGMENTS

Reviewer comments and study of the L. C. Dow Collection, Largo, Florida, are ap-
preciated. Contribution No. 640, Bureau of Entomology, Florida Department of Agri-
culture & Consumer Services.

LITERATURE CITED

CapusE, I. 1981. Sur les représentants de la famille des Ethmiidae (Lepidoptera: Ge-
lechioidea) de Cuba. Result. Exped. Biospeologiques Cubano-Romaines 4 Cuba (Bu-
charest) 3:125-143.

KIMBALL, C. P. 1965. The Lepidoptera of Florida, an annotated checklist. In Arthropods
of Florida and neighboring land areas. Vol. 1. Florida Department of Agriculture,
Gainesville. 363 pp.

POWELL, J. A. 1973. A systematic monograph of New World ethmiid moths (Lepidop-
tera: Gelechioidea). Smiths. Contr. Zool. 120:1-302.

Received for publication 12 January 1988; accepted 5 July 1988.

Journal of the Lepidopterists’ Society
42(4), 1988, 285-290

LARVAE OF NORTH AMERICAN LEUCONYCTA (NOCTUIDAE)

KENNETH A. NEIL

Agriculture Canada, Research Station,
Kentville, Nova Scotia B4N 1J5, Canada

ABSTRACT. Mature larvae of Leuconycta diphteroides (Guenée) and L. lepidula
(Grote), the only known members of Leuconycta, are illustrated, described, and diagnosed
based on eight specimens of the former reared from ova on Solidago sp. and two specimens
of the latter reared from ova on Taraxacum sp. Although resembling one another in
coloration and structure, larvae of the two species can be distinguished by characters in
the hypopharyngeal complex.

Additional key words: Acontiinae, Leuconycta diphteroides, L. lepidula, hypopha-
ryngeal complex.

The North American noctuid genus Leuconycta Hampson (Acon-
tiinae) contains two species, L. diphteroides (Guenée) and L. lepidula
(Grote). Larval systematic and life history information has been pre-
sented by Dyar (1898), Forbes (1954), and Crumb (1956). Crumb was
unable to find color or structural differences by which to separate larvae
of the species of Leuconycta. The purpose of this paper is to more fully
describe and diagnose mature larvae, especially with respect to mouth-
parts and chaetotaxy, which have been shown by Godfrey (1972) to be
of taxonomic value.

Leuconycta diphteroides and L. lepidula are common and widely
distributed in North America, both ranging from Nova Scotia S to North
Carolina, and W to Manitoba, Kansas, and Colorado (Forbes 1954).
Larvae of L. diphteroides have been recorded feeding on Solidago sp.
(Dyar 1898), and those of L. lepidula on Taraxacum sp. (Forbes 1954).

Terminology and abbreviations here follow Godfrey (1972). Specific
collecting localities and dates are provided in individual descriptions.

Genus Leuconycta Hampson
(Figs. 1-14)

Diagnostic description (diagnostic characters in italic). Head 1.6-2.5 mm wide, total
body length 25.8-32.2 mm (N = 10). Head and body smooth. Body broad at middle,
tapering slightly anteriorly and posteriorly. Prolegs present on abdominal segments
(Ab) 3-6, size increasing posteriorly; those of Ab6 twice size of those on Ab3. Crochets
uniordinal. All setae simple. Coloration of living material. Head green, no lines or
markings present. Body green, darker at edges of mid-dorsal and subdorsal lines and
ventral edge of subdorsal area; ventral area lighter green; mid-dorsal and subdorsal lines
white, the latter wider and more irregular; spiracular line greenish white, more whitish
on dorsal and ventral edges and bordered dorsally by a narrow red line on thoracic
segments (T)1 and 2. Cervical and anal shields concolorous with trunk, the latter with a
white medial and two white lateral lines. Pinacula white, the dorsal pinacula larger than
lateral and ventral pinacula. Spiracles yellow with black peritremes. True legs greenish,
slightly brown distally. Proleg shields concolorous with trunk. Coloration of preserved
material. Head and body light cream color. Lines and pinacula concolorous with body.

286 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1 eu 5 0.5mm

Distal

Medial Transverse
Region Cleft

Proximomedial

Region
Labial

Palpus \_ é a 2, ——
ars me ert Ao)

Proximolateral
Region

Proximolateral

5 —*
Stipular Seta Spines

2 0.5 mm

Fics. 1-6. Leuconycta diphteroides larval structures. 1, Head, frontal view. 2, Hy-
popharyngeal complex, left lateral view. 3, Labial palpus, lateral view. 4, Left mandible,
oral surface. 5, Left mandible, outer surface. 6, Anal shield, dorsal view.

VOLUME 42, NUMBER 4 287

—E—E—————
7 2.0mm 1 1 eo WU
Distal Medial Transverse
Region Cleft
Proximomedial
\ Region
ee, | ig eigenstate
alpus \.
wid i Ape Proximolateral
- Region

Proximolateral
Spines

Stipular Seta ——

8 0.5 mm

(a

9 01mm

Fics. 7-12. Leuconycta lepidula larval structures. 7, Head, frontal view. 8, Hypo-
pharyngeal complex, left lateral view. 9, Labial palpus, lateral view. 10, Left mandible,
anal surface. 11, Left mandible, outer surface. 12, Anal shield, dorsal view.

288 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

i 4 -
| | | se om ane (

SIE) MeL Sead TS Lenape

eee f. fs ~ sv3 Se oA [ mie +
Ab1

“=P Ab7 Ab8 Ab9 Ab10

2.0mm

Fic. 13. Leuconycta diphteroides. Dorsal and lateral chaetotaxy of prothoracic (T1),
mesothoracic (T2), and abdominal segments (Ab1-3, Ab6-10).

Spiracles white, peritremes dark brown. Head (Figs. 1, 7). Cervical indentation shallow;
adfrontal sutures terminating at epicranial suture; epicranial suture longer than height
of frons; frons slightly higher than its basal width. Adfrontal setae (AF)2 above, and
posterior head setae (P)1 below, even with, or slightly above apex of frons; anterior head
puncture (Aa) below a straight line between posterior head puncture (Pa) and anterior
head setae (A)3. P2, Pl, and AF1 in a straight line, Aa closer to A3 than to A2, Al-A8
forming an obtuse angle at A2; lateral head seta (L) even with or slightly below juncture
of adfrontal ecdysial lines. Distance between ocelli (Oc)2—Oc3 greater than Ocl—Oc2 and
Oc3-—Oc4. Mouthparts. Hypopharyngeal complex (Figs. 2, 8): Spinneret short and broad.
Stipular setae varying from extremely short to slightly less than % length of Ist segment
of labial palpus (Lps1). Labial palpi (Figs. 3, 9) with length of segments variable. Distal
and proximal regions of hypopharyngeal complex separated by shallow medial transverse
cleft; distal region of hypopharyngeal complex covered with spines which are long,
slender, shorter proximally; proximolateral region with spines small, triangular, numbers
variable. Mandible (Figs. 4, 5, 10, 11). Two well separated outer setae present; inner
surface with 3 distinct ridges, the last short; outer margin with 12 teeth, the Ist small,
2nd to 4th well developed and angular, 5th to 12th small and angular.

Thorax. Prothoracic segment (T1) (Figs. 13, 14): Shield smooth and weakly sclerotized,
subdorsal body setae (SD)1 and 2 separated from prothoracic shield, setae SD1 and lateral
body seta (L)2 fine, hairlike, and with a thickened sclerotized annulus at base; major axis
of prothoracic shield passing behind SD1 and SD2, and between subventral body setae
(SV)1 and 2; spiracle broadly elliptical, height less than twice its width. T2-3 (Figs. 13,
14): SD1 fine, hairlike, with a thickened sclerotized annulus at base. Tarsal claws with
basal angles acute. Metathoracic coxae contiguous. Abdomen. Dorsal and lateral chae-
totaxy of Ab2-6 with 3 SV setae; SV1 and SV2 setal insertions well separated. Ab7-8 with
1 SV seta. Ab9: SD1 fine, hairlike, with a thickened annulus at base. Ab10: Anal shield
as in Figs. 6 and 12. Dorsal margin convex, posterior margin entire; subanal setae widely
separated.

Leuconycta diphteroides (Guenée)
(Figs. 1-6, 13)

Head 2.2-2.5 mm wide, total length 25.8-31.0 mm (N = 8). Larva as described above
except: Hypopharyngeal complex (Fig. 2) with spinneret about % length of 1st segment
of labial palpus (Lps). Labial palpus (Fig. 3) with Lpsl about 3 times length of seta borne
by 1st segment of labial palpus (Lp), 13 times length of Lps2, 2 times length of Lp2.
Lps2 less than % length of Lp1. Stipular setae very short, about % length of Lpsl, less
than 4 length of Lp1, subequal to Lps2. Proximolateral spines small, 10-14 small triangular
spines.

Material examined. 8 specimens: New Minas, Kings Co., Nova Scotia, reared on Sol-
idago sp. from ova obtained from a female collected 19 June 1985. Moth collected,

VOLUME 42, NUMBER 4 289

Fic. 14. Leuconycta lepidula. Dorsal and lateral chaetotaxy of prothoracic (T1),
mesothoracic (T2), and abdominal segments (Ab1-3, Ab6—-10).

determined, and larvae reared by author. All specimens in Nova Scotia Museum collectio.,,
Halifax, Nova Scotia.

Leuconycta lepidula (Grote)
(Figs. 7, 12, 14)

Head 1.6-1.8 mm wide, total length 31.3-32.2 mm (N = 2). As described for genus
except: Hypopharyngeal complex (Fig. 8) with spinneret slightly less than % length of
Lps1; labial palpus (Fig. 9) with Lps1 slightly less than 5 times length of Lp], 12 times
length of Lps2, less than 2 times length of Lp2. Lps2 about % length of Lpl. Stipular
seta slightly less than 4 length of Lpsl, longer than Lpl, and about twice length of Lps2;
proximolateral region with 10-14 triangular spines; spines larger than in L. diphteroides.

Material examined. 2 specimens: Chicago, Illinois, reared on Taraxacum sp., 24 June
1934, A. K. Wyatt. Specimens in U.S. National Museum, Washington, D.C.

Discussion

Larvae of L. diphteroides and L. lepidula resemble one another
closely, but can be separated by the following mouthpart characters:
In L. diphteroides, the stipular seta is ¥,, the length of Lpsl1 compared
with L. lepidula in which this seta is slightly less than % length of Lps1,
and proximolateral spines are larger in L. lepidula than in L. diphte-
roides.

Franclemont and Todd (1983) placed Leuconycta in Acontiinae,
whereas Forbes (1954) and Crumb (1956) placed it in Amphipyrinae.
The latter two authors considered larval characters; those such as the
open silk pore on the spinneret, presence of five pairs of abdominal
prolegs and two SV setae on Abl strongly indicate Amphipyrinae, but
resolution of the discrepancy should be based on a wider range of
material than examined here.

ACKNOWLEDGMENTS

I thank D. C. Ferguson, Systematic Entomology Laboratory, U.S. National Museum,
Washington, D.C., for supplying specimens of L. lepidula, and Arthur Lightfoot of
Agriculture Canada, Kentville, Nova Scotia, for photographing the plates.

290 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

LITERATURE CITED

Crump, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agr. Tech. Bull. 1135,
356 pp.

Dyar H. G. 1898. Microcoelia diptheroides, Grote. Can. Entomol. 30:16.

ForBEs, W. T. M. 1954. Lepidoptera of New York and neighboring states. Pt. 3. Cornell
Univ. Agr. Expt. Sta. Mem. 329, 433 pp.

FRANCLEMONT, J. G. & E. L. Topp. 1983. Noctuidae, pp. 120-159. In Hodges, R. W.,
et al. (ed.), Checklist of the Lepidoptera of North America north of Mexico. E. W.
Classey and Wedge Entomological Research Foundation, London. 284 pp.

GopFREY, G. L. 1972. A review and reclassification of larvae of the subfamily Hadeninae
(Lepidoptera, Noctuidae) of America north of Mexico. U.S. Dept. Agr. Tech. Bull.
1450, 265 pp.

Received for publication 16 March 1987; accepted 20 May 1988.

Journal of the Lepidopterists’ Society
42(4), 1988, 291-292

GENERAL NOTES

NEW DISTRIBUTION RECORDS AND A PROBABLE NEW LARVAL HOST
PLANT FOR PHILOTES SONORENSIS (LYCAENIDAE) IN
KERN AND TULARE COUNTIES, CALIFORNIA

Additional key words: Sierra Nevada, Dudleya calcicola, Crassulaceae.

Philotes sonorensis (Felder & Felder) is of limited distribution, known only from Sierra
Co. in north-central California S to Cedros Island, Baja California Norte, Mexico (John
W. Brown pers. comm.). Published distribution records do not include Kern and Tulare
counties, California (Langston, R. L. 1963, J. Lepid. Soc. 17:201-228; 1965, J. Lepid. Soc.
19:95-102; 1969, J. Lepid. Soc. 23:49-62; Shields, O. 1978, Bull. Allyn Mus. 15:1-16;
Shapiro, A. M. 1974, Pan-Pac. Entomol. 50:442-443). Recently, P. sonorensis was dis-
covered at scattered locations in these two counties and in the southern Sierra Nevada.
New distribution records (Fig. 1) are as follows:

Kern Co.: Pleito Creek and Canyon (located nr. Mt. Pinos at S end of San Joaquin
Valley), 1 6, 16 IV 81 (W. D. Patterson). Laura Peak (“Rock Tip” on some maps) in Piute
Mts. E of Lake Isabella; adults found by author in canyon on S slope nr. rocky outcrops
at 3600-4600 ft (1097-1402 m) elev. on this peak of 5260 ft. (1603 m), 2 4, 28 III 87; 3

| aes
\ S) satus r

\ National
Park

1
aes | Sierra

im
fa

|
sure Wiiguccs
COUNTY 2
yee: { cous
San Joaquin \ Greenhorn
Valley \ Mts.
Bakersfield Oe

—_——
wee

AA
WF

2 KERN COUNTY

LS
nouechant fi |
15 30 Pleito fe
mele ean eaoven
scale in km

apt cine Pinos

Fic. 1. Map of Kern and Tulare counties, California, showing new localities for
Philotes sonorensis (black dots). Triangles represent nearby mountain peaks important
as reference points. Cross marks represent major cities.

292 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

4, 29 III 87, 2 6, 14 IV 87. The latter locality is dry grassland with some junipers on steep
rocky slopes. The dry streambed probably carries water for short periods following rainfall.

Tulare Co.: Canyon 1.6 km N of Lamont Peak S of Chimney Peak Road, 3 6, 10 IV
85 (K. Davenport). Canyon N of Sherman Pass Road 5.6 km E of Kern Canyon Road, 5
6, 2 IV 87 (Davenport); 1 6, 5 IV 87 (R. Meyer); 2 6, 1 9, 14 IV 87 (Davenport & K.
Richers). Kern River Canyon 1 km N of Roads End at dam, in side canyon E of road, 4
6, 12,2 1V 87 (Davenport); 1 4, 1 2, 5 IV 87 (Meyer); 2 6, 14 IV 87 (Davenport & Richers),
8 4, 1 2, 9 III 88 (Davenport).

The Tulare Co. colonies are in rocky canyons of limestone or granitic composition with
small streams between 3000 and 5000 ft (914-1524 m) elev., in chaparral and foothill
woodland.

The host plant of P. sonorensis in the southern Sierra Nevada (including Laura Peak
in the Piute subrange) is likely Dudleya calcicola Bartel & Shevock (Crassulaceae), which
occurs locally “on pre-Cretaceous limestones within chaparral or pinyon-juniper woodland
at 850-1700 m”’ (Bartel, J. A. & J. R. Shevock 1983, Madrono, 30:210-216), and is limited
in distribution to Kern, Tulare, and extreme SW Inyo counties (J. A. Bartel pers. comm. ).
Adults are closely associated with calcicola (no other Dudleya spp. present) at the new
localities. All known hosts are in the genus Dudleya (Shields, O. 1973, Bull. Allyn Mus.
15:9-11). Collections and identifications of Dudleya at the new localities were made by
J. F. Emmel, J. A. Bartel, and J. R. Shevock. No larvae were collected or reared, and
oviposition was not observed. The discovery of Philotes sonorensis on Laura Peak was
made using herbarium records of Dudleya calcicola provided by Emmel.

Eight voucher specimens of Philotes sonorensis representing each of the four new
Sierran localities have been deposited in the Natural History Museum of Los Angeles
County, Los Angeles, California. Remaining specimens are in private collections.

I thank J. A. Bartel, J. W. Brown, J. F. Emmel, R. Meyer, K. Richers, J. R. Shevock,
O. Shields, and W. D. Patterson for records and assistance.

KENNETH E. DAVENPORT, 6601 Eucalyptus Dr. #325, Bakersfield, California 93306.

Received for publication 22 June 1987; accepted 16 May 1988.

Journal of the Lepidopterists’ Society
42(4), 1988, 292-294

A MELANIC MALE OF ANTHERAEA POLYPHEMUS POLYPHEMUS
(SATURNIIDAE)

Additional key words: Canada, New Brunswick, light trap.

W. B. Preston and W. B. McKillop (1979, J. Lepid. Soc. 33:147-148) summarized the
known information about melanic Antheraea polyphemus polyphemus (Cramer), and
published the first illustration of a melanic specimen. Our specimen appears to be the
fourth ever collected.

On the night of 30 June-1 July 1986, we collected a melanic male of A. p. polyphemus
on a white sheet illuminated by a 250-W M-V bulb, at the edge of a sphagnum bog in
the Acadia Forest Experiment Station, 20 km E of Fredericton, New Brunswick. Dorsally,
this male (Fig. 2) differs from typical A. p. polyphemus (Fig. 1) by having a dark chocolate
ground color. Prothorax and costal edge of upper forewings are black so that the apical
and subapical forewing spots are not discernible as distinct spots. However, the grayish
lilac dash from the subapical spot toward the forewing apex is still present. In typical
specimens, prothorax and costal edge are whitish gray. The blackish component of the
submarginal band on fore- and hindwings is exaggerated, obliterating the pinkish shading
beyond it on the hindwings but leaving it just discernible on the lower half of the forewings.

VOLUME 42, NUMBER 4 293

Fics. 1-4. Males of Antheraea polyphemus polyphemus (Cramer) from New Bruns-
wick. 1, Typical dorsum. 2, Melanic dorsum. 3, Typical venter. 4, Melanic venter.

The discal spots on both fore- and hindwings are typical, with the transparent center
surrounded by an ocher-yellow ring, edged outwardly with a black ring. The whitish-
blue semicircle that outwardly edges the black ring on the proximal side is still discernible
on the forewing and very extensive on the hindwing. The black component of the hindwing
submarginal band is conspicuously recurved along the outer margin of the wing. The
outer margin of both wings beyond the submarginal band is of the usual bright ocherous
tawny brown, giving the specimen a distinctive bicolored appearance.

On the underside, the wings are uniformly dark chocolate (Fig. 4), and the markings

294 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

contrast less than those of a typical specimen (Fig. 3). Costal regions of both fore- and
hindwings are dark, as is the basal area of both wings.
The specimen is in the senior author's private collection.

ANTHONY W. THOMAS, Canadian Forestry Service-Maritimes, P.O. Box 4000, Fred-
ericton, New Brunswick E3B 5P7, Canada, and WAYNE L. FAIRCHILD, Department of
Biology, University of New Brunswick, Fredericton, New Brunswick E3B 6E1, Canada.

Received for publication 7 December 1987; accepted 21 June 1988.

Journal of the Lepidopterists’ Society
42(4), 1988, 295-296

BOOK REVIEWS

THE MONARCH BUTTERFLY: INTERNATIONAL TRAVELER, by Fred A. Urquhart. 1987. xxii
+ 232 pp., 24 pp. color photographs. Nelson-Hall, Chicago. Hard cover. $31.95.

To his everlasting credit Fred A. Urquhart showed that the monarch butterfly migrates
from Canada to overwintering sites in Mexico. In his most recent book Urquhart gives
an enlightening account of his pioneering investigations of North America’s most famous
butterfly, including its morphology, development, behavior and ecology.

Urquhart documents the migration of the monarch butterfly in historical and personal
terms. He writes in the first person and selects information intended for a popular audience.
The result is informative and exciting. He describes how he developed the tagging
technique for following monarchs. He reveals the clues that ultimately led to the discovery
of the Mexican overwintering sites. His conversational, informal style allows him to include
anecdotal digressions that enrich the factual account.

Often when writing about the biology of the butterfly, Urquhart poses a simple question
and then describes an experiment which he conducted looking for answers. What deter-
mines which trees monarch butterflies choose for roosting? Do pheromones help monarch
butterflies aggregate? How does a monarch caterpillar which has fallen off a milkweed
plant find its way back to that plant or another? The question-and-experiment format
generates a certain suspense and captures the reader’s attention. The experiments are
intriguing even when inconclusive.

Few entomological authors dare to write about the emotions which their love of insects
arouses. Scientific papers require a tone of objectivity, and expressions of pleasure come
as a surprise. In this book Urquhart overcomes the inhibitions which desiccate more
scholarly treatises. Urquhart understands that his audience wants to know why a person
would go to the extremes he did in pursuit of a common butterfly (p. 153):

Those who have had a dream and have lived to see that dream come true will have
some conception of my feelings when I first entered the Mexican forest and there,
before my eyes, was the realization of a dream that had haunted me since I was a
lad of sixteen. :

On the negative side, Urquhart’s personal approach overemphasizes his own work at the
expense of others. For example, in the opening chapter about the monarch butterfly’s
foodplant, he ignores the ecological chemistry of milkweeds. Later he alludes to the
transfer of milkweed toxins to monarch butterflies, but only to dismiss its importance.
Urquhart’s failure to recognize the work of others leads him to farfetched speculation.
His attempt to explain control of sexual maturation of migrants is an example (p. 121):

As our sinall planet earth travels in its elliptical orbit around the sun, it is possible
that twice each year it passes through an area rich in some sort of radiation that
impinges upon animal life. The radiation cycle might affect in some manner the
cells of the body, causing reproductive organs to abort in the fall and develop in the
spring and to initiate the migratory response. Perhaps our astronomy researchers
may add a missing part to the migration puzzle. Perhaps animal life on our earth
is being controlled by what is happening in outer space more than we now consider
feasible. ...

Urquhart could have avoided this fantasy if he had discussed hormonal control of monarch
butterfly development, and photoperiodic regulation of the timing of this development
(reviewed in Rankin, M. A., M. L. McAnelly & J. E. Bodenhamer 1986, The oogenesis-
flight syndrome revisited, pp. 27-28 in Danthanarayana, W. (ed.), Insect flight: Dispersal
and migration, Springer-Verlag, Berlin, 289 pp.).

Urquhart makes generalizations that violate basic biological principles. He writes (p.
194): “The characteristics of size, shape and color that we now see in different species of
butterflies were indelibly fixed in the hereditary gene complex millions of years ago and
have persisted to the present time.” It is doubtful whether any trait can be “indelibly

296 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

fixed,” since genes are neither indelible nor fixed. Urquhart implies that evolution of
butterfly coloration eons ago stopped. The fossil record does not answer the question, but
industrial melanism demonstrates that, at least for some Lepidoptera, such evolution
continues. When Urquhart does address the literature, he becomes polemical (p. 190):

The scientific literature abounds with attempts to justify the mimicry theory as it
applies to birds feeding on butterflies. These papers contain an impressive array of
tables, charts and graphs resulting from experiments carried out in the crowded
confines of cages in a laboratory. By the use of abstruse terminology the research
assumes an aura of highly qualified investigations, but, when carefully analyzed,
contains nothing of real value and no meaningful conclusions.

Much of the information Urquhart presents may be found in his earlier book (Urquhart,
F. A. 1960, The monarch butterfly, Univ. Toronto Press, Canada, 361 pp.). That work
contains an extensive bibliography, which the current book lacks. More recent findings
are described in Urquhart’s other publications, which he lists in his current book.

Fred A. Urquhart made perhaps the most spectacular discovery in the field of lepi-
dopterology this century. This book will interest anyone who wonders what he has to say
about monarch butterflies and his studies of them. However, to find out what others have
to report about this species, readers will have to consult sources other than Urquhart’s

book.

KENNETH D. FRANK, 2508 Pine St., Philadelphia, Pennsylvania 19103.

Journal of the Lepidopterists’ Society
42(4), 1988, 296-297

THE GEOMETROID MOTHS OF NORTH EUROPE (LEPIDOPTERA: DREPANIDAE AND GEO-
METRIDAE), by Peder Skou. Translated from Danish edition by Elisabeth Folino. Ento-
monograph Vol. 6. 1986. 348 pp., 24 color pls., 358 figs. E. J. Brill/Scandinavian Science
Press, Leiden & Copenhagen. 17 x 25 cm, hard cover. $100.00.

This book covers all moths of the families Drepanidae, Thyatiridae, and Geometridae
known from Norway, Denmark, Sweden, and Finland. After a brief section introducing
categories of information to follow, the author moves directly to species treatments. These
consist of scientific name, author and year citation, plate and figure references, description,
range, habitat, flight period, and biology. Descriptions are usually brief, with emphasis
on variation. The color plates are among the best I have seen in sharpness and color value,
comparing favorably with those in Skinner’s 1984 Moths of the British Isles. Color plates
include both sexes and sometimes additional varieties; they accomplish well the identi-
fication of most species. Genitalia drawings for some species are included, especially in
difficult genera such as Eupithecia. Similar species are discussed when separation is
difficult, and the author has added text figures showing useful body parts such as wing
patterns, heads, and abdomens, with arrows pinpointing diagnostic features.

The worldwide range for each species is given, followed by detailed locality information
for the four countries featured. The habitat section gives variably detailed characteristics
of known sites, with black-and-white photos of typical habitats for many species. Flight
periods are general (“From late April until mid-May.”), and the biology section features
larval foodplants, time of year in larval stage, place of pupation, and other information.
Larvae and pupae are not described, but the book is generously illustrated with large
black-and-white photos of the caterpillars, usually on their foodplants. A final line tells
how the adult is best collected (at light, usually).

Following the species treatments are a selected bibliography and a table of distribution
for all species in the four northern countries.

The arrangement of taxa anticipates a new catalogue of European moths in preparation
by K. Schnack. Thus Thyatiridae are treated as a subfamily of Drepanidae. The subfamilies
of Geometridae are named as we now recognize them in the North American fauna, but

VOLUME 42, NUMBER 4 297

they are arranged in the same order as in the McDunnough 1938 Check List: Archiearinae,
Oenochrominae, Geometrinae, Sterrhinae, Larentiinae, and Ennominae. The Hodges et
al. 1983 Check List is the same except that Ennominae are moved to a position between
Oenochrominae and Geometrinae.

This book builds on several previous works, and appears to be an excellent identification
guide for species. Taxa above species are not described or defined, and there are no keys.
Recent expansions of European species to North America were missed (the establishment
of Hemithea aestivaria (Hiibner) in Canada was published in 1979); so range information
outside Scandinavia and Finland is questionably thorough.

The English composition is awkward in only a few places—forgivable, considering the
nationalities of the book’s producers. Some typographical errors were iound; and a number
of words were broken in mid-syllable—irritating to a former English teacher! The print
on coated paper is generally sharp, but there are numerous poorly impressed or broken
characters which marr an otherwise lavishly produced book.

The expense of this book will unfortunately preclude its addition to the bookshelves
of many amateur lepidopterists in North America and other parts of the lepidopterological
world outside Europe. This is sad because it is a first-rate work, and is just one of many
fine works on European moths that have recently come out. Those who specialize in
Geometroidea should certainly find it a valuable investment.

CHARLES V. COVELL JR., Department of Biology, University of Louisville, Louisville,
Kentucky 40292.

Journal of the Lepidopterists’ Society
42(4), 1988, 297

NOCTUELLES ET GEOMETRES D’ EUROPE. DEUXIEME PARTIE. GEOMETRES. Volume IV—
1919-1920. Jules Culot. Reprint edition, 1987. Apollo Books, Svendborg, Denmark. Order
from: Apollo Books, Lundbyvej 36, DK-5700 Svendborg, Denmark. Vols. IIJ-IV, DKK
1380.00; Vols. I-IV, DKK 2550.00.

This is the fourth and last volume of the set, with 167 pp. and color plates 38-70 (Figs.
772-1403). It covers part of Larentiinae (beginning with Eupithecia) and Ennominae,
although neither of these subfamily terms are used, much less defined. Having worked
with Eupithecia of North America and Chile, I found the 45 pages and 140 figures
devoted to this group particularly frustrating, as there are no descriptions or figures of
genitalia; to me, a study of these structures is almost a necessity to correctly name many
of the species. The same can be said about the species grouped together in Boarmia; in
this case a number of different generic names are in use today.

Having reviewed Vol. III (J. Lepid. Soc. 41:239), I need not repeat comments made
there, except to add that the text is in French in the entire set. This volume can be useful
to determine some of the more obvious and distinct species, but the scientific names date
from 1901. Much more up-to-date works are available and, to me, they could very well
prove more useful than the volumes of this set.

FREDERICK H. RINDGE, Department of Entomology, American Museum of Natural
History, New York, New York 10024.

Journal of the Lepidopterists’ Society
42(4), 1988, 298-300

INDEX TO VOLUME 42

(New names in boldface)

Acentria ephemerella, 247

Acontiinae, 285

acuta, Hylesia, 132

adaptation, 184

adunca, Viola, 184

alabamae, Catocala, 116

alfalfa butterfly, 196

amatus, Colotis, 57

Anaea andria, 263

andria, Anaea, 263

angustifolium, Vaccinium, 120

Animal Evolution in Changing Environ-
ments ... (book review), 146

announcement, 203

Antheraea polyphemus, 292

Apaturidae, 269

Archaeoprepona, 269

arcigera, Schinia, 144

Arizona, 204

asterius, Papilio polyxenes, 94

atlantis, Speyeria, 1

Azeta versicolor, 14

Becker, V. O., 149

betularia cognataria, Biston, 213

bina, Schinia, 144

biogeography, 19, 37, 269

biology, 46, 120, 182, 144, 184

Biston betularia cognataria, 213

Blanchard, R., 94

book reviews, 59, 146, 147, 149, 240, 244,
245, 295, 296, 297

bremnerii, Speyeria zerene, 184

Brou, V. A., Jr., 116

Brown, K. S., Jr., 240

butterflies, 19

Butterflies of Costa Rica ..
review), 240

Butterflies of North Dakota (book review),
244

butterfly

alfalfa, 196
monarch, 32
Butterfly Garden, The (book review), 147
Butterfly Gardener, The (book review), 147

., The (book

calcicola, Dudleya, 291
California, 291
capitatus, Croton, 263
Catocala

alabamae, 116

charlottae, 116
Catocalinae, 218
Charaxinae, 263, 269

charlottae, Catocala, 116
Chew, F. S., 59
chrysella, Schinia, 144
cibriani, Rhyacionia, 236
cladogram, 247
Claussen, C. L., 196
coastal habitat, 184
cognataria, Biston betularia, 218
cohabita, Ocalaria, 218
Colias eurytheme, 196
Colorado, 1, 46
Colotis

amatus, 57

vestalis, 57
conservation, 63
Costa Rica, 14
courtship, 1
Covell, C. V., Jr., 296

cover illustrations (announcement), 203

Crassulaceae, 291
Croesia curvalana, 120
Croton capitatus, 263
curvalana, Croesia, 120
Cyanotricha necyria, 103

Danaus plexippus, 32

Davenport, K. E., 291

Descimon, H., 94, 269

Dioptidae, 103

diphteroides, Leuconycta, 285
Dirig, R., 147

dispar, Lymantria, 213

distribution, 291

dos Passos, C. F. (bibliography), 168
dos Passos, C. F. (obituary), 155, 164
Dudleya calcicola, 291

egg weight, 138
Eichlin, T. D., 231
ephemerella, Acentria, 247
Ethmia

farrella, 281

powelli, 281
Ethmiinae, 281
Euphydryas gillettii, 37
Euptychia rubrofasciata, 276
Euptychiini, 276
European corn borer, 188
eurytheme, Colias, 196
evolution, 63
extinction, 37

Fabaceae, 14
Fairchild, W. L., 292

VOLUME 42, NUMBER 4

farrella, Ethmia, 281

faunus, Polygonia, 46

feature photographs (announcement), 203
fecundity, 138

Finkelstein, I. L., 60

Florida, 116, 281

Frank, K. D., 63, 295

Gazoryctra wielgusi, 204

Geometridae, 213

Geometroid Moths of North Europe ...,
The (book review), 296

Georgia, 19

gillettii, Euphydryas, 37

Glassberg collection, 31

Gliricidia sepium, 14

gracilis, Polygonia, 46

grassland, 184

Grey, L. P., 164

gynandromorph, 94

gypsy moth, 213

Hammond, P. C., 184

Hayes, J. L., 196

Hepialidae, 204

Hepialus, 204

Heppner, J. B., 281

Herman, W. S., 32

Hevel, G. F., 31

hippolyta, Speyeria zerene, 184
Holarctic, 204

Hyatt, J. A., 19

hybrid, 94

Hylesia acuta, 132
hypopharyngeal complex, 285

immature stages, 46, 132, 247
involucrata, Lonicera, 37

Johnson, K., 269
Josiinae, 103

Kitching, I. J., 218

arsen, ff. B., 57
larva, 46, 182, 247, 285
larval photoperiod, 263
Lehman collection, 31
lepidula, Leuconycta, 285
Leuconycta
diphteroides, 285
lepidula, 285
light pollution, 63
Lives of Butterflies, The (book review), 59
Lonicera involucrata, 37
Louisiana, 116

299

Lycaenidae, 291
Lymantria dispar, 213

machaon, Papilio, 94

Manley, T. R., 213

manuscript reviewers (1987), 152, 230

marking, 196

McCorkle, D. V., 184

meiosis, 94

melanic, 213, 292

Mexico, 132, 236, 276

Michigan, 231

migration, 32

Miller, J. S., 103

Miller, J. Y., 276

Miller, L. D., 276

Miller, W. E., 138, 203, 230, 236

mimicry, 276

Minnesota, 32

monarch butterfly, 32

Monarch Butterfly ... , The (book review),
295

morphology, 103, 218, 247

moth, gypsy, 213

moths, 63

Moths of Borneo ..
245

., The (book review),

necyria, Cyanotricha, 103

Neil, K. A., 285

Neotropics, 132, 236, 269, 276

New Brunswick, 292

New Mexico, 204

nigrozephyrus, Polygonia progne, 46

Noctuelles et Géométres d’ Europe . . . (book
review), 297

Noctuidae, 14, 116, 144, 213, 218, 285

Noctuoidea, 103

Notodontidae, 103

nubilalis, Ostrinia, 138

Nymphalidae, 1, 32, 37, 46, 184, 247, 263

obituaries, 60, 155, 164
Ocalaria

cohabita, 218

oculata, 218

quadriocellata, 218
oculata, Ocalaria, 218
Oecophoridae, 281
Olethreutinae, 236
Ostrinia nubilalis, 138
oviposition, 57

Panama, 218

Papilio
machaon, 94
polyxenes asterius, 94

300

Papilionidae, 94
parasitoid, 144
Passoa, S., 247
Peigler, R. S., 144, 245
Pellmyr collection, 31
Pennsylvania, 213
Peterson, R. D., II, 244
Philotes sonorensis, 291
phylogeny, 218, 247
Pieridae, 57, 196
plexippus, Danaus, 32
Polygonia

faunus, 46

gracilis, 46

progne nigrozephyrus, 46

satyrus, 46
polymorphism, 1, 213
polyphemus, Antheraea, 292
polyxenes asterius, Papilio, 94
Ponder, B. M., 120
population dynamics, 14
powelli, Ethmia, 281
Prepona werneri, 269

progne nigrozephyrus, Polygonia, 46

pupa, 46, 132, 247
Pyralidae, 138, 247
Pyraustinae, 138

quadriocellata, Ocalaria, 218

reproduction, 138
reproductive diapause, 268
Rhyacionia
cibriani, 236
rubigifasciola, 236
Riley, T. J., 263
Rindge, F. H., 297
Robbins, R. K., 31
rubidium, 196
rubigifasciola, Rhyacionia, 236
rubrofasciata, Euptychia, 276

Salvadoraceae, 57
Saturniidae, 132, 292
Satyridae, 276
satyrus, Polygonia, 46
Schinia

arcigera, 144

bina, 144

chrysella, 144
Schoenobiinae, 247
Scott, J. A., 1, 46
Seabrook, W. D., 120
seasonal dimorphism, 263
Selaginella, 276

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

sepium, Gliricidia, 14
Sesia

spartani, 231

tibialis, 231
Sesiidae, 231
Shapiro, A. M., 146
Sierra Nevada, 291
Solidago, 285
Solis, M. A., 149
sonorensis, Phylotes, 291
spartani, Sesia, 231
Speyeria

atlantis, 1

zerene

bremnerii, 184
hippolyta, 184
Sphingidae Mundi . .. (book review), 149
survey, 19, 213
systematics, 116, 204, 218, 236, 247, 269,
276, 285

Pantie, \hy, Jal, Oeil
Taraxacum, 285

Tennessee, 19

Texas, 144

Thomas, A. W., 292
tibialis, Sesia, 231

Tindale, N. B., 204
Tortricidae, 120, 236
Towers, A. A. (obituary), 60
trapping, 120, 213, 231

underwings, 116
urban ecology, 63

Vaccinium angustifolium, 120
versicolor, Azeta, 14

vestalis, Colotis, 57

Vinson, S. B., 144

Viola adunca, 184

Virginia, 19

Wagner, D. L., 204
Watson, C. N., Jr., 19
werneri, Prepona, 269
wielgusi, Gazoryctra, 204
Wilkinson, R. S., 155, 168
Williams, E. H., 37
Wolfe, K. L., 132

Young, A. M., 14
zerene, Speyeria

bremnerii, 184
hippolyta, 184

Date of Issue (Vol. 42, No. 4): 6 December 1988

|

EDITORIAL STAFF OF THE JOURNAL
WILLIAM E. MILLER, Editor

Dept. of Entomology
University of Minnesota
St. Paul, Minnesota 55108 U.S.A.

Associate Editors and Editorial Committee:
M. DEANE BOWERS, BOYCE A. DRUMMOND III, DOUGLAS C. FERGUSON,
LAWRENCE F. GALL, ROBERT C. LEDERHOUSE, THOMAS A. MILLER,
THEODORE D. SARGENT, ROBERT K. ROBBINS

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories
are Articles, General Notes, Technical Comments, Book Reviews, Obituaries, Feature
Photographs, and Cover Illustrations. Journal submissions should be sent to the editor at
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Abstract: An informative abstract should precede the text of Articles.

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Text: Manuscripts should be submitted in triplicate, and must be typewritten, entirely
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Literature Cited: References in the text of Articles should be given as Sheppard (1959)
or (Sheppard 1959, 1961a, 1961b) and listed alphabetically under the heading LITERATURE
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SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London.
209 pp.

196la. Some contributions to population genetics resulting from the study of

the Lepidoptera. Adv. Genet. 10:165-216.

In General Notes and Technical Comments, references should be shortened and given
entirely in the text as P. M. Sheppard (1961, Adv. Genet. 10:165-216) or (Sheppard, P.
M., 1961, Sym. R. Entomol. Soc. London 1:23-80) without underlining.

Illustrations: Only half of symmetrical objects such as adults with wings spread should
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Voucher specimens: When appropriate, manuscripts must name a public repository
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CONTENTS

SYSTEMATIC POSITIONS OF ACENTRIA EPHEMERELLA (DENIS &
SCHIFFERMULLER), NYMPHULINAE, AND SCHOENOBIINAE
BASED ON MORPHOLOGY OF IMMATURE STAGES (PYRA-
LIDAE). | Steven Passoa 20000

EFFECT OF LARVAL PHOTOPERIOD ON MATING AND REPRODUCTIVE
DIAPAUSE IN SEASONAL FORMS OF ANAEA ANDRIA (NYMPH-
ALIDAE). Thomas J. Riley 0000

SYSTEMATIC STATUS AND DISTRIBUTION OF THE LITTLE-KNOWN
CHARAXINE PREPONA WERNERI HERING & Hopp. Kurt

Johnson & Henri Descimon 0
A NEW EUPTYCHIA SPECIES FROM NORTHWESTERN MEXICO
(SATYRIDAE). Lee D. Miller & Jacqueline Y. Miller
A NEW SPECIES OF ETHMIA FROM THE FLORIDA KEYS (OECOPH-
ORIDAE: ETHMIINAE). J. B. Heppner _.
LARVAE OF NORTH AMERICAN LEUCONYCTA (NOCTUIDAE). Ken-
neth A, Neisseria

GENERAL NOTES

New distribution records and a probable new larval host plant for Philotes
sonorensis (Lycaenidae) in Kern and Tulare counties, California. Ken-
neth E. Davenport

A melanic male of Antheraea polyphemus polyphemus (Saturniidae). An-
thony W. Thomas & Wayne L. Fairchild

onan nen nee een en eens en an cen ennnnenn sa naeweneesenseswensenan

BooK REVIEWS
The Monarch Butterfly: International Traveler. Kenneth D. Frank -eccccco0

The Geometroid Moths of North Europe (Lepidoptera: Drepanidae and
Geometridae). Charles V. Covell Jr: 0

Noctuelles et Géométres d’Europe. Deuxiéme Partie. Géométries. Vol. IV—
1919-1920 (reprint). Frederick H. Rindge

INDEX TO VOLUME 42

291

292

Volume 48 1989 Number 1

ISSN 0024-0966

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

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A. Atkins 148

3 March 1989

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Cover illustration: Male of the primitive ithomiine Tellervo zoilus zoilus (Fabricius)
feeding from Stachytarpheta cayennensis (Rich.) Vahl (Verbenaceae). Illustration based
on observation of three of the butterflies persistently attracted to this plant which was
growing on a pebbled stream-bank in a clearing of dense rainforest at Mission Beach,
northern Australia. Feeding was confined to leaf surfaces and stems. Submitted by Andrew
Atkins, 45 Caldwell Ave., Dudley, N. S. W. 2290 Australia.

JOURNAL OF

THe LeEpPIDOPTERISTS’ SOCIETY

Volume 43 1989 Number 1

Journal of the Lepidopterists’ Society
43(1), 1989, 1-10

SUNDRY ARGYNNINE CONCEPTS REVISITED
(NYMPHALIDAE)

be RAUIA GREY
Rt. 1, Box 1925, Lincoln, Maine 04457

ABSTRACT. Suggestions for revisions in the Argynninae section of the 1981 Miller
and Brown checklist are presented, the taxa principally discussed being Semnopsyche,
Boloria, Proclossiana, and the Speyeria species nokomis, zerene, adiaste, callippe, hy-
daspe, atlantis, and mormonia. For the genera, hitherto undescribed characters are noted
as reasons for retaining Boloria while synonymizing Proclossiana and Semnopsyche.
Within the Speyeria species, several type locality changes are recommended and new
synonymies proposed. While most Speyeria “subspecies” intergrade extensively, the cat-
egory has appealed to many as a useful one, providing convenient tags for geographically
localized color forms. Despite lack of definitives, suggesting need for further studies, no
immediate drastic curtailing of subspecific listings is recommended.

Additional key words: Speyeria, Semnopsyche, Boloria, Proclossiana, North Amer-
ica. =

Comments, corrections, and suggested emendations to the Argyn-
ninae section in the Miller and Brown (1981) checklist are detailed
under the following headings. References for taxa mentioned herein
are available in that checklist if not found in the literature citations
appended.

Speyeria versus Semnopsyche

Miller and Brown (1981) correctly place Semnopsyche as a synonym
of Speyeria although giving no reason for doing so. I recently made
dissections that settle this matter unambiguously. To my embarrassment
I find that the species idalia (Drury) has a “secondary” bursal sac in
female genitalia. This was the character used by dos Passos and Grey
(1945) in delimiting Semnopsyche. But since idalia is the generotype
of Speyeria, Semnopsyche perforce becomes a junior synonym thereof.
Perhaps, then, some future splitter will want to categorize separately
those speyerians which have a simple, long ovate bursa, as described
by dos Passos and Grey (1945), a refinement which would seem un-

2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

desirable since that type of bursa is usual in argynnines worldwide.
Thus, it is an exception worth noting that a bursa almost exactly like
that of idalia may be seen in the Eurasiatic Mesoacidalia charlotta
(Haworth) (described by Haworth 1803). The latter’s bursa is longer
and pointing more dorsad than usual in argynnines, terminating in a
definite constriction, followed by enlargement to a small round sac.
Comparison of the Speyeria generotype with charlotta should be of
incidental interest since it adds hitherto unpublished evidence sup-
porting the idea that Nearctic argynnines placed in Speyeria are closest
in phylogeny to Mesoacidalia, as various students have speculated when
judging by wing facies and as Warren (1944) deduced from features
of male genitalia.

Boloria and Proclossiana

Studies of bolorian genitalia have lead me to conclude (reluctantly)
that generic restriction of Boloria Moore to pales (Denis & Schiffer-
miller) and the other species placed in that genus by Warren (1944)
probably should stand. This, however, is only because Miller and Brown
do not use subgenera. It is fairly certain that many, given the oppor-
tunity to study the surprisingly little-varied genitalia of females, plus
the invariably bifid uncus of males, would want to place all of the world
bolorians in a single category. Nevertheless, in the pales group as defined
by Warren (1944), the generic diagnosis fails to mention a heavily
spiculate uncus, and this, so far as I have seen, is a unique feature in
the bolorians, an extreme divergence of probable phylogenetic signif-
icance further supporting Warren’s categorical treatment.

There is less chance for divided opinions when reviewing Proclos-
siana Reuss. The variety of characters in male genitalia of bolorians
may be seen in dos Passos and Grey (1945:figs. 1-21). It would seem
to be in violation of consistency and parsimony to accord the single
species eunomia (Esper) a separate category when the male genitalia
appear no more distinctive than in the group now lumped in Clossiana.
Genitalia of eunomia, accredited to the then-prevailing taxon aphirape
Hiibner, are illustrated by dos Passos and Grey (1945:fig. 10). Addi-
tionally, when reviewing female genitalia of world bolorians (unpubl.
studies), I found only slight distinctions in eunomia, nothing to suggest
any degree of phylogenetic divergence above the species level. There-
fore, I recommend placing Proclossiana in synonymy under Clossiana.

Neotype and Type Locality of
Speyeria nokomis nokomis (W. H. Edwards)

A neotype for S. n. nokomis was fixed by dos Passos and Grey (1947).
It was a specimen purportedly collected by Oslar in the Mt. Sneffels

VOLUME 48, NUMBER 1 3

area of Ouray Co., Colorado. That action was criticized both in Miller
and Brown (1981) and in Brown’s (1965) monumental work on W. H.
Edwards types. The historical improbability of Mt. Sneffels as the exact
type locality of nokomis may be granted, but one source of doubt has
been removed: the butterfly does occur there, as has been verified by
Richard L. Klopshinske, of Olathe, Colorado. Vouchers, five pairs taken
at Mt. Sneffels Jeg. Klopshinske, are in the American Museum of Natural
History. I think our neotype designation meets even the rigid Code
requirements of today, since the very muddled history of this taxon, as
related by Brown (1965), has to be taken into account. The true type
locality promises to remain forever obscure, and, therefore, objections
could be raised against any other fixation whatsoever. As it stands, the
name is tied satisfactorily to all essential requirements of the original
description, namely, the neotype is from the “Rocky Mountains” and
it has a “cinnamon brown” disk. I therefore reaffirm the earlier (1947)
designation of neotype and type locality as having been an acceptable
solution to an admittedly murky problem.

Revisions Required in Speyeria zerene (Boisduval)

The “Yosemite” type locality chosen for S. z. zerene by dos Passos
and Grey (1947) is invalid in view of Lorquin’s itinerary as traced by
Masters (1979). Masters designated a type locality to conform thereto,
namely, Agua Fria, which is just west of Mariposa and about 56 km
from Yosemite. Evidently Masters concluded that the regional variation
was such that taxonomic concepts would remain unchanged.

The taxon gunderi (Comstock) is incorrectly placed as a subspecies
under S. coronis (Behr). Field evidence was discussed by Grey (1975)
to this effect: Intensive collecting in the Warner Mountains of California
reveals a massive regional phenotypic fluctuation in the species zerene
because of a collision between a yellow and a red subspecies in a ““Basin-
Sierran” tension zone. Some resulting individuals have yellow disks,
others have pallid greenish disks, yielding very close matches with the
type material of gunderi, which was beautifully depicted by Comstock
(1927:plate 27). In contrast, the species coronis, although here strictly
sympatric, is relatively little-varied and never appears to verge toward
the facies of gunderi. This appears to be sufficient proof for the com-
bination S. zerene gunderi.

Removal of gunderi from coronis to zerene necessitates putting cyn-
na dos Passos & Grey as a junior synonym of gunderi. Both taxa apply
to the same concept, that is, to a pallid yellow-disk subspeciation of
zerene.

Speyeria z. pfoutsi (Gunder) is a junior synonym of S. 2. platina
(Skinner), and should be so listed. The reasons why Gunder became
confused in this instance are detailed by Grey (1969).

4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Specific Recognition for
Speyeria adiaste (W. H. Edwards)

Contemporary students, including the Emmels (1973) and Howe
(1975), recognize adiaste as a distinct species. This change should be
made in the Miller and Brown checklist. Evidence for the subspecific
association was circumstantial, and is now outweighed by other con-
siderations, particularly the electrophoretic study by Brittnacher et al.
(1978). Students will have to continue to marvel at the narrow distri-
bution of the adiaste subspeciation in southern California, which is a
huge anomaly in Nearctic argynnine speciations, and something of a
world wonder in Argynninae.

Subspeciation of Speyeria callippe (Boisduval)
in the Sierra Nevada

Reexamination of the lectotype of S. c. juba (Boisduval), and com-
parison with the holotype of S. c. sierra dos Passos & Grey suggests
treating sierra as a junior synonym of juba. Variation of callippe in the
California mountains near Lake Tahoe has bewildered many collectors;
the name sierra was advanced to be descriptive of the yellowish and
greenish-disk variants, associating them with the proper species, cal-
lippe. The lectotype of juba appears to be within bounds assignable to
the diversity in the region from whence sierra derived. With the ap-
parent need to cut back on subspecific nomenclature, as discussed later
on, this would be a good place to start. Variation usually assigned to S.
c. inornata, centering more southerly in the Sierra, now appears to me
to be very distinct from juba. Perhaps inornata (W.H. Edwards) should
be resurrected from synonymy. Despite recent work by Arnold (19838,
1985), the whole Sierran callippe subspeciation badly needs further
study. Earliness of its flight season can be allowed for, and it appears
that colonies are far more numerous than might appear from available
records.

Type Locality of Speyeria hydaspe rhodope (W. H. Edwards)

Brown (1965) asserted that the type locality of rhodope should be
restricted to the “Fraser River Lowlands’, rejecting the dos Passos and
Grey (1947) restriction to the Cariboo District of British Columbia.
Three of the four recognized syntypes bear ““Cariboo” labels. But Brown
found a letter to Edwards from Crotch, the original collector, stating
that “... the small Argynnis with purple beneath . . .”’ was taken in an
area that Brown interprets to have been in a westerly direction from
100-Mile House, whereas the Cariboo District lies easterly from there.

Based on my visits in 1973 and 1975 to the approximate region

VOLUME 43, NUMBER 1 5

suggested by Brown, the habitat appears unsuitable to support any
hydaspe subspecies. Remaining undisturbed areas are mostly in dry
lodgepole pine forest. Going easterly, however, the foothill spruce-fir
forests of the Cariboo District present suitable habitat, and specimens
of rhodope were collected. To my knowledge there are no records of
this insect from the Frazer River lowlands, and very few from the
Cariboo area, suggesting that it is quite locally restricted.

Discrepancies between Brown’s conclusions and my findings could
be eliminated by postulating that the reference in Crotch’s letter was
not to rhodope but to Clossiana titania (Esper). In the terminology of
that day this bolorian would have been called a “‘small Argynnis’’ and
it also displays “purple beneath”. The habitat preferences of titania
vs. rhodope would support that alternative, titania being locally abun-
dant in the region where Brown would place rhodope. A return to the
“Cariboo” type locality for rhodope would make syntype labeling con-
sistent with field evidence.

Type Locality and Status of
Speyeria mormonia mormonia (Boisduval)

The lectotype of mormonia is from the Boisduval Collection leg.
Lorquin, and bears a “Lac Sal’ notation on a label. This, conjoined
with the name, plus the impression from facies that the specimen might
have derived from Utah, led dos Passos and Grey (1947) to designate
Salt Lake City as type locality.

A key bit of data, then unpublicized but now well known, is that
Lorquin did cross the Sierra from somewhere in northern California,
and probably collected as far east as extreme western Nevada.

My recent reinspection of the mormonia lectotype suggests that this
specimen originated in or closely adjacent to the Sierra Nevada of
California. That conclusion would be hard to prove because, as is so
often the case in Speyeria, it comes down to subtle nuances in color
and pattern. But aside from the ipse dixit, others whose opinions |
value, such as John Emmel and Paul Hammond, apparently have con-
curred that the specimen obviously is “‘sierran’”’.

Placement of the original mormonia in Utah resulted in S. m. arge
Strecker being applied to the California subspeciation. To accord with
the revised status of S. m. mormonia, arge becomes a junior synonym
thereof.

Miller and Brown (1981) recognized both m. arge and m. mormonia
as valid subspecies, and for the latter, proposed a type locality restriction
to Pyramid Lake, Nevada. Some historical justification was adduced
for that action, and undoubtedly it is close to the mark in a geographical
sense. Since I have never seen mormonia material from Pyramid Lake,

6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

I would be curious, as others might be, to learn what is available and
where it is deposited, and especially how well it matches the lectotype.

Status of Speyeria mormonia opis (W. H. Edwards)

This taxon is known from three syntypes, two described by Brown
(1965:322), and another in the Smithsonian which, like the first two,
appears derived from “Bald Mt.” leg. Crotch. These specimens support
the concept of opis as a subspecies of mormonia, and all three are
similarly characterized by small size and dorsal melanic pattern, being
ventrally sordid yellowish and unsilvered.

The Bald Mountain upland is in the Cariboo District of British Co-
lumbia, south of Barkerville (in the same area where I think rhodope
probably originated). In 1981, Edward Peters collected the first con-
temporary series of topotypical opis, 40 specimens, which he kindly
allowed me to examine and select 23 examples for deposit in the Amer-
ican Museum of Natural History. Variation in this sample is far greater
than in the above-noted syntypes, proving that in the Bald Mountain
population there are individuals which, compared to the syntypes, are
larger, smaller, lighter, darker, are silvered and unsilvered in about
equal proportions, and thus encompass the whole range of mormonia
variation in British Columbia.

This extensive variation at the type locality necessitates broadening
the concept of opis, and one result must be to synonymize jesmondensis,
described by McDunnough (1940), and considered in Miller and Brown
(1981) as validated by and attributable to dos Passos and Grey (1947).
The population represented by jesmondensis overlaps opis extensively
in variation, and also yields brown-disk forms reminiscent of the Oregon
subspecies m. erinna (W. H. Edwards). In addition, occasional speci-
mens are like m. washingtonia (Barnes & McDunnough), less melanic
and light yellowish to pale greenish discally, this being a form dominant
in the Okanagan region of British Columbia as well as in Washington.

It was a welcome surprise to find that the legendary opis is similar
to jesmondensis in being a hodgepodge of color forms, thus further
justifying suspicions that expanding nomenclature is not likely to pro-
mote better understanding of northwestern mormonia, which, in itself,
is a sharply discrete entity.

Type Locality of Speyeria mormonia bischoffii (W. H. Edwards)

The involved history of the taxon bischoffii was exhaustively sum-
marized by Brown (1965:316-321), who recommended that Sitka, Alas-
ka, be regarded as type locality. Nothing in the original description
supports that conclusion, and a dissenting criticism by dos Passos and
Grey (1965) has been reinforced by subsequent events. A colony of
mormonia has been discovered at Anchorage, far north of records extant

VOLUME 48, NUMBER 1 7

in 1965, at a logical spot for a mainland landfall by a sailing vessel
operating in the vicinity of Kodiak Island, that is, substantially where
dos Passos and Grey had conjectured. Even more persuasive, the An-
chorage melanics match the Edwards Kodiak neotype better than any
other Alaskan material I have seen. I therefore propose that the Sitka
type locality restriction be withdrawn in favor of Anchorage. This will
bring the original description, the neotype, and extant material into
better agreement. Vouchers, taken in the “Ski Bowl” near Anchorage,
leg. Bond Whitmore, are presently in the collection of Donald Eff,
Boulder, Colorado.

Subspecies in Speyeria

In earlier days it was easier to define subspecies of Speyeria. For the
most part they were distinctive in facies and well separated geograph-
ically. Advent of the automobile changed all that: road networks ex-
panded, collectors travelled more, and geographic coverage burgeoned.
Consequently, gaps between named subspecies have been partly or
wholly bridged by intermediates, giving rise to much name-shuffling
and even to questions about the validity of subspecies as a category.

Speyerian populations are notoriously varied in single localities, in-
cluding type localities. From this it follows that judgments made as to
what is “typical” of particular taxa, if based on samplings from type
localities, are subjective in presuming a local norm, or are inadequate
if based on a single holotype specimen. Still worse, most speyerian
subspecies are insufficiently isolated to prevent occasional straying. The
variety of local color forms in topotype populations can thus disperse,
mingle, and blend with others similarly afflicted. Where then, and how,
should subspecific lines be drawn?

A comment by Rindge (1987) gives one answer, and it carries the
weight of having resulted from surveying 37,500 specimens of Speyeria
during geographical rearrangement of series in the American Museum
of Natural History. He says: “... it quickly became apparent to me
that the majority of subspecific names proposed in this genus are, at
best, but random points on or at the end of clines, and hence are of
little or no scientific value. There appear to be very few completely
allopatric populations to which legitimate names might be attached.”

In a similar vein, Arnold (1983, 1985) recognized only 3 callippe
subspecies of 16 accepted in Miller and Brown (1981). I heartily sub-
scribe to the idea that the majority of subspecific names in Speyeria
could be dropped; they are essentially undefinable. However, I have
one major question concerning Arnold’s methodologies: As one well
acquainted with callippe variation in all the geographical regions Ar-
nold sampled, I can only wonder how through any mathematical leg-
erdemain the large brown-disk callippe callippe of the San Francisco

8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Peninsula can be directly associated with, say, the smaller, often yellow-
disk “‘sierra”’ of Plumas Co.? Or the red-disk silvered elaine of southern
Oregon with the sordid yellowish and unsilvered /aurina of the Green-
horns? Also, retention of semivirida as one of the three recognized
subspecies invites the criticism that semivirida in itself is a catchall,
beginning in the Tahoe region with creeping intrusion of brown into
the green-disk series, and culminating in British Columbia (for example
near Jesmond) with individuals nearly black discally. So why is semi-
virida singled out, except in a vague regional sense, from the other
intergrading forms that were synonymized?

These are relatively minor quibbles. A measure of how far I agree
in principle with Arnold is that I think ambiguities will persist until all
trinomials are discarded, and callippe is allowed to stand alone as a
distinct, nonoverlapping entity. But even the “‘rigidly definable species”
is by no means easily attained, as confirmed in a paper by Ferris (1983)
mentioned below. Before ending discussion of Arnold’s callippe study,
however, a serious objection must be stated, namely, that his method-
ology would seem destined to fail where most needed, namely in de-
termining what are to be accepted as valid Speyeria species. The prob-
lem here would be with the many local parallels that blur superficial
distinctions among species. In many places the differences in facies
between species widely agreed to be valid, can be, and often are, fewer
and more subtle than among the callippe subspecies Arnold synony-
mized.

This allied and more vital problem of delimiting species amid the
welter of subspecific variations is exemplified in Ferris (1983). No better
statement of the often confusing impressions conveyed by field-collected
series can be found in the literature; this reference should be consulted
by everyone interested in sympatry as a means of defining species. It
would be hard to dispute Ferris’s tentative hypothesis that two sibling
species may be involved in the Colorado “atlantis” material; this would
apply even more certainly to some of the Canadian series. On the whole,
however, it probably would require a hand-pairing breakthrough such
as Ferris envisions to decide among alternatives. The moral of Ferris’s
work perhaps is not to worry unduly about subspecies until we can say
more precisely what species to recognize. A recent paper by Scott (1988)
suggests a fairly objective and practicable way to assess conflicting data
in sympatrisms. He studied the situations in atlantis described by Ferris.
By rearing broods from areas where hesperis predominated over at-
lantis and vice versa, Scott obtained enough intermediates to incline
him toward the single polytypic entity theory.

The one thing certain from all this is that speciation and subspeciation
in Speyeria will continue to fuel debate and taxonomic disagreements.
Revising this section of Miller and Brown will be an unenviable chore.

VOLUME 438, NUMBER 1 9

So far as I would venture recommendations, I think the species, aside
from the elevation of adiaste, are standing the test of time.

The status of presently listed subspecies in Miller and Brown (1981)
is problematical. While I would retain most of the subspecific taxa, it
seems to me a distinction should be made between the utility of these
names versus their reality in nature as definable biological units. La-
fontaine (1987) retained certain Euxoa “‘subgenera”’ by the device of
a distinct typeface; strictly speaking they are synonyms but practically
speaking they are helpful in classifying that difficult genus. The situation
in Speyeria is analogous: variation is huge and not well understood,
sure to be further exploited since furnishing so many exciting possi-
bilities for geneticists and other students of evolution. It would be
convenient, then, to have discriminant tags available, and indeed there
is something to be said for their ‘reality’ —they enable succinct ref-
erence to color forms which students can see actually do prevail in
certain geographical areas. For that reason, if for no other, I suspect
they will refuse to die even if formally synonymized.

ACKNOWLEDGMENTS

J. F. Gates Clarke helped me study the National Museum of Natural History Speyeria
material. C. D. Ferris was consulted often and gave substantial aid.

LITERATURE CITED

ARNOLD, R. A. 1983. Speyeria callippe (Lepidoptera: Nymphalidae): Application of
information-theoretical and graph-clustering techniques to analyses of geographic
variation and evaluation of classifications. Ann. Entomol. Soc. Am. 76:929-941.

1985. Geographic variation in natural populations of Speyeria callippe (Bois-
duval) (Lepidoptera: Nymphalidae). Pan-Pac. Entomol. 61:1-23.

BRITTNACHER, J. G., S. R. Sims & F. J. AYALA. 1978. Genetic differentiation between
species of the genus Speyeria (Lepidoptera: Nymphalidae). Evolution 32:199-210.

BROWN, F.M. 1965. The types of the nymphalid butterflies described by William Henry
Edwards. Part I. Argynninae. Trans. Am. Entomol. Soc. 91:233-350.

ComsTock, J. A. 1927. Butterflies of California. Published by author, Los Angeles,
California. 334 pp.

Dos Passos, C. F. & L. P. GREY. 1945. A genitalic survey of Argynninae (Lepidoptera,
Nymphalidae). Am. Mus. Nat. Hist. Novit. 1296, 29 pp.

1947. Systematic catalogue of Speyeria (Lepidoptera, Nymphalidae) with des-

ignation of types and fixations of type localities. Am. Mus. Nat. Hist. Novit. 1370,

30 pp.

1965. Notes on certain lectotypes and neotypes designated by the authors in
their systematic catalogue of Speyeria (Lepidoptera: Nymphalidae). Trans. Am. Ento-
mol. Soc. 91:357-358.

EMMEL, T. C. & J. F. EMMEL. 1973. The butterflies of southern California. Science Ser.
26, Nat. Hist. Mus. of Los Angeles Co., Los Angeles. Pp. 29-30.

FERRIS, C. D. 1983. Speyeria atlantis phenotypes in the southern Rocky Mountains
(Lepidoptera: Nymphalidae: Argynninae). J. Res. Lepid. 22:101-114.

Grey, L. P. 1969. On the Gunder collection of argynnids (Lepidoptera: Nymphalidae).
J. Res. Lepid. 8:55-64.

1975. Argynnis gunderi: A many-splendored snafu. News Lepid. Soc. No. 4

(August).

10 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Haworth, A. H. 1803. Lepidoptera Brittanica, sistens digestionen novam Lepidopto-
rum. Part 1. J. Murray, London. P. 32.

Howe, W. H. 1975. The butterflies of North America. Doubleday & Co., Garden City,
New York. 442 pp.

LAFONTAINE, J. D. 1987. The moths of America north of Mexico. Fascicle 27.2. Noc-
tuoidea. Noctuidae (part). Wedge Entomol. Res. Found., Washington. 237 pp.
MASTERS, J. H. 1979. The type locality of Argynnis zerene Boisduval (Nymphalidae):

_ A correction. J. Lepid. Soc. 33:137-138.

McDuNNouGH, J. 1940. The argynnids of the Cariboo Region of British Columbia.
Can. Entomol. 72:23-25.

MILLER, L. D. & F.M. BROWN. 1981. A catalogue/checklist of the butterflies of America
north of Mexico. Mem. Lepid. Soc. 2, 280 pp.

RINDGE, F. H. 1987. Speyeria collection of Paul Grey to the American Museum of
Natural History. J. Lepid. Soc. 41:123.

ScoTT, J. A. 1988. Speyeria atlantis in Colorado: Rearing studies concerning the relation
between silvered and unsilvered forms. J. Lepid. Soc. 42:1-138.

WARREN, B.C. S. 1944. Review of the classification of the Argynnini: With a systematic
revision of the genus Boloria (Lepidoptera; Nymphalidae). Trans. Roy. Entomol. Soc.
London 94:1-101.

Received for publication 2 February 1988; accepted 31 August 1988.

Journal of the Lepidopterists’ Society
43(1), 1989, 11-32

PHYLOGENY AND ZOOGEOGRAPHY OF THE BIGGER AND
BETTER GENUS ATALOPEDES (HESPERIIDAE)

JOHN M. BURNS

Department of Entomology, NHB 169, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560

ABSTRACT. What makes Atalopedes bigger and better is the addition of two tropical
species, A. clarkei, new species and A. bahiensis (Schaus), and the subtraction of another,
nabokovi (Bell & Comstock), which belongs in Hesperia. Comparison of genus Atalopedes
with its sister Hesperia, using characters of size, antenna, facies, stigma, and, especially,
male and female genitalia, precedes comparisons among the species of Atalopedes, using
these same characters. The five species form three highly distinct groups, whose phylo-
genetic sequence is (1) A. campestris (Boisduval), which ranges from equator to USA;
(2) the mesogramma group—A. mesogramma (Latreille), on most Greater Antilles, Isle
of Pines, and some Bahama Islands including New Providence, and A. carteri Evans,
New Providence Island; and (3) the clarkei group—A. clarkei, Margarita Island, Vene-
zuela, plus Cartagena, Colombia, and A. bahiensis, coastal central Brazil. The far-out
clarkei group has switched its ecologic niche to seashore grass; habitat is very restricted.
The older the species of Atalopedes, the wider its geographic range.

Additional key words: genitalia (male and female), Hesperia, H. nabokovi, taxonomy,
evolution.

What makes Atalopedes bigger and better is the addition of two
tropical species, an undescribed one plus its misplaced sister, and the
subtraction of another, nabokovi (Bell & Comstock), which belongs in
Hesperia (Burns 1987).

Because the five resulting species form three highly distinct clusters,
Atalopedes seems riddled by extinctions—far more than sister genus
Hesperia, which, with four times as many species, is still relatively
compact.

Atalopedes is American; Hesperia, mostly so—but it also spans the
Palearctic, an extension of range considered rather recent (Scudder
1874, MacNeill 1964). Though basically northern in modern distribu-
tion, Hesperia turns out, with the inclusion of H. nabokovi, to be lowland
Hispaniolan as well as Holarctic, which raises questions about area of
origin, particularly since nabokovi is among the oldest species of Hes-
peria (Burns 1987). The idea that Hesperia may have arisen in the
Neotropics becomes less astonishing in light of the fact that sister Ata-
lopedes occurs from the middle of South America to the middle of North
America and in much of the West Indies.

As currently known, the few species of Atalopedes tend to replace
one another geographically. The only species in the continental United
States is widespread and weedy and thus (for a skipper) familiar. Ata-
lopedes campestris (Boisduval) ranges from about the equator, through
northern South America (up to at least 3100 m in Colombia), through
Central America, and through Mexico, to most of the United States

12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

below Canada. Across the southern United States, this multivoltine grass-
eater flourishes, especially in disturbed open habitats, becoming scarcer
toward higher latitudes and altitudes. It commonly invades various
northern states in which it fails to overwinter: records typically reflect
the mid- to latter warm season. To many temperate lepidopterists (such
as-Clark & Clark 1951, Shapiro 1966, 1974, Opler & Krizek 1984), this
mobile skipper is a classic immigrant.

Three of the four remaining species of Atalopedes are emigrants,
wholly or partly on islands. Only one has much of an insular distribution:
A. mesogramma (Latreille) extends through all the Greater Antilles
except Jamaica, as well as south of Cuba to the Isle of Pines and north
of it to some of the Bahama Islands. By contrast, A. carteri Evans occurs
on New Providence Island in the Bahamas; and A. clarkei, new species,
on Margarita, an island (68 km east to west) roughly 25 km north of
the Venezuelan mainland and 250 km west of Trinidad.

Beyond that, this new skipper hails from the Caribbean coast of
Colombia (Cartagena); and its sister, A. bahiensis (Schaus), from the
Atlantic coast of central Brazil (Bahia and Espirito Santo)—though,
surely, neither is quite so localized.

The last reviser of Atalopedes (Evans 1955) saw one species, A.
mesogramma, as polytypic. Two subspecies, A. m. mesogramma and
A. m. apa Comstock, are real, but apparently minor, differentiates of
no special concern here: the latter (with broader light markings that
make it brighter overall) occurs on Puerto Rico and Hispaniola; the
former (with narrower light markings that leave it darker), to the west.
But what Evans (1955:339) described as a third subspecies, A. m. carteri,
differs more sharply from the others in both size and facies; and, to
quote Evans, it “occurs [at Nassau on New Providence in the Bahamas]
with mesogramma, which probably is a visitor from Cuba, while carteri
breeds locally.”’ Presumably taking this as evidence of sympatry be-
tween differentiates without breakdown of their sizeable differences,
Riley (1975:186) called carteri a full species. The situation points to
double invasion, with complete speciation on the part of the first invader
(see Mayr 1963:504-507 for discussion of multiple invasions).

The sistership of Atalopedes and Hesperia (Burns 1987) has mostly
been missed. Lindsey (1921) and Lindsey et al. (1931), in treating the
skippers of North America north of Mexico, inserted one genus (Hy-
lephila) between Atalopedes and Hesperia; and Evans (1955), in treat-
ing the entire New World fauna, five (Appia, Linka, Polites, Wallen-
grenia, and Pompeius), though all these workers, in attempting to
characterize Atalopedes, compared it with Hesperia, Evans (1955:338)
even going so far as to say, “Palpi as Hesperia... . Resembles Hesperia
in facies.” MacNeill (1975) set the two genera side by side, noting a

VOLUME 43, NUMBER | 13

relation. Yet, in the subsequent spate of North American butterfly books
and checklists, only Stanford (1981) and Pyle (1981) followed his ar-
rangement (both consulted MacNeill).

Atalopedes vis-a-vis Hesperia

On the whole, Atalopedes (Figs. 1-18, 42, 43) is very like Hesperia,
but less sexually dimorphic with respect to wing shape (in Hesperia,
females have much more rounded wings than males). The few depar-
tures in Atalopedes from an intergenerically shared pattern are on the
ventral secondary, which is not surprising since this is the surface a
resting individual shows the world (Figs. 42, 43): most extreme is a
vertical pale stripe down the middle of a dark wing from vein 8 to
mid-space lc in A. mesogramma (Figs. 8, 18) and A. carteri (Fig. 10).
Atalopedes is more variable in size, with A. mesogramma (Figs. 7, 8,
17, 18) averaging larger than any species of Hesperia (except H. na-
bokovi!) and A. clarkei (Figs. 3, 4, 18, 14) and A. bahiensis (Figs. 5, 6,
15, 16), smaller. In both genera, the antenna and its apiculus are rel-
atively short, and the club, stout; but the apiculus is a bit longer in
Atalopedes.

The stigma (Figs. 19-23)—an elaborate communicative device
spreading over the dorsal primary of the male from the junction of
veins 3 and 4 inward and downward to vein 1—resembles that of
Hesperia but clearly differs from it. The central, dustlike microandro-
conial mass (terminology of MacNeill 1964) is more or less open to
view, not neatly enclosed by flanking rows of large, wide, silvery-gray
scales as it is in Hesperia. The dark brown apical and lower brush
patches are more developed, more conspicuous, than they are in Hes-
peria, although the scales comprising them are narrower. The poststig-
mal patch, too, is well developed and dark, contrasting with adjacent
areas of the wing. Altogether, the stigma of Atalopedes looks less linear
and more massive than that of Hesperia.

Parts of the male (though not the female) genitalia of Atalopedes
hint at those of Hesperia. In both genera the valva ends in two large,
pointed, more or less dorsally-directed projections whose bases join on
the lateral valval surface by way of a smooth, U-shaped edge. This U,
narrow in Atalopedes (Figs. 24-33), varies from narrow to wide in
Hesperia. The more distal projection is the more complex, almost always
extending forward, medial to the proximal projection. In both genera
the aedeagus is slender and comparatively simple—quite unlike the
formidable one bristling with bizarre, often outsized, titillators and
cornuti in such related genera as Yuretta, Polites, Ochlodes, Poanes,
and Paratrytone.

Many genitalic features divide Atalopedes from Hesperia (Burns

14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

?

Fics. 1-18. Adults of Atalopedes (all x1) (in USNM unless otherwise indicated): 1-
10, males; 11-18, females; odd numbers, dorsal views; even numbers, ventral views. 1,
2, campestris, Charleson St., Annandale, Fairfax Co., Virginia, USA, 1 September 1979,
J. M. Burns; 3, 4, clarkei, El] Morro, Margarita Island, VENEZUELA, 12 February 1985,
J. F. G. Clarke; 5, 6, bahiensis, holotype, Bahia, BRASIL; 7, 8, mesogramma, Tanamo,
CUBA, March 1902; 9, 10, carteri, Nassau, BAHAMAS, 1 February 1898, H. G. Dyar;
11, 12, campestris, Austin, Travis Co., Texas, USA, 3 June 1967, J. M. Burns; 13, 14,
clarkei, Cartagena, COLOMBIA, 14 July 1969, J. Herrera (collection of C. D. MacNeill);
15, 16, bahiensis, Conceicao da Barra, Espirito Santo, BRASIL, 25 March 1969, C. & C.
T. Elias (collection of O. H. H. Mielke); 17, 18, mesogramma, Matanzas, CUBA, October.

VOLUME 43, NUMBER 1 13,

Fics. 19-23. Stigmas on the dorsal left primaries of the Atalopedes males in Figs. 1-
10. 19, campestris; 20, clarkei; 21, bahiensis; 22, mesogramma, 23, carteri.

1987). The aedeagus is longer than the rest of the intact genitalia in
Atalopedes (Figs. 25, 27, 29, 31, 33) but not in Hesperia; and it bears
either two multidentate cornuti (Figs. 24, 25) or none (Figs. 26-33) in
Atalopedes compared with a single bidentate cornutus in Hesperia.
Paired prongs projecting forward from the front end of the juxta are
short and stout in Atalopedes (Figs. 25, 27, 29, 31, 33) but long and
delicate in Hesperia. The valva is elongate and its top and bottom about

16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 24,25. Male genitalia of Atalopedes campestris from Skippers, Greensville Co.,
Virginia, USA, 25 July 1983, J. M. Burns (genitalic dissection no. X-2107). 24, Tegumen,
uncus, gnathos, distal ends of valvae, and distal end of aedeagus in dorsoposterior view;
25, Complete genitalia (minus right valva) in left lateral view.

parallel in Atalopedes (Figs. 25, 27, 29, 31, 33), whereas the top of the
valva is usually quite humped in Hesperia; and the distal projection of
the valva, apart from its one (Figs. 24, 25) or two (Figs. 26-33) big
points, is smooth along its dorsal edge in Atalopedes (Figs. 24-33) rather

VOLUME 43, NUMBER 1 17

than serrate, as it is in Hesperia. The dorsal tegumen forms a more or
less raised sac in Atalopedes (Figs. 24-33) but not in Hesperia. The
uncus, always short and blunt in Atalopedes (Figs. 24-33), ranges from
a roughly similar state to long and pointed in Hesperia. The gnathos,
which may be well developed (Figs. 24, 25), reduced (Figs. 30-33), or
vestigial (Figs. 26-29) in Atalopedes, is always well developed in Hes-
peria. (For figures of male genitalia of Hesperia, see Scudder 1874,
1889, Skinner & Williams 1924, Lindsey et al. 1931, Lindsey 1942,
MacNeill 1964, and Burns 1987.)

Female genitalia of the two genera are thoroughly distinct. A mid-
ventral prong projecting backward and downward from the back part
of the lamella postvaginalis (Figs. 34-41) marks Atalopedes. The body
of the lamella postvaginalis in Atalopedes consists of midventral scler-
otization (extending the roof of the ductus bursae to the base of the
prong) flanked by surfaces (curving upward and outward) that may be
entirely (Figs. 34, 35, 40, 41) or scarcely (Figs. 36-39) sclerotized. In
ventral outline the simpler lamella postvaginalis of Hesperia approaches
a rectangle. The ductus bursae is more or less symmetric in Atalopedes
(Figs. 34-41) but asymmetric in Hesperia, where it begins with a caudal
chamber on the right and then slants to the left. Sclerotization of the
ductus bursae stops before the junction of the ductus seminalis in Ata-
lopedes (Figs. 35, 37, 39, 41) but after its junction in Hesperia. And
the ductus bursae connects with the corpus bursae by means of a dorsal
jog in Atalopedes (Figs. 35, 37, 39, 41) but not in Hesperia. The corpus
bursae itself is essentially cylindrical in Atalopedes (Figs. 34-41), spher-
ical in Hesperia. (For figures of female genitalia of Hesperia, see mainly
MacNeill 1964 but also Gillham 1954 and Burns 1987.)

Additions to Atalopedes

Full and formal treatment of all included species (which would swell
an ordinary taxonomic text) is not essential. I have introduced them all
already by way of their peculiar distributions and gone on, in con-
nection with strict definition of the genus, to provide comparative
figures of their facies, stigmas, and genitalia—a wealth of information
useful at the specific level, as well as above. The species are few enough
and diverse enough to stand out. Their characters flood the upcoming
discussion of phylogeny anyway. Ritual is therefore restricted to the
two species in nominal need.

These, the South American sisters, are the most southern in distri-
bution and the smallest in size. Because the sample of the new species
is far larger than that of the old—87 males and 3 females as opposed
to 1 male and 1 female—and because a larger sample affords a better
description, the new species comes first.

18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Atalopedes clarkei, new species

(Figs. 3, 4, 13, 14, 20, 26, 27, 36, 37, 42, 43)

Length of right primary (mm).

Sample Sex N Range Mean + SE SD CV
Margarita I, Venezuela 4 28 11.0-13.4 12.28 + 0.09 0.46 3.70
Q 1 2a
Cartagena, Colombia 3 8 LNO=130" ess) 40:27 0.78 6.53
Q 2; 13.2-14.0

Antenna. Club shorter and thicker in males than in females; anterodorsally, from base
to (or close to) start of apiculus, club scaled orange in males, blackish brown in females.
(This marked sexual dimorphism involving size, shape, and color of the antennal club is
of general occurrence in Atalopedes—not to mention Hesperia and some other genera—
though details of expression can vary between species: for instance, the anterodorsal
orange stripe of males extends farther out the club in A. clarkei and A. campestris than
it does in A. mesogramma and A. carteri.) Nudum usually 7/5: of 34 specimens with at
least one antenna intact, 3 are 7/4; 27, 7/5; and 4, 7/6. (Nudum usually 7/7 in campestris,
8/8 in mesogramma. )

Facies. Much as in campestris (compare Figs. 8, 4 with 1, 2; and 18, 14 with 11, 12)
except for the ventral secondary, where a yellow ray from the cell through most of spaces
4 and 5 divides two yellow spots in spaces 2 and 8 from a third (not always present) in
space 6 (Figs. 4, 14, 42, 43). This distinctive ventral secondary relates readily to that of
many campestris males (compare Figs. 4 and 2) but not to that of campestris females
(compare Figs. 14 and 12). (It suggests the ventral secondary of marsh-dwelling species
of Poanes.) Spots of the primary (which become hyaline in females of campestris and
mesogramma) opaque in both sexes.

Stigma. Well developed (Fig. 20) but not hypertrophied as it is, slightly, in mesogramma
(Fig. 22) and, grossly, in campestris (Fig. 19).

Male genitalia. Valva similar to that of mesogramma and carteri (compare Figs. 26,
27 with 30-33), with the more distal of the two large, terminal projections expanded into
two major, dorsally directed points just mediad of the one-pointed proximal projection.
Tip of uncus broadly notched (Fig. 26) and roughly textured (Figs. 26, 27). Gnathos
vestigial (Figs. 26, 27). No cornuti, but tooth present on left side of aedeagus before the
single, backward pointing, terminal tooth (Figs. 26, 27).

Female genitalia. Midventral prong projecting from back of lamella postvaginalis short
(compare Figs. 36, 37 with 34, 35 and 40, 41). Much of lamella postvaginalis unsclerotized
(Figs. 36, 37). Simple, sclerotized ductus bursae modestly and rather evenly tapered from
ostium bursae forward (Fig. 36).

Material examined. Holotype: Male. VENEZUELA, [Nueva Esparta], Margarita I[slan]d,
El Morro, [ca. 4 km E Porlamar], on seashore grass, 12 Feb[ruary] 1985, J. F. G. Clarke.
Deposited in National Museum of Natural History, Smithsonian Institution (USNM).

Paratypes (39): 27 males with same data as holotype, plus 7 genitalia dissections (USNM).
1 male, 1 female, VENEZUELA, Nueva Esparta, Margarita Island, near Pampatar, be-
tween Playa Morefio and Playa El Angel, 19 August 1987, J. Glassberg & J. Scott, plus 2
genitalia dissections (USNM). 2 females, COLOMBIA, Cartagena, 14 July 1969, J. Herrera,
plus 2 genitalia dissections (collection of C. D. MacNeill). 8 males with same data except
15 July 1969, plus 4 genitalia dissections (collection of C. D. MacNeill).

Atalopedes bahiensis (Schaus), new combination
(Figs. 3,6, 1a, 6, 205928, 2938759)

Thymelicyus [sic] bahiensis Schaus (1902:436).
[The brief, verbal original description of nothing but facies and wing-

VOLUME 48, NUMBER 1 19

spread is so vague that Evans (1955:337) questioningly assigned ba-
hiensis to the synonymy of Pompeius amblyspila (Mabille).]

Length of right primary (mm). Male, 11.8; female, 13.2; so probably about as in A.
clarkei.

Antenna. [Male, missing.] Female, club scaled blackish brown anterodorsally; nudum
7/5 or 7/6 (count equivocal owing to incomplete suture).

Facies. Reminiscent of clarkei but darker, without the pale ray (parallel to veins through
the middle of the ventral secondary) characteristic of that species (compare Figs. 5, 6
with 8, 4; and 15, 16 with 18, 14); chiefly in female, brown background encroaches upon
pale pattern elements (compare Figs. 15, 16 with 18, 14). Spots of the primary opaque
in female as well as in male, as in clarkei (but not campestris and mesogramma).

Stigma. Similar to that of clarkei (Fig. 20), but lower brush patch perceptibly larger
and poststigmal patch, smaller (Fig. 21).

Male genitalia. Overall, very like those of clarkei, but with scattered differences (com-
pare Figs. 28, 29 with 26, 27). In lateral view (Fig. 29 versus 27), one-pointed, laterally
placed, proximal terminal projection of valva more nearly horizontal, extending farther
back to about end of valva; two-pointed, medially placed, distal terminal projection of
valva with its proximal point higher than its distal point, rather than the reverse; posterior
edge of valva not curved prominently backward the way it is in clarkei; anterior end of
tegumen more angular, less rounded. In dorsoposterior view (Fig. 28 versus 26), tegumen-
uncus usually narrower; left lateral tooth near end of aedeagus longer and basally much
broader.

Female genitalia. Similar to those of clarkei, but midventral prong projecting from
back of lamella postvaginalis even shorter, lamella postvaginalis still more narrowly scler-
otized, and ductus bursae more (and more abruptly) flared around level of ostium bursae
(compare Figs. 38, 39 with 36, 37).

Material examined. Male. Holotype: Bahia, Brazil; Collection W. Schaus; Thymelicus
Bahiensis Sch{au]s Type [handwritten in black]; Type No. 5999 U.S.N.M. [label red];
Genitalia No. X-2357 J. M. Burns 1987.

Female. Conceicao [da] Barra, E[spirito] S[anto], Brasil, 25 March 1969, C. & C. T.
Elias; DZ 3081; Genitalia No. X-2390 J. M. Burns 1987; [collection of O. H. H. Mielke].

Niche Switch to Seashore Grass

Atalopedes is a genus of multivoltine grass-eaters. One or, more likely,
both additions to it have made a striking ecologic shift analogous to
those of marsh-dwelling species of Euphyes, Poanes, Ochlodes, Prob-
lema, and Panoquina. Atalopedes clarkei lives in a peculiar and rela-
tively simple community dominated by short grass growing in sand
behind sandy marine beach (Figs. 42-44). This physically harsh envi-
ronment may offer fewer biotic pressures. At any rate, as in marsh-
dwelling skippers whose foodplant is local but sometimes locally abun-
dant, populations can be extremely local (and thus hard to find), but
the density of a population can be high. It was certainly high at El
Morro on Margarita when J. F. G. Clarke found A. clarkei (see Material
examined). In that area, man is the worst enemy of A. clarkei now
because he is rapidly wrecking its limited seashore habitat with hotels
and such (J. Glassberg pers. comm.).

The habitat must have been basically similar, and the population
density more or less high, when J. Herrera found A. clarkei at the
Colombian seaport of Cartagena (see Material examined). During his

20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 26, 27. Male genitalia of Atalopedes clarkei from El Morro, Margarita Island,
VENEZUELA, 12 February 1985, J. F. G. Clarke (X-2104). 26, Tegumen, uncus, distal
ends of valvae, and distal end of aedeagus in dorsoposterior view; 27, Complete genitalia
(minus right valva) in left lateral view.

brief time in Cartagena, Herrera collected at the airport, which is right
by the water (C. D. MacNeill pers. comm.).

Indirect evidence suggests that seashore grass is also the habitat of
A. clarkei’s close Brazilian sister. The female of A. bahiensis, taken
rather recently, is without question from a town on the coast. The male,
collected before 1902, is labelled “Bahia Brazil’? which may mean the
large coastal state of Bahia but probably refers to its capital, a seaport

VOLUME 43, NUMBER 1 eS

Fics. 28, 29. Male genitalia of Atalopedes bahiensis, holotype, from Bahia, BRASIL
(X-2357). 28, Tegumen, uncus, distal ends of valvae, and distal end of aedeagus in
dorsoposterior view; 29, Complete genitalia (minus right valva) in left lateral view.

now called Salvador but formerly called Bahia. Restriction to special
habitat could explain the scarcity of the skipper in collections.

Phylogeny of Atalopedes

The species of Atalopedes form three obvious groups: (1) campestris,
(2) mesogramma and carteri (the mesogramma group), and (3) clarkei
and bahiensis (the clarkei group). Between these groups, differences
are large; within them, small—so small that, when I think about the

22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 30, 31. Male genitalia of Atalopedes mesogramma from Guantanamo Bay,
CUBA, 14 September 1943, W. H. Wagner (X-2115). 30, Tegumen, uncus, gnathos, distal
ends of valvae, and distal end of aedeagus in dorsoposterior view; 31, Complete genitalia
(minus right valva) in left lateral view.

genus as a whole, it comes down to a trio of widely spaced points,
which beg for connection. Drawing those real but unseen lines of phy-
logenetic relationship (Fig. 45) is trying.

Though extinction seems to have obliterated most of the record so
that the surviving species groups stand out and apart from one another,

VOLUME 43, NUMBER 1 23

Fics. 32, 33. Male genitalia of Atalopedes carteri from Nassau, BAHAMAS, 1 Feb-
ruary 1898, H. G. Dyar (X-2116). 32, Tegumen, uncus, gnathos, distal ends of valvae,
and distal end of aedeagus in dorsoposterior view; 33, Complete genitalia (minus right
valva) in left lateral view.

there is no problem uniting them, using a diversity of characters, in a
well-defined, monophyletic higher group distinct from Hesperia and
sister of it (see Atalopedes vis-a-vis Hesperia). But within Atalopedes,
it is unfortunate (from a phylogenetic viewpoint) that many character
states are unique to each group and that, where they are shared, it is
hard to peg them as primitive or derived.

The gnathos of the male genitalia becomes critical. Despite a hy-
pertrophied stigma (Fig. 19), the most primitive species of Atalopedes
must be campestris because its gnathos is fully developed (Figs. 24, 25)
as it is in every species of sister genus Hesperia. In the remaining species
of Atalopedes, the gnathos is reduced—less severely in mesogramma

24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

34

Fics. 34, 35. Female genitalia of Atalopedes campestris from Skippers, Greensville
Co., Virginia, USA, 30 September 1981, J. M. Burns (X-2169). 34, Sterigma and bursa
copulatrix in ventral view; 35, The same, plus part of the ductus seminalis, in right lateral
view.

VOLUME 43, NUMBER 1 p25)

and carteri (Figs. 30-33) than in clarkei and bahiensis (Figs. 26-29)—
so that these two groups must be derived, and in that order.

The notion of a gradual and progressive trend toward elimination of
the gnathos is entirely different from abrupt total suppression of sec-
ondary sex characters like the costal fold, metatibial tufts plus meta-
thoracic pouch, or stigma of male skippers. I hypothesized long ago
that presence or absence of those elaborate odor-disseminating struc-
tures could have a simple genetic basis such that they might easily
reappear in a descendant species after having been switched off in an
ancestor (Burns 1964:196-197). Where reversals of that kind are likely,
establishing a sequence can be difficult. But in Atalopedes, parsimonious
interpretation of its three discrete levels of gnathos expression yields
the main lines of Fig. 45.

Two more features of the male genitalia lend what may be flawed
support to the argument that campestris is the most primitive species.
First, cornuti occur in campestris (Figs. 24, 25) but in no other species
of Atalopedes (Figs. 26-33). The weakness here is that even though all
species of sister Hesperia have a cornutus, it is always single and usually
bidentate and hence may not be strictly homologous with the paired
multidentate cornuti in campestris. Second, the distal terminal projec-
tion of the valva comes to a single, dorsally directed point in campestris
(Figs. 24, 25) but broadens to two conspicuous, dorsally directed points
(medial to the proximal projection) in all other species of Atalopedes
(Figs. 26-33). However, since the latter configuration more nearly re-
sembles the distal, terminal valval projection in all species of Hesperia,
it may be primitive and the one-pointed projection of campestris, de-
rived.

Other male genitalic characters bolster the sequence that puts the
mesogramma group between campestris and the clarkei group. The
strangely protuberant uncus of the mesogramma group (Figs. 30-33)
relates clearly to that of campestris (Figs. 24, 25), though differing in
many details and appearing, as a whole, rather less extreme. (Among
the figures just cited, the protuberance shows better in the dorsoposterior
views.) The protuberance is wanting in the clarkei group (Figs. 26-29).
On the other hand, the posterior tip of the uncus is more deeply notched
in the mesogramma group (Figs. 30, 32) than it is in campestris (Fig.
24); and the deep notch persists in the clarkei group where, moreover,
its sides diverge widely (Figs. 26, 28). The very tip of the aedeagus is
finely dentate in campestris (Figs. 24, 25) and more coarsely bidentate
in the mesogramma group, where, in addition, one tooth is decidedly
more anterior than the other (Figs. 30-33); in the clarkei group, the
anterior tooth appears to have moved upward and forward along the
left side of the aedeagus and to have grown bigger still (Figs. 26-29).

26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

36

Fics. 36, 37. Female genitalia of Atalopedes clarkei from Cartagena, COLOMBIA,
14 July 1969, J. Herrera (X-2267) (collection of C. D. MacNeill). 36, Sterigma and bursa
copulatrix in ventral view; 37, The same, plus part of the ductus seminalis, in right lateral
view.

All parts considered, the male genitalia set the clarkei group farthest
out (which is reasonable from an ecologic perspective, considering the
shift to seashore grass). Admittedly, some far-out facies mark the me-
sogramma group (compare Figs. 7-10, 17, 18, particularly the ventral
secondaries, with Figs. 1-6, 11-16, and the entire genus Hesperia); but
facies can be much more labile even than genitalia. When such data
conflict, favor the genitalia.

Size is another labile character of little value in working out low-
level skipper phylogeny. In Atalopedes size varies from medium in

VOLUME 48, NUMBER 1 OF

Fics. 38, 39. Female genitalia of Atalopedes bahiensis from Conceicao da Barra,
Espirito Santo, BRASIL, 25 March 1969, C. & C. T. Elias (X-2390) (collection of O. H.
H. Mielke). 38, Sterigma and bursa copulatrix in ventral view; 39, The same, plus part
of the ductus seminalis, in right lateral view.

campestris (Figs. 1, 2, 11, 12) to large in mesogramma (Figs. 7, 8, 17,
18) but small in carteri (Figs. 9, 10) and, independently, to very small
in clarkei (Figs. 8, 4, 18, 14) and bahiensis (Figs. 5, 6, 15, 16).
Female genitalia neither help nor hurt the case built from male
genitalia, except that campestris does seem to reflect a more generalized
morphology from which the disparate expressions of the mesogramma
and clarkei groups could readily come. In campestris (Figs. 34, 35) the
midventral prong projecting backward and downward from the back
of the lamella postvaginalis is of medium length, the whole sterigma is

28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

40

Fics. 40, 41. Female genitalia of Atalopedes mesogramma from Guantanamo Bay,
CUBA, 15 September 1948, “caught laying eggs on Poa lawn,” W. H. Wagner (X-2172).
40, Sterigma and bursa copulatrix in ventral view; 41, The same, plus part of the ductus
seminalis, in right lateral view.

VOLUME 43, NUMBER 1 29

42

Fics. 42, 43. Fresh and worn individuals of Atalopedes clarkei perched, larger than
life, on flower heads amid the seashore grass in Fig. 44 on 19 August 1987.

well sclerotized, and the sclerotized ductus bursae is short, wide, and
evenly tapered. In mesogramma (Figs. 40, 41) the midventral prong
is hypertrophied, sclerotization of the sterigma is reduced medially but
not laterally, and the sclerotized ductus bursae is fairly short, narrow,
and parallel-sided. Conversely, in the clarkei group (Figs. 36-39) the
prong is atrophied, sclerotization of the sterigma is much reduced lat-

Fic. 44. Seashore grass habitat of Atalopedes clarkei between Playa Moreno and Playa
El Angel, near Pampatar, Margarita Island, Nueva Esparta, VENEZUELA, 19 August
1987. Caribbean Sea shows at upper right through crescentic gap in barrier dune.

30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Hesperia campestris mesogramma__ carteri clarkei bahiensis

Atalopedes

Fic. 45. Phylogeny of the bigger and better genus Atalopedes.

erally but not medially, and the sclerotized ductus bursae is long, wide,
and tapered.

Within each group of two species, which species is primitive and
which derived? Geographic distribution indicates that mesogramma
must have given rise to carteri: mesogramma is complexly widespread
(Puerto Rico, Hispaniola, Cuba, Isle of Pines, and some of the Bahamas,
including New Providence) whereas carteri, so far as known, is limited
to one small, low island (New Providence); furthermore, mesogramma
is somewhat differentiated across its discontinuous range. I should note
tangentially that mesogramma may be even more widely distributed:
Evans (1955) lists one female in the British Museum (Natural History)
from Costa Rica, and I find in the National Museum of Natural History,
Smithsonian Institution, one male labelled ““Mex’’ and three males la-
belled “Yucat.’’—but all of these mainland records need verification.
Except for its small size and reduced pattern elements, daughter species
carteri (Figs. 9, 10, 23) is very like mother mesogramma (Figs. 7, 8,
17, 18, 22). I have seen only one example of carteri, a male whose
genitalia (Figs. 32, 33) are essentially those of mesogramma (Figs. 30,
31; Comstock 1944:606, pl. 1, fig. 4)—the slight differences between
figures may reflect nothing more than individual variation.

Ancestral-descendant relations in the clarkei group (whose species
differ more from each other) are not obvious. To judge from genitalic
form, clarkei (Figs. 26, 27, 36, 37) probably preceded bahiensis (Figs.
28, 29, 38, 39). In the male the narrow tegumen-uncus and the enlarged
anterior tooth of the aedeagus that mark bahiensis (Fig. 28) appear
more derived. In the female the more atrophied midventral prong of
bahiensis (Figs. 38, 39) seems farther out.

Zoogeography

Geographically, too, it is the derived species of each two-species group
which is farther out—in this case from some generic center of distri-

VOLUME 438, NUMBER 1 31

bution at about the north end of South America: carteri is to the far
side of Antillean mesogramma, on a small island on the northern edge
of the latter’s range; bahiensis, on the central coast of Brazil, is far to
the southeast of coastal Colombian and Venezuelan clarkei.

The older the species of Atalopedes (Fig. 45), the wider its geographic
range: campestris, the oldest, has much the widest range (equator to
USA), even when the vast area temporarily taken by immigrants is
discounted; mesogramma, the second oldest, is second most widespread
(Greater Antilles plus); clarkei (Cartagena to Margarita) is third; ba-
hiensis (coastal central Brazil), fourth; and young carteri (New Prov-
idence Island), a distant fifth.

ACKNOWLEDGMENTS

Thanks to J. F. G. Clarke who provoked this study by collecting a long series of a
Venezuelan skipper that kept reminding me of North American kinds while continually
eluding placement; to C. D. MacNeill, O. H. H. Mielke, and J. B. Sullivan for lending
or giving crucial South American material; to Jeffrey Glassberg and Jane Scott for donating
not only a pair of specimens but also the color photographs of live adults and their habitat
which became Figs. 42-44; to Lee-Ann Hayek for calculating statistics; to Adrienne
Venables for meticulously KOH-dissecting numerous genitalia; to Young Sohn for taste-
fully rendering genitalia, as well as mounting drawings and photographs; to Victor Krantz
for photographing dead adults and their stigmas and making black-and-white prints from
color slides; and, again, to Don MacNeill for just plain seeing.

MacNeill and A. M. Shapiro thoughtfully reviewed the manuscript.

LITERATURE CITED

BurRNS, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif.
Publ. Entomol. 37:1-217.

1987. The big shift: nabokovi from Atalopedes to Hesperia (Hesperiidae). J.
Lepid. Soc. 41:173-186.

CLARK, A. H. & L. F. CLark. 1951. The butterflies of Virginia. Smithsonian Misc. Coll.
116(No. 7):vii + 239 pp.

Comstock, W. P. 1944. Insects of Porto Rico and the Virgin Islands—Rhopalocera or
butterflies. New York Acad. Sci., Scientific Survey of Porto Rico and the Virgin Islands
12:421-622, 12 pls.

Evans, W. H. 1955. A catalogue of the American Hesperiidae indicating the classifi-
cation and nomenclature adopted in the British Museum (Natural History). Part IV.
Hesperiinae and Megathyminae. British Museum, London. 499 pp., pls. 54-88.

GILLHAM, N. W. 1954. The type of Hesperia horus Edwards (Lepidoptera: Hesperidae).
Psyche 61:162.

LinpDsEy, A. W. 1921. The Hesperioidea of America north of Mexico. Univ. Iowa Stud.
Nat. Hist. 9(No. 4):1-114.

1942. A preliminary revision of Hesperia. Denison Univ. Bull., J. Sci. Lab. 37:
1-50, pls. 1-6.

LinpsEy, A. W., E. L. BELL & R. C. WILLIAMS, JR. 1931. The Hesperioidea of North
America. Denison Univ. Bull., J. Sci. Lab. 26:1-142.

MACNEILL, C. D. 1964. The skippers of the genus Hesperia in western North America
with special reference to California (Lepidoptera: Hesperiidae). Univ. Calif. Publ.
Entomol. 35:1—280.

1975. Family Hesperiidae, pp. 423-578. In Howe, W. H. (ed.), The butterflies

of North America. Doubleday & Co., Inc., Garden City, New York.

32 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Mayr, E. 1963. Animal species and evolution. Belknap Press of Harvard Univ. Press,
Cambridge, Massachusetts. 797 pp.

OpLER, P. A. & G. O. KRIZEK. 1984. Butterflies east of the Great Plains. Johns Hopkins
Univ. Press, Baltimore, Maryland. 294 pp., 54 pls.

PyLe, R. M. 1981. The Audubon Society field guide to North American butterflies.
Alfred A. Knopf, New York, New York. 916 pp.

RiLEy, N. D. 1975. A field guide to the butterflies of the West Indies. Collins, London.
224 pp., 24 pls.

ScHaus, W. 1902. Descriptions of new American butterflies. Proc. U.S. Nat. Mus. 24:
383-460.

SCUDDER, S. H. 1874. The species of the lepidopterous genus Pamphila. Mem. Boston
Soc. Nat. Hist. 2(Part 3, No. 4):341-358, pls. 10-11.

1889. The butterflies of the eastern United States and Canada with special
reference to New England. Publ. by the author, Cambridge, Massachusetts. Vol. 3,
pp. vii + 1775-1958, pls. 1-89, 3 maps.

SHAPIRO, A. M. 1966. Butterflies of the Delaware Valley. Special publ. Am. Entomol.
Soc. 79 pp.

1974. Butterflies and skippers of New York State. Search, Agr. (Cornell Univ.)
4(No. 3):60 pp.

SKINNER, H. & R. C. WILLIAMS, JR. 1924. On the male genitalia of the Hesperiidae of
North America, Paper VI. Trans. Am. Entomol. Soc. 50:177-208.

STANFORD, R. E. 1981. Superfamily Hesperioidea Latreille, 1802 (skippers), pp. 67—
108, 117-144. In Ferris, C. D. & F. M. Brown (eds.), Butterflies of the Rocky
Mountain states. Univ. Oklahoma Press, Norman, Oklahoma.

Received for publication 18 July 1988; accepted 27 September 1988.

Journal of the Lepidopterists’ Society
43(1), 1989, 33-49

TERRITORIAL BEHAVIOR AND DOMINANCE IN SOME
HELICONUNE BUTTERFLIES (NYMPHALIDAE)

WOODRUFF W. BENSON, CELIO F. B. HADDAD!
AND MARCIO ZIKAN

Departamento de Zoologia, IB, Universidade Estadual de Campinas,
C. P. 6109, 13081 Campinas, Sao Paulo, Brazil

ABSTRACT. By marking and systematically observing activities of focal individuals
of Heliconius sara, H. leucadia, and Eueides tales at six trailside sites at Serra dos Carajas,
Para, Brazil, we found that resident male butterflies returned daily during 1-3-h periods
to patrol and defend fixed 10-15-m-long sunny corridors against conspecific males. De-
fenders expelled intruders about once every 5-20 min, and unoccupied territories were
taken over by vagrant males in about the same time interval. Marked primary residents
of the two Heliconius species won all 149 combats observed with encroachers, and could
evict newcomers settling on territories left temporarily vacant. Resident E. tales were
more than 95% victorious. Besides fleeing vigorously from residents, trespassing H. sara
and H. leucadia frequently departed slowly from territories when accompanied by the
resident from below and behind. Resident H. sara flew erratic blocking patterns under-
neath slowly departing invaders, although a trespasser sometimes avoided immediate
expulsion by diving to soil level and flying in circles too close to the ground for the
accompanying resident to get under it. These ground-circling flights of H. sara appear
to be contests to decide territory ownership whereas the peculiar slow exits of desisting
H. sara and leucadia apparently function as appeasement behavior that bridles territorial
aggression. Eueides tales sometimes followed one another through several steep glides
(interpreted as ritualized chases) during territorial encounters. Territories seem to be
rendezvous sites attractive to receptive females, although E. aliphera may defend emer-
gence sites.

Additional key words: appeasement behavior, Eueides tales, Heliconius leucadia,
H. sara, mate location.

Territorial behavior gains its advantage by permitting the prefer-
ential use of resources in restricted areas (Brown & Orians 1970). Ter-
ritorial defense has been reported repeatedly in temperate-zone but-
terflies (Powell 1968, Baker 1972, Wellington 1974, Douwes 1975, Davies
1978, Bitzer & Shaw 1980, 1983, Lederhouse 1982, Dennis 1982, Alcock
1988, 1985, Knapton 1985, Wickman 1985a, Dennis & Williams 1987),
where, so far as known, males defend probable mate encounter sites
against other males (Baker 1983).

Defended encounter sites are frequently defined by landmarks, such
as hilltops (Shields 1968, Alcock & O'Neill 1986) and other landscape
features that reliably bring the sexes together (Parker 1978). Thus,
butterfly territories may occur along flyways (Baker 1972, Bitzer &
Shaw 1983) or occupy sheltered sites offering favorable conditions until
matings occur (Knapton 1985, Wickman 1986). Oviposition sites fre-
quented by gravid females (Baker 1972) and female emergence sites

' Present address: Departamento de Zoologia, IB, Universidade Estadual Paulista, 13500 Rio Claro, Sao Paulo, Brazil

34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

(Dennis 1982) may also be defended. Territorial males often return to
defend the same place over a period of days or weeks (L. E. Gilbert
in Maynard-Smith & Parker 1976, Lederhouse 1982, Alcock 1983,
Knapton 1985, Wickman 1985a).

Territorial interactions in butterflies may be characterized by their
greater duration (Wickman & Wiklund 1983, Wickman 1985a) and by
the peculiar combat behavior of residents (Fitzpatrick & Wellington
1983, Wickman & Wiklund 1983). Territorially related dominance
hierarchies and appeasement behavior, although present in other non-
social insects (Ewing 1972, Raw 1976), are apparently unreported for
Lepidoptera.

Recently Baker (1983) suggested that tropical Heliconius butterflies
are territorial. Indeed, Seitz (1913) reported seeing male heliconiines

. showing some characteristic defect, daily during four weeks flying
at about the same place... up and down in that characteristic fashion

. called ‘promenading,’” and added, “‘this habit of flying for hours
or half days at a time up and down for a certain distance, turning
sharply around at a certain point and returning the same way .. . is
nowhere quite so distinct as in the genera Eueides and Helene:
On the other hand, Crane (1957) found no evidence for territorial
behavior or social hierarchies during insectary studies of six species of
Trinidad heliconiines. Murawski (1987), however, observed territory-
like stationary defense of flowers by Heliconius when floral resources
were scarce.

The “large scale promenading’”’ reported for several Heliconius (Brown
& Mielke 1972, Brown 1972, Cook et al. 1976, Mallet & Jackson 1980)
refers to the repeated use of flyways within daily activity ranges, and
does not correspond to the behavior reported by Seitz (1913).

We report here observations on male Heliconius sara thamar (Htib-
ner), H. leucadia pseudorhea Staudinger, Eueides tales pythagoras
Kirby and E. aliphera (Godart) patrolling and expelling conspecifics
from territories. Results show that defense is often achieved through
specialized ejection behavior, and that invaders rapidly occupy vacant
territories. Notes are given for other heliconiines indicating that similar
behavior may occur widely in these insects.

STUDY SITES AND METHODS

Systematic observations were undertaken during the austral dry sea-
son (July) of 1986 and 1987 in the Serra dos Carajas near Serra Norte,
Para, Brazil (6°03'S, 50°07’ W), at sites occupied by promenading (sensu
Seitz 1913) heliconiine butterflies. Heliconius sara (Fabr.), H. leucadia
Bates and Eueides tales (Cramer) were observed along trails near Cal-
deirao (5°53'S, 50°27’W), an abandoned mineralogical camp at 210 m

VOLUME 43, NUMBER 1 35

elev. by State Road PA-275 where it crosses the Rio Itacaitnas. A second
site, with only H. sara, was at 650 m on an abandoned spur of PA-275,
2 km N of the iron ore outcrop called “N-1”’ (5°59’S, 50°16’W). In 1987,
males of H. sara held territories at three points 100-200 m apart at
Caldeirao, here referred to as Areas 1, 2, and 3, and 3 others—4, 5,
and 6—arranged linearly 75-100 m apart at the N-1 site. Only Caldeirao
was worked in 1986 where H. sara territories were observed at Areas
1 and 2. Heliconius leucadia was observed only in 1987 defending
territories at Areas 1 and 3. Eueides tales defended in both years at
Area 1. Observations on territorial H. sara totaled 1194 min, on H.
leucadia 664 min, and on E. tales 627 min. Notes on territorial Eueides
aliphera were mostly taken in Costa Rica.

The climate at Carajas is moist tropical with mean annual rainfall
2100 mm (CV = 23%) and mean temperature between 24° and 26°C,
depending on altitude. Temperatures and relative humidities during
observation periods at Caldeirao were typically near 24°C and 90-95%
at 0930 h and 30°C and 75% at 1130 h. Most days were cloudless or
with scattered clouds only appearing at the ends of observation periods.

Net-captured butterflies were marked using colored porous-point
pens or by cutting notches from wing margins. Individually recogniz-
able animals were merely netted to establish positive species identifi-
cation. Sex, wing length, and wing damage and wear indicative of age
were noted. Behavioral observations were made from trailsides with
the aid of a stopwatch and binoculars. In territorial interactions, sex of
unmarked nonresident butterflies was inferred from their behavior us-
ing Crane (1957) and observations on marked intruders of known sex.
Male Eueides tales were distinguished from females by their narrower
wings. During 1987 observation periods, specific activity of focal in-
dividuals (Altmann 1974) was monitored continuously or noted every
15 sec for time budgets. For measured variables we give arithmetic
means and use standard deviations (SD) to describe data variation.

RESULTS

Territoriality in Heliconius sara, H. leucadia, and Eueides tales is
broadly similar. Defending males divided their time between perching
and promenading over 10-15-m-long territories situated in sunlit vege-
tation corridors. Territory cores were normally delimited by conspic-
uous features such as overhanging limbs or jutting bushes, and were
adjoined by less frequently visited peripheral areas of 5-30 m at one
or both ends. Males did not feed when engaged in territorial activity
nor were host-plants or other resources consistently present on terri-
tories. Heliconius leucadia and H. sara flew irregular paths 1-2 m wide,
usually staying within 1-2 m of neighboring vegetation whereas E. tales

36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

tended to occupy the center of the approximately 5-m-wide trail clear-
ing. A complete circuit of a core area took approximately 10-15 sec
when butterflies did not tarry in localized circling. In all three species,
conspecific males were challenged when approaching within 2-3 m of
a resident. Pursuing residents normally followed intruders well beyond
the patrolled area where they broke off the chases.

Territorial behavior. Discounting courtships and matings, approxi-
mately 100 conspecific interactions were observed in each of the three
main heliconiine species studied (Table 1). These were almost always
lengthy, and continued until one butterfly was either driven from the
territory or was able to evade the other.

In Heliconius sara, a resident male often followed, rather than chased,
a male encroacher, and attempted to get below it. Eighteen of 32
interactions observed in 1986 began with downward flight that tended
to bring the two butterflies to ground level (the other 14 were rapid,
straightforward chases). In five of these chases, the insects descended
almost to the ground, and in two, terminating in relatively open un-
dergrowth, the butterflies flew 5 to 15 cm above the ground, and circled
and weaved back and forth over contiguous areas 30-40 cm in diam.

We have observed such ground circling behavior on many additional
occasions. Circling may last from a few seconds to a minute or more,
after which normally the interloper begins flying upwards with the
resident joining to accompany it from below and behind. Low flight
apparently prevents the dominant butterfly from getting under and
expelling the subordinate. On occasion, and despite apparent attempts
to block it, an intruder may slip past a resident and initiate another
bout of circling. One series of interactions of this type observed in 1987,
involving several individuals, went on for 15 min.

Once the resident succeeded in getting under the trespasser, a char-
acteristic ascending ejection generally followed. Initially, when flying
through vegetation, if the intruder became separated from the resident
by more than 30-40 cm, the latter usually dashed in the interloper’s
direction until again immediately below and behind. Once free of
confining foliage, an intruder tended to fly slowly upwards, almost
hovering, with the resident darting erratically back and forth almost
directly underneath it. Ejections gave the impression of the invader
being driven upwards by the resident, and in one instance the lower
butterfly was seen to dart several times at a subordinate that was as-
cending at an abnormally slow pace. Butterfly pairs habitually rose 15
m or more to pass over vegetation bounding the territory, and could
wander as far as 40 m laterally before the resident disengaged. Ex-
pulsions sometimes ended with the intruder bolting away with the
resident in pursuit. Territory holders returned from such excursions

37

VOLUME 43, NUMBER 1

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38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

\

BPA

Fic. 1. Typical territorial interaction in Heliconius sara at Serra dos Carajas, Para,
Brazil. Resident H. sara (solid line) promenades (A) or perches on territory while intruder
(broken line, segment length approximately proportional to intruder velocity) patrols
along forest margin (A’). One or both butterflies attack, with resident attempting to get
below invader (B) and both butterflies sometimes diving to ground where they circle
adjacent to one another (C). Intruder starts flying slowly upwards with resident darting
back and forth below and behind it (D) until reaching tree-top level where invader may
dash away with resident in pursuit (E). Resident returns to territory where it flies briskly
over core and peripheral areas as it resumes promenading (F).

generally after a few 10s of seconds, flying briskly over the core and
peripheral areas before resuming usual patrolling. Fig. 1 schematizes
an ejection sequence in H. sara.

Evasive diving in intruding H. sara typically led to ground circling,
which we interpret as an endurance contest to determine territorial
possession. Although in the sequences we observed, territorial residents
seemed always to expel invaders, extended contests, sometimes involv-
ing several butterflies, were witnessed just before and after changes in
ownership. Twice in 1986 and once in 1987, an intruding H. sara was
seen to attack a promenading resident which dove into the undergrowth.
In each of these cases, the trespasser shortly left the area, and the
submissive individual resumed patrolling, suggesting that downward
dives may also aid less capable males in retaining territories, at least
temporarily.

In Heliconius leucadia, agonistic territorial behavior seems less com-
plex than in H. sara. After the initial rush at an intruder, the resident
may expel it by simply following it off the territory from approximately
2 m below and behind. These tandem flights were often leisurely, and

VOLUME 48, NUMBER | 39

TABLE2. Behavior of Eueides tales during territorial interactions at Serra dos Carajas,
Para, Brazil during 1986 (n = 44) and 1987 (n = 31).

Behavior of intruder

Interaction initiated Interaction initiated
by resident by intruder
Slow Rapid Slow Rapid
Behavior of resident departure departure departure departure
Chase or following 9 49 I 9
No chase or following 0 i ] 5

normally passed over bounding vegetation, sometimes attaining a height
of 15-20 m before the resident suddenly and spontaneously disengaged.
At Area 3 one resident often returned to the vicinity of its territory by
means of spectacular 40-m-long glides.

Although more or less rapid, apparently aggressive expulsions were
common in each of the three main species studied here, we did not
observe spiraling pursuits of the type reported for other Lepidoptera
(Baker 1983, Fitzpatrick & Wellington 1983). In H. leucadia, vigorous
circular chases were accompanied by sounds of wing contact indicative
of physical combat.

The leisurely exit of trespassing H. sara and leucadia, when being
conducted from a territory by its owner, is almost certainly a form of
appeasement behavior, behavior functioning to “inhibit or reduce
aggression . . . where escape is impossible or disadvantageous” (Mc-
Farland 1981:17). Territorial defense in butterflies typically consists of
direct combat, with the dominant positioning itself above its opponent
and striking at it with its wings (Fitzpatrick & Wellington 1983, Wick-
man & Wiklund 1983). In the Heliconius studied by us, rapidly flying
invaders may be vigorously pursued and perhaps hit by territory hold-
ers. In contrast, the slow ejections of heliconiines involve neither ag-
gressive pursuit nor physical combat, and slow intruder movements,
perhaps in concert with other behavior, seem to signal submission and
stimulate “‘escort’”’ behavior. Serious challenges are apparently resolved
by endurance contests (within a context of appeasement) in H. sara
and by brief but violent combats in H. leucadia.

Trespassing Eueides tales usually fled from a territory with the res-
ident in pursuit, although often an intruder left slowly with the resident
merely following (Table 2). Less commonly, intruders initiated inter-
actions by flying at residents, but these were usually attacked in return
or withdrew rapidly without being chased or followed. When a tres-
passer being followed from a territory got more than about 2 m ahead
of the resident, or entered into foliage, the owner normally dashed after
it, which at times provoked a high-speed chase.

40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Intruding male E. tales sometimes evaded pursuit by alighting on
leaves. In the seven instances of landing by escorted butterflies, the
resident flew agitatedly around the point of last contact. However, only
once did the resident find the perched intruder, and in this case dislodge
it, apparently by butting and landing on it. In two instances, landing
intruders succeeded in fleeing unmolested, and in four, they returned
to the territory where the resident found and expelled each once again.

On five occasions an attacked butterfly, rather than flee or retaliate
against its aggressor, assumed a descending gliding flight with its wings
partially folded. This posture, apparently starting with the attacked
individual, seemed to be copied by the attacker which trailed about
20-40 cm above and behind in descending flight. As the butterflies
drifted downwards, the lead individual sometimes switched places by
darting swiftly behind the trailing one, or one chased the other back
up to patrolling altitude, initiating another descent, or chased it off the
territory in an expulsion. Although sample sizes are small, data from
1987 suggest that glide chases may occur more frequently when in-
vaders challenge residents (3 times in 7 attacks) than the converse (2
times in 25 attacks). One of the two observed cases of a resident E.
tales losing its territory to an intruder followed an intruder-initiated
attack and 4-5 glide sequences. Glide chases in E. tales seem to be
ritualized territorial pursuits and, like ground circling in H. sara, may
constitute assessment behavior that helps resolve disputes in lieu of
potentially injurious combat.

Territorial defense. Territorial defense in the heliconines studied at
Carajas was concentrated in the late morning (Table 1) with Eueides
tales and exceptional Heliconius sara continuing as late as 1245 h. At
Area | where territorial males of all three species flew, H. leucadia
promenaded somewhat lower (about 3-4.5 m from the ground) than
H. sara (4-5 m) and E. tales (5-6 m). The percentage of time spent in
promenade flight ranged from less than 50% in sara to more than 90%
in tales (Table 1). Eueides tales seemed to glide more than the two
Heliconius species, perhaps assisted by its smaller size and the generally
stronger breezes higher up and around mid-day.

During defense, territory owners typically clashed with conspecifics
from 3 to 12 or more times/h, depending on the frequency of intrusions
(Table 1). Intruders were almost always successfully intercepted; we
have only one record of a probable intruder H. leucadia crossing a
territory apparently unseen by the perched resident.

Each species had territory holders that rarely lost contests. In H. sara,
one territory owner marked in 1986 and two marked in 1987 were
observed to win all of their 52 conflicts with intruders (Table 1). One
of these (1987, Area 4) additionally ejected another male that had set

VOLUME 48, NUMBER 1 4]

up a territory during a temporary absence. A resident briefly observed
at Area 5 also dispossessed a newcomer that took over its territory after
it was captured for marking. The two individually recognizable H.
leucadia won the 97 clashes with trespassers, and in addition, each twice
evicted newcomers that took up residence when the owner was away.
A marked E. tales won 33 of its 34 territorial clashes, and of the 23
conflicts involving unmarked butterflies in 1987, only 1 was for certain
lost by the resident butterfly. Reconquest of territories from subsidiary
residents was not observed in E. tales.

The two Heliconius species usually returned to defend the same
territory on successive days. The marked H. leucadia at Area 1 was
observed defending on all nine visits over 19 days, and the Area 3
resident on all six visits over the last eight days we were at Carajas.
The latter stayed in Area 3 at least 20 days after marking. On the last
day of observation, the Area 3 resident patrolled for only 7 min, and
the Area 1 butterfly did not appear. The H. sara at Area 3 defended
on all seven visits made over the 11 days following marking. An H.
sara at Area 4 apparently abandoned the territory after being marked,
but returned in the role of owner 10 and 11 days later. Similarly, a
butterfly at Area 6 took up residence 15 days after it was marked. The
marked E. tales that returned to defend the Area | territory in 1987
was among the defenders present during visits made on three of the
four days following marking. On day 4 it was evicted by an intruder
and did not reappear on day 5.

The limited nature of territories is indicated by the rapidity with
which newcomers reoccupied those left vacant. Three Heliconius sara
removed from territories at Areas 4, 5 and 6 at N-1 were replaced in
3, 11, and <80 min, respectively. Three others taken from territories
at Areas | and 2 were replaced by unmarked H. sara on the same or
following days. On 7 of the 15 occasions in which an identifiable H.
leucadia left or was removed from a territory (including final daily
departures), another individual took up patrolling within 10 min. On
the five occasions in 1987 that a Eueides tales territory became vacant
before noon, a new resident took up promenading within 7 min. Ter-
ritorial E. tales netted in 1986 were also rapidly replaced by newcomers.

The similarity in time intervals between expulsions and that required
to reoccupy a vacant territory suggests that most intruders are floating
males seeking territories. Intrusion rates in H. sara and H. leucadia
seemed to decline during the morning (Figs. 2 and 3), perhaps because
floaters took up searching for mates in other habitats.

Territorial heliconiines usually dashed after any large butterfly pass-
ing by. With non-conspecifics, chases normally terminated when the
owner came within 10-30 cm of the intruder. Butterflies pursued in

42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

this manner included a number of heliconiines, including the other
territorial species, and pierids. Heterospecific interactions between sim-
ilarly patterned H. sara, H. leucadia and H. wallacei Reakirt often
included mutual “on-and-off” chasing that, in the first pair of species,
sometimes occurred across a territorial boundary in Area 3. We interpret
this case of interspecific territoriality as a nonadaptive result of imper-
fect species recognition (Murray 1971). The H. sara and H. leucadia
at Area | simultaneously occupied broadly overlapping territories with-
out markedly interfering with one another. Both E. tales and H. sara
darted after falling leaves, and one of the latter pursued a large bee,
and another flew 8-10 m upwards eight times in succession in the
direction of lesser yellow-headed vultures (Cathartes burrovianus Cas-
sin) gliding low overhead. Apparently a combination of color, apparent
size, form, and movement stimulates inspection flights in H. sara with-
out the intervention of physical proximity or chemical stimuli.

In 1987, the marked territorial H. sara at Areas 3 and 4 each once
courted and was rejected by a female, and a marked male patrolling
Area 5 was seen to chase a probable female. Before systematic obser-
vations were begun, a recently emerged female H. sara was found
copulating in undergrowth immediately adjacent to Area 5. The marked
H. leucadia at Area 3 was also observed courting a female. In one
encounter, which may represent a courtship flight, the other marked
H. leucadia at Area 1, rather than follow the intruder from behind,
flew 30 cm almost directly below it with rapid wing beats and darting
flight reminiscent of the ascending expulsion flight of H. sara. The pair
rose approximately 20 m directly overhead before drifting out of sight
behind trees. The resident returned 14 min later, ousted an unmarked
individual that had taken up promenading in the meantime, and re-
sumed patrolling. In 1986 a courtship involving a male E. tales of
unknown status was observed at Area 1.

Territoriality in other heliconiines. Territorial behavior in Eueides
aliphera was noted in 1967 in a weedy coffee field 5 km S of San Vito,
Puntarenas Province, Costa Rica. One of the several E. aliphera present
on 27 April, flying about 1 m above the vegetation, seemed especially
pugnacious, and dashed after Heliconius charitonia (L.), Hypanartia
lethe (Fabr.), and on three occasions after other E. aliphera, in 18 min
of observation. A male E. aliphera color-marked at the site 5 days later
was re-encountered defending a territory at the same place on the six
visits made over the next 19 days. This E. aliphera won all combats
with natural intruders (Table 1) in addition to one with an apparently
territorial male that was experimentally herded onto the marked but-
terfly’s territory. The resident attacked and chased this insect into the
underbrush. In encounters with other species, the territorial E. aliphera

VOLUME 438, NUMBER 1 43

16 EXPULSION RATE e
(EVENTS /HOUR)
14
iG Caldertrae, Area 3
10

: pene
6 a eu

6 aaa.
e IN=IL- Meee. 5

0920 0940 1000 1020 1040 1100 1120 1140 1200
lal CO)" Whe

Fic. 2. Expulsion rates for territorial Heliconius sara at Serra dos Carajas, Para, Brazil.
Results grouped for intervals of 20 min. Numbers above abscissa show minutes of obser-
vation in time interval.

immediately turned away; however, conspecifics were vigorously pur-
sued with pairs usually rising 10-15 m into the air and speeding off
the territory. After flying well outside the patrolled area—pairs some-
times flew out of sight—the resident disengaged and returned to the
territory. All chases were energetic, and “escorting” analogous to that
recorded in other heliconiines was not observed.

The marked butterfly arrived on the territory as early as 0830 and
0910 h, and departures were as late as 1433 h during sunny weather.
On one afternoon, the E. aliphera flew off the territory and over the
canopy of neighboring forest about 50 m away three times during
cloudy periods, and returned to patrol during intervening sunny spells.

The territory of the marked E. aliphera contained larval food-plant
(Passiflora oerstedii Mast. in Mart.) with immature stages. An E. ali-
phera observed in August 1985 near Serra Norte, Brazil, promenaded
above a roadside tangle of Passiflora vines on which E. aliphera larvae
were also feeding. The territories of other heliconiines studied at Cal-

44 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

2. EXPULSION RATE
(EVENTS /HOUR)

Caldeimao, Aeeaws

“\,
12
Callderrac, nea:
10
e
8
° e
6 Ne
e e
4 “se ia

0920 0940 1000 1020 1040 1100 ILA) 1140
st © WR

Fic. 3. Expulsion rates for territorial Heliconius leucadia at Caldeirao, Serra dos
Carajas, Para, Brazil. Results grouped for intervals of 10 or 20 min. Numbers above
abscissa show minutes of observation in time interval.

deirdo lacked larval food-plants, although at N-1 an H. sara host, Pas-
siflora (Astrophea) sp., was abundant next to Area 5. This situation
seems fortuitous.

Territoriality probably occurs in other heliconiines. In January 1968
near Huixtla, Chiapas, Mexico, two well separated Eueides isabella
(Cramer) were briefly observed while tracing 10-15-m-long paths over

VOLUME 43, NUMBER 1 45

low vegetation in a roadside ravine. Each shared its space with a single
similarly behaving E. aliphera, although the latter flew somewhat lower
(1 m vs. 2 m) above the vegetation. On approaching within 2-3 m of
each other, one butterfly would occasionally dash at the other without
chasing it. Heliconius ricini (L.) promenades in Trinidad (W. W. Benson
field notes), and at Caldeirao appears to defend spaces over the forest
canopy. Heliconius antiochus (L.) at Caldeirao is both aggressive and
site-tenacious, and may be territorial. However, Eueides vibilia (Godart)
promenading near N-1 did not fight or expel closely approaching con-
specifics. Seitz (1913) reported promenading to be especially well de-
veloped in E. aliphera, E. isabella and Philaethria dido (L.). We have
detected no sign of promenading or area defense in Heliconius erato
(L.), H. melpomene (L.), H. wallacei, Eueides lybia (Fabr.), or Dryas
iulia (Fabr.) at any locality.

DISCUSSION

The heliconiines studied here are clearly territorial. Conspecific males
rarely remained together in a promenade area longer than the few
seconds necessary for the resident to find and expel the encroacher. In
the absence of an owner, territories were rapidly taken over, and the
time for this to occur was comparable to the average interval between
intrusions, suggesting that most intruders are territory-seekers. In three
species, both residents and invaders seem to possess a repertory of
species-specific behaviors for use during territorial confrontations.

Territoriality in butterflies seems based on male defense of encounter
sites where chances of mating are high (Baker 1983, Wickman 1985b,
Courtney & Anderson 1986). The territorial heliconiines studied by us
probably also defend rendezvous points. Although we observed few
courtships and no matings by known territorial males, female helico-
niines seem often to mate only once (Crane 1957, Gilbert 1976), and
their rarity is expected in long-lived insects that mate infrequently
(Alcock 1983). The places defended by Heliconius sara, H. leucadia
and Eueides tales are humid, sunny, and seem protected from wind,
and possibly attract receptive females. Eueides aliphera defends more
exposed sites, and its territoriality may be in part based on the despotic
control of host-plant patches where females are likely to emerge.

Territorial combat in heliconiines contrasts greatly with courtship.
In Heliconius sara and related species, courting males hover above and
in front of females (Crane 1957, pers. obs.). Territorial males fly below
intruders or harass or “escort”” them from behind. Owner behavior
seems adaptive since intruders are denied searching foliage for receptive
mates, taking the profit out of trespassing. However, trespassing and
skirmishing may still benefit interlopers in assessing vacancies and dis-

46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

covering weak residents (Lederhouse 1982, Wickman & Wiklund 1983,
Grafen 1987).

Promenading heliconiines fly slowly and seemingly with little effort
through their territories. Butterflies are conspicuously exposed under
such circumstances, and aposematic unpalatability in H. sara (Brower
et al. 1963), and probably the other species studied, must compensate
for much of the added risk of predator attack. Butterflies that are
profitable prey may be selected to reduce conspicuousness by perching
more, or to reduce catchability by patrolling territories using energet-
ically costly acrobatic flight, or both.

Recent works on territoriality (Bitzer & Shaw 1983, Wickman 1985a,
Dennis & Williams 1987, Shreeve 1987) equate Scott’s (1974) terms
“perching” and “patrolling” with territorial and nomadic mating strat-
egies, respectively. Although this terminology is misleading when ap-
plied to a species such as Eueides tales, which spends more than 90%
of its time in patrolling flight and combats, it is probably too intrenched
to change. Promenading, sensu Seitz (1913), may be a useful alternative
to designate site-faithful patrolling.

Given the prowess of resident Heliconius sara and H. leucadia in
rivalries, unrestrained fights may be risky. The fact that intruders often
flee, flying at what seems to be maximum speed, suggests that some
chance of injury exists. The submissive stance of many intercepted
intruders is noteworthy, and, combined with a less aggressive domi-
nance of the resident, results in slow but safe and effective expulsions.
To our knowledge these are the first reports of stereotyped appeasement
behavior in butterflies, and one of the few among nonsocial insects
(Fitzpatrick & Wellington 1983). They are also apparently the only
known cases of relatively pacific dominance relations inserted in ter-
ritorial behavior sequences.

The ground-circling behavior of H. sara seemed to help submissive
individuals avoid ejection, and may be, as appears with the glide chases
of E. tales, a ritualized contest used to decide territorial ownership.
Whatever their precise origins and functions, these behaviors are clearly
tied to presumably adaptive defense of territory.

Territoriality in Heliconius sara and H. leucadia is not simply ex-
plained by current hypotheses. In general, female attraction to en-
counter sites (and competition for them by males) is thought to stem
from male rarity and the increased speed of mating permitted by their
use (Shields 1968, Lederhouse 1982, Alcock & O'Neill 1986). However,
nonterritorial male H. sara and H. leucadia were common in our study,
and sara is frequently abundant throughout its range, suggesting that
isolated females would be quickly found and mated and that rendezvous
sites may be superfluous. On the other hand, the known territorial

VOLUME 43, NUMBER 1 47

Heliconius lay eggs in batches that give rise to synchronously developing
groups of larvae. It seems likely that emergence of large broods of
females in these species may commonly overtax mating capacity of
local males and result in pulses of virgin females that might profitably
seek mates at encounter sites. The importance of intrasexual competition
among females is manifest in the use of what is apparently an aggre-
gating sex pheromone by female pupae of H. sara (W. W. Benson field
notes), similar to that of the related H. charitonia (Edwards 1881,
Gilbert 1975) and H. hewitsoni Staudinger (J. T. Longino in DeVries
1987). Pupal pheromone production may aggravate mate shortages by
concentrating males at pupation sites well ahead of mating. We believe
that common heliconiines that lay solitary eggs would tend to have a
more uniform production of, and a more assured rapid mating of,
receptive females, and thus tend not to be territorial. Territoriality in
Eueides tales and E. aliphera may be explained by current theory.

ACKNOWLEDGMENTS

We acknowledge logistic support by the Organization for Tropical Studies in Costa
Rica. Observations in Brazil were made during field courses of the Universidade Estadual
de Campinas, for which transport was furnished by the Brazilian Air Force (FAB) and
the Coordenacao de Aperfeicoamento de Nivel Superior (CAPES) and logistic support
by the Companhia Vale do Rio Doce and Rio Doce Geologia e Mineracaéo SA under
Contract #220/82 with the Conselho Nacional de Desenvolvimento Cientifico e Tec-
nologico (CNPq). Haddad and Zikan received fellowships from CAPES and CNPq,
respectively. We thank Joao Semir and Neuza Taroda for aid in plant identifications, and
R. R. Baker, Ivan Sazima, Eleonore Setz, Thomas Lewinsohn, Felipe Costa, and an
anonymous reviewer for suggestions concerning the manuscript.

LITERATURE CITED

ALCOCK, J. 1983. Territoriality by hilltopping males of the great purple hairstreak,
Atlides halesus (Lepidoptera: Lycaenidae): Convergent evolution with a pompilid
wasp. Behav. Ecol. Sociobiol. 13:57-62.

1985. Hilltopping in the nymphalid butterfly Chlosyne californica (Lepidop-
tera). Am. Midl. Nat. 113:69-75.

ALCOCK, J. & K. M. ONEILL. 1986. Density-dependent mating tactics in the grey
hairstreak, Strymon melinus (Lepidoptera: Lycaenidae). J. Zool. 209:105-113.
ALTMANN, J. 1974. Observational study of behavior: Sampling methods. Behavior 49:

227-267.

BAKER, R. R. 1972. Territorial behaviour of the nymphalid butterflies, Aglais urticae
(L.) and Inachis io (L.). J. Anim. Ecol. 41:453-469.

1983. Insect territoriality. Ann. Rev. Entomol. 28:65-89.

BITzZER, R. J. & K. C. SHAW. 1980. Territorial behavior of the red admiral, Vanessa
atalanta (L.) (Lepidoptera: Nymphalidae). J. Res. Lepid. 18:36-—49.

1983. Territorial behavior in Nymphalis antiopa and Polygonia comma (Nym-
phalidae). J. Lepid. Soc. 37:1-18.

BROWER, L. P., J. v. Z. BROWER & C. T. COLLINS. 1963. Experimental studies of
mimicry. 7. Relative palatability and Miillerian mimicry among neotropical butterflies
of the subfamily Heliconiinae. Zoologica (N.Y.) 48:65-84.

Brown, J. L. & G. H. ORIANS. 1970. Spacing patterns in mobile animals. Ann. Rev.
Ecol. Syst. 1:289-262.

48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Brown, K. S., JR. 1972. The heliconians of Brazil (Lepidoptera: Nymphalidae). Part
Ill. Ecology and biology of Heliconius nattereri, a key primitive species near ex-
tinction, and comments on the evolutionary development of Heliconius and Eueides.
Zoologica (N.Y.) 57:41-69.

Brown, K. S., JR. & O. H. H. MIELKE. 1972. The heliconians of Brazil (Lepidoptera:
Nymphalidae). Part II. Introduction and general comments, with supplementary
revision of the tribe. Zoologica (N.Y.) 57:1-40.

Cook, L. M., E. W. THOMASON & A. M. YOUNG. 1976. Population structure, dynamics
and dispersal of the tropical butterfly Heliconius charitonius. J. Anim. Ecol. 45:851-
863.

COURTNEY, S. P. & K. ANDERSON. 1986. Behaviour around encounter sites. Behav. Ecol.
Sociobiol. 19:241-248.

CRANE, J. 1957. Imaginal behavior in butterflies of the family Heliconiidae. Changing
social patterns and irrelevant actions. Zoologica (N.Y.) 42:135-145.

Davies, N. B. 1978. Territorial defense in the speckled wood butterfly (Pararge aegeria):
The resident always wins. Anim. Behav. 26:138-147.

DENNIS, R. L. H. 1982. Mate location strategy in the wall brown butterfly, Lasiommata
megera L. (Lep: Satyridae). Wait or seek? Entomol. Rec. J. Var. 94:209-214.

DENNIS, R. L. H. & W. R. WILLIAMS. 1987. Mate location behavior of the large skipper
butterfly Ochlodes venata: Flexible strategies and spatial components. J. Lepid. Soc.
4]:45-64.

DEVRIES, P. J. 1987. The butterflies of Costa Rica and their natural history. Princeton
Univ. Press, Princeton, New Jersey. 327 pp.

DouwEs, P. 1975. Territorial behaviour in Heodes virgaureae (Lep., Lycaenidae) with
particular reference to visual stimuli. Norw. J. Entomol. 22:143-154.

EDWARDS, W. H. 1881. On certain habits of Heliconia charitonia Linn., a species of
butterfly found in Florida. Papilio 1:209-215.

EwInc, L. S. 1972. Hierarchy and its relation to territory in the cockroach Nauphoeta
cinerea. Behaviour 42:152-174.

FITZPATRICK, S. M. & W. G. WELLINGTON. 1983. Insect territoriality. Canad. J. Zool.
61:471-486.

GILBERT, L. E. 1975. Ecological consequences of a coevolved mutualism between
butterflies and plants, pp. 210-240. In Gilbert, L. E. & P. H. Raven (eds.), Coevolution
of animals and plants. Univ. Texas Press, Austin. 246 pp.

1976. Postmating female odor in Heliconius butterflies: A male contributed
antiaphrodisiac? Science 193:419—420.

GRAFEN, A. 1987. The logic of divisively asymmetric contests: Respect for ownership
and the desperado effect. Anim. Behav. 35:462-467.

KNAPTON, R. W. 1985. Lek structure and territoriality in the chryxus arctic butterfly
(Satyridae). Behav. Ecol. Sociobiol. 17:389-395.

LEDERHOUSE, R. C. 1982. Territorial defense and lek behavior in the black swallowtail
butterfly, Papilio polyxenes. Behav. Biol. Sociobiol. 10:109-118.

MALLET, J. L. B. & D. A. JACKSON. 1980. The ecology and social behaviour of the
neotropical butterfly Heliconius xanthocles in Colombia. Zool. J. Linn. Soc. 70:1-13.

MAYNARD-SMITH, J. & G. A. PARKER. 1976. The logic of asymmetric contests. Anim.
Behay. 24:159-175.

MCFARLAND, D. (ed.). 1981. The Oxford companion to animal behaviour. Oxford Univ.
Press, New York. 657 pp.

MURAWSKI, D. A. 1987. Floral resource variation, pollinator response, and potential
pollen flow in Psiguria warscewiczii. Ecology 68:1273-1282.

Murray, B.G., Jr. 1971. The ecological consequences of interspecific territorial behavior
in birds. Ecology 52:414—423.

PARKER, G. A. 1978. Evolution of competitive mate searching. Ann. Rev. Entomol. 23:
173-196.

POWELL, J. A. 1968. A study of area occupation and mating behavior in Incisalia iroides
(Lepidoptera: Lycaenidae). J. N.Y. Entomol. Soc. 76:47-57.

VOLUME 43, NUMBER 1 49

Raw, A. 1976. The behaviour of males of the solitary bee Osmia rufa (Megachilidae)
searching for females. Behaviour 56:279-285.

ScoTT, J. A. 1974. Mate-locating behavior of butterflies. Am. Mid]. Nat. 91:103-117.

SEITZ, A. 1913. Heliconidae, p. 377. In Seitz, A. (ed.), Macrolepidoptera of the world.
Vol. 5. Kernen, Stuttgart. 1139 pp.

SHIELDS, O. 1968. Hilltopping. J. Res. Lepid. 6:69-178.

SHREEVE, T. G. 1987. The mate location behaviour of the male speckled wood butterfly,
Pararge aegeria, and the effect of phenotypic differences in hind-wing spotting.
Anim. Behav. 35:682-690.

WELLINGTON, W. G. 1974. A special light to steer by. Nat. Hist. 83(10):46-53.

WICKMAN, P.-O. 1985a. The influence of temperature on the territorial and mate
locating behaviour of the small heath butterfly, Coenonympha pamphilus (L.) (Lep-
idoptera: Satyridae). Behav. Ecol. Sociobiol. 16:223-238.

1985b. Territorial defence and mating success in males of the small heath

butterfly, Coenonympha pamphilus L. (Lepidoptera: Satyridae). Anim. Behav. 33:

1162-1168.

1986. Courtship solicitation by females of the small heath butterfly, Coeno-
nympha pamphilus (L.) (Lepidoptera: Satyridae) and their behavior in relation to
male territories before and after copulation. Anim. Behav. 34:153-157.

WICKMAN, P.-O. & C. WIKLUND. 1988. Territorial defence and its seasonal decline in
the speckled wood butterfly (Pararge aegeria). Anim. Behav. 31:1206-1216.

Received for publication 13 January 1988; accepted 16 November 1988.

Journal of the Lepidopterists’ Society
43(1), 1989, 50-58

REPRODUCTIVE TRACT DEVELOPMENT IN
MONARCH BUTTERFLIES OVERWINTERING
IN CALIFORNIA AND MEXICO

WILLIAM S. HERMAN

Department of Genetics and Cell Biology, University of Minnesota,
St. Paul, Minnesota 55108

LINCOLN P. BROWER AND WILLIAM H. CALVERT

Department of Zoology, University of Florida,
Gainesville, Florida 32511

ABSTRACT. Reproductive organs of male and female monarch butterflies captured
in December, February, and March in overwintering colonies in both Mexico and Cali-
fornia were examined as soon as possible after capture. In addition, response of such
organs to incubation in summer-like conditions was determined for animals from both
locations in all three months. Results demonstrated numerous similarities between the
two populations, indicating comparable stages of reproductive tract development in the
two locations. However, a higher percentage of mating was recorded in Californian
females, and data obtained after incubation indicated that diapause might last longer in
both sexes of Mexican monarchs.

Additional key words: Nymphalidae, Danaus plexippus, diapause, overwintering.

North American monarch butterflies (Danaus plexippus L.) aggre-
gate in two major overwintering locations. One is California, where
several colonies form each winter (Lane 1984). The other is in the
mountains of the states of Michoacan and Mexico in Mexico, where
multiple overwintering colonies have now been located (Calvert &
Brower 1986). The Mexican colonies are principally aggregations of
monarchs that emerge in the eastern United States and Canada, while
the Californian colonies are the major overwintering sites for monarchs
originating west of the Rocky Mountains. Although it might be rea-
sonably assumed that overwintering monarchs from both locations were
in similar reproductive states, direct comparative evidence concerning
the reproductive status of such animals is not available. To obtain such
evidence, we weighed reproductive organs in both sexes collected in
December, February and March from colonies in both locations. Ad-
ditionally, we compared such weights to those found at eclosion, and
to those found when animals from both locations were exposed to
summer-like environments. Our data demonstrate striking similarities,
and some differences, between the two populations.

MATERIALS AND METHODS

Butterflies came from three localities. Those providing data on eclo-
sion values and prediapause (Herman 1981) response to summer-like
conditions were obtained from larvae reared outdoors on Asclepias
syriaca L. in June and July in Minnesota. Larvae were collected as first

VOLUME 438, NUMBER 1 Bl

instars immediately after hatching from eggs laid by wild-caught fe-
males. Eclosion data came from adults dissected on the day of emer-
gence. Prediapause response to summer-like conditions was measured
by holding newly emerged adults, fed daily with 30% honey, in in-
cubators at 25°C on a 16-h photophase for 10 days before dissection.
Californian monarchs were airmailed to Minnesota from the Natural
Bridges colony near Santa Cruz. These insects were either dissected
immediately upon arrival or incubated as above before dissection. An-
imals from California were examined in three separate years (1977-
79), and all results were pooled. Diapause values (Herman 1981) were
obtained by holding animals captured during the principal portion of
the diapause period (September—November) in summer-like conditions
for 10 days before dissection. Diapause data were also obtained over a
3-yr period from animals captured in both Minnesota (September) and
California (October-November), and the results were pooled. Mexican
animals were collected during 1983-84 at the Chincua and Herrada
colonies in Michoacan and Mexico, respectively, and airmailed (in three
separate shipments) to Minnesota as soon as possible (within eight days)
after capture. These adults were either dissected immediately or in-
cubated as above before dissection.

Anatomy of the reproductive tracts of both monarch sexes, and ter-
minology applied to the tracts, is discussed elsewhere (Urquhart 1960,
Herman 1975). Reproductive organs were dissected, cleaned of fat
body, blotted to remove excess saline, and weighed to the nearest 0.01
mg. Mature oocytes, defined as oocytes with chorionic ridges, were
counted in both ovaries in all females. Mated females were those with
sperm in the spermatheca, which in monarchs is that portion of the
receptable gland proximal to the common oviduct. Rear-wing maximal
length was measured to the nearest 0.56 mm with a ruler. Data were
analyzed with Student’s t-test. In this report “significant” refers to
statistical significance at the P < 0.05 level. All data are presented as
mean + SE. Some of the reproductive tract weight data obtained at
eclosion, in prediapause, and in diapause have been reported earlier
(Herman 1981, 1985, Herman et al. 1981) but this report includes new
data from additional animals.

RESULTS

Wing lengths. Females had rear wing lengths of 37.7 + 0.1 mm (n
= 145) and 37.8 + 0.1 mm (n = 201) in the Minnesotan and Mexican
populations, while males had significantly larger wings (38.1 + 0.1, n
= 145 and 38.2 + 0.1, n = 188) in the two populations, respectively.
Monarch wings examined at eclosion were not significantly different
from those of the Minnesotan and Mexican populations. Animals ob-

52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

tained from California in October-November had significantly smaller
wings than those from Mexico or Minnesota: 37.5 + 0.1 mm (n = 161)
and 37.9 + 0.1 mm (n = 141) for females and males, respectively. In
addition, Californian adults of both sexes exhibited significant declines
of wing length in February-March, to 37.0 + 0.2 mm (n = 114) and
37.6 + 0.1 mm (n = 201) for females and males, respectively. These
latter values were the lowest recorded during this study.

Reproductive organ weights. Mexican and Californian females showed
no significant weight differences in the ovaries (OV) and colleterial
glands (CG) on arrival in either December or February, but both organs
were slightly and significantly heavier in Californian females in March
(Fig. 1). The OV and CG weights of Mexican animals on arrival never
exceeded the eclosion values, but those of the Californian adults in
March were significantly heavier than at emergence. Mature oocytes
(MO) were never observed on arrival in females from either location.

Response of the OV and CG to summer-like conditions was quali-
tatively similar, but quantitatively different, in the two groups of fe-
males (Fig. 1). Incubated Californian females in all three months had
final organ weights significantly above the diapause value, generally
comparable to or above those in prediapause females, and larger than
those of incubated Mexican females. Mexican females exhibited OV
and CG weights after incubation that were close to the diapause values
in December, but well above those values in February and March.
Mating was more often observed in Californian animals, with 29%,
37%, and 96% mated in December, February and March, respectively,
while Mexican females exhibited only 17%, 8%, and 15% mating, re-
spectively, in the same months.

Accessory glands (AG), tubular glands (TG), and ejaculatory ducts
(ED) were typically heavier than at eclosion in males examined on
arrival (Fig. 2). There were no significant differences in gland weights
on arrival in December or March between the Californian and Mexican
males, but all three glands of Californian males were significantly heavier
in February. After incubation, all three glands of Californian males
exhibited responses significantly above those of diapause males, and
similar to prediapause animals, in all three months (Fig. 2). Mexican
male glands showed lesser responses only in December and February,
and the December response of TG and ED approached the diapause
value. All three male glands exhibited the same level of development
in both groups of incubated males in March.

On arrival, testes (TE) in both populations were always smaller than
at eclosion, while seminal vesicles-vas deferens (SV) complexes were
always larger (Fig. 3). In addition, Californian males had comparable
TE, but significantly smaller SV, in all three months. Incubated Cali-

VOLUME 438, NUMBER | 53

ON ARRIVAL INCUBATED

ORGAN WET WEIGHT (mg)

O rau,
OOCYTES/ANIMAL
120
80
40
7
eee) Sg PAD 2 i imey ls

STAGE OR MONTH EXAMINED

Fic. 1. Wet weights of colleterial glands (CG) and ovaries (OV), and total number
of mature oocytes (MO) in females collected in Californian (solid lines) and Mexican
(dashed lines) overwintering colonies. On-arrival data obtained from at least 17 animals/
data point, and incubated data obtained from at least 18 animals/data point. E = organ
weights at eclosion (n = 26), P = organ weights from prediapause animals (n = 47), and
D = organ weights from diapause animals (n = 85); 12 = December, 1 = January, 2 =
February, and 3 = March. Vertical lines indicate SE; negligible values are omitted.

fornian males consistently had both TE and SV near the diapause level,
while Mexican animals exhibited slightly smaller TE and significantly
larger SV (Fig. 3).

Receptacle glands (RG) and bursae copulatrix (BC) showed little
variation from eclosion level in Mexican females on arrival, as did RG
and BC of December females from California (Fig. 3). Both organs

54 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ON ARRIVAL INCUBATED

ORGAN WET WEIGHT (mg)

,

Eyal 1 2 3 Poe ya2 1 2 3
STAGE OR MONTH EXAMINED

Fic. 2. Wet weights of accessory glands (AG), tubular glands (TG), and ejaculatory
ducts (ED) from monarch males collected in Californian and Mexican colonies. On-arrival
data obtained by dissection of at least 27 animals/data point, and incubated data obtained
from at least 24 animals/data point. Data presentation and other abbreviations as in Fig.
1. Values of n for E, P, and D were 35, 59, and 79, respectively.

SS

exhibited weights significantly above eclosion values in Californian
animals examined on arrival in both February and March. Incubation
reduced RG size to similar values in both groups of females (Fig. 3),
and caused little BC weight change.

DISCUSSION

Our data show maior similarities and some differences in the repro-
ductive tracts of monarch butterflies from the Mexican and Californian

VOLUME 438, NUMBER 1 55D

ON ARRIVAL INCUBATED
ORGAN WET WEIGHT (mg)

ee
ee

See 1 2 3 Pa new 1 2 3
STAGE OR MONTH EXAMINED

Fic. 3. Wet weights of testes (TE) and seminal vesicles-vas deferens complexes (SV)
from males, and of receptacle glands (RG) and bursae copulatrix (BC) from females,
collected in Californian and Mexican colonies. Other abbreviations, data presentation,
and n values in Figs. 1 and 2.

overwintering colonies. Females from both populations possess OV and
CG that are nearly identical in December and February, and only
slightly different in March. In addition, these two organs are indistin-
guishable in December and February, in both groups of monarchs,
from those of insects emerging in summer in Minnesota. Female RG
and BC are also indistinguishable in December and similar in February

in the two populations, but both are larger in Californian females in
March.

56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

We believe the differences noted in weights of the female repro-
ductive organs of the two groups in March may be due to a higher
percentage of mating in the Californian animals. This conclusion is
supported by reports that mating increases juvenile hormone levels in
female monarchs (Herman & Barker 1977), and that juvenile hormone
stimulates the development of all four organs in this species (Herman
1985). Moreover, the considerable difference in BC weights appears
to be due principally to the greater number of Californian females
carrying spermatophores in their BC.

Female OV and CG also exhibit qualitatively similar responses to
incubation in summer-like conditions, that is, both groups of animals
exhibit their lowest response in December and their highest in March.
Again, we believe the best explanation for the quantitatively greater
response of these organs in Californian females is the higher percentage
of mated females in the Californian colonies. Response of female RG
to incubation mimics that of posteclosion animals in both groups, that
is, the glands diminish in weight after exposure to summer-like con-
ditions for 10 days (Herman et al. 1981). Response of BC in both groups
is comparable to that of both prediapause and diapause monarchs: the
weights of these organs after incubation are usually slightly above those
recorded at eclosion. The relatively high weight of BC in incubated
Californian females we again attribute to a higher proportion of mated

females.
For most of the overwintering period, the tracts of both groups of

females resemble those of females at eclosion in Minnesota. Both groups
of monarchs also show remarkable similarity in the response of the
female reproductive tract to incubation. Our data do suggest, however,
that diapause may last longer in Mexican females than in Californian,
since a response of the OV and CG to incubation similar to that of
diapause animals is found only in Mexican females in December.

Males from the two colonial sites also exhibit strong similarities and
some differences. Both groups of animals have AG, TG and ED of
similar size and significantly above eclosion values in both December
and March. However, the increased weight noted in Californian males
in February was delayed until March in Mexican males. On arrival,
both groups also possessed TE that were comparable and smaller than
those of newly emerged males. The SV of Californian animals were
significantly smaller on arrival, perhaps resulting from the additional
mating occurring in the Californian colonies.

The response of male organs to incubation was also similar in both
groups. AG, TG and ED exhibited pronounced responses that were
comparable in February and indistinguishable in March. Mexican males,

VOLUME 438, NUMBER 1 Pili

however, exhibited a reduced response of those three organs in Decem-
ber, suggesting that male diapause might also last somewhat longer in
the Mexican colonies. No striking changes were observed in the TE or
SV from either group after incubation. Thus, male monarchs from the
two colonial locations exhibited only minor differences in the weight
of their reproductive organs on arrival, and in the response of those
organs to incubation. The organs were, with the notable exception of
the SV, frequently indistinguishable on arrival and they normally ex-
hibited similar responses to summer-like conditions.

The above differences in Californian and Mexican monarchs do not
appear to be due to size differences of monarchs in the two populations.
As indicated by our data on wing length, Mexican monarchs were
somewhat larger than Californian, but exhibited similar or smaller
reproductive organs on arrival.

We conclude that overwintering monarchs of both sexes in both
California and Mexico maintain similar and low levels of reproductive
tract development through most of the overwintering period, and that
monarchs from both populations become more responsive to summer-
like conditions as the overwintering period progresses. The data suggest
that the postdiapause response of the reproductive tract to summer-
like conditions may be delayed in both sexes in Mexico, that is, repro-
ductive diapause may last longer in monarchs in the Mexican colonies.
The observed quantitative differences in the condition of the tracts,
and of their responses to incubation, may be due principally to the
greater percentage of mated females observed in Californian animals.
Why such a difference in mating exists in the two locations remains to
be determined.

ACKNOWLEDGMENTS

We are grateful for the use of the Northfield, Minnesota, estate of Alden and Ann
Mikkelsen for capturing and rearing monarchs, and for the efforts of Jeremy Criswell
and Claude Herman in that regard. This work was supported by the University of
Minnesota Graduate School and NSF Grant BSR-8500416 (to L.B.).

LITERATURE CITED

CALVERT, W. H. & L. P. BROWER. 1986. The location of monarch butterfly (Danaus
plexippus L.) overwintering colonies in Mexico in relation to topography and climate.
J. Lepid. Soc. 40:164-187.

HERMAN, W. S. 1975. Endocrine regulation of posteclosion enlargement of the male
and female reproductive glands in monarch butterflies. Gen. Comp. Endocrinol. 26:
034-540.

1981. Studies on the adult reproductive diapause of the monarch butterfly,

Danaus plexippus. Biol. Bull. (Woods Hole) 160:89-106.

58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1982. Endocrine regulation of the bursa copulatrix and receptacle gland of

Danaus plexippus L. (Lepidoptera: Danaidae). Experientia 38:631-632.

1985. Hormonally mediated events in adult monarch butterflies. In Rankin, M.
A. (ed.), Migration: Mechanisms and adaptive significance. Contrib. Mar. Sci. 27:
799-815.

HERMAN, W.S. & J. F. BARKER. 1977. Effect of mating on monarch butterfly oogenesis.
Experientia 22:688-689.

HERMAN, W.5S., C. A. LESSMAN & G. D. JOHNSON. 1981. Correlation of juvenile hormone
titer changes with reproductive tract development in the posteclosion monarch but-
terfly. J. Exp. Zool. 218:387-395.

LANE, J. 1984. The status of monarch butterfly overwintering sites in Alta, California.
Atala 9:17-20.

URQUHART, F. A. 1960. The monarch butterfly. Univ. of Toronto Press, Toronto. Pp.

242-249.

Received for publication 7 June 1988; accepted 4 November 1988.

Journal of the Lepidopterists’ Society
43(1), 1989, 59-65

INSTAR NUMBER AND LARVAL DEVELOPMENT IN
LYCAENA PHLAEAS HYPOPHLAEAS (BOISDUVAL)
(LYCAENIDAE)

GREGORY R. BALLMER AND GORDON F. PRATT

Department of Entomology, University of California,
Riverside, California 92521

ABSTRACT. The arctic-alpine butterfly Lycaena phlaeas hypophlaeas (Boisduval)
may have either four or five larval instars, the number apparently being fixed at ovipo-
sition. Factors affecting instar number were investigated in a laboratory colony of L. p.
hypophlaeas from the White Mountains of California. Adults in oviposition cages were
subjected to outdoor ambient conditions of day-length and temperature, but larvae were
reared indoors under nearly constant conditions (ca. 16 h light, 25°C). Larvae with five
instars predominated when oviposition occurred during short days (<11 h light) and low
maximum diurnal temperatures (ca. 22°C). When oviposition occurred during longer days
(>12 h light) and higher mean diurnal temperatures (ca. 33°C) most larvae had four
instars. Larvae having five instars required about 70% longer to mature than larvae having
four instars. Although diapause is not obligate, overwintering probably occurs as larvae,
which are more resistant to cold than are pupae and adults.

Additional key words: diapause, Lycaeninae, Oxyria digyna.

The primarily holarctic lycaenid butterfly Lycaena phlaeas (L.) in-
habits a wide range of habitats from sea level to ca. 4000 m elevation.
Various subspecies of L. phlaeas in Asia, Europe, and eastern North
America are multivoltine, polyphagous (primarily on Rumex species),
and common at low elevations. However, L. p. hypophlaeas (Boisduval)
of western North America is univoltine, apparently monophagous on
Oxyria digyna (L.) Hill, and confined to arctic-alpine habitats (Ferris
1974). This subspecies occurs in isolated colonies above 3000 m in the
central Sierra Nevada (Bishop Pass to Sonora Pass) and White Mountains
of California; collection records indicate a flight period from mid-July
to early September (Shields & Montgomery 1966, Ferris 1974). Its
habitat is one of extreme (especially cold) and unpredictable weather
for much of the year; even in summer there may be frost and occasional
snow. The restricted range of this subspecies is puzzling since suitable
hosts (Rumex spp.) are widespread at lower elevations in California
where they are utilized by other Lycaena species (Ballmer & Pratt,
1988). Also puzzling is the fact that, unlike other California Lycaeninae,
L. p. hypophlaeas may have either four or five instars (Ballmer & Pratt,
1988). Investigations reported here concerning the biology of L. p.
hypophlaeas were undertaken primarily to clarify instar number under
controlled environmental conditions. Additional observations on growth
rate and cold tolerance of stages may help explain the ability of this
butterfly to survive in the arctic-alpine zone.

60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

MATERIALS AND METHODS

A laboratory culture of L. p. hypophlaeas was derived from progeny
of a single female captured in the White Mountains (California, Mono
Co., White Mt., elev. ca. 4000 m, 26 July 1987) by J. F. Emmel. A
single mature larva was also found on Oxyria digyna (same data) by
G. F. Pratt. No other likely host was encountered at the collection site,
although at a lower elevation (3300 m) a few km away, Rumex pau-
cifolius Nutt. ex Wats. was abundant and utilized as a larval host by
Lycaena cupreus (W. H. Edwards) and L. editha (Mead). One of us
(G.F.P.) has also found larvae of L. p. hypophlaeas on Oxyria digyna
in the nearby Sierra Nevada (Mono Co., Mt. Dana, elev. 3600 m, 7
August 1985). Oxyria digyna is also a host for other arctic-alpine pop-
ulations of L. phlaeas in North America (Ferris 1974, Harry 1981).

In captivity, adult butterflies were confined with Rumex crispus L.
and R. acetosella L. in a screened cage (0.8 m X 0.3 m X 0.3 m) for
mating and oviposition. The cage was outdoors in a sheltered location
with partial sun exposure. Ova were brought indoors and larvae were
reared in quart (0.95-1) plastic food containers on leaves of both R.
acetosella and R. crispus. Pupae were transferred to the screened cage
for eclosion.

Nineteen neonatal larvae from ova produced during the first week
of September (long-day ova) were placed individually in 7-dram (25-
ml) plastic vials and reared on leaves of R. crispus. Leaves were replaced
as needed (usually every 2-3 days for young larvae and daily for last
instars). A second group of 23 neonates from ova produced at the end
of October (short-day ova) was reared under slightly different condi-
tions. These larvae were kept individually in 25-dram (90-ml) plastic
vials with two 25-mm-diam. screened ventilation openings, and fed
leaves of R. crispus. A small hole in each lid allowed the leaf stem to
protrude for immersion in water contained in a second vial; this per-
mitted leaves to remain fresh longer while the ventilation prevented
mold which occasionally appeared in the smaller nonventilated vials
used earlier. All larvae were reared indoors at 25 + 38°C (brief tem-
perature fluctuations resulted from operation of indoor heating and
cooling equipment). Larvae were inspected daily for signs of ecdysis.
Dates of ecdysis were recorded for each larva, and head capsules were
measured using a microscope and ocular micrometer.

Total illumination from indirect natural daylight and artificial light-
ing from overhead fluorescent lamps exceeded 16 h per day. Only
adults and ova were exposed to natural (outdoor) diurnal photoperiods
and temperatures. There were 12.5 h of daylight (sunrise to sunset) on
10 September, the mean eclosion date for long-day ova, and 10.75 h

VOLUME 43, NUMBER 1 61

of daylight on 3 November, mean eclosion date for short-day ova; the
effective period of daylight on both dates may have been somewhat
longer. Mean maximum and minimum diurnal temperatures for the
seven days preceding mean eclosion dates were 33.38°C and 13.3°C,
respectively, for long-day ova and 21.5°C and 12.8°C, respectively, for
short-day ova.

Four mature larvae from long-day ova were preserved and the re-
mainder allowed to pupate. Four pupae were refrigerated at 5°C for
28 days to test the effect of mild but prolonged chilling. All larvae from
short-day ova were reared to adults without chilling.

Other experiments tested the effect of extreme chilling on additional
larvae, pupae, and adults. Ten second instars and 12 fourth instars (all
destined to have 5 instars) were removed from the colony during De-
cember (from short-day ova) and placed in 25-dram (90-ml) ventilated
vials, as described above, with fresh host leaves.

Vials were refrigerated (5°C) for 21 days, then wrapped in damp
paper towels and placed inside larger sealed jars which were kept at
—7°C for 28 days. After the freezing treatment, jars were allowed to
thaw at 5°C for 24 h; the vials were then removed, larval condition was
assessed, and survivors were given fresh host leaves; rearing continued
at 25°C. Ten pupae were similarly treated (7 days at 5°C followed by
28 days at —7°C).

While refrigerated at 5°C, second instars fed considerably, but fourth
instars did not feed. Leaves damaged by feeding were dried in a press,
weighed, and photocopied. The paper images were cut out and weighed;
then their feeding-damaged portions were cut out and weighed to
determine percent of feeding damage. The latter values were then used
to calculate quantity of leaf tissue eaten per larva.

On several occasions it was noted that brief periods (1-6 h) of exposure
to —7°C were not lethal to adults; but death usually occurred after 2-
3 consecutive exposures of such duration. The effect of milder but more
prolonged exposure to cold was tested by refrigerating 13 freshly eclosed
adults at 5°C for 30 days. Adults were placed individually in 25-dram
(90-ml) ventilated vials which were then placed inside plastic bags with
damp paper towels and refrigerated.

Statistical significance of differences in head size and instar duration
was determined by t-tests.

RESULTS

Instar number and duration. Mean duration of each larval instar
and overall larval and pupal stages are presented in Table 1. Only one
male and six females from long-day ova are included owing to loss of

62 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

gender data for the remainder; therefore, discussion of sex-correlated
differences in development rates is restricted to larvae from short-day
ova. In general, females developed more rapidly than males, especially
in the third and ‘extra’ instars (fourth instar of five-instar larvae). Sex-
correlated differences in development times among larvae from short-
day ova were most significant for third instars (P = 0.02 and 0.051 for
four- and five-instars, respectively); for other larval instars P ranged
from 0.13 to 0.84. Mean larval stage duration of five-instar larvae was
greater than that of four-instar larvae; the difference for males (ca. 35%
greater) is not significant (P = 0.08), but for females (ca. 49% greater)
is (bh = D008).

Most larvae from long-day ova (17 of 19) had four instars, and
required a mean of 23 days to pupate (both sexes combined); one larva
had five instars, and another, which died of a whitish fungal infection
in the fourth instar, would have molted again judging from its head
size had it survived. Short-day ova produced mostly five-instar larvae
(17 of 23), and required a rounded mean of 27 days to pupate (both
sexes‘ combined); remaining larvae had five instars and required a
rounded mean of 41 days to pupate. Nevertheless, there was consid-
erable individual variation and some overlap in developmental time
ranges. It is remarkable that among both four- and five-instar larvae,
some individuals remained as larvae about twice as long as others; range
of larval duration for all larvae was 13-59 days. No significant differ-
ences in pupal duration were found with respect to sex, number of
larval instars, or day length.

Head size. Measurements of head widths indicate no significant sex-
related differences (P = 0.47, 0.21, 0.40, and 0.55 for instars 1, 2, 3,
and 4, respectively, of five-instar larvae from short-day ova). There
were also no significant differences in mean head size between four-
and five-instar larvae from short-day ova for instars 1, 2, and 3 (P =
0.39, 0.54, and 0.78, respectively). Therefore, data were pooled for all
larvae in comparing head sizes of first, second, third, and ‘extra’ instars
of larvae conceived under long- and short-day conditions (Table 2).
Since most larvae were reared to pupation, and the last-instar head
capsule was invariably deformed at pupation, the head widths of last-
instar larvae included in Table 2 are based primarily on preserved
larvae reared concurrently. The mean first-instar head width of larvae
from short-day ova was slightly but significantly (P = 0.015) larger than
that of long-day ova. The head size of the ‘extra’ (fourth) instar of five-
instar larvae was intermediate between that of third and last instars;
thus, some growth occurred in all instars.

Values presented here for head size should not be considered typical
of all populations of L. p. hypophlaeas. The mean last-instar head width

63

VOLUME 438, NUMBER 1

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64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

(1.46 mm) of 10 L. p. hypophlaeas larvae from Mount Dana in the
Sierra Nevada is 17% larger than that of 13 larvae from the White
Mountain colony (1.24 mm); this difference is highly significant (P =
0.00002).

Cold exposure. The mortality rate from freezing was similar for
second- and fourth- (‘extra’) instar larvae. Six (of 10) second instars
exposed to —7°C for 30 days survived, as did 7 (of 12) fourth instars
exposed to the same conditions. During 21 days of exposure to 5°C
(before freezing), second instars consumed a mean 0.47 cm? (0.002 mg,
dry wt.) of leaf tissue.

Adults and pupae were less tolerant of cold. Although all 4 pupae
exposed to 5°C for 28 days survived and produced adults, only 1 of 10
pupae frozen (—7°C) for 28 days eclosed, and it was unable to properly
expand its wings. All 13 adults refrigerated at 5°C for 30 days perished.

CONCLUSIONS

Instar number in Lycaena p. hypophlaeas is variable; four instars are
prevalent under warm, long-day (late summer) conditions while five
instars predominate when days are cooler and shorter in fall. Number
of instars is apparently fixed at oviposition. Developmental time is
greater for five-instar larvae than for four-instar larvae. Similar length-
ened larval development and extra molts, but without apparent growth,
in response to short day-length have also been reported in the multi-
voltine L. p. daimio Seitz of Japan (Sakai & Masaki 1965, Endo et al.
1985).

The greater development time required for L. p. hypophlaeas larvae
produced under short-day conditions is probably important in winter
survival. Unlike at least most other California Lycaena species, L. p.
hypophlaeas does not have an obligate diapause. However, an extended
larval duration induced by short day-length and further promoted by
reduced activity due to cold fall and winter temperatures reduces the
possibility of premature maturation and subsequent exposure of the less
cold-tolerant stages to winter conditions. The great variability in de-
velopment time probably also contributes to survival, since it ensures
that some individuals are likely to be in the most cold-tolerant (larval)
stage at all times of the year.

ACKNOWLEDGMENTS

We thank J. F. Emmel for providing the ova of L. p. hypophlaeas that began our
colony. The Riverside office of the National Weather Service provided local ambient
temperature and day-length information. David M. Wright graciously reviewed the
manuscript.

VOLUME 43, NUMBER 1 65

LITERATURE CITED

BALLMER, G. R. & G. F. PRATT. 1989. A survey of the last instar larvae of the Lycaenidae
of California. J. Res. Lepid. 27:1-80.

ENpo, K., Y. MARUYAMA & K. SaKal. 1985. Environmental factors controlling seasonal
morph determination in the small copper butterfly, Lycaena phlaeas daimio Seitz.
J. Insect Physiol. 31:525-532.

FERRIS, C. D. 1974. Distribution of arctic-alpine Lycaena phlaeas L. (Lycaenidae) in
North America with designation of a new subspecies. Bull. Allyn Mus. 18:1-13.

Harry, J. L. 1981. A new foodplant for Lycaena phlaeas. Utahensis 1:5.

SAKAI, T. & S. MASAKI. 1965. Photoperiod as a factor causing seasonal forms in Lycaena
phlaeas daimio Seitz (Lepidoptera: Lycaenidae). Kontyti 33:275-283.

SHIELDS, O. & J. C. MONTGOMERY. 1966. The distribution and bionomics of arctic-
alpine Lycaena phlaeas subspecies in North America. J. Res. Lepid. 5:231-242.

Received for publication 12 August 1988; accepted 14 November 1988.

Journal of the Lepidopterists’ Society
43(1), 1989, 66-68

GENERAL NOTES

STATUS OF THE PAPILIONID TYPES PAPILIO STEWARTI AVINOFF
AND P. MORRISI EHRMANN

Additional key words: taxonomy, Neotropics.

Recently we reviewed the types and some newly acquired specimens of several pa-
pilionid taxa of uncertain taxonomic status known only from extremely small samples
(Johnson, K., R. Rozycki & D. Matusik 1985, J. N.Y. Entomol. Soc. 93:99-109, 1986, 94:

Pmctis Shur Te ds Pmctis Chm TPe as
heen. Hogcle Ce. a0. JY y trea. Motcle Ck, V3.0. /Y/4

Loja SG, PRLCuK Lope 5:6. BeOum

dann. Carn, Mas.

wr

Die Vo/ 1927.
me: PLEX, fig. 3 @

Fic. 1. Papilio holotype males. A, P. stewarti, upper surface on right, under on left;
forewing expanse, base to apex, 50.0 mm; B, P. morrisi, as above; forewing expanse 40.0

mm.

VOLUME 438, NUMBER 1 67

Fic. 2. _Papilionid male genital valves, inner lateral view. A, Papilio stewarti holotype
male; B, P. scamander joergenseni, Tucuman, Argentina (David Matusik Collection); C,
P. morrisi holotype male; D, Protesilaus xenaides, Rio Pastaza, Ecuador (Am. Mus. Nat.
Hist., New York).

383-393; Johnson, K. & R. Rozycki 1986, J. N.Y. Entomol. Soc. 94:516-525; Johnson, K.,
R. Rozycki & D. Matusik 1986, J. Lepid. Soc. 40:65-66; Johnson, K. & D. Matusik 1987,
J. Lepid. Soc. 41:65-69, 108-118; Johnson, K., D. Matusik & R. Rozycki 1987, J. Res.
Lepid. in press). The status of two other papilionid taxa, P. stewarti Avinoff and P. morrisi
Ehrmann, are of interest to South American colleagues preparing a study of Neotropical
Papilionidae (K. S. Brown Jr. pers. comm.). These taxa, originally described as species
from one, or very few, specimens (types at Carnegie Museum of Natural History, Pitts-
burgh, CMNH), have had little subsequent report in the literature, and their genitalia
have hitherto not been examined.

Papilio stewarti (Avinoff, A. 1926, Ann. Carnegie Mus. 16:355-375, type locality, TL,
Samaipata, Bolivia). The holotype male (Fig. 1A) indicates P. stewarti belongs to the
“scamander Group’ of Pterourus (tribe Papilionini) (sensu Hancock, D. L. 1983, Smith-
ersia 2:1-48), and is a synonym of the tailed subspecies P. scamander joergenseni Rober
(Rober, J. K. M. 1925, Entomol. Mitteil. 14:85) which occurs commonly southward in
Bolivia and northwestern Argentina (D’Almeida, R. F. 1965, Catalogo dos Papilionidae
Americanos, Sociedade Brasileira de Entomologia, 366 pp.). Male genitalia of P. stewarti
(Fig. 2A) differ negligibly from P. s. joergenseni (Fig. 2B) and nominate P. scamander
(Johnson, Matusik & Rozycki 1985, above:fig. 2A).

Papilio morrisi (Ehrmann, G. A. 1921, Lepidoptera 5(2):17, TL of original description
“Peru” but TL of holotype “Loja, S.E. Peru” [sic]). The holotype male (Fig. 1B) indicates
P. morrisi belongs to the “lysithous Group” of Protesilaus (tribe Leptocircini) (sensu
Hancock 1983, above), and, more specifically, the “harmodius cluster” (sensu Johnson,
Rozycki & Matusik 1986, J. N.Y. Entomol. Soc. 94:383-393; 1987, above). Wing characters
(Fig. 1B), genitalia (Fig. 2C), and locality data (other P. morrisi specimens labelled “Rio
Bamba, Ecuador’’) indicate P. morrisi is a synonym of Protesilaus xenaides (Hewitson)
(Fig. 2D) (Johnson, Rozycki & Matusik 1986, J. N.Y. Entomol. Soc. above:fig. 4F).

W. J. Holland (1927, Ann. Carnegie Mus. 17:299-365) noted that Ehrmann, in de-
scribing many (often synonymic) taxa, sometimes made clerical errors. Holland, in his
entry concerning P. morrisi, repeats Ehrmann’s citation of “Peru” as the type locality,

68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

but in text cites Ehrmann’s “notebook” as stating “Laja, Peru” [sic]. Holland questioned
this as possibly “Loja” [Ecuador]. The holotype’s labels, not figured by Holland (but shown
here in Fig. 1B) appear to say “Loja” [Ecuador], compatible with data on two paratype
males (CMNH) labelled “Rio Bamba, Ecuador’.

KurRT JOHNSON, Department of Entomology, American Museum of Natural History,
Central Park West at 79th Street, New York, New York 10024; DAviD MATusIk, De-
partment of Entomology, Field Museum of Natural History, Roosevelt Road at Lake
Shore Drive, Chicago, Illinois 60605; AND RICK ROZYCKI, 5830 South McVicker Avenue,
Chicago, Illinois 60638.

Received for publication 8 December 1987; accepted 19 September 1988.

Journal of the Lepidopterists’ Society
43(1), 1989, 68-71

REPEATED COPULATION IN AN ORANGE HAIRSTREAK,
SHIROZUA JANASI. A CASE OF MATE GUARDING?

Additional key words: Lycaenidae, mating, behavior.

In butterflies, multiple copulations are common not only in males (Svard, L. & C.
Wilkund 1986, Behav. Ecol. Sociobiol. 18:325-330) but also in females (Burns, J. M. 1968,
Proc. Nat. Acad. Sci. U.S.A. 61:852-859; Ehrlich, A. H. & P. R. Ehrlich 1978, J. Kans.
Entomol. Soc. 51:666-697; Thornhill, R. & J. Alcock 1983, The evolution of insect mating
systems, Harvard Univ. Press, Cambridge, Massachusetts, 547 pp.; Drummond, B. A. 1984,
pp. 291-370 in Smith, R. L. (ed.), Sperm competition and the evolution of animal mating
systems, Academic Press, Orlando, Florida, 687 pp.). However, within-a-day repeated
copulations are very rare in both sexes (Svard & Wilkund, above; Fujii, H. unpubl. data).

Recently, Tanaka and Unno (in Fukuda, H., E. Hama, K. Kuzuya, A. Takahashi, M.
Takahashi, B. Tanaka, H. Tanaka, M. Wakabayshi & Y. Watanabe 1984, The life histories
of butterflies in Japan, Vol. 3, Hoikusha, Osaka, 373 pp., Japanese, English summary)
observed that females of an orange hairstreak, Shirozwa janasi (Janson) soon copulated
with other males after preceding copulations. Such immediate remating seems to be
exceptional in butterflies.

In the summer of 1986, I observed repeated within-pair copulations in S. janasi. This
paper describes mating behavior in S. janasi and suggests that mate guarding is a possible
consequence of remating.

Shirozua janasi is the only omnivorous species in the tribe Theclini. Like other Theclini,
it has one generation per year, and imagines are on the wing from late July to September
(Fukuda et al., above).

Field observations were made in secondary forest including Quercus serrata Murray
(Fagaceae), Pinus densiflora Sieb. et Zucc. and Larix Kaempferi (Lamb.) (both Pinaceae),
at Sakai village, Nagano, Japan in August 1986.

The male of S. janasi flies 3-10 m above the ground and alights just behind the female.
This has been called a patrolling-type mate-locating strategy (Scott, J. A. 1973, J. Res.
Lepid. 11:99-127; Fujii, H. 1982, Yadoriga (107/108):1-37, Japanese). Then the male’s
wings are held open about 30° apart and fluttered. The male moves slowly to the side of
the female, bends its abdomen towards the tip of the female’s abdomen, and copulates
(Fig. 1). This courtship sequence usually ends in successful copulation within 5 sec.

During the survey, five courting pairs were found, and all copulated thereafter. At
intervals after copulation began, I disturbed these pairs by approaching or touching them
with my fingers until they separated or flew away in copula. As shown in Table 1, most

VOLUME 43, NUMBER Il 69

approaching
mate-searching

SS flight

courtship

Wa flying away

F
ing

flying away =

&
patrolling ie =

copulation

Fic. 1. Sequence of repeated copulations in S. janasi. M: male, F: female. Further
details in text.

pairs (No. 1; No. 2, lst & 2nd copulations; No. 8, lst & 2nd; No. 4, lst; No. 5, 2nd) were
separated easily when disturbed within 10 min after they initiated copulation. In contrast,
the pair (No. 5, 1st copulation) that had been copulating more than 30 min was not easily
separated. Instead, it usually flew away in copula, during which the female always carried
the male. Possible mate guarding was observed when the pair was separated as a result
of my disturbance: an uncoupled male flew away but returned immediately to where
the male had copulated just before. An uncoupled female from a disturbed pair also flew
away from the place where it had copulated (usually a leaf), but the female rarely moved
so far. Therefore, a returned male could usually find its previous partner, and then the
male courted and mated the same partner again (Fig. 1). Such behavior was observed in
four of the seven separated pairs, including not only pairs that had copulated for less
than 10 min but one pair that had copulated more than 150 min and then remated twice
(Table 1).

According to Tanaka and Unno (Fukuda et al., above), copulation in S. janasi usually
starts within about 10 sec and ends within 10 min of first contact. In this study, most
pairs ended copulations within 10 min as a result of my disturbances. It should be noted,
however, that the lst copulation of pair No. 5 lasted about 2.5 h in spite of my intensive
disturbances (Table 1). Further observations are needed to determine how long a bout
of copulation lasts under undisturbed conditions.

Although my data are insufficient to say how long a time is necessary for the male to
inseminate the female, it seems that 10 min is too short for successful insemination because

70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Effects of artificial disturbances on in copula pairs. Asterisk indicates oc-
currence of remating. C: first copulation. R: remating. U: uncoupling. F: flight (flying
female always carried male). L: uncoupled male lost previous partner even though he
seemed to search for her. X: I could not follow uncoupled individuals because of rapid
flights. S: I stopped observing.

Distance Time after copulation began (min)

from previous
Pair no. copulation (m) 0 1 3 5 10 30 150
1 — C UX
2 — C FF FU
2 I! R F IONE,
3 _ C FU
3F ] R UL
4 _— C F U
4* 0 R F FX
) — C FF F FF FF FFFU
On 0) R FF U
ia 2 R S

duration of copulation in almost all butterflies is known to last over 30 min (Scott, above;
Shields, O. & J. F. Emmel 1973, J. Res. Lepid. 12:25-64; Fukuda et al. 1982-1986, The
life histories of butterflies in Japan, Vol. 1, Hoikusha, Osaka, 277 pp., Vol. 2, 325 pp.,
Vol. 3, above, Vol. 4, 373 pp., Japanese, English summary). If so, any male that uncouples
within 30 min after copulation begins should remate with the previous partner to insure
successful insemination. If this male does not find the previous partner, she will be
inseminated by another male. In fact, Unno and Tanaka observed that such a female
copulated again with another male.

Pair No. 5 remated twice after the lst copulation, which lasted about 2.5 h. The male
of this pair is likely to have transferred its sperm to the female’s bursa copulatrix during
the Ist copulation, because in butterflies most successful copulations are known to finish
within 1-2 h (Scott, above; Shields & Emmel, above). If insemination did occur, the 2nd
and 3rd copulations of pair No. 5 may be copulatory mate guarding behavior by the
male. Copulatory mate guarding has not been reported in Lepidoptera previously (Thorn-
hill & Alcock, above; Drummond, above), but Drummond considered that lepidopteran
males might also guard their mates from the advances of other males while still in copula.
However, in some cases where the male successfully copulates several times within 1 or
2 days, a bout of copulation may last several hours after the 2nd copulation (Svard &
Wilkund, above; Fujii unpubl.). Additional studies are needed to know whether or not
repeated copulations in S. janasi are truly copulatory mate guarding.

Although in copula pairs of S. janasi were separated very easily by my disturbances,
this is not true in other butterflies (Fujii, H. 1975, Gekkan-Mushi [52]:14-19, Japanese).
Why do in copula pairs of S. janasi separate so easily? Longer copulations are probably
more dangerous than shorter copulations, because in copula pairs are more conspicuous
and less mobile and should therefore suffer higher predation. Moreover, both sexes of S.
janasi are reddish orange in color, so they are very conspicuous on green leaves. Therefore,
the easy-to-separate copulation behavior of S. janasi may have evolved in response to
predation pressure. In favor of the hypothesis is the fact that two other orange hairstreaks,
Japonica lutea (Hewitson) and J. saepestriata (Hewitson), copulate at dusk (Fujii, above;
Fukuda et al., above), while S. janasi copulates during the day when bird predation seems
much heavier (Fujii, above; Saigusa, T. 1983, 30th annual meeting of the Lepidopte-
rological Society of Japan).

VOLUME 43, NUMBER 1 7Ay

I thank B. A. Drummond and an anonymous referee for valuable comments on the
manuscript.

HisasHI Fuji, Department of Zoology, Kyoto University, Sakyo, Kyoto, 606 Japan.

Received for publication 24 September 1987; accepted 2 September 1988.

Journal of the Lepidopterists’ Society
43(1), 1989, 71

AN EMENDED SPECIFIC NAME IN EUPITHECIA (GEOMETRIDAE)
Additional key words: Chile, Eupithecia taracapa, E. tarapaca.

Prof. Raul Cortés, of the Instituto de Entomologia, Universidad Metrolitan de Ciencas
et la Educacion, Santiago, Chile, called my attention to an incorrect geographical name
and a resulting incorrect species-group name in my 1987 paper “The Eupithecia (Lep-
idoptera, Geometridae) of Chile,” Bull. Am. Mus. Nat. Hist. 186:269-363. On p. 325 I
gave the type locality of the new species as being in “Taracapa” Province and Region,
and proposed for it the specific name Eupithecia taracapa, a noun in apposition taken
from the type locality. The correct geographic term is Tarapaca, and so I am emending
the name of the species to Eupithecia tarapaca, thus replacing the incorrect E. taracapa
Rindge 1987; both names have the same holotype. This emendation is in conformity with
Articles 32(d) and 33(b)(ii) of the International Code of Zoological Nomenclature.

FREDERICK H. RINDGE, Department of Entomology, American Museum of Natural
History, New York, New York 10024.

Received for publication 8 September 1988; accepted 8 September 1988.

Journal of the Lepidopterists’ Society
43(1), 1989, 72

THE VALID GENERIC PLACEMENT FOR
“CALOTHYSANIS” AMATURARIA (WALKER)
(GEOMETRIDAE, STERRHINAE)

Additional key words: taxonomy, Timandra amaturaria.

~The common eastern North American sterrhine geometrid moth described by Walker
in 1866 as Timandra amaturaria has often been placed in the genus Calothysanis Hiibner
1823. Examples are A. S. Packard (1876, Monograph of the geometrid moths or Pha-
laenidae of the United States, in Hayden, F. V., Report of the United States Geological
Survey of the Territories 10:317), L. B. Prout (1934, Lepidopterorum catalogus, Part 61:
51), Prout in A. Seitz (1936, Macrolepidoptera of the world, Vol. 8:94), and W. T. M.
Forbes (1948, Lepidoptera of New York and neighboring states, Part 2:119).

Timandra, on the other hand, was used in the 1917 check list of Barnes and Mc-
Dunnough and the 1938 one of McDunnough (numbers 3913 and 4205, respectively), as
well as in earlier works by A. Guenée, C. F. Gumppenberg, and Prout himself (1913, in
Seitz, A., Macrolepidoptera of the world, Vol. 4:47). Both combinations have appeared
in other literature, and on the head labels of collections, creating considerable confusion.

Since Calothysanis Hiibner 1823 predated Timandra Duponchel 1829, and had been
applied by Forbes and by Prout in his most recent works, I used Calothysanis in my
Sterrhinae section of the R. W. Hodges (ed.) (1983) Check list of the Lepidoptera of
America north of Mexico (p. 100) and my Field Guide to Moths of Eastern North
America (Covell 1985, p. 377; pl. 46, fig. 14).

Prout (1913) chose Timandra over Calothysanis on the basis of Butler’s selection of
Acidalia imitaria Hubner as the type of Calothysanis (Butler, A. G. 1881, Trans. Entomol.
Soc. London 1881:342). D. S. Fletcher (1979, in Nye, I. W. B., Generic names of the
moths of the world, Vol. 3:34) verified that selection.

The type of Timandra was originally designated as Phalaena amataria innaane 1761.
Fletcher, in his treatment of Timandra (p. 206), pointed out that Phalaena amataria
Linnaeus is an unjustified emendation of P. amata Linnaeus, and therefore an objective
synonym of amata. The original description of amata was based not on specimens but
on two figures in another work, which turn out to be two other species. An unnamed
series of moths left by Linnaeus were misidentified by later workers as P. amataria.
Fletcher concluded that Timandra griseata Petersen 1902 is the earliest available name
for those moths which Linnaeus had misidentified as P. amataria, and is therefore the
type of Timandra.

Since imitaria Hubner is in the genus Scopula Schrank 1802, Calothysanis must be
considered a junior synonym of Scopula as it was thus first published by Prout (1934:
169).

The generic and species treatments in Hodges (above) should therefore read as follows:

TIMANDRA Dup., 1829
BRADYEPETES Steph., 1831
7147 amaturaria Wlk., 1866
effusaria (Prout, 1936)

The other 14 species of Timandra, including griseata, occur in Eurasia (Prout 1934).
The author thanks an anonymous reviewer for helpful criticism.

CHARLES V. COVELL Jr., Department of Biology, University of Louisville, Louisville,
Kentucky 40292.

Received for publication 8 September 1988; accepted 18 October 1988.

Journal of the Lepidopterists’ Society
43(1), 1989, 73-78

BOOK REVIEWS

THE MOTHS OF AMERICA NORTH OF MEXICO. The Wedge Entomological Research
Foundation, Washington, D.C. (Distributed by the Wedge Entomological Research Foun-
dation, % National Museum of Natural History, MCR-127, Washington, D.C. 20560; E.
W. Classey Ltd., P.O. Box 98, Faringdon, Oxfordshire SN7 7DR, England; Bioquip
Products, 17808 LaSalle Ave., Gardena, California 90248; Entomological Reprint Spe-
cialists, P.O. Box 77224, Dockweiler Station, Los Angeles, California 90007.)

Fascicle 18.1. Geometroidea, Geometridae (Part), by Douglas C. Ferguson. 1985. 131
pp., 4 color pls. Soft cover. $55.

The appearance of this fascicle of “MONA,” as Richard B. Dominick affectionately
dubbed it, is unique. It is a memorial fascicle in which intimate details of Dick’s life,
personality, and contributions to lepidopterology as founder of this series are presented
most touchingly by his widow, Tatiana. This, with a full-page portrait of Dick, precedes
the paginated body of the work.

Also, this fascicle is the first covering “macros” since the Lymantriidae volume in 1978,
and is also the first to treat a subfamily of Geometridae.

Ferguson's treatment consists of nomenclatural and descriptive introduction to the
subfamily Geometrinae, leaving superfamily and family material as headings followed
by “(continued),” thus anticipating placement of Archearinae, Oenochrominae, and En-
nominae ahead of the greens in phylogenetic order. We can expect superfamily and
family treatments in a later fascicle.

Tribal and generic descriptions follow, each with a key to the next lower category.
Species, and where appropriate, subspecies, are painstakingly described, with illustrations
of wing venation and genitalia adding greatly to the usefulness of descriptions. In addition
to the bibliography, there are appended abbreviations for contributing collections and
individuals, an animal-name index, and a plant-name index.

This work is based primarily on Ferguson (1969, A revision of the moths of the subfamily
Geometrinae of America north of Mexico [Insecta, Lepidoptera], Bull. 29, Peabody Mu-
seum, Yale University)—a publication based on his doctoral dissertation. Since publishing
that work, Ferguson has made some changes, most of them introduced in R. W. Hodges,
ed. (1983, Checklist of the Lepidoptera of America north of Mexico, The Wedge Ento-
mological Research Foundation, Washington, D.C., 284 pp.). These include Synchlora
albolineata and S. liquoraria treated as subspecies of S. aerata; three new synonyms for
S. frondaria; S. frondaria denticularia reduced to synonymy of S. frondaria frondaria;
S. xysteraria (Hulst) applied to the Florida moths treated as S. gerularia, a similar species
reaching North America only in southern Texas; S. herbaria hulstiana reduced to syn-
onymy of S. herbaria; Merochlora synonymized to Chetoscelis (not indicated as new
synonymy in the Checklist); exchange of position of Xerochlora and Chloropteryx (the
1969 work had Xerochlora first); addition of Hemithia aestivaria (Hbn.), a European
introduction discovered in Canada in 1979; and elevation of Hethemia pistasciaria
insecutata from synonym to subspecies status, with auranticolorata as its synonym. In
addition, the 1985 fascicle elevates former synonym remotaria (Wlk.) to replace the name
latipennis Hulst—a correction from the 1969 treatment in which remotaria was attributed
to Grossbeck.

The text abounds in small refinements and improvements over the revision, and re-
duction in details that a formal revision normally includes. Ranges and other information
are improved for some species. I found partial life history information available for 8
species of the 76 in our fauna for which none appeared in the earlier work. An example
of range extension is that of Nemoria tuscarora Ferguson (1969:61), once known only
from Appalachian North Carolina, Virginia, and West Virginia, now known from north-
central Kentucky with flight date extending into August from the 27 July limit stated
earlier. Likewise, the ranges of N. saturiba Ferguson and N. elfa Ferguson are extended
northward by addition of Kentucky records in the fascicle.

Genitalia and other line drawings are copious and well rendered, and the delicate
patterns and pastel colors of the moths on the four plates are appealing. Several years

74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

elapsed between photography of the plates and their production for the book, however;
some moths appear more grayish green or duller than specimens with which I compared
their published likenesses. Having had similar disappointments with color registry, I am
sure Ferguson must be equally disappointed that the lovely gee colors did not come
out as well in production as one would wish.

This is a well written and illustrated book which enables one to identify usually by
superficial features the North American Geometrinae. It also contains considerable in-
formation additional to that in Ferguson’s earlier revision, plus variation represented in
the color plates by multiple illustrations of some species (six of N. elfa, for example). It
is a worthy addition to the MONA series, and a fitting fascicle to commemorate the life
and contributions of Dick Dominick.

CHARLES V. COVELL JR., Department of Biology, University of Louisville, Louisville,
Kentucky 40292.

Fascicle 7.1. Gelechioidea, Gelechiidae (Part), Dichomeridinae, by Ronald W. Hodges.
1986. 195 pp., 4 color & 34 monochrome pls. Soft cover. $70.

This volume presents the first revision of any large group of North American Gelechiidae
in contemporary times, and as such, it brings welcome order to part of a family of small
moths whose classification is chaotic at best. The fauna covered is small, however, in
relation to the size of the family: 84 species out of possibly 1500+ on this continent.
Three genera are recognized (one is monobasic), with most species (74) placed in Di-
chomeris. How confused the group was previously is reflected in the 81 generic synonyms
under Dichomeris, 60 of which are new or revised. The generic synonymy will prove
especially useful because it is worldwide in scope. Also noteworthy in the treatment of
one genus, Helcystogramma, is a list of extralimital (non-North American) species. Un-
fortunately, a similar list is not included for the larger genus Dichomeris, presumably for
reasons of length (it includes several hundred species worldwide). The number of new
species, 42 or 50% of taxa treated, is a fair reflection of how poorly North American
gelechiids are known.

Because this is the first MONA fascicle to treat gelechiids, family and subfamilies are
defined. Only three subfamilies are recognized, with Gelechiinae being vastly enlarged
to include the majority of our gelechiids. It is quite probable that this assemblage of taxa
comprising several thousand species worldwide is defined by primitive character states,
and that it will eventually be broken up into monophyletic units. Nevertheless, Hodges
must be praised for attempting to delineate precisely the notoriously ill-defined higher
categories of gelechiids.

Keys based on external features are given for Dichomeris and Helcystogramma species.
They do not permit the separation of all species, however, because several species are
distinguished with certainty by genitalia only. This is an unavoidable fact of many
microlepidoptera groups, at least until distributions and natural histories become better
known. For Dichomeris species, there are also keys based on male and female genitalia.

Species descriptions are lengthy and detailed. They could have been shortened to
conserve space and improve readability by deleting unnecessary details of color. For
many species, genitalia receive only a brief reference to a figure. It would have been
more useful to give distinctive, comparative features because of their importance for
species separation. Perhaps this was omitted on account of lepidopterists who dislike or
are unable to make genitalia preparations. However, it is likely that whoever is interested
in these small moths will also get involved in the techniques required for their study.
This notwithstanding, omission of genitalia comparisons partly defeats the purpose of
including plates showing genitalia of all species treated because the reader is often left
trying to figure out what detectable differences in the figures have taxonomic value.
Systematists with a phylogenetic bent will be pleased to find a table of character states
that covers 38 characters, albeit nonpolarized, for the 20 species groups of Dichomeris.

The four color plates are stunningly sharp—an improvement over previous fascicles

VOLUME 48, NUMBER 1 75

on oecophorids and cosmopterigids in which the pictures were slightly fuzzy. The line
drawings are of fine quality, although several genitalia illustrations lack contrast between
membranous and sclerotized parts, giving the impression that these structures are some-
what uniformly sclerotized. The 34 monochrome plates illustrating male and female
genitalia are generally of excellent quality. Hodges is to be praised for doing such a fine
job at the very difficult task of microphotography. However, the shortcomings of using
photographs to illustrate genitalia are apparent in discrepancies in slide quality, mostly
brought out by different staining intensities. Some genitalia are too dark, and their details
obscured. In the aedeagus illustrations, one wonders whether visible differences in the
photos represent taxonomic differences or artifacts of preparation. Differences are even
more tenuous in female genitalia where one can only guess at the important characters.
Loss of resolution has been minimized by using contact prints, which accounts for the
large format of photographs and the fact that many are composites of two prints.

As in previous “micro’’ fascicles, the style resembles more that of a taxonomic revision
than a general manual, but this seems to be an unavoidable aspect of treating groups
where lack of previous revisionary work and large numbers of undescribed taxa preclude
more popular-style treatment.

The author is to be commended for this fine treatment of little known, small moths.
Like previous fascicles on microlepidoptera, this one should be on the shelf of any serious
student of moths, but given its price, it is hardly a manual for the general lepidopterist.

J.-F. LANpDry, Agriculture Canada, Biosystematics Research Centre, Ottawa, Ontario
K1A 0C6, Canada.

Fascicle 15.2. Pyralidae (Part), Phycitinae (Part—Acrobasis and Allies), by H. H.
Neunzig. 1986. 114 pp., 5 monochrome & 6 color pls. Softcover. $45.

This is the sixth fascicle on Pyralidae and the first to be written by H. H. Neunzig. In
this fascicle, Neunzig does an exhaustive study of the large and complex genus Acrobasis,
and allied genera Cryptoblabes, Trachycera, Anabasis, and Hypargyria. There has been
much difficulty with the identification of species in this group, in particular those of
Acrobasis. Using primarily C. Heinrich (1965, U.S. Natl. Mus. Bull. 207:1-581) as a basis,
Neunzig incorporated his own studies on biology and immature stages. In his more
comprehensive approach, he examined all available type specimens, studied in detail
male antennae, shape of male forewings, color pattern of the undersurface of wings and
thorax of males, ventral scale tufts of the eighth abdominal segment of males and females,
and male and female genitalia.

Neunzig made significant taxonomic changes, including description of three new species,
and placement of 16 new synonyms and 2 new combinations. The 38 species of Acrobasis
are divided into 10 species-groups based on adult and immature morphology and biology.
A convenient table is provided which gives host plant relations and geographic distri-
butions of the species-groups. Future taxonomic work will be facilitated also by the
designation of 18 lectotypes in this fascicle.

Accurate identification is aided by detailed species descriptions, excellent illustrations,
and keys to genera, to adults and larvae of Acrobasis, and to adults of Trachycera. Many
characters of larvae and adults are nicely shown with 61 line drawings. Included also are
drawings of larval frass tubes and pupal chambers for 17 Acrobasis species. Four mono-
chrome plates have very good scanning electron micrographs of male antennal characters.
In addition, another monochrome plate shows the black scaling of the undersurface of
the wings of six species. As with preceding fascicles of MONA, the fine color plates are
a strong contribution to this work. Neunzig shows the variation within species by using
6 color plates and 258 specimens to show 44 species photographed at twice natural size.
The specimens are in good to excellent condition (few have missing abdomens). A minor
inconvenience is the carryover of the same species to subsequent plates, probably to
economize on space and reduce costs.

I noticed one error in the text on page 11: “ZMHB” was used for the Museum Alexander

76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Humboldt, Berlin, instead of “HUMB,” the standard found in “Notes” on page vii. I did
not find “ZMHB” in type specimen data or elsewhere in text.

This fascicle will be a valuable addition to the library of those who curate collections,
and especially those who are interested in Pyralidae. Those concerned with economic
species such as the cranberry fruitworm, leaf crumpler, pecan nut casebearer, pecan leaf
casebearer, walnut shoot moth, and the birch tubemaker will find it especially useful to
have the known biological information, keys for identification, and color photographs in
one publication. Neunzig has made a significant contribution to the knowledge of Ac-
robasis and its allies through a more comprehensive approach, and is to be congratulated
on his work.

EVERETT D. CAsHatrt, Illinois State Museum, Springfield, Illinois 62706.

Journal of the Lepidopterists’ Society
43(1), 1989, 76

A TAXONOMIC REVISION OF THE NEW WorRLD MOTH GENUS PERO (LEPIDOPTERA:
GEOMETRIDAE), by Robert W. Poole. 1987. U.S. Dept. Agric., Agric. Res. Serv., Tech.
Bull. 1698. 257 pp., 1116 figs. No price given.

This work is but one of a small handful of major revisionary papers on the New World
Geometridae—in fact, for any large family of New World moths. As such, it is an
invaluable aid for determining the members of this genus, which have been in utter
taxonomic chaos. That this genus has proven to be a problem over the years is indicated
by the list of 10 generic synonyms given, with 6 being placed in synonymy in this paper.

Pero is one of the largest genera in Ennominae; it makes up, by far, the largest portion
of the Azelini. Members are restricted to the New World, and occur almost everywhere
except in the far northern and southern regions. Pero includes 294 species, of which Poole
described 119 as new, and there are 74 junior synonyms for the genus. (One omission is
the four subspecific names I proposed in my 1955 paper on this genus in western North
America, even though my paper is cited in the text.) With this many species, it is not
surprising that there are some that exhibit sexual dimorphism, polymorphism, extreme
geographic variation, and a high degree of individual variation. This means that genitalic
dissections are often necessary to place the correct name on a species; in fact, I prefer to
base determinations on study of genitalia rather than pattern and color of an individual
specimen.

This work is a condensation of Poole’s doctoral thesis. Descriptions have been reduced
to diagnoses, as the author uses them to supplement illustrations of the adults (photographs)
and genitalia (drawings). One item I believe should have been included is length of
forewings, as specimens range from about 10 to nearly 30 mm; there is no indication in
text or photographs, as to specimen size. Each species has a listing of localities for the
specimens examined.

For anyone interested in New World moths, especially the Geometridae, this paper is
a necessary addition to his or her library.

FREDERICK H. RINDGE, Department of Entomology, American Museum of Natural
History, New York, New York 10024.

Journal of the Lepidopterists’ Society
43(1), 1989, 77-78

IMMATURE INSECTS, Volume 1, Frederick W. Stehr (ed.). 1987. Kendall-Hunt, Dubuque,
Iowa. 754 pp. Quarto. Hard cover. $69.95.

Had the finger of zoological fate pointed to larvae as the “perfect’’ insect stage instead
of adults, entomology and lepidopterology might be different today. The applied branch,
so concerned with larvae, might include systematics; the Code might outlaw adults for
naming purposes; dermestids might matter less; and visionaries like Alvah Peterson, and
now Fred Stehr, might be drawing our attention to the neglected adult stage instead of
to the neglected larval stage.

A decade in the making, Immature Insects doubtless had its origin in the summer of
1957 when Stehr took a field course in immatures from Alvah Peterson at the Itasca
Biological Station of the University of Minnesota. Stehr and a generation of Ohio State
University students taking Peterson’s immature insects course (including me) duly keyed
the collected or prescribed material, but sometimes without relish. The main tool then
was Peterson’s Larvae of Insects, a plesiomorphic ancestor; Stehr and company’s Im-
mature Insects is an apomorphic, streamlined descendant.

Volume 1 of Immature Insects deals with 24 orders, but is dominated by Lepidoptera,
to which more than 300 pages are devoted; when supporting sections are considered,
easily half of this big book concerns Lepidoptera. The Lepidoptera section, coordinated
by general editor Stehr, contains contributions by 19 specialists, a number large enough
to greatly thin the ranks of candidates to review the book. Volume 2 of Immature Insects,
covering the 10 remaining orders, including worldwide coverage of Coleoptera, should
appear in 1989.

The book’s focus is larvae rather than eggs or pupae. Additional sections of integral
interest to lepidopterists include Introduction (6 pages), Techniques for Collecting, Rear-
ing, Preserving, and Studying Immature Insects (12 pages), Key to Orders of Immature
Insects and Selected Arthropods (28 pages), Glossary (11 pages), Host Plant and Substrate
Index (6 pages), and the overall Index (26 pages). The key to orders includes immature,
brachypterous, and wingless adult insects as well as other terrestrial and freshwater
invertebrates that might be confused with immature insects.

The Lepidoptera section provides coverage basically to family level. Treated families
number 75 whose larvae occur north of Mexico. Classification largely follows the 1983
Check List. Backbone of the section is the dichotomous key to families developed by
Stehr and P. J. Martinat. With a whopping 225 couplets, the key is essentially in two
parts; it may be entered at couplet 39 for larvae with normal numbers of thoracic legs
and prolegs. By comparison, Peterson’s three-part key had 99 couplets. The authors did
not attempt to make the key reflect phylogeny or reveal all family characters.

Refreshingly, the key was designed with uncommon consideration for the user. It is
sprinkled with helpful hints, reminders, cautions (“may be very small, look carefully”),
as well as italic and boldface type for extra emphasis (“... distinctly closer.”’). It suc-
cessfully decreases conditional statements, the bane of many a key user: there is 1 such
statement per 9 couplets compared with Peterson’s 1 per 6, and the ones left are not very
convoluted.

The new and larger key resulted from use of more characters as well as decomposition
of complex couplets. The authors rightly claim that key paths are not necessarily longer:
I found the ratio of number of key-out points to number of couplets to be high, 227/225
or 1.00 compared to Peterson’s 100/99 or 1.01. The authors say it is possible to make
wrong choices in the key and still arrive at the correct family. Such robustness is likewise
supported by my checking: there are more key-out points for five diverse moth and
butterfly families than in Peterson’s key: 8 vs. 2 for Noctuidae; 4 vs. 1 for Pyralidae; 3
vs. 1 for Geometridae; 7 vs. 5 for Nymphalidae (broad sense); and 6 vs. 1 for Lycaenidae.
Most key characters are illustrated with serviceable line drawings which occupy part of
every page of the key for ready accessibility. The ratio of such illustrations to number
of key couplets exceeds 0.80. When introductory illustrations are added, the ratio becomes
0.95; the introductory illustrations are part of some 15 pages of larval description preceding
the key that thoroughly review external anatomy, and include setal maps as well as
chaetotaxic tables.

78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Following the key, every family is individually discussed. These discussions are uni-
formly presented under the headings Relationships and Diagnosis, Biology and Ecology,
Description, Comments, and Selected Bibliography. Most are accompanied by instructive
line drawings and photographs ranging from structural details to whole larvae. The
Selected Bibliographies provide easy entry to pertinent literature for each family. The
family information is up to date and insightful, a result of the specialist expertise of the
various authors. I found the family discussions an unexpected highlight of the book; they
form an encyclopedic source of current information on North American Lepidoptera.

There are no keys to genera of Lepidoptera. However, for Noctuidae, Pyralidae, and
Tortricidae, three of the five most speciose families, there are keys to selected species.
The pyralid keys treat stored product and corn-sugarcane pests; the tortricid keys treat
pine feeders, soybean-alfalfa-cultivated legume feeders, and pome-fruit feeders; and the
noctuid key, representative last instars. The value of such keys seems equivocal to me.
At worst, they mislead the unwary; at best, they provide a starting point from which
comprehensive keys can be built. Fortunately, the keyed species of Pyralidae and Noc-
tuidae are illustrated or described to help confirm key results.

Physically, the big green book is sturdily manufactured and attractively designed. An
eye-catching color photograph of a limacodid caterpillar adorns the front cover.

Immature Insects delivers a solid background for an interest in lepidopterous and
other larvae; I venture it will also inspire much new interest in larvae. It will surely build
a following among a new and more demanding generation of students and devotees.
Hardly anyone could fail to get a good lepidopterological return on its purchase price.

WILLIAM E. MILLER, Department of Entomology, University of Minnesota, St. Paul,
Minnesota 55108.

Journal of the Lepidopterists’ Society
43(1), 1989, 79

FEATURE PHOTOGRAPH

A rare and perhaps unique interfamily mating between a female Glaucina erroraria
Dyar.(Geometridae) (upper) and a male Protorthodes melanopis (Hampson) (Noctuidae)
(lower) shown 2.8x natural size. Taken in copulo at 5131 Bannock St., Pueblo del Sol,
Huachuca Mts., Cochise Co., Arizona, at UV light, 25 March 1988. Collected and mounted,
still attached as shown, by Ronald S. Wielgus of the above address. Specimens deposited
in the National Museum collection under G. erroraria. Identified and photographed by
Douglas C. Ferguson, Systematic Entomology Laboratory, U.S. Dept. Agric., % U.S.
National Museum of Natural History, Washington, D.C. 20560.

Date of Issue (Vol. 48, No. 1): 8 March 1989

s
un)
-
p iy

EDITORIAL STAFF OF THE JOURNAL

BoycE A. DRUMMOND, Editor WILLIAM E. MILLER, Retiring Editor
Natural Perspectives Dept. of Entomology
P.O. Box 9061 University of Minnesota
Woodland Park, Colorado 80866 U.S.A. St. Paul, Minnesota 55108 U.S.A.

Associate Editors and Editorial Committee:
_ M. DEANE BOWERS, BOYCE A. DRUMMOND III, DOUGLAS C. FERGUSON,
LAWRENCE F. GALL, ROBERT C. LEDERHOUSE, THOMAS A. MILLER,
THEODORE D. SARGENT, ROBERT K. ROBBINS

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories
are Articles, General Notes, Technical Comments, Book Reviews, Obituaries, Feature
Photographs, and Cover Illustrations. Journal submissions should be sent to the editor at
the above address. Short manuscripts concerning new state records, current events, and
notices should be sent to the News, June Preston, Editor, 832 Sunset Drive, Lawrence,
Kansas 66044 U.S.A. Journal contributors should prepare manuscripts according to the
following instructions, and submit them flat, not folded.

Abstract: An informative abstract should precede the text of Articles.

Key Words: Up to five key words or terms not in the title should accompany Articles,
General Notes, and Technical Comments.

Text: Manuscripts should be submitted in triplicate, and must be typewritten, entirely
double-spaced, with wide margins, on one side only of white, letter-sized paper. Titles
should be explicit, descriptive, and as short as possible. The first mention of a plant or
animal in the text should include the full scientific name with author, and family.
Measurements should be given in metric units; times in terms of the 24-hour clock (0930 h,
not 9:30 AM). Underline only where italics are intended.

Literature Cited: References in the text of Articles should be given as Sheppard (1959)
or (Sheppard 1959, 1961a, 1961b) and listed alphabetically under the heading LITERATURE
CITED, in the following format without underlining:

SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London.
209 pp.

196la. Some contributions to population genetics resulting from the study of

the Lepidoptera. Adv. Genet. 10:165-216.

In General Notes and Technical Comments, references should be shortened and given
entirely in the text as P. M. Sheppard (1961, Adv. Genet. 10:165-216) or (Sheppard, P.
M., 1961, Sym. R. Entomol. Soc. London 1:23-80) without underlining.

Illustrations: Only half of symmetrical objects such as adults with wings spread should
be illustrated, unless whole illustration is crucial. Photographs and drawings should be
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secutively in Arabic numerals; “plate” should not be employed. Figure legends must be
typewritten, double-spaced, on a separate sheet (not attached to illustrations), headed
EXPLANATION OF FIGURES, with a separate paragraph devoted to each page of illustrations.

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for
tables should not be capitalized. Tabular material must be typed on separate sheets, and
placed following the main text, with the approximate desired position indicated in the
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Voucher specimens: When appropriate, manuscripts must name a public repository
where specimens documenting identity of organisms can be found. Kinds of reports that
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Proofs: The edited manuscript and galley proofs will be mailed to the author for
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PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A.

CONTENTS

SUNDRY ARGYNNINE CONCEPTS REVISITED (NYMPHALIDAE). L.
Paul Grey oe Oe ee

PHYLOGENY AND ZOOGEOGRAPHY OF THE BIGGER AND BETTER
GENUS ATALOPEDES (HESPERIIDAE). John M. Burns ...... ef.

TERRITORIAL BEHAVIOR AND DOMINANCE IN SOME HELICONIINE
BUTTERFLIES (NYMPHALIDAE). Woodruff W. Benson, Célio
F, B. Haddad & Mércio Zikan 2... eee

REPRODUCTIVE TRACT DEVELOPMENT IN MONARCH BUTTERFLIES
OVERWINTERING IN CALIFORNIA AND MExiIco. William S.
Herman, Lincoln P. Brower ¢ William H. Calvert ...........

INSTAR NUMBER AND LARVAL DEVELOPMENT IN LYCAENA PHLAEAS
HYPOPHLAEAS (BOISDUVAL) (LYCAENIDAE). Gregory R.
Ballmer & Gordon F. Pratt 00 0

GENERAL NOTES

Status of the papilionid types Papilio stewarti Avinoff and P. morrisi Ehr-
mann. Kurt Johnson, David Matusik d+ Rick POZY CK: oc.ccccccccccccsssscceeeeneene

Repeated copulation in an orange hairstreak, Shirozua janasi: A case of mate
guarding? Hisashi Fujtt (0000

An emended specific name in Eupithecia (Geometridae). Frederick H.
Rimd ge iain PU OI UN EON ON a a a

The valid generic placement for “Calothysanis” amaturaria (Walker) (Geo-
metridae, Sterrhinae). Charles V. Covell Jr...)
Book REVIEWS

The moths of America North of Mexico (three fascicles). Charles V. Covell
Jr., J.-F. Landry & Everett D. Cashatt

A taxonomic revision of the New World moth genus Pero (Lepidoptera:
Geometridae). Frederick H. Rindge

Immature insects, Vol. 1. William E: Miller’ ee

FEATURE PHOTOGRAPH
Douglas C. Ferguson

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

59

66

Volume 43 1989 Number 2

ISSN 0024-0966

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

18 May 1989

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

JULIAN P. DONAHUE, President GEORGE W. GIBBs, Vice
JERRY A. POWELL, Immediate Past President

President RONALD LEUSCHNER, Vice
JOHN N. ELIOT, Vice President President
RICHARD A. ARNOLD, Secretary JAMES P. TUTTLE, Treasurer

Members at large:

JOHN W. BROWN M. DEANE BOWERS FREDERICK W. STEHR
MOocGENS C. NIELSEN RICHARD L. BROWN JOHN E. RAWLINS
FLOYD W. PRESTON PAUL A. OPLER Jo BREWER

EDITORIAL BOARD

PAUL A. OPLER (Chairman), FREDERICK W. STEHR (Member at large)
Boyce A. DRUMMOND (Journal), WILLIAM E. MILLER (Memoirs), JUNE PRESTON (News)

The object of the Lepidopterists’ Society, which was formed in May 1947 and for-
mally constituted in December 1950, is “to promote the science of lepidopterology in all
its branches, .... to issue a periodical and other publications on Lepidoptera, to facilitate
the exchange of specimens and ideas by both the professional worker and the amateur
in the field; to secure cooperation in all measures’ directed towards these aims.

Membership in the Society is open to all persons interested in the study of Lepi-
doptera. All members receive the Journal and the News of the Lepidopterists Society.
Institutions may subscribe to the Journal but may not become members. Prospective
members should send to the Treasurer full dues for the current year, together with their
full name, address, and special lepidopterological interests. In alternate years a list of
members of the Society is issued, with addresses and special interests. There are four
numbers in each volume of the Journal, scheduled for February, May, August and
November, and six numbers of the News each year.

Active members—annual dues $25.00
Student members—annual dues $15.00
Sustaining members—annual dues $35.00
Life members—single sum $500.00
Institutional subscriptions—annual $40.00

Send remittances, payable to The Lepidopterists’ Society, to: James P. Tuttle, Treasurer,
3838 Fernleigh Ave., Troy, Michigan 48083-5715, U.S.A.; and address changes to: Julian
P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, California
90007-4057 U.S.A. For information about the Society, contact: Richard A. Arnold, Sec-
retary, 50 Cleaveland Rd., #3, Pleasant Hill, California 94523-3765, U.S.A.

For back issues of the Journal and News, write the Publications Coordinator at the
address below about availability and prices. Prices of The Lepidopterists’ Society com-
memorative volume 1945-19783 are $12.00 ($8.00 to members and subscribers); of A
catalogue/checklist of the butterflies of America north of Mexico, clothbound, $19.00
($12.00 to members and subscribers), paperbound, $10.50 ($7.00 to members and sub-
scribers). Order from the Publications Coordinator, Ronald Leuschner, 1900 John St.,
Manhattan Beach, California 90266-2608, U.S.A. Make remittance payable to “The Lep-
idopterists’ Society.”

Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for
$40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’
Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los
Angeles, California 90007-4057. Second-class postage paid at Los Angeles, California and
additional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’
Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, California 90007-
4057.

Cover illustration: Larvae of the small-eyed sphinx, Paonias myops (J. E. Smith) (Sphin-
gidae), resting “leaf-like” on a shoot of black cherry, Prunus serotina Ehrh. (Rosaceae).
Submitted by Gerald P. Wykes, 2569 Reinhardt Road, Monroe, Michigan, 48161.

a

JouRNAL OF
Tue LeEpipoprTrerRistTs’ SOCIETY

Volume 43 1989 Number 2

Journal of the Lepidopterists’ Society
43(2), 1989, 81-92

ELECTROPHORETIC COMPARISONS OF VICARIANT
VANESSA: GENETIC DIFFERENTIATION BETWEEN
V. ANNABELLA AND V. CARYE (NYMPHALIDAE)
SINCE THE GREAT AMERICAN INTERCHANGE

ARTHUR M. SHAPIRO AND HANSJURG GEIGER!
Department of Zoology, University of California, Davis, California 95616

ABSTRACT. Vanessa carye and V. annabella are very similar species found in South
America and North + Central America, respectively; they probably differentiated in the
three million years since the Great American Interchange. Electrophoretically they are
differentiated at a level typical of animal morphospecies (Nei’s I = 0.855, D = 0.157)
and are much more unlike than small samples of V. cardui from California vs. France.
Using the Sarich method of estimating time of divergence, we date their speciation at
roughly 2.97 million years ago, suggesting that Vanessa was an early crosser of the Panama
land bridge. Our results support continued recognition of V. carye and V. annabella at
the species, rather than the subspecies, level.

Additional key words: systematics, biogeography.

Since the emergence of protein electrophoresis as a technique in
population genetics, it has been applied widely in systematics as well
(Burns 1975, Ayala 1983). Several attempts have been made to compare
levels of electrophoretic differentiation to conventional (morphologi-
cally-based) taxonomic judgment (Avise 1974, Ayala 1983, Ayala &
Powell 1972, Ayala et al. 1974, Nevo et al. 1974, Mickevich & Johnson
1976, Thorpe 1982). Burns and Johnson (1967) first suggested that
enzyme variation might offer a powerful tool for recognizing sibling
species; Webster and Burns (1973) demonstrated its value in a pioneer-
ing study with lizards. Despite the widespread use of electrophoresis
in systematic investigations of other taxonomic groups, it has seldom
been brought to bear on butterflies (Geiger & Scholl 1985). The present
study attempts to resolve the status of two putative vicariant mor-

‘ Present address: Zoologisches Institut, Universitat Bern, Baltzerstrasse 3, Bern CH 3012, Switzerland.

82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

phospecies of the genus Vanessa Fabricius (Nymphalidae) by electro-
phoretic means, and interprets the data in historical-biogeographical
terms to estimate the antiquity of their speciation event.

Hamadryas carye was described by Hibner in 1812 with no specific
type locality. The name was subsequently applied to both North and
South American, superficially similar populations later placed in the
genus Vanessa. As early as 1951, W. D. Field had noted color and
pattern differences between North and South American specimens,
which he communicated personally to his Chilean collaborator J. Her-
rera. Herrera et al. (1958) then asserted that “‘after studying the genitalia
of the examples which we possess from the United States (Oregon and
California), Mexico, Argentina and Chile we are able to affirm that we
are dealing with two quite different species.’”” Once Field was able to
establish from Hiibner’s figure that his (lost) type must have been South
American, it was now possible to fix that usage; Field (1971) named
the (newly-nameless) North American entity Cynthia annabella. Al-
though his generic judgment has not been generally accepted, the spe-
cific epithet annabella continues in use for material from Central Amer-
ica northward.

Vanessa carye, sens. str.,and V. annabella have sufficient phenotypic
differences (in both habitus and genitalia) that if they co-occurred
without intergrading there would be no hesitation in calling them dif-
ferent, though very closely related, species. However, they are appar-
ently completely allopatric; carye ranges from southern Patagonia to
Colombia, annabella from British Columbia to Guatemala. Neither
species is recorded from montane or lowland Costa Rica (DeVries 1986).
Such allopatric sister-species were called “‘vicars’” by Udvardy (1969)
and are commonly known as “‘vicariants’’ or “vicariant species” in the
literature; they are often considered to be relatively recently-differ-
entiated. In the absence of genetic data, and sometimes in the presence
of such data, taxonomists’ judgments as to how to rank such entities are
often controversial. Thus, the suggestion by Higgins and Riley (1970)
that several Palearctic-Nearctic pairs of pierid taxa were conspecific
has remained unresolved despite laboratory hybridizations and elec-
trophoretic studies (Shapiro 1980, Shapiro 1983, Shapiro & Geiger 1986
for Pontia, in which compatibility studies were done between popu-
lations far-removed from one another on the alleged Holarctic cline).
In Vanessa carye and annabella there is near-unanimity in usage; doubts
as to the validity of a species-level distinction have remained largely
unpublished, appearing only in one major work (Scott 1986:283—284
treats them as subspecies). Such doubts are sure, however, to be exac-
erbated by the recent demonstration by Herrera (1987) of wing-pattern
overlap between the taxa, and his forthcoming publication of laboratory

ERRATUM

A printing error resulted in the omission of the numbers on Figures 1-18 in John M.
Burns article, “Phylogeny and zoogeography of the bigger and better genus Atalopedes
(Hesperiidae), which appeared in Volume 43, Number 1 of the Journal of the Lepi-

dopterists’ Society, pp. 11-32. The original version of this figure is reprinted here. To
correct this error in your copy, cut around the figure below, peel off the back, and affix
it above the caption on page 14 of Volume 43, Number 1.

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VOLUME 43, NUMBER 2 83

hybridization data (Herrera pers. comm.). Although we had only limited
local samples available, we considered it worthwhile to attempt to assess
the level of electrophoretic differentiation between the taxa. Since there
are no published taxic comparisons within the Nymphalini, we at-
tempted to place the data in context by comparing these two entities
to members of different species-groups in Vanessa, to European vs.
American V. cardui L., and to a few other nymphalines to which we
had access at the time the study was done. Once in hand, the data
permit a very crude estimate of the time since gene flew was inter-
rupted, that is, the time of speciation—an estimate which is particularly
interesting in cases such as this one, in which very different models of
the history and biogeography of the situation may be advanced.

MATERIALS AND METHODS

The sources of our samples are listed in Table 1. All animals were
collected from the field, transported alive and immediately stored at
—70°C until electrophoresis. Only autumn 1985 through 1986 catches
were used, except for European V. cardui; we had only a handful of
old frozen specimens (1979) of these, but they had conserved most of
their activity such that the zymograms were completely satisfactory for
comparison with recent American material. All wings were retained
by HJG. The head and thorax of each butterfly were homogenized in
four volumes of Tris-HCl buffer (0.05 M, pH 8.0). We used horizontal
starch gel electrophoresis procedures slightly modified from Ayala et
al. (1972) (Geiger 1981). Twenty-two enzymes were scored:

adenylate kinase (AK-1, AK-2) hexokinase (HK)

aldolase (ALD) indophenol oxidase (IPO)

arginine kinase (APK) isocitrate dehydrogenase (IDH-1, IDH-2)
fumarase (FUM) malate dehydrogenase (MDH-1, MDH-2)
glutamate-oxaloacetate transaminase malic enzyme (ME-1, ME-2)

(GOT-1, GOT-2) phosphoglucomutase (PGM)
glutamate-pyruvate transaminase (GPT) 6-phospho-gluconate dehydrogenase
glyceraldehyde-phosphate dehydrogenase (6-PGD)

(GAPDH) phosphoglucose isomerase (PGI)
a-glycerophosphate dehydrogenase pyruvate kinase (PK)

(a-GPDH)

There are no studies known to us of the heredity of any of these loci
in Nymphalini, and we made the usual assumption by treating elec-
tromorphs as alleles. “Allelic” distributions were generally in good ac-
cord with Hardy-Weinberg expectations in samples large enough to
warrant such a test. The most frequent “allele” in carye was arbitrarily
given the standard index 100 in all cases; electromorphs with different
mobilities are designated with relation to it, such as an “allele 105” for
an enzyme that migrates 5 mm faster than the commonest carye allelic
product.

84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Localities and dates of samples. Altitudes are given only for mountain
samples.

Vanessa carye, sens. str.

ARGENTINA: Prov. Salta: Abra Molina, 4000 m, i.2.86 (n = 3); Valle Encantado, 2700
m, i.22.86 (n = 2); Salta, i.22.86 (n = 2). Prov. Tucuman: San Javier, i.18.86 (n = 8);
Abra Infiernillo, 3300 m, i.20.86 (n = 1); Tafi del Valle, 2100 m, i.28-27.86 (n = 7);
San Miguel de Tucuman, i.29-iii.9.86 (n = 5 + 1 reared ex Sida).

V. annabella

CALIFORNIA, USA: Siskiyou Co.: Ball Mt., 2200 m, viii.28.86 (n = 7); Yolo Co.: Da-
vis, ix.7.86 (n = 10); Solano Co.: Suisun City, ii.6.86 (n = 1), Fairfield, ii.6.86 (n =
2); Nevada Co.: Donner Pass, 2100 m, ix.25.85 (n = 9), Lang Crossing, South Yuba
River, 1750 m, ix.25.85 (n = 1).

V. cardui

CALIFORNIA, USA: Nevada Co.: Donner Pass, ix.4.86 (n = 5). FRANCE: Dept. Vau-
cluse: Bolléne, vi.4.79 (n = 1); Dept. Bouches du Rhéne: Le Grau du Roi, vi.4.79 (n
= 1); Dept. Herault: Oppidium d’Enserune, vi.2.79 (n = 1).

V. virginiensis
CALIFORNIA, USA: Nevada Co.: Donner Pass, ix.4.86 (n = 8).
Polygonia zephyrus W. H. Edwards
CALIFORNIA, USA: Nevada Co.: Donner Pass, ix.25.85 (n = 2).
Nymphalis milberti Godart
CALIFORNIA, USA: Nevada Co.: Donner Pass, ix.25.85 (n = 2).

The statistic I (Nei 1972) was used to estimate genetic similarity
between samples over all loci. Calculated I values were then used to
construct a dendrogram (Fig. 1) by cluster analysis (UPGMA method,
Ferguson 1980). Because the set of loci is very similar and the same
statistic has been used, direct comparisons may be made to earlier studies
from our laboratories (Geiger & Scholl 1985 for example), while com-
parisons to others must be made with more caution. For estimating
time of divergence of the taxa, a different statistic (D or I, not I) was
employed, as explained below.

RESULTS

There are genetic differences at several loci between Vanessa carye
and V. annabella (Tables 2, 3). At most loci they consist of moderately
to strongly divergent allelic frequencies; at only one locus (HK) is there
an apparent fixed difference. In sympatry these data would be un-
equivocal evidence for speciation. In allopatry they must be compared
to similar data for entities in other groups, using the same statistic and
more or less similar procedures, to determine what constitutes a “species-
level’ difference. It is now well-established that some groups are much
more conservative electrophoretically than others, and that the relation

85

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86 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

between rates of morphological and electrophoretic differentiation can
be extrapolated among taxa only with great caution. The classic study
of the Drosophila willistoni (Diptera: Drosophilidae) complex by Ayala
et al. (1974) established I values of 0.970 for conspecific, geographic
populations; 0.795 for subspecies; 0.873 for semispecies; 0.517 for sibling
species and 0.352 for morphospecies (recalculated from the original
figures, which were given for Nei’s I). The corresponding values for
the same taxonomic levels are considerably higher in pierid butterflies,
a very conservative group at the level of electrophoretic genetics (Geiger
1981, Geiger & Scholl 1985, Shapiro & Geiger in prep.). Thus, within
the genus Pieris, sens. lat., the European and Japanese subspecies of
Pieris rapae L. cluster at 0.989; these with the morphospecies P. mannii
Mayer at 0.902; these three with P. canidia L. at 0.874; the European
and North American groups of “napi’’-taxa with each other at 0.748
and the napi and rapae species-groups in toto at 0.546 (23 loci). The
I value for V. carye and V. annabella, being in the mid-0.8 range,
would indicate very well-differentiated species in Pieris and in Pieridae
generally, but only infraspecific status in the D. willistoni group.

By Field’s (1971) classification, the other two Vanessa used in this
study (cardui and virginiensis Drury) belong to different species-groups
(or splitter’s genera). Thus the degree of differentiation in the dendro-
gram (Fig. 1) is not surprising. The lack of differentiation between
Californian and French V. cardui mirrors their phenotypic similarity
but is still somewhat surprising, especially given the small samples which
would tend to amplify any differences purely probabilistically. Vanessa
cardui is migratory in both Europe and America, with a huge summer
breeding range (whence come our samples from both continents) but
a much smaller overwintering one. This situation would tend to swamp
out any tendency to local population differentiation, as in the Monarch,
Danaus plexippus L. (Danaidae) (Eanes & Koehn 1979, Kitching 1985).
But migration between Europe and America is neither known nor
suspected for Vanessa cardui; nor is it a recent introduction in North
America—at least Boisduval (1868) and Scudder (1889) treat it as native
on the Pacific and Atlantic coasts, respectively. The possible stability
of gene frequencies over its vast range deserves further study.

Both Vanessa carye and V. annabella are highly vagile, though
neither is documented as a seasonal mass-migrant as is V. cardui. Our
samples are drawn in both cases from more or less contiguous lowland
and montane sites. There are hints in both species (Shapiro unpubl.) of
a disorganized, individual altitudinal migration in mountainous terrain,
tracking the seasonal availability of hosts. The virtual identity between
nearby high- and low-elevation populations is not surprising. The bi-
ology of V. carye in Argentina is largely unpublished, but like V.

VOLUME 43, NUMBER 2 87

annabella carye
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9 ee eee er gens
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Fic. 1. Dendrogram illustrating clustering among three populations of Vanessa an-
nabella, two of V. carye, and small samples of V. cardui, V. virginiensis, Polygonia
zephyrus and Nymphalis milberti, using Nei's statistic I.

annabella it is a “weedy,” often urban species, and its behavior is nearly
identical to V. annabella. There is very pronounced rainfall seasonality
at Tafi del Valle, while hosts are available all year at San Miguel de
Tucuman. Other butterflies—several Pierini and Coliadini at least—
appear to undergo regular seasonal up- and downslope movements in
the Province of Tucuman (Shapiro & R. Eisele pers. obs.).

DISCUSSION

Instances in which speciation can be associated with a specific geo-
historical event afford the opportunity to time-calibrate rates of bio-
chemical evolution in particular groups, which is intrinsically superior
to existing procedures for estimating time of divergence from genetic
similarity or distance data (Nei 1971, Sarich 1977, Carlson et al. 1978,
Thorpe 1982, Menken 1982). When biogeography suggests a specific
time of speciation, this can be cross-checked using these procedures;
agreement does not necessarily validate the scenario, nor disagreement
falsify it, but such results are always suggestive.

88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The ranges of V. carye and V. annabella can be viewed as products
of strict vicariance (a formerly continuous range divided by the ap-
pearance of a barrier) or of dispersal followed by differentiation (the
classic geographic-speciation model). Herrera (1987) takes a vicariance
position and attributes this case to continental drift, the “vicariance
event” leading to speciation being the breakup of Pangaea. Such a
scenario makes the last common ancestor of carye and annabella as old
as the Triassic (some 200 million years ago), which seems unlikely for
many reasons. However, the geography of this case strongly suggests
dispersal across the Isthmus of Panama during the Great American
Interchange which commenced roughly three million years ago when
that corridor emerged, and which is very thoroughly documented for
mammals (Marshall 1988, Stehli & Webb 1985). It resulted in coloni-
zation of each continent by faunal elements from the other, with a
higher percentage of successful colonizations from North into South
America than the reverse. Until fairly recently there was a tendency
to attribute virtually all High Andean occurrences of otherwise Hol-
arctic groups to this event (compare Mani 1968), regardless of the
amount of evolutionary differentiation observed in the Andean biota
(which by this view must have occurred since the Interchange). A
consensus is now emerging to the effect that insects have evolved more
slowly than mammals, at least in the Quaternary (Brown 1982, Coope
1978, 1979; D. W. Jenkins & L. D. Miller pers. comm.)—such that
evolutionary origin of taxa above the species level in the Quaternary
seems unlikely in Lepidoptera. Indeed, most butterfly evolution in the
Quaternary seems to have been at the subspecies level, despite great
geoclimatic dynamism. We suspect that the level of differentiation
shown by V. carye and V. annabella, if fairly represented here, lies
near the high end of the range to be expected once many candidates
for Quaternary trans-Isthmian differentiation have been investigated.

The Panama land bridge was not only a corridor for migration and
colonization by terrestrial organisms; it also formed a barrier to marine
ones at the same time (Woodring 1966), and several speciation events
have been attributed to it as a result. The genetic differentiation of
sister species of marine organisms in the tropical eastern Pacific vs. the
Caribbean has been quantified and cross-checked using dating estimates
from electrophoretic data (Lessios 1979, 1981, Vawter et al. 1980).
There is no reason in principle why the same should not be possible
for terrestrial species. Like Vawter et al., we used Sarich’s (1977) pro-
cedure, as modified by Carlson et al. (1978), to convert Nei’s distance
measure for V. carye and V. annabella (I = 0.855, D = 0.157) to an
estimate of time of divergence, which is 2.97 million years (discussion
of significant figures below). Such estimates entail many assumptions

VOLUME 438, NUMBER 2 89

and should not be taken unduly seriously, even when they give re-
markably close agreement to estimates derived from biogeography.
[Sarich’s method was developed using vertebrate data, and we are aware
of the dangers in extrapolating among taxonomic groups—as were
Vawter et al. (1978)]. But this number is in fact very consistent with
speciation consequent on the Great American Interchange and with an
early dispersal across Panama, perhaps even before a continuous land
corridor was available. What is most important is that it is wildly
inconsistent with Herrera’s (1987) invocation of the breakup of Pangaea,
250 to 100 million years ago depending on how far the animals could
still disperse over water.

It is premature to state the direction of dispersal before a careful
phylogenetic analysis of Vanessa is completed. There are more species-
groups represented in North than South America, but more species on
the latter continent. Herrera (1987) provides no explicit rationale for
his claim that “The origin of carye is indubitably in Gondwanaland.”’

We have successfully resisted the temptation to generate scenarios
for the history of V. carye and annabella, such as the proximate cause
of the interruption of gene flow after invasion of one continent from
the other, or for their failure to re-establish contact in montane Central
America. Such exercises of the imagination are not in any sense testable
with the tools used in this investigation.

Summing over many studies, Thorpe (1982) concludes that in general,
“If allopatric populations of dubious status have genetic identities below
about 0.85 it is improbable that they should be considered conspecific,
while nominate species with I values above 0.85 should be considered
doubtful if there is no other evidence of their specific status.’”’ He goes
on to chide geneticists for violating common sense and the rules of
significant figures by treating three-digit decimal I values as givens.
Thorpe’s rule of thumb for species status is inappropriate for Pierini
but may be appropriate in Nymphalini and various other butterflies;
time (and more studies) will tell. Nymphalini seem to undergo very
slow morphological differentiation: Nearctic and Palearctic populations
of Nymphalis and Vanessa species do not differ phenotypically; the
genera are so uniform morphologically that generic splitting and lump-
ing are a chronic problem in the group; even different genera show
homologous responses to temperature shock during development (Sha-
piro 1984); and an Oligocene fossil attributed to Vanessa by Miller and
Brown (1988) demonstrates morphological near-stasis over geologic time.
On the other hand, Nymphalini seem to be more normal animals elec-
trophoretically than Pierini are, that is, more labile, at least to judge
by our work.

We agree with Thorpe’s comments on significant figures; slight dif-

90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ferences in electrophoretic data must be interpreted reasonably in the
context of overall patterns of variation in the group, and calculations
such as Sarich’s estimator—one more manipulation removed from real-
ity—must be used with still more caution. When one is primarily in-
terested in orders of magnitude, as we are here, they are still quite
valuable. Although our findings are very preliminary and we are cog-
nizant of the limitations of our study, including small sample sizes and
the use of samples from arbitrary locations within very large ranges,
we are pleased with the outcome—and still comfortable with Field’s
decision to treat Vanessa annabella as a species distinct from V. carye.

ACKNOWLEDGMENTS

We thank F. J. Ayala and A. Scholl for advice and for permitting use of their facilities;
R. Eisele (Tucum4an, Argentina) and A. H. Porter (Davis) for assistance, J. Herrera G. for
sharing his unpublished data with us, and NSF grant BSR-8306922 (Systematic Biology
Program, to AMS) for supporting HJG’s work in Davis. This study forms part of California
Agricultural Experiment Station Project CA-D*-AZO-3994-H, “Climatic Range Limita-
tion of Phytophagous Lepidopterans” (AMS, Principal Investigator).

LITERATURE CITED

AVISE, J. C. 1974. Systematic value of electrophoretic data. Syst. Zool. 23:465—481.

AYALA, F. J. 1983. Enzymes as taxonomic characters, pp. 3-26. In Oxford, G. S. & D.
Rollinson (eds.), Protein polymorphism: Adaptive and taxonomic significance. Aca-
demic Press, New York.

AYALA, F. J. & J. R. POWELL. 1972. Allozymes as diagnostic characters of sibling species
of Drosophila. Proc. Natl. Acad. Sci. U.S.A. 69:1094-1096.

AYALA, F. J., J. R. POWELL, M. L. TRACEy, C. A. MOURAO & S. PEREZ-SALAS. 1972.
Enzyme variability in the Drosophila willistoni group. IV. Genetic variation in natural
populations of D. willistoni. Genetics 70:113-139.

AYALA, F. J.. M. L. TRACcEy, D. HEDGECOCK & R. C. RICHMOND. 1974. Genetic
differentiation during the speciation process in Drosophila. Evolution 28:576-592.

BOISDUVAL, J. B. A. 1868. Lepidoptéres de la Californie. Ann. Soc. Entomol. Belg. 12:
o-94.

Brown, K. S., JR. 1982. Historical and ecological factors in the biogeography of apo-
sematic Neotropical butterflies. Am. Zool. 22:453—471.

BurRNsS, J. M. 1975. Isozymes in evolutionary systematics, pp. 49-62. In Markert, C. L.
(ed.), Isozymes IV. Genetics and evolution. Academic Press, New York.

BURNS, J. M. & F. M. JOHNSON. 1967. Esterase polymorphism in natural populations
of a sulfur butterfly, Colias eurytheme. Science 156:93-96.

CARLSON, S. S., A. C. WILSON & R. D. MAxson. 1978. Do albumin clocks run on time?
A reply. Science 200:1183-1185.

Coope, G. 1978. Constancy of insect species versus inconstancy of Quaternary envi-
ronments, pp. 176-187. In Mound, L. A. & N. Waloff (eds.), Diversity of insect
faunas. Blackwell, Oxford.

1979. Late Cenozoic fossil Coleoptera: Evolution, biogeography, and ecology.
Ann. Rev. Ecol. Syst. 10:247-267.

DEVRIES, P. J. 1986. The butterflies of Costa Rica and their natural history: Papilionidae,
Pieridae, Nymphalidae. Princeton Univ. Press, Princeton, New Jersey. 327 pp.
EANES, W. F. & R. K. KOEHN. 1979. An analysis of genetic structure in the monarch

butterfly, Danaus plexippus L. Evolution 32:784-797.

FERGUSON, A. 1980. Biochemical systematics and evolution. Blackie, Glasgow & London.

194 pp.

VOLUME 438, NUMBER 2 91

FIELD, W. D. 1971. Butterflies of the genus Vanessa and of the resurrected genera
Bassaris and Cynthia (Lepidoptera: Nymphalidae). Smiths. Contrib. Zool. 84:1-105.

GEIGER, H. J. 1981. Enzyme electrophoretic studies on the genetic relationships of
pierid butterflies. I. European taxa. J. Res. Lepid. 19:181-195.

GEIGER, H. J. & A. SCHOLL. 1985. Systematics and evolution of holarctic Pierinae: An
enzyme electrophoretic approach. Experientia 41:24—29.

HERRERA, J. 1987. Biologia de Cynthia carye Hiibner 1812, especie criptica de C.
annabella Field 1971 (Lepidoptera: Nymphalidae). Acta Entomol. Chilena 14:65-
116.

HERRERA, J.. M. ETCHEVERRY & R. BARRIENTOS. 1958. Los Nymphalidae Chilenos.
Ediciones Anales Univ. Chile (ser. azul). 39 pp.

HiccIns, L. G. & N. D. RitEy. 1970. A field guide to the butterflies of Britain and
Europe. Houghton Mifflin, Boston. 380 pp.

KITCHING, I. J. 1985. Allozyme variation in the milkweed butterflies (Lepidoptera:
Danaidae). Zool. J. Linn. Soc. 86:367-389.

Lessios, H. A. 1979. Use of Panamanian sea urchins to test the molecular clock. Nature
280:599-601.

1981. Divergence in allopatry: Molecular and morphological differentiation
between sea urchins separated by the Isthmus of Panama. Evolution 35:618-634.

ManI, M. S. 1968. Ecology and biogeography of high-altitude insects. W. Junk, The
Hague. 526 pp.

MARSHALL, L. G. 1988. Land mammals and the Great American Interchange. Am. Sci.
76:380-388.

MENKEN, S. B. J. 1982. Biochemical genetics and systematics of small ermine moths.
Z. Zool. Syst. Evol.-forsch. 20:131-148.

MIcKEvIcH, M. F. & M. S. JOHNSON. 1976. Congruence between morphological and
allozyme data in evolutionary inference and character evolution. Syst. Zool. 25:260-
270.

MILLER, J. Y. & F. M. BROWN. 1988. The fossil Vanessa butterflies with the description
of a new species from the Florissant formation (Oligocene). 39th Ann. Meeting Lepid.
Soc., 14-17 July. (Abstract.)

Nel, M. 1971. Interspecific gene differences and evolutionary time estimated from
electrophoretic data on protein identity. Am. Nat. 105:385-398.

1972. Genetic distance between populations. Am. Nat. 106:283-292.

NEvo, E., Y. Kim, C. R. SHAW & C. S. THAELER. 1974. Genetic variation, selection and
speciation in Thomomys talpoides pocket gophers. Evolution 28:1-23.

SARICH, V. 1977. Rates, sample sizes, and the neutrality hypothesis for electrophoresis
in evolutionary studies. Nature 265:24—28.

Scott, J. A. 1986. The butterflies of North America: A natural history and field guide.
Stanford Univ. Press, Stanford. 583 pp.

SCUDDER, S. H. 1889. The butterflies of the eastern United States and Canada with
special reference to New England. Vol. I. Publ. by author, Cambridge, Massachusetts.
Pp. 430-487.

SHAPIRO, A. M. 1980. Genetic incompatibility between Pieris callidice and P. occiden-
talis nelsoni: Differentiation within a periglacial relict complex. Can. Entomol. 112:
463-468.

1983. Taxonomic uncertainty, the biological species concept, and the Nearctic

butterflies: A reappraisal after twenty years. J. Res. Lepid. 21:212-218.

1984. The genetics of seasonal polyphenism and the evolution of “general
purpose genotypes’ in butterflies, pp. 16-30. In W6hrmann, K. & V. Loeschke (eds.),
Population biology and evolution. Springer, New York.

SHAPIRO, A. M. & H. J. GEIGER. 1986. Electrophoretic confirmation of the species status
of Pontia protodice and P. occidentalis (Pieridae). J. Res. Lepid. 25:39-47.

STEHLI, F. G. & S. D. WEBB, (eds.). 1985. The Great American Biotic Interchange.
Plenum, New York. 532 pp.

THORPE, J. P. 1982. The molecular clock hypothesis: Biochemical evolution, genetic
differentiation and systematics. Ann. Rev. Ecol. Syst. 13:139-168.

92 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Upvarpy, M. D. F. 1969. Dynamic zoogeography. Van Nostrand Reinhold, New York.
445 pp.

VAWTER, A. T., R. ROSENBLATT & G. C. GORMAN. 1980. Genetic divergence among
fishes of the eastern Pacific and the Caribbean: Support for the molecular clock.
Evolution 34:705-711.

WEBSTER, T. P. & J. M. BuRNS. 1973. Dewlap color variation and electrophoretically
detected sibling species in a Haitian lizard, Anolis brevirostris. Evolution 27:368-
377.

WOoODRING, P. W. 1966. The Panama land bridge as a sea barrier. Proc. Am. Phil. Soc.
110:425-434.

Received for publication 26 August 1988; accepted 21 December 1988.

Journal of the Lepidopterists’ Society
43(2), 1989, 93-101

EVALUATION OF SPERMATOPHORE COUNTS IN STUDYING
MATING SYSTEMS OF LEPIDOPTERA

ROBERT C. LEDERHOUSE, MATTHEW P. AYRES,
AND J. MARK SCRIBER
Department of Entomology, Michigan State University, East Lansing, Michigan 48824

ABSTRACT. Counts of spermatophores contained by field-collected females have
been widely used to investigate mating behavior of Lepidoptera. We present new counts
for Papilio glaucus L. females and reanalyze published data for this species to evaluate
the often implicit assumptions of this technique. In addition, we relate spermatophore
size and sequence to mating history of tiger swallowtail females captured in Wisconsin.
Number of spermatophores per female increased with both wear class and capture date.
Females that received small first spermatophores were significantly more likely to contain
one or more additional spermatophores than those that received large first spermatophores.
This suggests that more spermatophores per female result from inferior initial matings
and not necessarily from male mating preference.

Additional key words: spermatophore size, multiple-mating probability, Papilio glau-
cus, Papilionidae.

Because one spermatophore is passed during each copulation in most
Lepidoptera, and generally persists in recognizable form, counts of
spermatophores contained in females have been used to infer aspects
of mating systems in this taxon (Burns 1966, 1968, Taylor 1967, Pliske
1978, Ehrlich & Ehrlich 1978, Smith 1984). Burns (1968) documented
the validity of these two assumptions and warned of potential biases
from different aged samples. However, interpretations of spermato-
phore data are limited in additional ways that frequently have been
ignored.

Mating histories of female tiger swallowtails, Papilio glaucus L., as
revealed by spermatophore counts, probably are documented better
than any other species of butterfly (Drummond 1984). Over much of
its geographic range, P. glaucus females may be tiger-striped yellow
like males, or dark mimics of the distasteful Battus philenor (L.) (Brower
1958). Spermatophore counts were analyzed to evaluate the role of
sexual selection in maintaining this sex-limited color dimorphism in
female adults (Burns 1966, Makielski 1972, Pliske 1972, Platt et al.
1984). Drawing on this literature and additional studies in our labo-
ratory, we illustrate the strengths and limitations of using spermatophore
count data.

MATERIALS AND METHODS

Samples of Papilio glaucus canadensis females were collected from
five adjacent counties in north-central Wisconsin during the flight pe-
riod of the single generation in 1985. Of the 282 females collected, 152
were set up for oviposition; the other 130 were frozen until dissected.

94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Females were assigned to a condition class either when set up or dis-
sected. We classified 78% of the females for wing wear within a four-
day period, which reduced potential bias due to changing standards by
the classifier during the flight season as discussed below. Females were
carefully dissected so that both number of spermatophores and their
relative position in the bursa copulatrix were determined (Drummond
1984). Since the spermatophore of an additional mating forces the
previous spermatophore forward in the saclike, posteriorly opening
bursa copulatrix, and since spermatophores change from creamy white
to yellow and collapse with time, accurate sequences of spermatophore
deposition were determined for 98% of collected females. Volumes of
the nearly spherical P. g. canadensis spermatophores were estimated
by averaging the longest and shortest diameters and calculating the
volume of such a sphere. Spermatophore dimensions were not measured
for 52 females.

RESULTS AND DISCUSSION

Reliability. Spermatophore counts are generally reliable as a measure
of female mating history (Burns 1968, Lederhouse 1981, Drummond
1984). Deviations from the one copulation-one spermatophore assump-
tion are rare. Only occasionally will Papilio males not pass a sper-
matophore during coupling, and we have detected only one case of two
spermatophores being passed during a single copulation (n = 226 hand-
pairings of Papilio). This exceptional case occurred during a prolonged
coupling lasting over 24 h. The same sample also contained one female
that laid viable eggs but contained no detectable spermatophore. She
did have seminal material in her bursa. Spermatophores are persistent
in swallowtail females; even females that have been maintained for 20-
30 days in the laboratory have obvious spermatophores. However, sper-
matophores may disintegrate rapidly in the lower Lepidoptera (Taylor
1967) where the spermatophore lacks chitin. Also, spermatophores are
gradually absorbed in females of a variety of higher Lepidoptera (Burns
1968, Ehrlich & Ehrlich 1978). This is particularly true for species
where females use nutrients contributed by males at copulation for egg
production (Boggs & Gilbert 1979, Boggs 1981). Therefore, it seems
prudent to verify the one copulation-one spermatophore relation for
each species for which spermatophore counts are used.

Ageing females. Even when spermatophore counts reliably indicate
number of copulations, there remain a variety of factors that must be
considered in evaluating such data in the context of mating behavior
and sexual selection. The difficulties chiefly involve controlling for fe-
male age and spermatophore quality.

It is logical to assume that older females should carry more sper-

VOLUME 438, NUMBER 2 95

TABLE 1. Mean number of spermatophores contained by Papilio glaucus canadensis
females from Wisconsin in relation to wing condition and date of capture.

June 1985
Condition 6 13 20 Fei

Fresh 1.00 1.00 1.50 1.63
Slightly worn 1.20 1.18 55 1.85
Intermediate 1.56 1.61 1.84 2.33
Very worn 2.00 2.10 242 2.62
Mean 1.48 1.58 1.89 2.30
Sample size 2S 108 79 70

matophores than younger females. However, it is difficult to age field-
collected females accurately. Recapture rates of marked females are
usually too low to provide known age samples (Lederhouse 1982). Wing
wear is an estimate of age, but may reflect the quality of life (encounters
with predators, inclement weather, or other factors; Lederhouse et al.
1987) as much as its quantity. By their nature, estimates of age are
subjective and may vary from investigator to investigator. Since rep-
resentatives of all age classes are not present throughout a flight season,
our experience suggests a tendency to overestimate age early in a gen-
eration when very worn individuals are scarce and underestimate it
late in a generation when very fresh individuals are rare. Nevertheless,
spermatophore numbers carried by females of a variety of species have
been shown to increase with estimates of age such as wing wear (Burns
1968, Lederhouse 1981, Drummond 1984, Lederhouse & Scriber 1987).
This is illustrated within and across four sampling days for P. g. can-
adensis females (Table 1).

Accurate comparisons of samples rely on similar age structures or
the ability to control for age structure in analysis. However, female age
structures of natural butterfly populations are largely unknown. Sper-
matophore counts necessarily underestimate lifetime mating frequency.
Sampling removes females at an artificial point. Once sampled, a female
that would have remated the next day, and perhaps again the following
week, becomes equivalent to a female that would never have remated.
Since females may be singly-mated as a result of their mating system
or their young age, studies indicating female monogamy (Wiklund
1977, 1982, Wiklund & Forsberg 1985) must give age estimates for
their samples.

Comparisons of mating histories for different female morphs within
a population must control for female age. Pliske (1972) questioned the
importance of sexual selection in maintaining the frequency of the
yellow female phenotype in a Florida population of P. glaucus because

96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

he found no difference in number of spermatophores carried by yellow
or dark females. Our reanalysis of Pliske’s data indicates that his sample
of P. glaucus females cannot address effects of sexual selection on
differential remating of the two female morphs because females of both
morphs had mated too few times for a differential to be detectable.
Either females in his sample were too fresh (young) for many to have
mated more than once, or multiple-mating was rarer in central Florida
P. glaucus populations (Pliske 1972, 1973) than elsewhere (Burns 1966,
Makielski 1972, Platt et al. 1984, Lederhouse & Scriber 1987).

Spermatophore quality. Spermatophore quality may influence the
frequency with which a female mates. A common assumption is that
more spermatophores indicate superior mating, which is based on the
supposition that all spermatophores are equal. Yet young or frequently-
mating males produce smaller than average spermatophores (Sims 1979,
Svard & Wiklund 1986). Spermatophore volumes in early season P. g.
canadensis females suggest a bimodal distribution (Fig. 1). Hand-pair-
ing in our laboratory indicates that the larger spermatophores (about
7 mm*) result from males’ first matings and the smaller ones (about 4
mm) from subsequent matings (Lederhouse, Ayres & Scriber in prep.).
Spermatophore size distribution from a late-season sample shows a
significant increase in frequency of smaller spermatophores compared
with the early-season sample (Fig. 1; Kolmogorov-Smirnov test, P <
0.005). Small spermatophores may result in lower egg fertility or more
rapid fertility declines (Lederhouse & Scriber 1987). Size of spermato-
phores is therefore an important aspect of quality that should be con-
sidered.

Since stretch receptors in the female’s bursa copulatrix may deter-
mine female receptivity (Sugawara 1979), smaller than average sper-
matophores might produce a shorter mating refractory period or none
at all. Indeed, a multiway contingency analysis of the probability of P.
g. canadensis females carrying multiple spermatophores demonstrates
a significant effect of spermatophore size, in addition to date of capture
and female condition class (Fig. 2, Table 2). Only 45% of 160 females
that received a large first spermatophore (>4 mm?) carried more than
one spermatophore compared with 63% of 70 females that had a small
first spermatophore (<4 mm). This difference is significant (x2, P =
0.002, Table 2) although both samples had similar age distributions as
indicated by wing-wear classes (x?, P > 0.25). This suggests that females
mated again to replace a small spermatophore. We observed a similar
relation for P. g. glaucus from an Ohio population (Lederhouse &
Scriber 1987). Of 164 females that received a large first spermatophore,
45% had mated more than once compared with 67% of 165 females

VOLUME 43, NUMBER 2 97

FREQUENCY (%)

EARLY SEASON

1 3 5 7 9 11
SPERMATOPHORE VOLUME (mm’)

Fic. 1. Size distribution of Wisconsin Papilio glaucus canadensis spermatophores
found in early-season females (captured 6-13 June 1985) and in late-season females
(captured 20-27 June 1985). N = 166 and 232 spermatophores, respectively.

98 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

TABLE2. Results of multiway contingency analysis (SAS CATMOD) of the probability
of multiple-mating in P. g. canadensis. The null hypothesis tested for each source effect
was that proportion of remated females (carrying 2 or more spermatophores) did not
differ between categories of the independent variable. A graphical representation of the
analysis is in Fig. 2.

Source df Chi-square P
Condition 2 32.4 0.0001
Spermatophore size 1 10.2 0.002
Date of collection il 5.6 0.018
Condition x date 2 4.7 0.10
Other interactions 1-2 <3.0 =0)22

that had a small first spermatophore (x? = 15.1, P < 0.0001, where
condition class and year were other sources of variation in the model).

Male P. glaucus generally emerge before females (Berger 1986),
which provides time for them to reach full sexual maturity before
copulation. Nevertheless, some small spermatophores carried by early
females could come from immature males because emergence curves
of the sexes do overlap (Berger 1986). The significant relation between
date of capture and probability of remating (Tables 1, 2) may result
from an increased proportion of females mating with previously mated
males later in the generation. Since size of spermatophore passed by a
male decreases with additional matings in P. g. canadensis (Lederhouse,
Ayres & Scriber in prep.) and proportion of males that had mated at
least once appears to increase later in the generation (Fig. 1), later
mating females were more likely to receive an insufficient spermato-
phore and mate again after a short refractory period. Late in the
generation, it was not uncommon for even fresh females to carry three
or four smaller than average spermatophores.

Even the size of a spermatophore may be a poor indicator of its
quality (Greenfield 1983, Jones et al. 1986). In our study of 1985 P. g.
canadensis females, we could detect no significant relation between
spermatophore size and percent egg hatch, despite wide variation in
egg hatch (range 0.0-97.5%, x = 58.5, n = 23). Similar-sized sper-
matophores may vary in relative proportions of different constituents
(Marshall 1982, Alcock & Hadley 1987, Marshall & McNeil in press).
This may be particularly important for those species where the sper-
matophore and associated secretions contribute to the nutrient pool
available to females for reproduction. Selection could favor male sperm
delivery strategies that treat females of different reproductive value
differently (Boggs 1981), or that fool a female with large but inexpen-
sive, low quality spermatophores.

Persistence of courtship is related to mating history in some lepi-

VOLUME 438, NUMBER 2 99

PERCENT MULTIPLE MATINGS

| |

Y

i
/

Late Early La Late

1-2 3 4
CONDITION CLASS

Fic. 2. Probability of remating in Wisconsin Papilio glaucus canadensis females as
influenced by first spermatophore size, date of collection, and condition class. In 6 of 6
comparisons females receiving a small 1st spermatophore (hatched bars) were more likely
to remate than females receiving a large lst spermatophore (stippled bars). Associated
contingency tests are in Table 2. Early-season females were captured 6-18 June and late-
season females 20-27 June 1985.

dopteran species (Rutowski 1979, 1980). High selectivity by either a
male or a female could lead to passing of a larger than average sper-
matophore followed by a longer than average mating refractory period.
Thus, preferred females might carry fewer but larger spermatophores
on average. Larger Dryas julia Fabr. females received larger sper-
matophores (Boggs 1981). Less selective males might mate more fre-
quently but pass smaller than average spermatophores. Less selective
females might receive smaller spermatophores, remate at shorter in-
tervals, and as a result carry more spermatophores on average. Such
potential results run counter to the logic of Burns (1966) and others.

These various factors do not invalidate spermatophore counts but
suggest that more care must be taken in interpreting count data. Count-
ing spermatophores remains a valuable tool, but count data must be
integrated with that of other techniques to yield an accurate appraisal
of mating behavior in Lepidoptera.

100 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ACKNOWLEDGMENTS

This research was supported in part by the National Science Foundation (BSR 8306060,
BSR 8718448), USDA grants #85-CRCR-1-1598 and #87-CRCR-1-2851, and the Grad-
uate School and College of Agricultural and Life Sciences (Hatch 5134) of the University
of Wisconsin and the Colleges of Natural Science and Agriculture of Michigan State
University (MAES Project 8051). We are grateful to P. Christi, M. Evans, E. Schuh, J.
Sibenhorn, J. Thorne, and V. Viegut for their assistance in collecting butterflies. R. H.
Hagen provided helpful comments on this manuscript.

LITERATURE CITED

ALCOCK, J. & N. F. HADLEY. 1987. Test of alternative hypotheses for assortative mating
by size: A comparison of two meloid beetles (Coleoptera: Meloidae). J. Kansas Ento-
mol. Soc. 60:41-50.

BERGER, T. A. 1986. Habitat use and reproductive ecology of the eastern tiger swal-
lowtail, Papilio glaucus L. Ph.D. Dissertation, Rutgers University, Newark, New
Jersey. 109 pp. Diss. Abstr. Int. microfilm order No. 8616563.

Boccs, C. L. 1981. Selection pressures affecting male nutrient investment at mating in
heliconiine butterflies. Evolution 35:931—940.

Boccs, C. L. & L. E. GILBERT. 1979. Male contribution to egg production in butterflies:
Evidence for transfer of nutrients at mating. Science 206:83-84.

BROWER, J. VZ. 1958. Experimental studies of mimicry in some North American but-
terflies. Part II. Battus philenor and Papilio troilus, P. polyxenes and P. glaucus.
Evolution 12:123-136.

BURNS, J. M. 1966. Preferential mating versus mimicry: Disruptive selection and sex-
limited dimorphism in Papilio glaucus. Science 153:551-558.

1968. Mating frequency in natural populations of skippers and butterflies as
determined by spermatophore counts. Proc. Natl. Acad. Sci. 61:852-859.

DRUMMOND, B. A. 1984. Multiple mating and sperm competition in the Lepidoptera,
pp. 291-370. In Smith, R. L. (ed.), Sperm competition and the evolution of animal
mating systems. Academic Press, New York.

EHRLICH, A. H. & P. R. EHRLICH. 1978. Reproductive strategies in the butterflies: I.
Mating frequency, plugging, and egg number. J. Kansas Entomol. Soc. 51:666-697.

GREENFIELD, M. D. 1983. The question of paternal nutrient investment in Lepidoptera:
Male contributed proteins in P. interpunctella. Intern. J. Invert. Reprod. 5:323-330.

JONEs, K. N., F. J. ODENDAAL & P. R. EHRLICH. 1986. Evidence against the sper-
matophore as paternal investment in checkerspot butterflies (Euphydryas: Nym-
phalidae). Am. Mid]. Nat. 116:1-6.

LEDERHOUSE, R. C. 1981. The effect of female mating frequency on egg fertility in
the black swallowtail, Papilio polyxenes asterius (Papilionidae). J. Lepid. Soc. 35:
266-277.

1982. Factors affecting equal catchability in two swallowtail butterflies, Papilio
polyxenes and P. glaucus. Ecol. Entomol. 7:379-383.

LEDERHOUSE, R. C., S. G. CODELLA & P. J. COWELL. 1987. Diurnal predation on
roosting butterflies during inclement weather: A substantial source of mortality in
the black swallowtail, Papilio polyxenes (Lepidoptera: Papilionidae). J. New York
Entomol. Soc. 95:310-319.

LEDERHOUSE, R. C. & J. M. SCRIBER. 1987. Ecological significance of a postmating
decline in egg viability in the tiger swallowtail. J. Lepid. Soc. 41:83-93.

MAKIELSKI, S. K. 1972. Polymorphism in Papilio glaucus L. (Papilionidae): Maintenance
of the female ancestral form. J. Lepid. Soc. 26:109-111.

MARSHALL, L. D. 1982. Male nutrient investment in the Lepidoptera: What nutrients
should males invest? Am. Nat. 112:197-213.

MARSHALL, L. D. & J. N. MCNEIL. In press. Spermatophore mass as an estimate of
male nutrient investment: A closer look in Pseudaletia unipuncta Haworth (Lepi-
doptera: Noctuidae). Functional Ecol.

VOLUME 43, NUMBER 2 101

PLATT, A. P., S. J. HARRISON & T. F. WILLIAMS. 1984. Absence of differential mate
selection in the North American tiger swallowtail, Papilio glaucus, pp. 245-250. In
Vane-Wright, R. I. & P. R. Ackery (eds.), The biology of butterflies. Sympos. 11.
Academic Press, London.

PLISKE, T. E. 1972. Sexual selection and dimorphism in female tiger swallowtails, Papilio
glaucus L. (Lepidoptera: Papilionidae): A reappraisal. Ann. Entomol. Soc. Am. 65:
1267-1270.

1973. Factors determining mating frequencies in some New World butterflies
and skippers. Ann. Entomol. Soc. Am. 66:164—-169.

RUTOWSKI, R. L. 1979. The butterfly as an honest salesman. Anim. Behav. 27:1269-
1270.

1980. Courtship solicitation by females of the checkered white butterfly, Pieris
protodice. Behav. Ecol. Sociobiol. 7:113-117.

Sims, S. R. 1979. Aspects of mating frequency and reproductive maturity in Papilio
zelicaon. Am. Midl. Nat. 102:36-50.

SMITH, D. A. S. 1984. Mate selection in butterflies: Competition, coyness, choice and
chauvinism, pp. 225-244. In Vane-Wright, R. I. & P. R. Ackery (eds.), The biology
of butterflies. Sympos. 11. Academic Press, London.

SUGAWARA, T. 1979. Stretch reception in the bursa copulatrix of the butterfly, Pieris
rapae crucivora, and its role in behaviour. J. Comp. Physiol. A. 130:191-199.

SVARD, L. & C. WIKLUND. 1986. Different ejaculate delivery strategies in first versus
subsequent matings in the swallowtail butterfly Papilio machaon L. Behav. Ecol.
Sociobiol. 18:325-830.

TayLor, O. R. 1967. Relationship of multiple mating to fertility in Atteva punctella
(Lepidoptera: Yponomeutidae). Ann. Entomol. Soc. Am. 60:583-590.

WIKLUND, C. 1977. Courtship behaviour in relation to female monogamy in Leptidea
sinapis (Lepidoptera). Oikos 29:275-283.

1982. Behavioural shift from courtship solicitation to mate avoidance in female
ringlet butterflies (Aphantopus hyperanthus) after copulation. Anim. Behav. 30:790-
793.

WIKLUND, C. & J. FORSBERG. 1985. Courtship and male discrimination between virgin
and mated females in the orange tip butterfly Anthocharis cardamines. Anim. Behav.
34:328-332.

Received for publication 17 August 1988; accepted 2 December 1988.

Journal of the Lepidopterists’ Society
43(2), 1989, 102-107

BEHAVIOR OF THE TERRITORIAL SPECIES
LIMENITIS WEIDEMEYERII (NYMPHALIDAE) WITHIN
TEMPORARY FEEDING AREAS

RIisA H. ROSENBERG!

Ecology and Evolutionary Biology, Cornell University,
Ithaca, New York 14853

ABSTRACT. Behavior of a population of Limenitis weidemeyerii Edwards (Nym-
phalidae) was studied in central Colorado using mark-recapture and observations. In 1984,
individuals of both sexes fed on sap excreted from a willow via holes made by sapsuckers.
The following season, L. weidemeyerii of both sexes fed on honeydew excreted by aphids.
In addition, in both years, individuals fed at artificial high-quality food sources experi-
mentally placed within the habitat. Territorial behaviors (patrols, chases, and investiga-
tions) were not observed within the temporary feeding areas, possibly because high
intruder pressures affected the defendability of these sites. It is suggested that some studies
citing a lack of territoriality in lepidopterans may have been conducted within temporary
feeding areas.

Additional key words: admiral butterfly, territoriality, sap feeding.

Resource defense territoriality involves defense of resources that are
patchy, predictable, and economically defendable (Davies 1978a). In
lepidopteran territoriality, males generally defend locations where fe-
males reliably can be found: oviposition sites (Baker 1972), landmark
sites (Shields 1967, Davies 1978b, Lederhouse 1982), or routes used by
females for feeding or oviposition (Fitzpatrick & Wellington 1983,
Baker 1972). While food resources are commonly defended in other
taxonomic groups (Wittenberger 1981), reports of butterflies defending
areas around adult feeding sites are rare. This may stem from the
economic defendability of adult lepidopteran feeding sites. Because
nectar resources used by butterflies often are widely scattered (Rutowski
1984; but see Murphy 1983, Murphy et al. 1984), it might prove difficult
for a butterfly to maintain exclusive use of a patch of flowers, even
though other insects (especially bees) do defend floral resources. Male
mason bees (Hoplitis anthocopoides (Schenck): Megachilidae) for in-
stance, have been found defending patches of flowers (Eickwort &
Ginsberg 1980). In addition to floral resources, Lepidoptera often use
temporary food sources such as sap holes, puddles, animal excreta, and
carrion (Wilson & Hort 1926, Norris 1936, Downes 1973, Adler &
Pearson 1982) which might prove to be more economically defendable.

I describe here the behavior of individuals of a territorial species,
Limenitis weidemeyerii Edwards (Nymphalidae) during two flight sea-
sons when the population had access to an unpredictable, patchy food
supply in addition to its normally undefended floral foraging sites.
During one season (1984), individuals of both sexes were found feeding

' Present address: Department of Zoology, University of Stockholm, S-106 91 Stockholm, Sweden.

VOLUME 43, NUMBER 2 103

at holes on a willow made by yellow-bellied sapsuckers (Sphyrapicus
varius L.: Picidae). The following year this site was not used, presumably
because sap no longer flowed freely. Individuals of both sexes were
found feeding at another temporary food supply: honeydew on willow
leaves in a stand where there were abundant aphids (Chaitophorus
viminalis Koch: Aphididae). Neither location was used by this species
during the previous three years as territorial, feeding, or oviposition
sites. Artificial food sources also were placed in similar sites at random
locations and times, and presence or absence of territorial behavior was
recorded. |

METHODS

A population of Limenitis weidemeyerii was studied during July and
August 1984 and 1985 along Cement Creek in Gunnison Co., Colorado,
as part of a larger study of social and genetic organization of populations
of this species (Rosenberg 1987). All individuals seen were marked and
color-coded using the 1-2-4-7 system of Ehrlich and Davidson (1960),
so individuals could be easily identified on the wing. Territorial behavior
of this species is reported in greater detail elsewhere (Rosenberg 1987).
Briefly, territorial behavior consists of a perched male flying out to
investigate any passing object, resulting in either a spiral flight (indi-
viduals fly around one another), or a chase (flight directly towards an
intruder leading away from the perch site). Territorial behavior also
includes patrols: smooth flights from and back to the perch without
obvious stimulus. Feeding behavior also was recorded.

During the 1984 flight season individuals of both sexes were found
aggregating at a series of holes made by yellow-bellied sapsuckers on
a 1.5 m tall willow bush (Salix sp., hereafter called sapwillow) located
approximately 23 m from the nearest territorial site. Behavior of in-
dividuals at this location was recorded at various times through the day
(0900-1800 h), over the season (12 July-27 August), and observations
also were concentrated for a full day only on activities at this site.
Weather conditions (sun, cloud, rain), time of day, sex, identity and
behavior as described above were recorded. Ages were estimated by
wing-wear in increments of 0.5 from scores of 1 (newly emerged) to 4
(many scales missing), following the conventions of Watt et al. (1977).
For unmarked individuals sighted, weather, time of day, and behaviors
were recorded; ages were unknown, and sex of only a sample of in-
dividuals could be ascertained by noting approximate wing lengths. As
with most nymphalids (Howe 1975), females of this species are larger
than males.

During the 1985 flight season the sapwillow was no longer used by
the butterflies. Instead, they frequented a willow stand of 10 m® area
ca. 100 m away (and 90 m from the nearest territorial site) where

104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

approximately 60% of leaves contained aphids on their undersides.
Behavior and identity of individuals visiting this site, weather condi-
tions, time of day, sex, and ages of a sample of individuals were recorded
during the 1985 season (25 July-27 August). Observers could approach
within 0.25 m of individual butterflies at both sites.

Four times each season artificial food sources were placed for a
minimum of two days in arbitrary locations in the habitat, and identity,
sex and age of individuals feeding there were recorded. Artificial sources
contained fermented fruit, beer or wine, and were placed in cages
styled after Platt (1969).

RESULTS

In the 1984 season there were 70 sightings of L. weidemeyerii feeding
at the sapwillow: 20 marked males, 4 marked females, and 21 unmarked
individuals. The marked butterflies fed there 49 different times on 13
separate days, many of these individuals (42%) feeding there repeatedly
on different days. It is possible that unmarked butterflies also were
resighted on different days. Individuals of various ages were found at
this site feeding at all times of the day and in all weather conditions.
The majority of the marked butterflies (84%) were not newly emerged;
the average age-class was approximately 2. In more than 30 h of ob-
servation, only feeding was observed at the sapwillow; no territorial
behaviors (patrols, chases, investigations) were observed. Other taxa also
fed there, including unidentified species of Diptera and Hymenoptera,
and other Lepidoptera such as Vanessa atalanta L., Nymphalis antiopa
L., and birds such as yellow-bellied sapsucker, and broad-tailed hum-
mingbird.

In the 1985 season there were 62 sightings of L. weidemeyerii of
both sexes feeding at the willow stand containing aphids: 1 marked
female, 16 marked males, and 34 unmarked individuals (at least 4 of
the last were females). Marked butterflies were sighted there 28 times;
8 of the marked males fed there on multiple days. Some of the unmarked
butterflies also may have fed there on different days. The only marked
female sighted had previously mated (as evidenced by a sperm plug).
No newly emerged individuals were found there. On average, the
marked individuals were of age-class 3 (out of a maximum wing-wear
score of 4). Weather conditions were noted for 23 observations: only 4
sightings occurred during a cloudy period, the other 19 when there was
sunshine. Feeding was observed at all times of day. Individuals within
the site spent most of their time probing with their proboscides on sticky
spots on leaf surfaces. The butterflies apparently were feeding on the
honeydew flicked onto the top surfaces by the aphids on leaves above
(as in Wilson 1971). In the laboratory after feeding, I observed L.

VOLUME 438, NUMBER 2 105

weidemeyerii regurgitate and probe repeatedly at the regurgitant. Be-
cause in the field this species was observed to probe repeatedly on the
leaves, it is reasonable to assume they were imbibing fresh (or possibly
dissolved) honeydew. In more than 10 h of intensive observation, patrol
flights never were seen in this area. Interactions between individuals
were extremely brief and slow moving, and rather than involving chases
away from the site, always resulted in the individuals landing on leaves
there and feeding. Other taxa also were observed feeding on the hon-
eydew including Diptera (Sarcophagidae, Muscidae) and Hymenoptera
(Dolichovespula arenaria (Fabricius): Vespidae, and Dialictus sp.: Ha-
lictidae).

Five individuals were found at artificial food sources placed in the
field: four males and one female. These individuals on average were
scored as age 2 (out of a total wing-wear score of 4). In more than 6 h
of observation, no territorial behavior was observed at or near these
sources.

DISCUSSION

Patchy and predictable resources in nature often are defended via
territoriality (Davies 1978a). Unpredictable sources, even if high quality
often are not defended. Male territorial behavior (perching, patrolling,
investigating, chasing) was not observed at three temporary feeding
sites of a population of Limenitis weidemeyerii in central Colorado.
These feeding sites, at sapsucker sap holes, leaves with aphid honeydew,
and artificial sources, were high-quality sources rich in sugars and free
amino acids. Four other willow stands with evidence of previous sap-
sucker damage were found within the boundaries of this population,
suggesting that although this food source is unpredictable in time and
space, it had been encountered by this population of L. weidemeyerii
previously. Limenitis butterflies have been reported feeding at sap holes
(Flemwell 1914, Wilson & Hort 1926) and Platt (1969) successfully
traps Limenitis using baits. To date there have been only a few reports
of adult butterflies other than lycaenids feeding on aphid honeydew
(Kershaw 1907, Bingham 1907, Johnson & Stafford 1986).

Limenitis weidemeyerii males defend sites where they have good
vantage points of approaching conspecifics, generally either at locations
of emerging females or along flyways with an open central area bounded
on other sides by vegetation (Rosenberg 1987). Although feeding lo-
cations described here proved to be good rendezvous sites for a single
season, they were within wide open areas, and there is no guarantee
of their utility in the following generation. Males appear to mate with
females emerging within their territorial sites (Rosenberg 1987); thus,
ovipositing within a previously unused territorial site might lead to

106 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

offspring being unmated longer. Also, larvae in these sites might be
harmed because sap can attract adults of predatory and parasitic groups
(Stary 1970), and also can breed bacteria. Finally, females visiting these
sites were older ones, hence probably not receptive anyway (Rosenberg
1987), so defense of these locations may have been a waste of a territorial
male’s time and energy.

Butterflies were observed feeding at these sites under all weather
conditions. Finding males feeding there on sunny days is particularly
interesting because in most butterfly species territorial defense occurs
on sunny days (Baker 1972, Davies 1978b, Lederhouse 1982, Wickman
& Wiklund 1983, Rosenberg 1987). It is probable that individuals come
to the temporary sites to quickly stoke up with a high energy food
source to support other activities such as territorial defense, mating,
and oviposition.

Alternatively, these high energy sources may not be defended ter-
ritorially because it would be uneconomical on account of high intruder
pressures. A breakdown of territorial behavior at feeding sites also has
been noted for other butterfly species (Baker 1972, Fukuda 1974), as
well as for birds (Gill & Wolf 1975).

Before the recent acceptance of lepidopteran territoriality, there were
a number of reports on a “lack of territoriality” in butterflies (Ross
1968, Suzuki 1976, Scott 1974). One such report, on two species of
Hamadryas, seems to have been undertaken at a temporary feeding
area; Ross (1963) described tree sapholes within the study site. The
“lack of territoriality’ hence may only refer to this feeding location.
More detailed study of these species away from a potential high-quality
temporary feeding area may indicate these to be territorial species. If
so, it is unfortunate that Ross’s study has been so widely cited as negative
evidence for lepidopteran territoriality. Further studies of the behavior
of individuals with and without unpredictable high quality food sources
can help us to better understand territoriality in Lepidoptera.

ACKNOWLEDGMENTS

I thank G. Eickwort, P. Brussard, S. Levin, C. Aquadro, P. Sherman, R. Pudim, A.
Platt, M. Cain, F. Sperling, R. Lederhouse, R. Baker, and many wonderful field and
laboratory assistants for help during this project. Financial assistance was provided by
several grants in aid of research from local and national chapters of Sigma Xi, the Rocky
Mountain Biological Laboratory, and Cornell University.

LITERATURE CITED

ADLER, P. & D. PEARSON. 1982. Why do male butterflies visit mud puddles? Can. J.
Zool. 60:322-325.

BAKER, R. R. 1972. Territorial behaviour of the nymphalid butterflies Aglais urticae
(L) and Inachis io (L). J. Anim. Ecol. 41:453-469.

BINGHAM, C. T. 1907. The fauna of British India including Ceylon and Burma. Taylor
& Francis, London. 480 pp.

Davies, N. B. 1978a. Ecological questions about territorial behavior, pp. 317-350.

VOLUME 43, NUMBER 2 107

In Krebs, J. R. and N. B. Davies (eds.), Behavioral ecology: An evolutionary

approach. Sinauer, Sunderland, Massachusetts. 494 pp.

1978b. Territorial defence in the speckled wood butterfly (Pararge aegeria):
The resident always wins. Anim. Behav. 26:138-147.

Downes, J. A. 1973. Lepidoptera feeding at puddle margins, dung and carrion. J.
Lepid. Soc. 27:89-99.

EHRLICH, P. R. & S. DAVIDSON. 1960. Techniques for capture-recapture studies of
Lepidoptera populations. J. Lepid. Soc. 14:227-229.

EICKWokRT, G. C. & H. GINSBERG. 1980. Foraging and mating behavior in Apoidea.
Ann. Rev. Entomol. 25:421—446.

FITZPATRICK, S. & W. WELLINGTON. 1983. Insect territoriality. Cen. J. Zool. 61:471-
486.

FLEMWELL, G. 1914. Beautiful Switzerland: Lausanne and its environs. Blackie &
Son Ltd., London. 64 pp.

Fukuba, H. 1974. Life history of butterflies. Seibundo-shinkosha, Tokyo. 272 pp.

GILL, F. B. & L. L. WoLF. 1975. Economics of feeding territoriality in the golden-
winged sunbird. Ecology 56:333-345.

Howe, W. 1975. The butterflies of North America. Doubleday, New York. 633 pp.

JOHNSON, J. B. & M. P. STAFFORD. 1986. Adult Noctuidae feeding on aphid honeydew
and a discussion of honeydew feeding by adult Lepidoptera. J. Lepid. Soc. 39:321-
327.

KERSHAW, J. C. 1907. Butterflies of Hong Kong. Kelly & Walsh, London. 184 pp.

LEDERHOUSE, R. C. 1982. Territorial defense and lek behaviour of the black swal-
lowtail butterfly, Papilio polyxenes. Behav. Ecol. Sociobiol. 10:109-118.

Murpny, D. D. 1983. Nectar sources as constraints on the distribution of egg masses
by the checkerspot butterfly Euphydryas chalecedona (Lepidoptera: Nymphali-
dae). Environ. Entomol. 12:463-—466.

Murpny, D. D., M. MENNINGER & P. R. EHRLICH. 1984. Nectar source distribution
as a determinant of oviposition host species in Euphydryas chalecedona. Oecologia
(Berlin) 62:269-271.

Norris, M. J. 1936. The feeding habits of the adult Lepidoptera Heteroneura. Trans.
Roy. Entomol. Soc. London 85:61-—90.

PLATT, A. P. 1969. A lightweight collapsible bait trap for Lepidoptera. J. Lepid. Soc.
23:97-101.

ROSENBERG, R. H. 1987. The social and genetic organization of populations of Wei-
demeyer’s admiral butterfly. Ph.D. Thesis, Cornell University, Ithaca, New York.
85 pp. Diss. Abs. Int. 48:B2527-B2528. Order No. 8725765.

Ross, G. 1963. Evidence for lack of territoriality in two species of Hamadryas (Nym-
phalidae). J. Res. Lepid. 2:241-246.

RUTOWSKI, R. 1984. Sexual selection and the evolution of butterfly mating behavior.
J. Res. Lepid. 23:125-142.

ScoTT, J. A. 1974. Mate-locating behavior of butterflies. Am. Midl. Nat. 91:103-117.

SHIELDS, O. 1967. Hilltopping. J. Res. Lepid. 6:69-178.

STARY, P. 1970. Biology of aphid parasites (Hymenoptera: Aphidiidae) with respect
to integrated control. Dr. W. Junk, The Hague. 643 pp.

SUZUKI, Y. 1976. So-called territorial behaviour of the small copper, Lycaena phlaeas
daimio Seitz (Lepidoptera, Lycaenidae). Kontyi, Tokyo 44:193-204.

WatTT, W. B., F. CHEw, L. SNYDER, A. WaTT & D. ROTHSCHILD. 1977. Population
structure of pierid butterflies I. Numbers and movements of some montane Colias
species. Oecologia 27:1-22.

WICKMAN, P.-O. & C. WIKLUND. 1983. Territorial defence and its seasonal decline
in the speckled wood butterfly (Pararge aegeria). Anim. Behav. 31:1206-1216.

WILSON, E. O. 1971. The insect societies. Belknap Press of Harvard Univ. Press,
Cambridge, Massachusetts. 548 pp.

WILSON, G. & N. Hort. 1926. Insect visitors to sap-exudations of trees. Trans. Ento-
mol. Soc. London 74:243-254.

WITTENBERGER, J. F. 1981. Animal social behavior. Duxbury Press, Boston. 722 pp.

Received for publication 11 September 1987; accepted 6 September 1988.

Journal of the Lepidopterists’ Society
43(2), 1989, 108-113

A NEW NEOTROPICAL ANONCIA SPECIES
(COSMOPTERIGIDAE)

DAVID ADAMSKI

Department of Entomology, Drawer EM, Mississippi State University,
Mississippi State, Mississippi 39762

ABSTRACT. Anoncia crossi is described from 1 male and 15 females collected in
Guerrero, Mexico. Anoncia crossi is differentiated from A. diveni, the only other congener
to occur in the Neotropics, by structural differences in male and female genitalia, and
shape of the eighth tergum. A photograph of the imago and illustrations of wing venation,
modified eighth tergum and sternum of the male, and male and female genitalia are
included.

Additional key words: Gelechioidea, Cosmopteriginae, Mexico, Anoncia crossi, A.
diveni.

Anoncia is a New World genus with 31 species known from the SW
United States, Mexico, and Honduras (Hodges 1978, 1983). Larvae feed
on Labiatae, Loasaceae, and Verbenaceae as leaftiers and leafminers,
or in immature ovaries of developing fruit (Hodges 1978).

Clarke (194la) proposed Anoncia to include seven previously de-
scribed species. After the description of a new Anoncia and two species
transferrals into Anoncia (Clarke 1942), the genus was ignored in taxo-
nomic treatments until the studies of Hodges (1962, 1978, 1983). Hodges
(1962) described 9 Anoncia and made 2 synonymies, and in a review
of the genus (Hodges 1978) synonymized 2 species and added 16 species
including 3 new combinations. Hodges (1978) provided adequate de-
scriptions, and a generic key that places the species described here in
Anoncia on the basis of the following characters: hindwing with M,
and Cu, connate or stalked, rarely separate; forewing with Cu, and Cu,
slightly downcurved from cell then parallel with M;, ocelli absent;
aedeagus without dorsal projection from midregion.

Discovery of the species described here resulted from examination
of unidentified specimens during systematics study of North American
Blastobasidae (Gelechioidea). The new species is described here because
it is only one of two species of Anoncia known from the Neotropics,
and will undoubtedly contribute to future understanding of the evo-
lution of the genus.

Pinned specimens and genitalic preparations were examined using
a stereomicroscope and a phase-contrast microscope. Colors of vestiture
were described using Kornerup and Wanscher (1978) as a standard.
Genitalia were dissected as described by Clarke (1941b), except that
mercurochrome and chlorazol black were used as stains.

VOLUME 438, NUMBER 2 109

Fic. 1. Anoncia crossi, holotype female.

Anoncia crossi Adamski, new species

(Figs. 1-6)

Head. Scales on frontoclypeus and vertex basally and apically white with subapical
brown band, or white with brown apex; scape, pedicel, flagellomeres mostly brown
intermixed with white scales, antennal pecten concolorous with vertex scales; 2nd segment
of labial palpus with basal and dorsomedial scales brown, medial and subapical scales
white, terminal segment with basal and dorsomedial scales white, medial and subapical
scales brown, or with mostly brown scales on outer surface, and mostly white scales on
inner surface. Thorax. Tegulae and mesoscutum concolorous with vertex scales, or brown
intermixed with light-brown, or mostly brown scales, tegulae occasionally with white
marginal scales. Legs. Mostly brown intermixed with white scales, each segment and
tarsomere with white apical band, each tibia with median band (sometimes not expressed
on tibia 1), coxa 1 and femora 2-8 occasionally mostly white intermixed with brown
scales. Forewing (Figs. 1, 2). Length 6.7-8.4 mm (n = 16); ground color gray; basal wing
scales white with brown, light-brown, or brownish gray apex; submedial fascia with mostly
semierect brown scales, delimiting a subcircular patch of light-brown scales near middle
of discal cell, posterior portion of submedial fascia pale; scales on wing adjacent to inner
margin of submedial fascia mostly white, intermixed with white scales tipped with light-
brown; scales on distal portion of wing mostly white, tipped with light-brown, intermixed
with white scales tipped with brown; ventral surface uniform grayish brown; venation as
in Fig. 2 (n = 8). Hindwing (Figs. 1, 2). Dorsal and ventral surfaces uniformly light
grayish brown; venation as in Fig. 2 (n = 3). Abdomen. Anterior portion of segments
with grayish brown scales, posterior portion with light grayish brown scales; eighth tergum
and sternum modified as in Figs. 3-4, seventh segment unmodified (n = 1). Male genitalia
(Fig. 5) (n = 1). Uncus absent, gnathos with two highly sclerotized asymmetrical projec-
tions, apical setae absent, larger projection serpentine-shaped and apically blunt, smaller
projection apically pointed posteriorly, posterior margin deeply arched; tegumen and
aedeagus heavily sclerotized, aedeagus ankylosed with heavily sclerotized diaphragma,
partially setose on apical rim, cornuti absent; vinculum broad throughout; valvae asym-
metrical, right valva reduced laterally, with large basal lobe, broad at base, left valva
laterally broadened, modified basally into a long, thin, apically setose projection, slightly
expanded basilaterally. Female genitalia (Fig. 6) (n = 3). Ostium bursae asymmetrically

110 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fic. 2. Wing venation of Anoncia crossi. Scale line = 2.0 mm.

situated on eighth tergum; ductus bursae membranous throughout; corpus bursae mem-
branous with spinules on walls, with accessory bursae at base; ductus seminalis arising
from accessory bursa, signum absent; apophyses anteriores widely separated basally,
arising from lateral margin of eighth tergum; eighth tergum with pair of short submedial

4

Fic. 3. Eighth tergum of male Anoncia crossi (dorsal view). Scale line = 0.5 mm.
Fic. 4. Eighth sternum of male Anoncia crossi (dorsal view). Scale line = 0.5 mm.

VOLUME 43, NUMBER 2 AT

Fic. 5. Male genitalia of Anoncia crossi (posterior view). Scale line = 0.5 mm.

projections, posterior margin slightly emarginate medially; seventh abdominal segment
unmodified.

Types. Holotype (Fig. 1) female: Mex[ico], Guerrero, Zapilote C[an]y[o]n, 8 km S[outh]
Mezcala, IX-10-82, 550 m; [Collectors] J. A. Powell and J. A. Chemsak, at light. Holotype
not dissected, deposited in Essig Entomology Museum, University of California, Berkeley,
California. Paratypes (1 male, 14 females): 1 female same data as holotype; 4 females
Mex. Guerrero, 32 km W Eguala, IX-11-82, elev. 1350 m; J. A. Powell and J. A. Chemsak,
D. Adamski wing slide nos. 3029, 3129, 3130 and gen. slide nos. 3127 and 3128; 2 females
same as previous data except IX-15-87; 1 male, 5 females same except IX-15-82, D.
Adamski male gen. slide no. 3027; 2 females 10 km E Tixtla; [X-18/22-82; elev. 1770 m,
D. Adamski female gen. slide no. 3028. Two female paratypes in U.S. National Museum,
other paratypes in same depository as holotype.

Remarks. Anoncia crossi appears closely allied to A. diveni (Heinrich), and these are
the only Anoncia congeners known in the Neotropics. Each possess a light-brown patch
of scales within the submedial fascia of the forewing. Males of both species possess an
unmodified seventh abdominal segment, valvae are short and broad, and size and shape
of aedeagus are similar. Females of both species have a membranous ductus bursae, an
accessory bursae that arises from base of the corpus bursae, lack signa, and have apophyses
anteriores that arise from lateral margins of the eighth sternum.

112 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fic. 6. Female genitalia of Anoncia crossi (dorsal view). Scale line = 1.0 mm.

VOLUME 43, NUMBER 2 113

Males of crossi can be separated from those of diveni by the acute angle and broader
lobe on the right valva, presence of apical setae on basal projection of the left valva,
presence of expanded base of the basal projection of the left valva, broad vinculum
throughout, pointed apex of the left brachium, absence of a dorsal projection on base of
right brachium, presence of setae on apical rim of the aedeagus, and linear shape of the
eighth abdominal tergum. Females of crossi can be separated from those of diveni by
presence of a pair of small projections on the anterior margin of the eighth tergum
between apophyses anteriores.

Hodges’s (1978) key to species of Anoncia is modified below to include A. crossi. In
the part treating males, couplet 23 is modified, and couplet 23’ added; in the part treating
females, couplet 39 is modified and couplet 39’ added. The modifications read as follows:

QouValvae short, broadly joined basally (text fig. 7b) 00
— Valvae longer, narrowly joined basally (text fig. 18d) 0
23 eebette valva with basal_lobesetose apically diveni
=eewchavatva with, basal lobe without setae 200 crossi
39. Eighth abdominal tergum concavely excavated, genital plate at least % width

Haeoumabadominalsteroume~(text-fie 28a) ees aM a 39’
— Eighth abdominal tergum concavely excavated, genital plate less than % width

Oeste abdominal. tergum (Text-me, 26h) a esa Ee Ee diveni
39’. Anterior margin of 8th abdominal tergum with two short lobes _ crossi
— Anterior margin of 8th abdominal tergum without such lobes smogops

Etymology. The new species is named in honor and memory of Dr. William H. Cross,
naturalist, ecologist, taxonomist, and founder of the Mississippi Entomological Museum
at Mississippi State University, who died in Mexico in 1984 on a collecting expedition.

ACKNOWLEDGMENTS

I thank J. A. Powell, University of California, Berkeley, for loan of the specimens
examined, R. W. Hodges, U.S. Dept. of Agriculture, Systematic Entomology Laboratory,
Washington, D.C., for confirming the identity of the species, and Larry Corpus, Dept.
of Entomology, Mississippi State University, for preparation of Fig. 1. Research supported
in part by National Science Foundation Grant no. BSR-8501212 and Mississippi Agri-
cultural and Forestry Experiment Station Projection no. J-6933.

LITERATURE CITED

CLARKE, J. F. G. 194la. Revision of the North American moths of the family Oeco-
phoridae, with descriptions of new genera and species. Proc. U.S. Natl. Mus. 90:33-
286.

1941b. The preparation of slides of the genitalia of Lepidoptera. Bull. Brooklyn

Entomol. Soc. 36:149-161.

1942. A synopsis of the genus Anoncia, with one new species (Cosmopterygidae
[sic]; Lepidoptera). Can. Entomol. 74:17-19.

HopceEs, R. W. 1962. A revision of the Cosmopterigidae of America north of Mexico,
with a definition of the Momphidae and Walshiidae (Lepidoptera: Gelechioidea).
Entomol. Americana 42(N.S.):1—166.

1978. Gelechioidea (in part), Cosmopterigidae. In Dominick, R. B., T. Dominick

and C. R. Edwards (eds.), The moths of America north of Mexico. Fase. 6.1. E. W.

Classey Ltd. & the Wedge Entomol. Res. Found., London. 166 pp.

[ed.]. 1983. Checklist of the Lepidoptera of America north of Mexico. E. W.
Classey Ltd. & the Wedge Entomol. Res. Found., London. 284 pp.

KORNERUP, A. & J. H. WANSCHNER. 1978. The Methuen handbook of colour. 3rd ed.
Methuen, London. 252 pp.

Received for publication 23 May 1988; accepted 9 November 1988.

Journal of the Lepidopterists’ Society
43(2), 1989, 114-119

A NEW SUBSPECIES OF NEONYMPHA MITCHELLII
(FRENCH) (SATYRIDAE) FROM NORTH CAROLINA

DAVID K. PARSHALL
4424 Rosemary Pkwy., Columbus, Ohio 43214

AND

THOMAS W. KRAL
Box 349, Necedah, Wisconsin 54646

ABSTRACT. In 1983, a colony of Neonympha mitchellii was discovered in south
central North Carolina. Before 1983, mitchellii was known only from Michigan, Indiana,
Ohio, and New Jersey. The newly discovered population co-occurs with Neonympha
areolatus (J. B. Smith). Comparison of 60 male genitalia of mitchellii from Michigan
and New Jersey and 20 from North Carolina with 60 of areolatus showed that the two
have distinct valvae and are separate species. Comparison of 200 Michigan and New
Jersey mitchellii with 47 North Carolina mitchellii revealed several population differ-
ences. The North Carolina population is here named N. m. francisci. Both nominate and
new subspecies are in need of conservation.

Additional key words: Neonympha mitchellii francisci, systematics, biology, endan-
gered, habitat.

Neonympha mitchellii French (1889) is one of the most restricted
butterflies in the eastern U.S. Its known range before 1983 was Michigan,
Indiana, Ohio, and New Jersey (Opler & Krizek 1984). On 2 June 1983,
Kral discovered a small colony of mitchellii on Fort Bragg Military
Reservation, Fort Bragg, North Carolina. This discovery was both a
new State record and a significant extension of known range.

There has been confusion as to whether Neonympha mitchellii and
N. areolatus are distinct species (Scott 1986, Mather 1965). Wing mac-
ulation characters are not always reliable. There are several populations
in North Carolina where some N. areolatus have round hindwing ocelli
much like typical N. mitchellii. Such a population is adjacent to the
habitat of the North Carolina N. mitchellii population, and there are
similar phenotypes in other N. areolatus populations such as in Gates
Co., North Carolina. The problem of identification is obviously greatest
where the ranges of the two species overlap. This confusion is resolved
here by genitalic structure. Male genitalia of 60 nominate mitchellii
from Michigan and New Jersey and 20 from North Carolina were
compared with 60 of areolatus, with results as follows. The distal process
of the areolatus valva has a distal toothlike process that projects sharply
dorsad, while that of mitchellii has a distal process that projects laterad
and is denticulate (Fig. 1).

Comparison of North Carolina mitchellii with Michigan and New
Jersey mitchellii revealed several population differences (Table 1). Be-

VOLUME 43, NUMBER 2 1B 5)

_ 2 ages
es

Fic. 1. Top left and right lateral views of Neonympha male valvae. Left, N. mitchellii,
Jackson Co., Michigan; right, N. areolatus, Fort Bragg, North Carolina.

low we describe and name the North Carolina population. Parenthetical
color names are based on Ridgeway (1886).

Neonympha mitchellii francisci Parshall & Kral, new subspecies

(Fig. 2, Table 1)

Description (male holotype). Left forewing length 17.0 mm (for all males, mean 16.7
mm, range 15.0-18.0 mm, N = 85). Dorsal wing surfaces uniform chocolate-brown
(vandyke brown) except for apex and outer margins. Outer margins and forewing apex
have modified hairlike scales slightly lighter brown than rest of dorsal surfaces, resulting
in lighter brown fringe along outer margins, and much wider (1 mm) band of light-
brown buff scales (drab brown) forming a second submarginal band which follows contour
of forewing margin. Second submarginal band lighter brown than fringe scales or rest of
ground scales of dorsal wing surfaces. A line of dark brown scales identical to first completes
submarginal bands. Dorsal submarginal bands viewed together are widest on hindwing
surfaces, forming uniform width of nearly 2 mm. Hindwing inner margin covered with
light brown scales from inner margin vein 2A to submarginal band at anal angle. It then
flows basally, completing the triangular light brown area. Rest of dorsal surface uniformly
unmarked.

Ventral surfaces of both wings light brown, lighter than dorsal surfaces but not as light
as ventral surfaces in nominate N. mitchellii. Fringes of apex and outer margin much
darker than dorsal surfaces, slightly contrasting to lighter ground color of these surfaces.
Outer marginal fringe followed by a rufous (ochraceous-rufous) submarginal band. This
band, 0.75 mm wide, begins along costal margin, closely following contour of outer margin,
ending at inner margin. Proximad to this band is a very thin submarginal line of dark
brown scales which follows entire length of much wider rufous band. A second less rufous
band follows lighter band. Second proximad rufous band is thinner than first and follows
outer margin contour beginning subapically at vein R; and ending at inner margin. Three
submarginal bands together are ca. 1.5 mm wide.

Forewing postmedial area has row of 4 ocelli in cells M,, M;, M;, and Cu,. Ocelli in
M, and M, largest and most developed. Ocellus of cell M, only faintly present. All 4 ocelli
have silvered pupils which are a series of flat, clear scales with silver sheen. Each 8 fully
developed ocelli have a thin ring of yellow buff scales with interior ground of black with
silver pupils. Ocellus in cell M, is largest.

Forewing with 2 medial transverse bands, 1 barely extracellular, the other transcellular,
both darker brown than rufous. Extracellular transverse band begins subapically at vein
R, and meets 2nd submarginal band to form continuous band. Extracellular band then
flows diagonally to vein M, ca. 2 mm from junction of M, and discal cell, ending vertically
at inner margin at vein 2A. This medial line forms closure around postmedial row of
ocelli open at inner margin. Second or transcellular line parallels path of first, ending

116 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Comparison of Neonympha mitchellii subspecies.

m. francisci (North Carolina) m. mitchellii (Michigan & New Jersey)
Character P) iMale 0 | Siemalennne Male Female

Left forewing length

“Mean + SD IOs as Us) 18.8 + 0.8 164+ 0.75 18.3 = 05

Range 15.0-18.0 17.5-20.1 16.0-19.0 18.0-24.0

N 395 2 44 3]
No. forewing ocelli

Mean + SD 3.3 + 0.7 4.0 + 0.65 O.1) = 0 3.9 = 0:7

Range 2-5 3-5 2-4 5-6

N 35 12 44 31
No. hindwing ocelli

Mean + SD 5D +016) onfo=n04 5.5 + 0.55 S905

Range 4-6 5-6 5-6 5-6

N 30 12 44 31
8rd & 4th hindwing ocelli bifid

% 2

N 47 200
3rd & 4th ocelli set at oblique angle*

% 74 100 18 20

N 35 V2 100 100
Ocelli ring Thin, not very contrasting Thick, contrasting with

with ground color ground color
Ventral wing color Not brightly contrasting Brightly contrasting with
with dorsal surface dorsal surface

Medial transverse bands More rufous than brown,
thin, and contrasting with Dark brown, contrasting less

ground color (98% of with ground color (90%
sample) of sample)
N 200 47
Voltinism Bivoltine Univoltine
Habitat Treed fen Tamarack bog

* Frequencies underlying percentages differ between subspecies for both males and females (P < 0.01, 2 x 2 contingency
tables, adjusted G-test).

near inner margin. Distance between the two parallel bands is visually uniform width of
3.75 mm. Ventral forewing outer margin fringed as dorsally. Two submarginal rufous
bands separated by light band of ground scales. Rufous bands of hindwing larger and
more rufous than forewing.

Six postmedial hindwing ocelli arranged in curved pattern mirror contour of suis
margin. Cells R;, M,, Mz, M,, Cu,, Cu, have ocelli. Ocelli of cells R;, M, greatly reduced
but retain silvered pupils. Ocelli in cells M,, M, largest, best developed. All ocelli ovoid
and pointed distally. Third and 5th ocelli of cells M,, Cu, nearly bifid with double silvered
pupils. Fourth ocelli in cell M, double-pupiled but not bifid. Each ocellus with a thin
yellow buff circulus as in forewing. Ocelli of cells M,, M, set at oblique angle, pointing
distally away from each other at 60°; ocellus of M, pointing in anterior direction, ocellus
of cell M, pointing in posterior direction.

Types. Holotype male, Fort Bragg, North Carolina, 21 August 1984, in U.S. National
Museum, Washington, D.C.; 46 male and 13 female paratypes in collections of American

VOLUME 438, NUMBER 2 £57

Fic. 2. Neonympha mitchellii. A, N. m. francisci female paratype, Fort Bragg, North
Carolina, 31 August 1984. B, N. m. francisci holotype male. C, N. m. mitchellii female,
Jackson Co., Michigan, 5 July 1984 leg. D. K. Parshall. D, N. m. mitchellii male, Springdale,
New Jersey, 12 July 1970 leg. W. B. Wright Jr. E, N. m. francisci type locality. F, N.
m. francisci paratype showing hindwirg with oblique ocelli.

Museum of Natural History, New York; private collections of Thomas W. Kral, Richard
Anderson of Gainesville, Florida, and David Parshall.

Etymology. We name the new subspecies in honor of Saint Francis of Assisi, known
for kindness to animals and a love of natural beauty.

DISCUSSION

Paratype males differ little from the holotype male. Female paratypes
are larger and more variable than male paratypes, with ventral ground
color lighter in females (Table 1).

Subspecies francisci differs from the nominate in several ways (Table
1). Dorsal surfaces contrast less with ventral in francisci males and
females; sexual dimorphism in number of forewing ocelli is less pro-

118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

nounced in francisci; francisci females are darker ventrally, with less
round ocelli distally more pointed; the yellow ring of ocelli is thinner
and contrasts less with ground color; third and fourth ventral hindwing
ocelli are occasionally (2%) bifid; and ventral forewing and hindwing
transverse bands are more rufous and contrast more with ground color.

The habitat of N. m. francisci is quite different from that of nominate
mitchellii (Fig. 2). McAlpine (1986) described the nominate type lo-
cality in Cass Co., Michigan, as grassy glades along streams running
through a tamarack bog. Pallister (1927) and Rutkowski (1966) gave
much the same description for Ohio and New Jersey habitats.

The North Carolina habitat is a treed fenlike area surrounded on
three sides by thickly forested sandhills. The colony occurs along an
apparently spring-fed stream where succulent growth of sedges and
grasses has developed in openings of a few meters. The surrounding
sandhill forest is composed mainly of Magnolia grandiflora L., M.
virginiana L., Sassafras albidum (Nutt.) Nees, Carya cordiformis
(Wangenh.) K. Koch, and Pinus taeda L., with an understory of fern
and Arundinaria tecta (Walkt.) Muhl., Vaccinium coymbosum L., and
with thick alder often choking the water course.

Only a few butterfly species are on the wing in this habitat at the
same time as Neonympha m. francisci. The most notable is N. areolatus
which begins flight at the end of the flight of N. m. francisci and, like
the latter, is bivoltine. Flight dates for N. m. francisci are 5 May to 6
June, and 26 July to 21 August. Flight dates for N. areolatus at Fort
Bragg are 30 May to 28 June, and 15 August to 8 September (1983-
86).

Subspecies francisci is isolated from all known nominate mitchellii
populations. The North Carolina population is bivoltine; all nominate
populations are univoltine, peaking around the first week in July. Iso-
lation of the North Carolina population may represent a preglacial
distribution and adaptation, or a post-Wisconsin isolation and adapta-
tion. Other colonies may exist south of North Carolina, and might yield
evidence to support a southern preglacial origin of mitchellii.

The North Carolina population of mitchellii is small, with less than
100 adults produced per season, but seems secure for the short term
because of its isolation on Fort Bragg away from the public. The Ohio
population is likely extinct (Shuey et al. 1987), and the small New Jersey
population’s status is unclear. In Indiana, the known range is greatly
restricted but may be somewhat protected because it still occurs in a
few State parks and preserves (Shull 1987). In 1987, Michigan placed
N. mitchellii on the State list of threatened and endangered species,
making the collection of mitchellii unlawful without a permit (Michigan
Public Act 203 of 1974 rules as amended effective 4 Sept. 1987). While

VOLUME 43, NUMBER 2 119

this gives the species some protection in Michigan for the short term,
it limits study of the insect to those who seek a permit, and does not
protect its habitat for the long term.

There is not a more endangered butterfly population in the eastern
U.S. than N. m. francisci. Because of its small population and uncertain
future over the long term on military lands, this butterfly will need
conservation. We hope in naming this unique population that more
field research will be generated, and that this attention will lead to real
protection of not only the colony at Fort Bragg but colonies elsewhere
as well.

ACKNOWLEDGMENTS

We thank R. A. Anderson for sharing his knowledge and help in acquiring material
for study; L. D. Miller for the original determination of the North Carolina mitchellii
specimen; the late Leland Martin for his much valued companionship in the tick-infested
swamps of North Carolina; D. C. Ferguson of the U.S. National Museum for allowing us
to examine material; J. A. Scott, D. C. Iftmer and E. H. Metzler for critically reviewing
the manuscript; M. C. Nielsen for much interesting discussion and help; and Leni Wils-
mann of the Michigan Department of Natural Resources for help.

LITERATURE CITED

FRENCH, G. H. 1889. A new species of Neonympha. Can. Entomol. 21:25-27.

MATHER, B. 1965. Euptychia areolatus: Distribution and variation with special refer-
ence to Mississippi (Satyridae). J. Lepid. Soc. 19:139-160.

MCALPINE, W. S. 1936. Habitat of Cissia mitchellii in Cass County, Michigan. Bull.
Brooklyn Entomol. Soc. 31:110-221.

OPLER, P. A. & G. O. KRIZEK. 1984. Butterflies east of the Great Plains. Johns Hopkins
Univ. Press, Baltimore. 294 pp.

PALLISTER, J.C. 1927. Cissia mitchellii (French) found in Ohio with notes on its habitats.
Ohio J. Sci. 27:203-204.

RIDGEWAY, R. 1886. A nomenclature of colors for naturalists. Little Brown, Boston.
129 pp.

RUTKOWSKI, F. 1966. Rediscovery of Euptychia mitchellii in New Jersey. J. Lepid. Soc.
20:43-44.

ScoTT, J. A. 1986. The butterflies of North America. Stanford Univ. Press, Stanford,
California. 658 pp.

SHUEY, J. A., E. H. METZLER, D. C. IFTNER, J. V. CALHOUN, J. W. PEACOCK, R. A.
WATKINS, J. D. HOOPER & W. F. BABCOCK. 1987. Status and habitats of potentially
endangered Lepidoptera in Ohio. J. Lepid. Soc. 41:1-12.

SHULL, E. M. 1987. The butterflies of Indiana. Indiana Univ. Press, Bloomington.
262 pp.

Received for publication 10 February 1988; accepted 16 December 1988.

Journal of the Lepidopterists’ Society
43(2), 1989, 120-146

REVISION OF CHLOROSTRYMON CLENCH AND
DESCRIPTION OF TWO NEW AUSTRAL
NEOTROPICAL SPECIES (LYCAENIDAE)

KURT JOHNSON

Department of Entomology, American Museum of Natural History,
Central Park West at 79th Street, New York, New York 10024

ABSTRACT. Neotropical Chlorostrymon Clench is revised to comprise six species,
including the austral C. patagonia, new species (Patagonian Steppe biotic province,
Argentina); C. chileana, new species (Central Valley biotic province, Chile); and C.
kuscheli (Ureta), new combination (N Andean Cordillera-High Andean Plateau biotic
provinces, Chile). Three additional congeners are C. simaethis (Drury) (Thecla simaethis
jago Comstock & Huntington, new synonym, Antilles; C. s. sarita (Skinner), C. s. rosario
Nicolay, new synonym, mainland Neotropics), C. telea (Hewitson) (Central and South
America), and C. maesites (Herrich-Schaeffer) (Antilles). Differentiating characters in-
clude female genitalia. The Andes have produced three distinctive species isolated in
high montane and austral regions.

Additional key words: Eumaeini, systematics, biogeography.

Chlorostrymon was erected by Clench (1961) to include three fa-
miliar and widely distributed New World hairstreaks: C. simaethis
(Drury), C. telea (Hewitson), and C. maesites (Herrich-Schaeffer). Sub-
sequently, Nicolay (1980) elucidated the original generic description,
and Clench (1963) further distinguished the Antillean species. Chlo-
rostrymon species are distinctly marked, and aside from the naming
of some subspecies (Skinner 1898, Stallings & Turner 1947, Comstock
& Huntington 1943, Nicolay 1980), the genus has appeared to be one
of the best known and taxonomically stable in Eumaeini (Nicolay 1980).

I have assembled and studied eumaeine samples from the austral
Neotropics (Johnson et al. 1986, 1988, Johnson 1987, 1989, Johnson in
press). These specimens derived from unsorted and unincorporated
material principally at the British Museum (Natural History) (BMNH),
and Museum National d’Histoire Naturelle, Paris (MNHN). Specimens
were also provided by the Central Entomological Collection, University
of Chile, Santiago (CECUC), and the Museo Nacional de Historia Nat-
ural, Chile, Santiago (MNHNC).

Three distinctive austral South American members of Chlorostrymon
are apparent: Thecla kuscheli Ureta (1949), hitherto not examined by
northern workers, and two new species. Unique characters in these
austral populations require expansion of Nicolay’s (1980) redescription
of the genus. I revise Chlorostrymon to comprise six species, including
these newly discovered austral ones.

Because of peculiar intraspecific variation, there is little utility in
pursuing extensive subspecific division of the three familiar Chloro-
strymon species (Nicolay 1980). Accordingly, I synonymize some sub-

VOLUME 438, NUMBER 2 121

species. I reduce subspecies in C. simaethis to two (Antillean and main-
land, consistent with the distribution of C. maesites and C. telea), and
reduce subspecies in C. maesites to the nominate. I treat C. maesites
and C. telea as species based on their traditionally cited features (Com-
stock & Huntington 1948, Klots 1951, Clench 1961, 1964, Riley 1975,
Thorne 1975, Pyle 1981, Opler & Krizek 1986, Scott 1986) as well as
a statistically significant difference in their female genitalia.

METHODS AND MATERIALS

I follow Clench (1961) in abbreviating dorsal hind- and forewing to
DHW and DFW, respectively, and ventral hind- and forewing to VHW
and VFW,, respectively. I also use TL for type locality.

Distribution data derive from specimens at the Allyn Museum of
Entomology—Florida State Museum (AME), American Museum of Nat-
ural History (AMNH), BMNH, Carnegie Museum of Natural History
(CMNH), CECUC, Instituto Miguel Lillo (Tucuman, Argentina) (IML),
Milwaukee Public Museum (MPM), MNHN, and MNHNC. To study
consistency of morphological characters, I dissected genitalia of males
and females from localities spanning distribution of each taxon, as well
as more extensive series available from particular sites. Such material
is listed for each taxonomic entry.

Chlorostrymon Clench
(Figs. 1-6)

Chlorostrymon Clench (1961:189). Clench (1963:248; 1976:269; 1977:186), dos Passos
(1970:27), Brown & Heineman (1971:4; 1972:230), Emmel & Emmel (1973:51), Ferris
(1974:278), Riley (1975:100), Thorne (1975:277), Ross (1976:188), Nicolay (1980:253),
Miller & Brown (1981:99; 1983:54); Pyle (1981:464), Schwartz & Jimenez (1982:8),

Garth & Tilden (1986:189), Opler & Krizek (1986:88), Scott (1986:359), Llorente-
Bousquets et al. (1986:25), Schwartz (1989).

Diagnosis. In wings (Figs. 1-3), DFW and DHW are variously iri-
descent blue to violet like many Eumaeini (though lacking DFW male
androconia as in some Eumaeini), but Chlorostrymon is distinctive in
its brilliant green (often chartreuse) ventral ground color; VHW with
brilliant white to silver bands (usually across entire wing) and lavish
reddish brown or gray suffusion across limbal area; and VFW post-
median silver-white or blackish bands. These markings are distinctive
in overall pattern regardless of occasional reduction, and can be con-
fused only with Cyanophrys crethona (Kaye), as discussed later under
Chlorostrymon simaethis. Chlorostrymon genitalia (Figs. 5, 6) differ
from other Eumaeini by the male aedeagus having its terminus sepa-
rated from the rest of the shaft by a transparent juncture, but conjoined
internally by the elongate, pointed cornutus, as discussed further on.

Type species. Papilio simaethis Drury (1778) by original description.

122 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fic. 1. Adult male Chlorostrymon simaethis (left, upper surface; right, under surface)
(AMNH except as noted). A, C. s. simaethis, proximate topotype (TL given as generalized
locality), Basseterre, St. Kitts. B, Thecla s. jago, holotype. C, C. s. sarita, proximate
topotype, San Antonio, Texas, 29 October 1933. D, C. s. sarita, Caripito, Venezuela, 1
July 1913. E, C. s. sarita, Callao, Lima Department, Peru (BMNH). F, C. s. sarita, Arroyo
San Pedro, Jujuy Province, Argentina, 17 July 1978.

VOLUME 43, NUMBER 2 123

Fic. 2. Adult female Chlorostrymon telea and C. maesites (left, upper surface; right,
under surface) (AMNH except as noted). A, C. telea, proximate topotype, Obidos, Ama-
zonas State, Brazil. B, C. telea, Villa Ana, Santa Fe Province, Argentina, 26 February
1927 (BMNH). C, C. maesites, proximate topotype, Guantanamo Bay, Cuba. D, Thecla
m. clenchi, allotype.

Diversity. Previously comprising the species simaethis, telea, and
maesites; hereafter, these species, kuscheli, and the two new species.
All are distinguished in the following key. A key character is not con-
sidered “distinctive’’ if obscure.

Wing Character Key to Species

1 VHW postdiscal band distinctive across @mntire Wing eee eee eee 2
VHW postdiscal band distinctive only costad vein M, or caudad vein M, 3
2m EN Vepostmedian line: white or silver 2.00 5
VFW postmedian line black (without white) or faint to absent 4
3 VHW postdiscal band distinctive only costad vein M, —. chileana, new species
VHW postdiscal band distinctive only caudad vein M, and with the costal fold of
forewing extremely wide (=1 mm) and rufous colored ....... patagonia, new species
4 VHW limbal patch extending costad to M,; postmedian line forming a distinct
“W”; VFW postmedian line very faint to absert eee telea

VHW limbal patch extending costad to M,; postmedian line not forming a “W’;
MENWVepostmedianvlime iolack ... = as st) RN maesites

124 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fic. 8. Adults of austral South American Chlorostrymon (left, upper surface; right,
under surface). A, C. patagonia, allotype male. B, C. patagonia, holotype female. C, C.
chileana, allotype male. D, C. chileana, holotype female. E, C. kuscheli, paratype male.
F, C. kuscheli, female, data in text.

5 VFW, VHW band wide, silver-white; DHW with uniformly colored ground ........
siete ate PR OS ot CIM, hal) SOE SAE? OL IRED TOS PCAN) simaethis

VFW, VHW band extremely thin, white, DHW with bright rufous limbal patch
ee 2 Be EES ET GRY a 8 A eae eee Seen kuscheli, new combination

Wing characters are correlated with characters of the female genitalia. Since the latter
have not been previously treated, the following key uses features of the ductus bursae
and cervix bursae.

VOLUME 43, NUMBER 2 125

Female Genitalic Key to Species

1 Ductus bursae cephalically inclined 60—90°; cervix bursae with two dorsal sclero-
Bezecmpads (Ee OAL) sure a ht in ee ieee ee simaethis

Ductus bursae inclined <60° or not at all; cervix bursae without sclerotized pads
OOS onan @ C(O) eres eee rc Ny OM NC oneal lee An aes 2

2 Ductus bursae with cephalic tubular ductus and caudally flared antrum (Fig. 6G-
1) ecsercennieh ec vesceeainas Mat cd St.) Cian AC URMn Ue ane ueh ew aouoey, i rT ore aD Aa pallid aR 3

Ductus bursae with caudally flared antrum only, antrum connected directly to
eorgoseoulusaes (tig aN, (©) mares slat fo el i eel Peel lie Oe 4
Se Muewissounsae inelined 30-60% sao lah Ne ae 5

Ductus bursae not inclined or inclined <20°; length of antrum less than length of
Ghurestirsmaunesaen (Lig eG bar 1). tisrse ls Wl eee eee Ce be testi ee) maesites

4 Terminus of antrum only slightly flared; lamella postvaginalis parabolic (Fig. 6O)

vranccesoesamsnnt nena elle ten UO ET ale ear Al re aed SS ee chileana, new species
Terminus of antrum greatly flared; lamella postvaginalis distally lobate (Fig. 6N)
sreeoestolecs ent ah a Na RC SN ae RD RTOS OYTO SC OR patagonia, new species
Length of antrum (Fig. 6A) equalling or exceeding length of ductus bursae (Fig.
CO ear Heat ae ee tet ed eC a Ne telea
Length of antrum less than length of ductus bursae (Fig. 6M)
ere SWE Lc) Sets shales Neuer Seis aegis kuscheli, new combination

Ol

Distribution (Fig. 4). Extreme southern United States (S Florida, S
Texas to Baja California and neighboring areas), Greater and Lesser
Antilles and Mexico, S through South America to northern and central
Chile and northern Patagonia.

Characters. Along with the distinctive wing pattern, Nicolay (1980)
proposed several diagnostic genitalic characters for Chlorostrymon.
From my analysis of 121 male and 138 female genitalia, one character
of male genitalia appears common to all Chlorostrymon taxa: separation
of aedeagal terminus and shaft by a stripe of transparent sclerotization
conjoined internally by the single elongate cornutus (Nicolay 1980:225)
(Fig. 5). Because of great structural divergence of male genitalia in
Eumaeini, other male genitalic characters of Chlorostrymon appear
less diagnostic.

In the female genitalia, Nicolay distinguished Chlorostrymon from
other Eumaeini by the cephalic one-quarter of the ductus bursae dor-
sally inclining ca. 90°, and by two sclerotized pads occurring on the
dorsum of the cervix bursae (Fig. 6A-F). However, my samples indicate
that only C. simaethis, the type species, has these characters.

Phylogenetic position. Search for the sister group of Chlorostrymon
(Hennig 1966) appears difficult and will probably be resolved only by
further study of the many undescribed Eumaeini. Considering Chlo-
rostrymon’s distinctive wing characters, and the wide geographic range
of its two sympatric congeners, the genus is probably very old. The
distinctive Chlorostrymon aedeagus may occur in other as yet unde-
scribed or unstudied Eumaeini, and be a key to recognizing the out-
group. In other respects, male and female Chlorostrymon genitalia
resemble taxa of Electrostrymon Clench (type species Papilio endym-

126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

B

KEY

Numbered Circles--
sympatric localities
Cc. simaethis and Cc.
telea or C. maesites
(see text)

[4 c. simaethis simaethis
Cc. simaethis sarita

- maesites
- telea

- patagonia
- chileana
- kuscheli

onmeco#
220200

Fic. 4. Geographic ranges of Chlorostrymon species. A, United States and northern
Mexico distributions of C. simaethis (vertical hatches), C. telea (stippled), and C. maesites
(black), from localities assembled by Scott (1986) and Opler and Krizek (1986). B, Neo-
tropical and austral distributions (specimens in AMNH except as noted). Numbers 1-27:
sympatric Neotropical localities of C. simaethis and C. telea (mainland) or C. maesites
(Antilles). Only localities not mentioned in text are given here. 1, San Francisco, Ta-
maulipas State, Mexico. 4, Guatemala City, Guatemala. 5, Havana, Cuba. 6, Guantanamo,
Cuba. 9, Turrialba, Costa Rica. 10, Coamo Springs, Puerto Rico. 11, St. Vincent, Lesser
Antilles. 12, Port-of-Spain, Trinidad. 15, Barta District, Guyana. 17, Pernambuco, Brazil
(AMNH, BMNH). 19, Espirito Santo, Brazil (MPM, BMNH). 20, Callao, Lima Depart-
ment, Peru (BMNH). 22, Campo Grande, Mato Grosso State, Brazil (BMNH). 27, Obidos,
Amazonas State, Brazil. Numbers 28-30: austral species distributions and biotic provinces
(BP): 28 & cross hatches, C. chileana, Cental Valley BP. 29 & stippling, C. patagonia,
Patagonian Steppe BP. 30 & vertical hatches, C. kuscheli (dark hatches, Northern Andean
Cordillera BP; light, High Plateau BP).

VOLUME 438, NUMBER 2 127

ion Fabricius). However, Electrostrymon has not been revised and,
since Clench (1961, 1963) never listed the taxa it included, the genus
has been subject to various interpretations (Barcant 1970, Riley 1975,
Johnson & Matusik 1988). Johnson and Matusik (1988) suggested the
Barcant and Riley treatments of Electrostrymon were, at least, diphy-
letic. To complicate matters, the short, non-inclined female genitalia
of Chlorostrymon maesites and the new austral Chlorostrymon species
(Fig. 6H, I, M-O) resemble those of at least two groups of Eumaeini
whose wing patterns differ greatly: the taxa-rich “Thecla celmus’’ and
“Thecla phrutus” groups (Johnson 1986, Johnson & Matusik 1988), and
Parrhasius Hubner and Michaelus Nicolay (Nicolay 1979). Without a
designated outgroup, or a basis for describing outstate characters, phy-
logenetic inference concerning Chlorostrymon taxa is too speculative.

Conspecificity of Chlorostrymon telea and C. maesites. This has
been much debated (Riley 1975, Clench 1961, Opler & Krizek 1986,
Scott 1986). Some early workers, and recently Scott (1986), proposed
the synonymy of the two taxa. Genitalic study indicates that, along
with traditionally cited pattern differences, female genitalia of the two
taxa differ consistently and distinctly (Female Genitalic Key, Fig. 6,
and discussion under C. telea). Accordingly, these taxa are treated as
distinct species here.

Temporal and spatial distribution. Rarity of Chlorostrymon taxa,
compared to many other hairstreak butterflies (Opler & Krizek 1986),
is reflected in museum samples. There is a correlation between Chlo-
rostrymon occurrence (particularly sympatry of C. simaethis with C.
telea or C. maesites) and location of major collectors. Study of such
samples indicates Chlorostrymon taxa are local, but sometimes locally
common. Major historical sources of Chlorostrymon specimens warrant
mention because they explain the origin of most data on the genus, and
have allowed study of C. simaethis and C. telea or C. maesites from
numerous areas of sympatry (Fig. 4). Such collectors, common collecting
localities, and depositories are listed with Fig. 4 locality numbers as
follows:

(2, 3) Mexico: Presidio, Cordoba, Vera Cruz State, Colima, Colima State; N. Hoffman
(AMNH). (7) Jamaica: B. Heineman (AMNH). (8) Hispaniola: A. Schwartz (private, AME),
D. Matusik (private, FMNH, AMNH, CMNB), K. Johnson (AMNH, AME). (12) Trinidad-
Tobago: R. Rozycki (AMNH). (13) Panama: collections of AMNH Research Station (AMNH).
(15) Venezuela: Caripito; collections of New York Zoological Society (AMNH). (16) French
Guiana: northern localities; expedition collections of MNHN (MNHN). (18) Brazil: Minas
Gerais; collections of J. F. & W. Zikan (Instituto Oswaldo Cruz, Guanabara). (21) Bolivia:
eastern localities; J. Steinbach (CMNH, BMNH). (23) Paraguay: Santissia-Trinidad; B.
Podtiaguin (AMNH). (24) Brazil: Rio de Janeiro vicinity; P. Gagarin (MPM). (25, 26)
Brazil: Curitiba, Parana State, Pelotas, Rio Grande del Sol State; C. Biezanko (BMNH,

AMNH). (29) Argentina: northwestern localities; R. Eisele (AME, AMNH), B. MacPherson
(AMNH), K. Hayward and N. Giocomelli (BMNH, IML). Austral South America: Pata-

128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

gonia; C. Larsen (MNHN). (28) Chile (Valparaiso, Santiago); R. Martin (MNHN). General:
collections of CECUC, MNHNC.

Chlorostrymon simaethis (Drury)
(Figs. 1, 5A-D, 6A-F)

Papilio simaethis Drury (1773:3; 1770:pl. 1, fig. 3) (mame given in index published in
LAWS).

Mitoura simaethis: Dyar (1903:38).

Tmolus simaethis: Kaye (1914:567).

Chalybs simaethis: Kaye (1921:103), Barcant (1970:251).

Thecla simaethis: Kirby (1871:398), Dewitz (1877:233, pl. 1), Godman & Salvin (1879-
1901:720, pl. 81), Moeschler (1889:301), Stahl (1882:93), Gundlach (1887:622), Draudt
(1919:798, pl. 158,f), Barnes & Benjamin (1923:17), Hall (1925:188; 1936:276), Kaye
(1926:462), Holland (1931:232), Wolcott (1936:403), Hoffman (1941:716), Schweizer
& Webster Kay (1941:19), Comstock & Huntington (1943:58, 78; 1961:54; 1963:190),
Beatty (1944:157), Comstock (1944:488), Avinoff & Shoumatoff (1946:284), Zikan &
Zikan (1968:57), Hayward (1973:157).

Thecla lycus Skinner (1898:48) (takes authorship of “lycus Hiibner” Skinner 1898:48)
(misspelling, misattribution of author, not licus Hiibner 1807:pl. 150, not lydus Hub-
ner 1818:75, no. 753), Dyar (1903:36), Draudt (1931:798), Barnes & McDunnough
(1917:18) (all follow Skinner, in error), Barnes & Benjamin (1923:17), Comstock &
Huntington (1958-64 [1963]:190) (both cite Skinner 1898 as an error).

Thecla sarita Skinner (1895:112; 1898:48), Barnes & McDunnough (1917:3), Barnes &
Benjamin (1923:17), McDunnough (1938:24), Stallings & Turner (1947:40) (the last
make sarita a subspecies of simaethis).

Strymon simaethis: Barnes & McDunnough (1917:13), Bates (1935:194), McDunnough
(1938:24), Stallings & Turner (1947:40), Klots (1951:139), Ziegler 1961:22 (as “Stry-
mon’), Lipes (1961:56), dos Passos (1964:56), Lewis (1974:67).

Chlorostrymon simaethis: Clench (1961:189; 1964:248; 1976:269; 1977:192), dos Passos
(1970:27), Brown & Heineman (1971:230), Emmel & Emmel (1973:51), Riley (1975:
100), Thorne (1975:277), Ross (1976:188), Nicolay (1980:253), Miller & Brown (1981:
99; 1983:54), Pyle (1981:465), Schwartz & Jimenez (1982:8), Garth & Tilden (1986:
189), Opler & Krizek (1986:89), Scott (1986:359), Llorente-Bousquets et al. (1986:
25), Schwartz (1989).

Diagnosis. DFW, DHW brown, strongly suffused with iridescent
purple (much duller on females); VF W, VHW ground chartreuse, VHW
postdiscal band white to pearly white, distinctive across entire wing;
VFW postmedian line, thick, white or pearly white (Fig. 1). Male
genitalia with vincular arc, valvae, saccus and aedeagus all more elon-
gate than congeners (and aedeagus not ventroterminally declined) (Fig.
5A-D); brush organs moderately dense, articulated to small basal mem-
brane along each ventrocephalic edge of vincular arc (Fig. 5J); female
genitalia with ductus bursae cephalically inclined 90°, cervix bursae
with two dorsal sclerotized pads, papillae anales terminally lobate,
apophyses of papillae anales short (not extending entire length of ductus
bursae) (Fig. 6A—F).

Types. Papilio simaethis type reported lost (Miller & Brown 1981);
TL “St. Christopher’s”’ (=St. Kitts, Riley 1975). Chlorostrymon simae-
this is unambiguous in facies; there is no need for a neotype. Thecla
lycus type and TL unknown (Comstock & Huntington 1961).

VOLUME 43, NUMBER 2 129

Distribution. In United States, extreme S California, Arizona, Texas,
and Florida; Baja California, throughout Mexico and Central America,
most of Greater and Lesser Antilles; in South America, SW to central
Peru, E over entire continent except Amazon basin, SE along Brazilian
coast, and W from SE Brazil to Paraguay and E Bolivia, S to NW
Argentina (Fig. 4). Scott (1986) portrays the Baja California distribution
as transient. However, since Opler and Krizek (1986) document recent
establishment of the species in Florida, the large series of specimens
from numerous Baja California locales (AMNH, CMNH) may also rep-
resent resident populations. John Brown (San Diego Museum, pers.
comm.) suspects that marked genitalic variation between Baja Califor-
nia populations may reflect a flux of resident and transient populations.

Superficially similar noncongeners. As noted in Diagnosis, Cyano-
phrys crethona (Kaye) somewhat resembles C. simaethis because both
have lavish VH W limbal suffusion. The former is much larger (forewing
base to apex in the male 15 mm, in the female 17 mm, compared with
12 and 14 mm for simaethis; Riley 1975); its DFW and DHW are deep
iridescent blue with wide black borders; and its VHW and VFW are
deep lime green, with VHW postdiscal band continuous basad discal
cell, disjunct costad.

Variability. Klots (1951) and Nicolay (1980) noted that wing pattern
variability in Chlorostrymon simaethis caused most of its subspecies
to be ill-defined geographically. Except for two major allopatric seg-
regations, subspecies are dropped here by placement in appropriate
synonymies. Below, I summarize these synonymies and the character
variation on which they are based.

1. Nominate C. s. simaethis (Fig. 1A) and Jamaican C. s. jago (Thecla s. jago Comstock
& Huntington, 1943:74, pl. 1, fig. 7; holotype male, Fig. 1B, allotype female, both AMNH,
TL Dunrobin District, Mandeville, Manchester, Jamaica), NEW SYNONYMY.

By virtue of its type locality, nominate Chlorostrymon s. simaethis has historically
been considered restricted to the Antilles. However, while C. simaethis was still poorly
known from the Antilles, Comstock and Huntington (1948) described subspecies jago
from Jamaica. Later, Riley (1975) noted that wing characters of jago duplicated those
of C. simaethis occurring on Hispaniola.

Compared to mainland populations, Antillean C. simaethis display some homogenous
wing characters (Fig. 7A), but Jamaican specimens are no more distinct than other
Antillean populations. Female genitalia of Jamaican specimens have a widely flared,
elongate antrum (Fig. 6A). However, equally distinctive innovations appear in other
Antillean C. simaethis: Virgin Islands specimens show a distinctively narrow and elongate
ductus and antrum; Hispaniola specimens have a markedly constricted genital plate. Male
genitalia also have numerous localized innovations, including a cephalically elongated
vinculum in Jamaica (Fig. 5A), an elongate, narrow, valve in Puerto Rico, and a wide,
blunt-ended valve in Hispaniola. Genitalia of Hispaniolan females most resemble Cuban
females; genitalia of Hispaniolan males most resemble St. Vincent males. Such genitalic
variation in Antillean C. simaethis makes jago appear no more distinct than other An-
tillean populations. Further, there is no distinctive character correlation between C.

simaethis of Jamaica and that of southern Hispaniola, which, if present, would have
biogeographic significance (Schwartz 1989, Schwartz et al. 1984, 1986a, 1986b, Johnson

130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 5. Male genitalia (A-I, N) and brush organs (J-M) of Chlorostrymon. Genitalia
shown in ventral view at left without aedeagus, aedeagus in lateral view at right (specimens
in AMNH unless otherwise noted here or in text). A, Thecla simaethis jago, holotype. B,
C. simaethis sarita, proximate topotype, San Antonio, Texas, 29 October 1933. C, C. s.
simaethis, proximate topotype, Basseterre, St. Kitts. D, C. s. sarita, Caripito, Venezuela.

VOLUME 48, NUMBER 2 131

& Matusik 1988). Description of subspecies from Antillean populations would invite an
inflated trinomial nomenclature on the mainland. Dissections examined (all AMNH). C.
s. simaethis: CUBA: Santiago (2), Havana (6), Guantanomo Bay (6, 2). DOMINICAN
REPUBLIC: El] Numero, Barahona Province, 3 July 1986 (3 6, 2 2). HAITI: Petionville,
9 May 19380 (2), 24-29 January 1922 (¢). LESSER ANTILLES: St. Vincent (6, 2); Dominica,
Canefields, 1-8 December 1933 (4 6, 16 2), October 1919 (2); St. Kitts, Basseterre (6, 2).
UNITED STATES: Puerto Rico, Cuamo Springs, 26 December 1914 (6); Virgin Islands,
St. Croix, 14 March 1951 (5 6, 6 2). C. s. jago: JAMAICA: primary types, paratypes with
same data except 28 December 1919 (6), 28 January 1919 (2), 4, 22, 28 December 1919
(3 2), 4 November 1919 (2), Mt. Diablo, 5 March 1951 (6), Constant Springs, 4 January
1924 (8 Q).

2. C. s. sarita (Skinner 1895:112; holotype male, CMNH, TL Comal Co., Texas) (Figs.
1C, 5B, 6B) and C. s. rosario (Nicolay 1980:254; holotype male and 10 paratypes in AME,
TL La Kenedy, Pichincha, Ecuador; 1 paratype, S. Nicolay Collection, same locality;
allotype female, San Bartolo, Ecuador, AME), NEW SYNONYMY.

Subspecies sarita has usually been characterized by generally straight VHW band
(poised perpendicular or slightly slanted costad FW inner margin), and with discal area
of band sometimes distally produced (Fig. 1). Stallings and Turner (1947) presented data
recommending use of C. s. sarita for populations extending from the SW United States
into Mexico. Subsequently, C. s. sarita was applied southward into Central America
(Llorente-Bousquets et al. 1986) and South America (Nicolay 1980). Nicolay (1980) also
described a new subspecies (C. s. rosario) from then unique Ecuadorian specimens.
Subsequently, however, numerous variable series of C. simaethis have been accumulated
from Ecuador (Banos: AMNH, CMNH; Aguarico, Duran, Latas, Mishahualli: AMNH)
and the species taken southward in Peru (AMNH, BMNBH).

Mainland C. simaethis are generally distinct from Antillean populations (Fig. 7A), and
notably high frequencies of wing characters occur in some regional mainland populations
(Fig. 7A). Though northern populations usually have a more uniform hindwing band, a
few (notably S Texas and insular montane Vera Cruz and Guerrero in Mexico) show
extreme swelling of the discal area of the VHW band. This trait becomes much more
common in South America (Figs. 1D-F, 7A) but the distinction is obviated by blending
in the Panamanian isthmus region.

In contrast to haphazard local genitalic variation in Antillean C. simaethis, genitalic
characteristics in both sexes of mainland C. s. sarita are often regional. Homogeneity is
most common in contiguous lowland regions and appears more varied in disjunct or high
montane areas. Male and female genitalia are most uniform from S Texas S across Mexico
(Figs. 5B, 6B) with variation increasing in specimens from S$ California-Baja California,
and Guatemala S through Panama. Males from S California-Baja California exhibit an
elongate vinculum, and females an unusually wide ductus (width 1% that of lamella;
normally about 4). From Guatemala S through Panama, females are more locally variable
in antrumal width and ductus length, and males more variable in valval width, terminal
recurvature and saccus length. In South America, males have more distally shouldered
valvae which are less terminally elongate or recurved. The last trait is strongest in pop-
ulations from Colombia E across the Guyana shield (Fig. 5D). Populations S of the Amazon
basin, SE Brazil to NW Argentina, show more elongation in the valve but not increased
recurvation. In South American females the antrum is usually outstanding and greatly
flared (most often from about half the distance between the lamellal termini and the

_

E, Thecla telea, holotype. F, C. maesites, Nicoll’s Town, North Andros Island, Bahamas.
G, Thecla maesites clenchi, holotype. H, C. patagonia, allotype (MNHN). I, P. chileana,
allotype (MNHN), with saccal brush organ. Brush organs shown in diagrammatic lateral
view from vinculum to labides including abuttment of anchoring membrane. J, C. si-
maethis. K, C. telea and C. maesites. L, C. patagonia. M, C. chileana (including saccus
and saccal brush organ). N, Aedeagus of C. kuscheli paratype male (CECUC).

132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

-antrum-

5
2)
=)

oe)
16)
=)

%
J

Fic. 6. Female genitalia of Chlorostrymon. Unless otherwise indicated, lateral view
at left; ventral view at center (antrum indicated in A; dorsal suture line, d.s.]., and ductus
indicated in G); lateral view of papillae anales and their apophyses at right (specimens
in AMNH unless indicated otherwise here or in text). A, Thecla simaethis jago, allotype.
B-F, Chlorostrymon simaethis sarita, proximate topotype, San Antonio, Texas, 29 October
1933 (ventral only, with lateral view of cephalic inclination in C, E, F). C, C. s. simaethis,
proximate topotype, Basseterre, St. Kitts. D, C. s. sarita, Caripito, Venezuela. E, C. s.
sarita, Balsapuerto, Huanuco Department, Peru. F, C. s. sarita, La Rioja, La Rioja Prov-
ince, Argentina. G, C. telea, proximate topotype, Obidos, Amazonas State, Brazil. H, C.
maesites, proximate topotype, Guantanamo, Cuba. I, Thecla maesites clenchi, allotype

VOLUME 48, NUMBER 2 133

cephalic area where the ductus is dorsally inclined) (Fig. 6D-F). Variation from this
ground plan is usually limited to difference in ductal length before and after the dorsal
inclination. Most notably, isolated montane Peruvian populations have the ductus more
elongate on both sides of the dorsal inclination (Fig. 6E), and Colombian specimens have
a decreased dorsal inclination. The numerous high montane populations in Ecuador show
extreme local variation.

In conclusion, some mainland regions evidence certain distinctive characters, but overall
contiguity suggests validity of a single name: C. s. sarita. Unless numerous subspecies are
recognized in South America, C. s. rosario is best considered a synonym of C. s. sarita.
Dissections examined (all AMNH). C. s. sarita: ARGENTINA: La Rioja Province, La
Rioja (2 6, 1 2); Salta Province, Yariquarenda (1 6, 2 2), Agua Blanca (2 4, 1 2), Mosconi
(2 6, 2 9), Tartagal (1 6, 1 2), La Merced (1 4, 1 2); Jujuy Province, San Pedro (2 6, 2 9),
Rio Lazares (1 6, 1 2). BOLIVIA: Rio Surutu, 350 m, E Bolivia (1 4, 1 2). BRAZIL:
highlands nr. Massaranduba Blumenau (é); Annaburg, St. Catarina (¢). COLOMBIA:
Cauca Valley, 3200 ft (975 m), 25 January 1935 (3 2). COSTA RICA: Turrialba, 29 May
1946 (6, 2). ECUADOR: Banos, February 1939 (6 6, 2 2); Duran, 400 ft (122 m), 24 June
1914 (1 6, 1 2); Aguarico, November 1979 (8 4, 6 2), Mishahualli (4); Latas, Oriente (¢).
GUATEMALA: Guatemala City (6, 2). GUYANA: “British Guiana” (6); Bartica District,
Bartica (6). MEXICO: Baja California, Arroyo del Refugio, 5 May 1935 (2 6, 3 2); Arroyo
del Rosario, 21 March 1935 (3 2); Cape San Lucas, 24, 26 December 1938, 13 November
1938 (3 2); North End, San Jose Island, 12 December 1938 (2 4, 2 2); Vera Cruz State,
Presidio (6, 2), Jalapa (4 4, 2 2); Colima State, Colima, April 1918 (1 4, 3 2); Tamaulipas
State, San Francisco, August 1964 (6, 2). PANAMA: La Boca, Canal Zone, 25 January
1908 (6, 2). PARAGUAY: Santissima Trinidad, Cordillera Province, June-July (2 4, 2 9).
PERU: Balsapuerto, Paranapura River, Loreto, June 1938 (6); Callao (BMNH) (6, 9);
Chanchamayo, Huanuco (BMNH) (6); Chosica, 850 m, January 1900 (BMNH) (2). TRIN-
IDAD-TOBAGO: Port-of-Spain, 1-9 April 1929 (2 6). UNITED STATES: Texas, Browns-
ville, 30 October 1965 (4 6, 5 2), Pharr (4 2), San Antonio (Comal Co.; TL), 29 October
1938 (6, 2); Arizona, Portal, 10 June 1958 (6); California, San Diego Co., 193- [sic] (2 4,
4 2). VENEZUELA: Caripito (3 4, 3 2); San Felipe Venezuela, 6 May 1938 (8). C. s.
rosario: | saw C. s. rosario type series but did not dissect specimens; I rely on illustrations
of Nicolay (1980) and specimens variously identified as rosario listed above under EC-
UADOR (AMNB).

Chlorostrymon telea (Hewitson)
(Figs. 2A, B, 5E, K, 6G, J-L)

Thecla telea Hewitson (1868:4) (cited by Comstock & Huntington 1958-64 [1964]:123,
as “1868, Specimen of a Catalogue of Lycaenidae in the British Museum, p. 4’,
probably referring to text later published by Classey with preface by L. Higgins; see
Higgins 1972). Kirby (1871:398), Hewitson (1862-78 [1873, February], vol. 1:148,
vol. 2:pl. 57, figs. 350, 351), Dewitz (1877:233, pl. 1), Godman & Salvin (1879-1901:
720), Moeschler (1889:301), Stahl (1882:93), Gundlach (1887:622), Draudt (1919:798,
pl. 158,f), Kaye (1926:462), Barnes & Benjamin (1923:17), Holland (1931:232), Bates
(1935:190), Wolcott (1936:402), Hoffman (1941:462), Beatty (1944:157), Comstock
(1944:488), Comstock & Huntington (1943:73; 1958-64 [1964]:123) (last two citations
place telea as subspecies of maesites; Comstock 1944, however, makes telea a species),
Avinoff & Shoumatoff (1946:284), Hayward (1973:158).

—_—

(including entire corpus bursae). J-L, Chlorostrymon telea (ventral only, with lateral
view of cephalic inclination in L); J, Port-of-Spain, Trinidad; K, Villa Ana, Santa Fe
Province, Argentina (BMNH); L, Callao, Lima Department, Peru (BMNH). M, C. kus-
cheli, data in text (ventral view, cephalic inclination). N, C. patagonia, holotype (including
entire corpus bursae). O, C. chileana, holotype (including entire corpus bursae).

134 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Thecla maesites telea: Comstock & Huntington (1943:78; 1958-64 [1964]:128), Zikan &
Zikan (1968:57).

Eupsyche telea: Dyar (1902:36), Wolcott (1936:402), Grossbeck (1917:23).

Chalybs telea: Kaye (1921:108), Barcant (1970:85).

Strymon telea: Barnes & McDunnough (1917:13), McDunnough (1938:24), Rindge (1952:
11), Ziegler (1960:22) (as “Strymon’’), Lipes (1961:56), dos Passos (1964:55).

Strymon maesites telea: Young (1937:47), Klots (1951:139), Kimball (1965:47).

Chlorostrymon telea: Clench (1961:190; 1976:269; 1977:186), dos Passos (1970:27), Riley
(1975:100), Thorne (1975:278), Ross (1975:189), Miller & Brown (1981:99; 1983:54),
Pyle (1981:464), Opler & Krizek (1986:89).

Chlorostrymon maesites telea: Brown & Heineman (1972:229) (Brown, in Miller & Brown
1981, 1983 considers telea a species), Scott (1986:360).

Diagnosis. DFW, DHW brilliant iridescent azure blue; VF W post-
median line black (with only faint white borders, if any); VHW limbal
suffusion extending costad to M,; postdiscal band thin (“‘line’’), often
broken costad discal cell; line forming distinct ““W” basad limbal area
(Fig. 2A, B). Male genitalia with vincular arc, valvae, saccus, and ae-
deagus less elongate than C. simaethis (Fig. 5E), brush organs attached
as in C. simaethis but less dense (Fig. 5K). Female genitalia less elongate
and not cephalically inclined as in C. simaethis, papillae anales ter-
minally constricted, papillae anales apophyses long (usually extending
entire length of ductus bursae) (Fig. 6G, J-L). For genitalic comparison
to C. maesites, see below.

Types. Holotype male in BMNH (Fig. 5E); TL “Amazon’’.

Distribution (Fig. 4). From S Texas S across Mexico and Central
America; in South America from SW Colombia SE (except for Amazon
basin) along SE coast of Brazil, W across Uruguay and Paraguay to E
Bolivia and E Argentina. W from SW Colombia but only a few spec-
imens from coastal Peru, none from Ecuador. Reports of C. telea from
Florida are usually considered to be C. maesites (Klots 1951, Opler &
Krizek 1986, Scott 1986, and as discussed below).

Conspecificity of C. telea and C. maesites. Possible conspecificity of
C. telea and C. maesites has been often discussed, and favored by
several early authors, more recently by Scott (1986). The taxa have well
defined morphological characters (Figs. 5-7) which are homogeneous
in their respective ranges. For this reason I retain them as species.

The major difference occurs in female genitalia: C. maesites (Fig.
6H, I) has a much smaller antrum and lamellal configuration than C.
telea (Fig. 6G, J-L). As Nicolay (1980) noted, the lamellal area of
Chlorostrymon has a membranous ventral covering. This occurs in
various Eumaeini (Brown 1982), but is artifactual since the covering
strips away easily to expose underlying structures (Johnson 1976, 1978).
In Chlorostrymon, when this membrane is stripped away, the lamella
antevaginalis may be damaged. Thus, the best measure of visual dif-
ference between C. telea and C. maesites is the ratio of the “dorsal

VOLUME 43, NUMBER 2 135

suture line’ (Fig. 6G, extending from terminus of lamellae to base of
“antrum”, Fig. 6A) to the remaining length of ductus bursae. Samples
of C. telea and C. maesites (each spanning distributions characterized
in respective Dissections Examined sections) produced frequency dis-
tributions (Fig. 7B), whose means differ by t-test (P < 0.05). To be
sure that extreme morphology in C. m. clenchi (Fig. 7B, intervals 1.2-
1.4) did not prejudice the distribution of C. maesites, t was recomputed
without these specimens, and also proved significant (P < 0.05). Con-
version of the data to “meaningful pairs’ lacking intracorrelation re-
duced t-values, but they are still significant (P < 0.05). This difference
in female genitalia along with the long cited differences in characters
of the wing make these allopatric taxa distinctive. As discussed under
C. maesites, lesser differences are apparent in male genitalia. Chlo-
rostrymon simaethis shows no comparable major difference between
mainland and Antillean populations.

As with my treatment of C. simaethis, I did not subdivide C. telea
into subspecies.

Dissections examined (AMNH except as indicated). VENEZUELA: Caripito (6, 2).
TRINIDAD-TOBAGO: Port-of-Spain (?). BRAZIL: Parana State, Caviuna (?); Santa Ca-
tarina State, highlands above Massaranduba, Blumenau (6); Amazonas State, Obidos,
January 1936 (2). COLOMBIA: Caldaz, 14 May 1914 (6, 2). MEXICO: Vera Cruz State,
Presidio (6, 2); Colima State, Colima (6, 2); Tamaulipas State, San Francisco (6, 2). UNITED
STATES: Texas, Loredo (2). COSTA RICA: Turrialba (2 6, 2). GUATEMALA: Guatemala
City (2) (BMNH). PANAMA: La Boca, Canal Zone (4, 9).

Chlorostrymon maesites (Herrich-Schaeffer)
(Figs. 2C, D, 5F, G, 6H, 1)

Thecla maesites Herrich-Schaeffer 1864:165. Dewitz (1877:233, pl. 1), Moeschler (1889:
301), Stahl (1882:93), Gundlach (1887:623), Wolcott (1986:402), Comstock & Hun-
tington (1943:72; 1958-64 [1961]:158), Comstock (1944:487), Zikan & Zikan (1968:
Bil):

Thecla maesites clenchi Comstock & Huntington (1943:72) (holotype male, allotype
female [Fig. 2D], AMNH, Roseau Valley, Dominica, British West Indies, April). NEW
STATUS.

Thecla moesites [sic]: Kirby (1871:398), Draudt (1919:798) (misspelling; Comstock &
Huntington 1958-64 [1961]:158 incorrectly attribute error to Draudt).

Thecla moesites Draudt (1919:798). Comstock & Huntington (1958-64 [1961]:158, 171)
(incorrect nomen nudum attributed to Draudt).

Strymon maesites: Barnes & McDunnough (1917:13), Bates (1935:194), Young (1937:47),
McDunnough (1938:24), Klots (1951:140), Kimball (1965:47), Rindge (1952:11), Zieg-
ler (1960:22) (as “Strymon’’), dos Passos (1964:55).

Chlorostrymon maesites: Clench (1961:189; 1963:248; 1976:269; 1977:186), dos Passos
(1970:27), Riley (1975:100), Thorne (1975:277), Miller & Brown (1981:99; 1983:54),
Pyle (1981:464), Opler & Krizek (1986:88), Scott (1986:360), Schwartz (1988).

Diagnosis. DFW, DHW brilliant iridescent azure blue; VHW ter-
minal patch extending costad to M,, postmedian line not making a
“WwW”: VFW postmedian line black (Fig. 2C, D). Genitalia differing

136 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

FREQUENCY (%)

100

75 A

50

25
dus EL Hal i
12 3 4 5/1 2 3 4 Sere

simaethis(Ant.) | sarita(Cent.Am.)| sarita(S. Am.)
N=97 N=123 N=125

FREQUENCY (No. )

4 =I
=
3} ==
===
i
: S22:
22==
Sees
heals 3 lee Pel ES PN SINTON ean Bete
I 1 1
Xm Xm Xt

UNITY- FREQUENCY INTERVAL (.05)

Fic. 7. Frequencies of characters in certain Chlorostrymon populations. A, Wing
characters in C. simaethis simaethis (Antilles) and C. s. sarita (Central America, South
America). Characters: 1, VHW postdiscal band undulate (angles of band, along inner
surface at consecutive veins, changing at least 6 times costad of vein 2A). 2, band generally
straight (not as in 1, and in at least 3 of 5 cells costad of vein 2A, generally in same
plane). 3, band swollen distad in area adjacent to discal cell. 4, VF W with marginal and
submarginal areas, cells CuA, and M,, suffused red-brown and gray. 5, FW with basal
area of costa folded and colored orange. B, Female genitalic shape in C. telea (hatched)
and C. maesites (white). Shape expressed as ratio of length of dorsal suture line (d.s.1.)
to length of ductus (d) (d.s.1./d). xm is mean of C. maesites sample including C. m.
clenchi (0.95, N = 14); xml is mean of C. maesites excluding C. m. clenchi (0.81, N =
10); xt is mean of C. telea sample (0.58, N = 14).

VOLUME 438, NUMBER 2 137

from C. simaethis as in C. telea. For genitalic comparison to C. telea
see C. telea and below.

Types. Location of T. maesites type not known (cited as possibly
Havana, Cuba, by Miller & Brown 1981:note 357); TL Cuba. Comstock
& Huntington (1958-64 [1961]:171) cite a “species” ‘““moesites Draudt’’,
taking Draudt’s (1919) treatment of this name as a description. They
cite no type or type locality as Draudt gave none. Draudt’s treatment
of ““moesites’” was an incorrect repetition of an earlier misspelling by
Kirby (1871:398). Clearly, Kirby, and consequently Draudt, were treat-
ing T. maesites.

Distribution (Fig. 4). S Florida, Bahamas, Greater and Lesser Antilles
S to St. Vincent.

Specificity of C. maesites and C. telea. Considering C. telea and C.
maesites separate species, Clench (1961) stated, without elaboration,
“the two ... are different in many traits’. Such observations probably
resulted from Clench’s experience with Chlorostrymon species in the
field (Clench 1976, 1977).

Variability of C. maesites. As noted under C. telea, male genitalia
of C. telea and C. maesites are similar (Fig. 5E—G). They differ from
C. simaethis (Fig. 5A—-D) by a generally reduced vincular arc, shorter
valval configuration, and aedeagus (a) short, its length usually not ex-
ceeding 8X maximum width of vincular arc (in C. simaethis, 3.5-
4.0x), and (b) with terminal % greatly flared and ventrally inclined
about 60°. Female genitalia of C. telea (Fig. 6G, J-L) and C. maesites
(Fig. 6H, I) differ consistently in structure of ductus and antrum (Fig.
7B). In addition, papillae anales of C. maesites are not as terminally
constricted as in C. telea (Fig. 6G-I). In Antillean C. maesites, as with
Antillean C. simaethis, infraspecific variation is more extreme than in
mainland populations. For example, male genitalia from the Bahamas
and Puerto Rico show notable cephalic sculpturing along outer valval
margins. These do not occur in any other Chlorostrymon and are
probably a parallelism. Female genitalia vary most in ratio of lengths
of antrum and ductus bursae (Fig. 7B), and in degree of dorsal incli-
nation, if any, at the cephalic terminus of ductus bursae (Fig. 6G-L).
Dominican endemic C. m. clenchi (Fig. 61) and some specimens from
Jamaica have a somewhat reduced cephalic terminus on the ductus
bursae. In the two new austral species described here, this tendency is
so extreme that only the area of the antrum remains.

Subspecies of C. maesites. Comstock and Huntington (1943) noted that C. m. clenchi
lacked a tail at vein CuA,, and that both sexes had a dull DFW, DHW ground color, and
pronounced black apical borders (noticable in male; emphatic in female, obscuring almost
any DFW blue). The wing pattern in C. m. clenchi is distinctive, more so than degree

of local differentiation in other Antillean C. m. maesites. However, certain genitalic
features of C. m. clenchi are duplicated in other Antillean C. maesites (female discussed

138 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

under species treatment; in male, elongation of valval terminus in C. m. clenchi [Fig.
5G] duplicated in males from Puerto Rico and the Bahamas). As in C. simaethis, there
appear to be Antillean populations of C. maesites that might equally be considered worthy
of subspecific status. Nevertheless, I recommend that no further subspecies of C. maesites
be recognized, although the name clenchi might still be useful historically to note the
pattern morph typifying Dominica.

Dissections examined (all AMNH). C. m. maesites: BAHAMAS: Nicoll’s Town, Andros
Island (4, 2), North Caicos, 17-18 May 1983 (2 4, 1 2). CUBA: Guantanamo Bay. JAMAICA:
Port Antonio, 10 March 1954 (6), Baron Hill, Trelawny, 16 February 1931 (6), Montego
Bay, 3 January 1965 (6), Reading, St. James, 27 March 1939 (2), Sandy Gully, St. Andrews,
20 June 1951, 3 December 1951 (2 4), 8 July 1951 (?). UNITED STATES: Miami, Florida,
various dates (7 6, 7 2), Brickell Hammock, Florida, 6 August 1914 (6, 2), Coamo Springs,
Puerto Rico, 26-29 December 1914 (2 8, 2). C. m. clenchi: types. DOMINICA: Roseau,
11 April 1929 (2 4, 3 9).

Chlorostrymon kuscheli (Ureta), new combination

(Figs. 3E, F, 5N, 6M)

Thecla kuscheli Ureta (1949:98, pl. 1, fig. 4), Comstock & Huntington (1958-64 [1961]:
58), Rojas (1964:103).

Diagnosis. F W small, base to apex 8.0-9.5 mm (N = 2); DFW, DHW
iridescent lavender in male, dull brown in female, and DHW of both
sexes with bright rufous suffusion across limbal area. VFW, VHW with
white bands limited to thin lines, VHW limbal suffusion only barely
perceptible as silverish streaks. Female genitalia with cephalic end of
ductus dorsally inclined as in C. telea, but with ductus length far
exceeding that of lamellae, as in C. maesites; papillae anales with
apophyses elongate, extending entire length of ductus bursae.

Description. Male. DFW, DHW iridescent lavender. VF W, VHW
ground dull chartreuse; each wing with complete postmedial band, but
constricted as thin white lines. VF W with red-brown suffused discal
slash; VHW with limbal area vaguely suffused silverish. Length of
forewing: 8.0 mm (N = 1). Female. Similar to male but slightly larger,
with DFW, DHW duller brown. Length of forewing: 9.5 mm (N = 1).
Male genitalia (Fig. 5N). Only aedeagus remains of paratype genitalic
preparation; aedeagus typical of genus but angled at junction of shaft
and caecum, latter rather elongate for genus. Female genitalia (Fig.
6M). Cephalic ductal terminus inclined dorsally about 45°, ductus elon-
gate compared to length of dorsal suture line (ratio 0.99). Papillae anales
with apophyses extremely elongate, extending entire length of ductus.

Types. Holotype male, MNHNC, Larancagua, Tarapaca, Chile, 2800
m, 9 December 1946. Allotype female, MNHNC, same data except 25
February 1948. Paratype (Fig. 83E), CECUC, labelled “Thecla kuscheli;
Larancagua, 2700 m, Kuschel, 8 xii 1946; Paratypus; donada par E.
Ureta.”

Distribution (Fig. 4). Tarapaca State, Chile, near border with Bolivia
and Argentina.

VOLUME 43, NUMBER 2 139

Remarks. Ureta’s description, in Spanish, was not widely distributed,
and specimens of C. kuscheli have only recently been available to
northern workers. Though uniquely marked, the species clearly belongs
in Chlorostrymon by wing, male aedeagal, and female genitalic char-
acters. The DHW rufous coloration is unique for the genus; reduced
VFW, VHW bands, and limbal suffusions are common to all austral
Chlorostrymon (but differ in each species). Female genitalia do not
show marked reduction of ductus bursae as in the new austral species
described further on. Though wing pattern in C. kuscheli is extreme,
and somewhat reminiscent of C. simaethis, genitalia are more like C.
maesites and C. telea.

Biogeography. The species is apparently a high montane (2700-3650
m) isolate of the genus. Specimens are known only from the cusp of
the Northern Andean Cordillera and Andean High Plateau biotic prov-
inces (Irwin & Schlinger 1986, Davis 1986) in northern Chile, but may
also occur in adjacent high montane Bolivia and Argentina.

Dissections examined (all CECUC). Paratype 6. CHILE: Putre, Arica region, 3650 m,
25 February 1940, leg. Ureta (@).

Chlorostrymon patagonia, new species

(Figs. 3A, B, 5H, 6N)

Diagnosis. Male DFW, DHW iridescent red-violet; fuscous, basally
overlaid with dull blue-gray in female. Both sexes with FW costa basally
folded, colored bright rufous; VF W, VHW without bands, patterned
as short silver cellular streaks across VHW discal cell and cells CuA, to
caudal M, (limbal suffusion, dull rusty-red to grayish, generally re-
stricted to latter cells). Larger than C. telea and C. maesites: forewing
base to apex averaging 12.2 mm, range 10.0-13.0 mm (N = 8); in C.
telea 8.8 mm, range 7.5-10.0 mm (N = 19); in C. maesites 8.6 mm,
range 6.0-11.0 mm (N = 18). Female genitalia sclerotized only in the
terminal antrumal configuration (as only in C. chileana), corpus bursae
uniquely lacking signa (Fig. 6N); male resembling C. telea and C.
maesites except bilobed valval configuration wider, more shouldered,
saccus reduced to small terminal point, aedeagus with unique cephalic
inclination and marked terminal declination, and brush organs attached
to long membrane spanning ventral surface of vincular arc (Fig.
SE iL).

Description. Male. DFW, DHW bright iridescent red-violet, basal
area of costa widely folded and colored bright rufous. VF W chartreuse,
patterned only with occasional, hardly visible, light streaks in various
cells from costa to cell M3; VHW chartreuse, patterned only by light
slash through discal cell, and silvery zig-zag markings, basad dull rusty-

140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

red to slightly gray suffusion from cells CuA, to caudal M3. Stubby tail,
terminus of vein CuA,. FW length 12.0 mm (allotype). Female. Similar
to male, but DFW, DHW fuscous and suffused dull blue-gray on base
of FW and basal half of HW. FW length 12.0 mm (holotype). Male
genitalia (Fig. 5H). Similar to C. telea and C. maesites but differing
by wider, more shouldered bilobed valval configuration; reduced, fun-
nel-shaped saccus; aedeagus markedly inclined at caecum, declined at
terminus; and brush organs attached along entire ventral surface of
vincular arc. Female genitalia (Fig. 6N). Resembling only C. chileana,
with sclerotized components including only the antrumal structure.
Lamellae distally lobated as in C. telea and C. maesites; corpus bursae
lacking signa; papillae anales constricted terminad as in C. telea, but
apophyses of papillae anales short (about equal to length of antrumal
sclerotization).

Types. Holotype female, allotype male, Nahuel Huapi, Mendoza
Province, Argentina, 15 March 1911 (@), 3 December 1908 (6) (C. S.
Larsen Collection in MNHN). Paratypes: MNHN—same data as allo-
type (6), Mendoza, Argentina, 8 April 1907 (6), 14 March 1907 (6), 18
December 1906 (68), all C. S. Larsen Collection; AMNH—same data as
allotype (4); MPM—Patagonia, August 1939, P. Gagarin Collection (@).

Distribution (Fig. 4). Known only from N to central Patagonian
Steppe biotic province (Davis 1986) of Argentina.

Remarks. In facies, C. patagonia might be considered a C. telea
population of extremely reduced wing pattern if it were not for its
larger size, unique wing characters, and female genitalia resembling
only C. chileana. The southernmost record of C. telea is Villa Ana,
Santa Fe Province, Argentina (BMNH); the southwesternmost, Callao,
Peru (BMNH) (Fig. 4). These specimens are females and typical of C.
telea (Figs. 2B, 6L, M).

It should be noted that Clench (1961) called the upper surface iri-
descent color of C. telea “red-violet’’. This is unfortunate since this
surface in C. patagonia is truly red-violet and distinctive from C. telea,
generally characterized by other authors as brilliant blue. The widely
folded, rufous colored DFW costal fold is also obvious on all specimens
of C. patagonia. A «irvey of 38 C. telea from across its range shows
no such costal cha: -ter. An orangish costal fold occurs in occasional
specimens of C. simaethis (Fig. 7A). In genitalia, the sclerotized struc-
tures in female C. patagonia (and C. chileana) duplicate only the
antrumal structure of other Chlorostrymon species. The ductal area of
C. patagonia (and C. chileana) is wholely membranous. Male genitalia
of C. patagonia resemble those of C. telea and C. maesites most, but
differ as summarized in Diagnosis.

VOLUME 43, NUMBER 2 14]

I speculate that such unusual characters in C. patagonia and C.
chileana are autapomorphic, as discussed under C. chileana.

Biogeography. Chlorostrymon patagonia is found within the Pata-
gonian Steppe biotic province of Davis (1986). From 30°S latitude, this
province extends S in a thin strip E of the Andean Cordillera to en-
compass all of Patagonia S and E of 44°S latitude. Vegetation is xeric
grassland, compatible with known habitats of Chlorostrymon taxa. Sev-
eral other butterfly species have insular distributions like C. patagonia.
One is the distinctly marked Thecla thargelia Burmeister, found only
occasionally northward to Tucuman (IML, MNHN). Five others are
T. larseni Lathy, T. restricta Lathy (both described from MNHN C.
S. Larsen material), and three species of Eiseliana Ajmat de Toledo
located recently in Patagonian material at AMNH, BMNH, and MNHN.

Chlorostrymon chileana, new species

(Figs. 3C, D, 51, 60)

Diagnosis. DFW, DHW of both sexes, dull brown, male slightly
suffused purplish. VF W, VHW lacking bands, VHW patterned only
with vague postdiscal line from discal cell costad to margin. Limbal
area suffused only vaguely gray-brown and dusted basad with silver
from cells CuA, and CuA,. Female genitalia sclerotized only in terminal
antrumal configuration (as in C. patagonia); male genitalia resembling
C. simaethis most but with an enlarged, broad saccus, and an additional
brush organ occurring distally at each juncture of saccus and vinculum.

Description. Male. DFW, DHW dull fuscous slightly hued with pur-
plish blue. VFW, VHW ground dull chartreuse, VF W without pattern,
VHW with obsolescent postdiscal line, discal ceil costad to costal margin;
limbal area, cells CuA, and CuA, slightly suffused reddish to grey distad,
silver basad; stubby tail at terminus of HW vein CuA,. FW length 11.5
mm (allotype). Female. Similar to male except DFW, DHW dull brown.
FW length 11.0 mm (holotype). Male genitalia (Fig. 51). Similar to C.
simaethis but with saccus enlarged and broad (length & width nearly
equal and each equally about % length of vincular arc), a second brush
organ distally at juncture of saccus and vinculum, aedeagus short, its
shaft only slightly exceeding length of entire genitalia, and with caecum
somewhat laterally displaced. Female genitalia (Fig. 60). Resembling
C. patagonia, with sclerotization limited to antrum; lamellae parabolic
as in C. simaethis; signa reduced as small blunt spines; papillae anales
lobate; apophyses of papillae anales short (length barely exceeding that
of antrumal sclerotization).

Types. Holotype male, allotype female, Santiago, Chile, R. Martin,
deposited in MNHN. Paratypes: MNHN—same data as primary types

142 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

(4 4,19); BMNH—“‘Chili’’, Walker, J. J. Joicey Collection, “Thecla sp.
not in collection, $.G”’ (¢); AMNH—same data as primary types (6).

Distribution (Fig. 4). Known only from TL and “Chili’.

Remarks. Chlorostrymon chileana differs greatly from C. simaethis
in its nearly immaculate undersurface and unusual male and female
genitalia. Female genitalia superficially resemble C. patagonia while
male genitalia have a number of unique characters as summarized in
C. patagonia and C. chileana Diagnosis sections.

Biogeography. MNHN has substantial series of butterflies bearing
the labels “Santiago, Chile, R. Martin” and “Valpariso, Chile, R. Mar-
tin’. Chlorostrymon chileana occurs only in the Santiago samples. This
locality, if taken literally, is within the Central Valley biotic province
(Davis 1986, Irwin & Schlinger 1986)—relatively xeric, former thorn
forest now extensively replaced by cultivation. This province is quite
small, extending inland from the Central Coastal Cordillera from about
32-38°S latitude. Its ecology is typical of that associated with Chloro-
strymon taxa. These circumstances, along with unusual characters, sug-
gest that C. chileana is an insular species. Its present-day occurrence
may be severely restricted by land use, as noted for several central
Argentinean plains butterflies (Johnson et al. 1988). The Central Valley
biotic province lies directly opposite the distribution of C. patagonia
on E slopes of the Andes in Argentina. MNHN “R. Martin’ samples
include a number of butterflies previously unrecorded for Chile which
have congeners occurring directly eastward in Argentina’s Coquena
biotic province (Davis 1986). Examples include Calycopis Scudder
(Johnson et al. 1988), Femniterga Johnson (1987), the little known
hairstreaks Thecla americensis Blanchard and T. wagenknechti Ureta,
and others. From such diversity, and comparison with information from
more recent Chilean collections (such as J. Herrera’s, on loan to AME),
I suspect that MNHN “Santiago” and “Valpariso” labels include diverse
Chilean habitats.

ACKNOWLEDGMENTS

I am grateful to the following curators: P. Ackery (BMNH), D. Ajmat de Toledo (IML),
G. Bernardi and J. Pierre (MNHN), L. D. & J. Y. Miller (AME), J. E. Rawlins (CMNH),
F. H. Rindge (AMNH), A. Young, S. Borkin (MPM), and J. Herrera (CECUC, MNHNC).
Lee D. & J. Y. Miller, D. Matusik (FMNH), and E. Quinter (AMNH) assisted in obtaining
literature, and L. D. & J. Y. Miller in obtaining Chilean specimens. For help with statistical
analyses I thank R. F. Rockwell (City College, New York) and Paula Martin.

LITERATURE CITED

AVINOFF, A. & N. SHOUMATOFF. 1946. An annotated list of the butterflies of Jamaica.
Ann. Carnegie Mus. 30:263-295.
BARCANT, M. 1970. The butterflies of Trinidad and Tobago. Collins, London. 314 pp.

VOLUME 43, NUMBER 2 143

BARNES, W. & F. H. BENJAMIN. 1923. Checklist of the diurnal Lepidoptera of boreal
America. Bull. So. Calif. Acad. Sci. 25:1-17.

BARNES, W. & J. H. MCDUNNOUGH. 1917. Checklist of the Lepidoptera of North
America. Herald Press, Decatur, Illinois. 392 pp.

BATES, M. 1935. The butterflies of Cuba. Bull. Mus. Comp. Zool. 78:63-258.

BEATTY, H. A. 1944. Fauna of St. Croix, V.I. The insects of St. Croix, V.I., Lepidoptera.
J. Agric. Univ. Puerto Rico 28:156-158.

BRowN, F. M. & B. HEINEMAN. 1971. A checklist of butterflies of Jamaica. Mid-Cont.
Lepid. Ser. 2:1-9.

1972. Jamaica and its butterflies. E. W. Classey, London. 478 pp.

CLENCH, H. K. 1961. “Lycaenidae.” In Ehrlich, P. R. and A. H. Ehrlich (eds.), How
to know the butterflies. W. C. Brown, Dubuque, Iowa. 262 pp.

1963 (1965). A synopsis of the West Indian Lycaenidae with remarks on their

zoogeography. J. Res. Lepid. 2:247-270.

1976. Butterflies of the Carnegie Museum Bahamas expedition, 1976. Ann.

Carnegie Mus. 46(16):265-283.

1977. A list of the butterflies of Andros, Bahamas. Ann. Carnegie Mus. 46(12):
173-194.

ComsTocK, W. P. 1944. The insects of Porto Rico and the Virgin Islands. Rhopalocera
or butterflies. Scientific Survey of Puerto Rico and the Virgin Islands (New York
Academy of Sciences) 12(1):421-622.

Comstock, W. P. & E. I. HUNTINGTON. 19438. Lycaenidae of the Antilles (Lepidoptera:
Rhopalocera). Ann. New York Acad. Sci. 45:119-130.

1958-64. An annotated list of the Lycaenidae (Lepidoptera, Rhopalocera) of
the western hemisphere. J. New York Entomol. Soc. 66[1958]:103-118; 67[1959]:59-
95, 1638-212; 68[1960]:49-62, 105-122, 176-186, 232-240; 60[1961]:54-58, 105-118,
157-176, 191-200; 70[1962]:39-49, 100-118, 177-179; 71[1963]:45-57, 115-119, 189-
197, 262-264; 72[1964]:62-64, 120-130.

Davis, D. R. 1986. A new family of Monotrysian moths from austral South America
(Lepidoptera: Palaephatidae), with a phylogenetic review of the Monotrysia. Smiths.
Contr. Zool. 434:1-202.

DeEwitTz, H. 1877. Tagschmetterlinge von Portorico. Stett. Entomol. Zeit. 38:233-245.

pos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. Soc.
Mem. 1, 145 pp.

1970. A revised synonymic catalogue with taxonomic notes on some Nearctic
Lycaenidae. J. Lepid. Soc. 24:23-28.

DraupT, M. 1919. Thecla, pp. 794-811. In Seitz, A. (ed.), Macrolepidoptera of the
world. Alfred Kernen Verlag, Stuttgart. Vol. 5 (“Vol. IT’), pp. 5938-1139, Vol. 5 plates
(“Vol: II Plates’), 194 pls.

Drury, D. 1770-82 [1773]. Illustrations of natural history. Vol. 1 and index. White,
London. Vol. I [1770], 180 pp., 50 pls., index [1773].

Dyar, H. G. 1902 (1908). List of North American Lepidoptera. Bull. U.S. Natl. Mus.
52, 722 pp.

EMMEL, T. C. & J. F. EMMEL. 1973. The butterflies of southern California. Nat. Hist.
Mus. Los Angeles Co., Sci. Ser., 26, 148 pp.

Ferris, C. F. 1974. Review: Butterflies of the world by H. L. Lewis. J. Res. Lepid. 3:
278-280.

GaRTH, J. S. & J. W. TILDEN. 1986. California butterflies. Univ. California Press,
Berkeley. 246 pp.

GopmaN, F. D. & O. SALVIN. 1879-1901. Biologia Centrali-Americana. Rhopalocera.
Privately printed, London. Vol. I, xiv + 487 pp. (1879-86); Vol. 2, 782 pp. (1887-
1901); Vol. 3 (pls.), 112 pls. (1879-1901).

GROSSBECK, J. A. 1917. Insects of Florida, IV. Lepidoptera. Bull. Am. Mus. Nat. Hist.
37:1-147.

GUNDLACH, J. 1887. Fauna Puerto-Riquena. Ann. Soc. Espanola Hist. Nat. 16:347-658.

HALL, A. 1925. List of the butterflies of Hispaniola. Entomologist (London) 59:186-
190.

1936. The butterflies of St. Kitts. Entomologist (London) 69:274-278.

144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

HAYWARD, K. J. 1973. Catalogo de los Ropaloceros Argentinos. Opera Lilloana 23:1-
318.

HENNIG, W. 1966. Phylogenetic systematics. Univ. Illinois Press, Urbana. 263 pp.

HERRICH-SCHAEFFER, G. A. W. 1864. Die Schmetterlinges-fauna der Insel Cuba. Cor-
respondenzblatt des zoologischer-mineralogische Verein in Regensburg 18:159-172,

- 19:52-60.

HEWITSON, W. C. 1862-78 [1873]. Illustrations of diurnal Lepidoptera, Lycaenidae.

John van Voorst, London. Vol. I, 228 pp., suppl., 48 pp.; Vol. II (pls.), 92 pls., suppl.,

8 pls.
1868. Descriptions of some new species of Lycaenidae. John van Voorst, London.
36 pp.
Hiccins, L. G. 1972. Hewitson on butterflies 1867-77. Hampton, E. W. Classey, London.
210 pp.

HOFFMAN, C. C. 1941. Catalogo sistematico y zoogeografico de los Lepidopteros Mex-
icanos. Sobret. Ann. Inst. Biol., Mexico, D.F. I: Papilionoidea, pp. 640-739.

HOLLAND, W. J. 1931. The butterfly book. Doubleday, New York. 382 pp.

HUBNER, J. 1806-38 [1807]. Sammlung exotische [sic] Schmetterlinge 1. Privately print-
ed, Augsburg. 36 pp., 213 pls.

1816-26 [1818]. Verzeichniss bekannter Schmettlinge [sic]. Privately printed,
Augsburg. 431 pp. + 72 pp. index.

IRWIN, M. E. & E. I. SCHLINGER. 1986. A gazetteer for the 1966-1967 University of
California—Universidad de Chile Arthropod Expedition to Chile and parts of Ar-
gentina. Occ. Pap. California Acad. Sci. 144:1-11.

JOHNSON, K. 1976. Three new Nearctic species of Callophrys (Mitoura), with a diag-
nostic [sic] of all Nearctic consubgeners (Lepidoptera: Lycaenidae). Bull. Allyn Mus.
38, 30 pp.

1978. Specificity, geographic distribution, and foodplant diversity in four Cal-

lophrys (Mitoura) (Lycaenidae). J. Lepid. Soc. 32:3-19.

1986. A new species of Tmolus (Lycaenidae) with notes on the Euptychia

westwoodi (Satyridae) mimicry complex. Bull. Allyn Mus. 106, 11 pp.

1987. Tergissima and Femniterga, new sister genera of Calycopis Scudder and

Calystryma Field from the south-central Andes. Insecta Mundi 2(1):28—42.

1989. A revisionary study of the Neotropical hairstreak butterfly genus Noreena

and its new sister genus Contrafacia. J. New York Entomol. Soc. 97:11-46.

In press. Penaincisalia, a new genus of “Elfin’’-like butterflies from the high
Andes (Lepidoptera: Lycaenidae). Pan-Pacific Entomol.

JOHNSON, K., R. E. EISELE & B. MACPHERSON. 1988. The “Hairstreak butterflies”
(Lycaenidae: Theclinae) of northwestern Argentina. I. Calycopis, Calystryma, Ter-
gissima and Femniterga. Bull. Allyn Mus. 123:1—49.

JOHNSON, K., B. MACPHERSON & J. INGRAHAM. 1986. A new genus and species of
Eumaeini (Theclinae) from western Argentina (Lepidoptera: Lycaenidae). Bull. Allyn
Mus. 102, 7 pp.

JOHNSON, K. & D. MATUSIK. 1988. Five new species and one new subspecies of butterflies
from the Sierra de Baoruco of Hispaniola. Ann. Carnegie Mus. 57:221—254.

KAYE, W.J. 1914. Additions and corrections to my catalogue of Lepidoptera Rhopalocera
of Trinidad (1904). Trans. Entomol. Soc. London 1914:545-585.

1921. A catalogue of the Trinidad Lepidoptera Rhopalocera (butterflies). Mem.

Dept. Agric. Trinidad and Tobago 2, 163 pp.

1926. The butterflies of Jamaica. Trans. Entomol. Soc. London 1924:413—428.

KIMBALL, C. P. 1965. The Lepidoptera of Florida, an annotated checklist. Vol. 1.
Arthropods of Florida and neighboring land areas. Florida Dept. Agric., Gainesville.
363 pp.

KirBy, W. F. 1871. A synonymic catalogue of diurnal Lepidoptera. John van Voorst,
London. 192 pp.

Kxiots, A. B. 1951. A field guide to the butterflies of North America, east of the Great
Plains. Houghton Mifflin, Boston. 349 pp.

Lewis, H. L. 1974. Butterflies of the world. Follett, Chicago. 312 pp.

VOLUME 43, NUMBER 2 145

LipEs, J. E. 1961. Butterfly records from Brownsville, Texas. J. Lepid. Soc. 15:56.

LLORENTE-BOUSQUETS, J., A. GARCES MEDINA & A. L. MARTINEZ. 1986. Las Mariposas
de Jalapa-Teocelo, Veracruz. Revista Teocelo 4:14—37.

McDUNNOUGH, J. 1938. Checklist of the Lepidoptera of Canada and the United States
of America, part I: Macrolepidoptera. Mem. So. California Acad. Sci. 1, 175 pp.
MILLER, L. D. & F.M. BRowN. 1981. A catalogue/checklist of the butterflies of America

north of Mexico. Lepid. Soc. Mem. 2, 280 pp.

1983. Papilionoidea, pp. 49-65. In Hodges, R. W. (ed.), Check list of the
Lepidoptera of America north of Mexico. E. W. Classey and Wedge Entomol. Res.
Found., London. 284 pp.

MOESCHLER, H. B. 1889. Die Lepidopteran fauna von Porto Rico. Abhandlungen der
Seckenbergischen naturforschenden Gesellschaft [1889]:69-360.

NICOLAY, S. S. 1979. Studies in the genera of American hairstreaks. 5. A review of the
Hiubnerian genus Parrhasius and description of a new genus Michaelus (Lycaenidae:
Eumaeini). Bull. Allyn Mus. 56, 52 pp.

1980. The genus Chlorostrymon and a new subspecies of C. simaethis. J. Lepid.
Soc. 34:253-256.

OPLER, P. R. & G. O. KRIZEK. 1986. Butterflies east of the Great Plains, an illustrated
natural history. Johns Hopkins Univ. Press, Baltimore. 294 pp.

PyLE, W. M. 1981. The Audubon field guide to North American butterflies. Alfred A.
Knopf (Chanticleer), New York. 916 pp.

{ILEY, N. D. 1975. Field guide to the butterflies of the West Indies. Collins, London.
244 pp.

RINDGE, F. H. 1952. The butterflies of the Bahama Islands, British West Indies (Lep-
idoptera). Amer. Mus. Novit. 1563, 18 pp.

Rojas, E. U. 1964. Catalogo de Lepidopteros de Chile. Bol. Mus. Nac. Hist. Nat. Chile
28(2):67-140.

Ross, G. N. 1975-77. An ecological study of the butterflies of the Sierra de Tuxtla, in
Veracruz, Mexico. J. Res. Lepid. 14:103-124, 169-188, 233-252; 15:41-60, 109-128,
185-200, 225-240; 16:87-130.

SCHWARTZ, A. 1988. The butterflies of Hispaniola. Univ. Florida Press, Gainesville,
Florida. 604 pp.

SCHWARTZ, A. & J. C. CorREA. 1986. The status of Calisto hysius batesi (Lepidoptera,
Satyridae) with the description of a new species of Calisto from Hispaniola. Florida
Sci. (Biol. Sci.) 49:11-18.

SCHWARTZ, A. & F. GALI. 1984. Five new species of Calisto (Satyridae) from Hispaniola.
Bull. Allyn Mus. 5:1-18.

SCHWARTZ, A. & C. J. JIMENEZ. 1982. The butterflies of Montserrat, West Indies. Bull.
Allyn Mus. 66:1-18.

SCHWARTZ, A. & W. W. SOMMER. 1986. A new subspecies of Synapte malitiosa (Lep-
idoptera: Hepseriidae) from Hispaniola. Florida Sci. (Biol. Sci.) 49:18-22.

SCHWEIZER, F. & R. G. WEBSTER Kay. 1941. Lepiddpteros del Uruguay. II. Catalogo
sistematico, parte 1. Rhopalocera and Grypocera. Anal. Mus. Hist. Nat. Montevideo
2.a, Serie-Tomo V, No. 3:4—24.

Scott, J. A. 1986. The butterflies of North America, a natural history and field guide.
Stanford Univ. Press, Stanford, California. 583 pp.

SCUDDER, H. 1872. A systematic revision of some of the American butterflies. Ann.
Rep. Peabody Acad. Sci. 4:24—-83.

1876. Synonymic list of the butterflies of North America, north of Mexico. Bull.
Buffalo Soc. Nat. Sci. 3:98-129.

SKINNER, H. 1895. Notes on Rhopalocera, with descriptions of new species. Entomol.
News 6:112-114.

1898. A synonymic catalogue of the North American Rhopalocera. Philadelphia,
215 pp.

STAHL, A. 1882. Insects, pp. 82-102. In Stahl, A. (ed.), Fauna Puerto Rico. San Juan.

STALLINGS, D. B. & J. R. TURNER. 1947. Texas Lepidoptera (with description of a new
subspecies). Entomol. News 58:37-41.

146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

THORNE, F. 1975. Chlorostrymon, pp. 277-278. In Howe, W. H. (ed.), The butterflies
of North America. Doubleday, Garden City, New York. 638 pp.

UrRETA, R. E. 1949. Lepidopteros de Chile (Rhopalocera). IV Parte. Familia Lycaenidae.
Bol. Mus. Nac. Hist. Nat. Chile 24:93-123.

WoLcoTT, G. N. 1936. “Insectae borinquenses’’, a revised checklist of the insects of
Puerto Rico. J. Dept. Agric. Puerto Rico 20(1):1-600.

YOUNG, F. N. 1987. Notes on the occurrence of Strymon maesites (Herrich-Schaeffer)
in Florida. Entomol. News 48:80-81.

ZIEGLER, B. J. 1960, 1961. Preliminary contribution to a redefinition of the genera of
North American hairstreaks (Lycaenidae) north of Mexico. J. Lepid. Soc. 14:19-23.

ZIKAN, J. F. & W. ZIKAN. 1968. Insecto-Fauna de Itatiaia e da Mantiquieva. Sec. Entomol.
Fito Patulog. Pesq. Agropec. Braz. 3:45-109.

Received for publication 23 November 1987; accepted 18 November 1988.

Journal of the Lepidopterists’ Society
43(2), 1989, 147

GENERAL NOTES

PAPILIO TROILUS L. ON A NEW AND RARE LARVAL FOOD PLANT
Additional key words: Papilionidae, endangered, Lindera melissifolia.

Papilio troilus L. is a common swallowtail found over a broad geographic range. It is
known from southern Canada to Florida and W to Manitoba and Texas, becoming less
common W of the Mississippi River (Klots, A. B. 1951, Field guide to the butterflies,
Houghton-Mifflin, Boston, 349 pp.). At least 15 species have been reported as larval food
plants; these are mainly in the families Lauraceae, Rosaceae, and Kutaceae (Teitz, H. M.
1972, An index to the described life histories, early stages, and hosts of Macrolepidoptera
of the continental United States and Canada, Allyn Mus. Entomol., Sarasota, Florida,
1041 pp.; Opler, P. A. & G. O. Krizek 1984, Butterflies east of the Great Plains, Johns
Hopkins Univ. Press, Baltimore, 294 pp.). Species for which there is direct evidence of
complete larval development are Cinnamomum camphora Nees & Eberm., Lindera
benzoin (L.) Blume, Persea borbonia (L.) Spreng., and Sassafras albidum (Nutt.) Nees
(R. C. Lederhouse pers. comm. ).

In Mississippi, Lindera benzoin is the most common and widely distributed spicebush.
The related pondberry or swamp spicebush, L. melissifolia (Walter) Blume, is an en-
dangered species throughout its range in the SE United States (Kral, R. 1983, U.S. Dep.
Agr. Forest Service Tech. Publ. R8-TP2, 1305 pp.; Currie, R. 1985, Federal Register 50:
32581-32585). Pondberry is known in Mississippi only from the Delta Region in Bolivar,
Sharkey, and Sunflower counties.

On 18 June 1988, when the latest Mississippi population of L. melissifolia was discovered
in Sunflower Co., a larva of Papilio troilus was noticed in its weblike, longitudinally rolled
nest on a leaf of L. melissifolia. The preserved larva was given to the Mississippi Ento-
mological Museum at Mississippi State University, Mississippi State, Mississippi, and vouch-
er specimens of L. melissifolia are deposited in university herbaria at Florida, Michigan,
Vanderbilt, and other herbaria.

In Mississippi, Papilio troilus larvae are commonly found on Sassafras albidum and
Lindera benzoin, both in Lauraceae. Sassafras albidum is common in the Delta Region
of Mississippi, but Lindera benzoin is relatively rare there, being more frequent eastward
in the Loess Bluff Region. It is therefore logical that P. troilus utilizes another species in
this family and in the genus Lindera. This observation is unique because a common
butterfly seems to accept a rare food plant in the natural environment when other sources
are much more common. However, no individuals of sassafras were located in the im-
mediate area, and populations of the more common spicebush are kilometers away from
the collection site.

M. WAYNE Morris, Department of Botany, University of Florida, Gainesville, Florida
32611.

Received for publication 26 August 1988; accepted 14 December 1988.

Journal of the Lepidopterists’ Society
43(2), 1989, 147-148

SPENCER COLLECTION GIVEN TO SMITHSONIAN

The National Museum of Natural History (Smithsonian Institution) has received the
Spencer Collection of Western Butterflies. The Collection consists of over 4000 specimens,
primarily of the genus Speyeria, and represents eight western states, Mexico, and Canada.

148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Among the species are fine series of Speyeria nokomis nitocris (Edwards), a topotypical
series of S. n. coerulescens (Holland), and a series of S. cybele pugetensis F. Chermock
and Frechin. Two important butterflies were rediscovered by Spencer: S. nokomis ni-
grocaerulea (W. and T. Cockerell), near Taos, New Mexico, and Clossiana selene ne-
braskensis (Holland), near Valley, Nebraska. The Collection is rich in Nebraska material,
including the only known Nebraska specimen of Colias alexandra krauthii Klots, from
Sow Belly Canyon, Sioux County, and the only recent eastern Nebraska specimen of
Speyeria aphrodite alcestis (Edwards). Nearly all of the Speyeria specimens were reared,
and each species series displays rich coloration and individual variation.

Mr. Orville D. Spencer and his wife Eunice of Lincoln, Nebraska spent 40 years amassing
their collection of Lepidoptera. Spencer’s interest in butterflies began when he was a boy
in Lincoln. Later he developed a highly successful technique for collecting eggs from
butterflies and rearing them at home. Mr. Spencer’s background is engineering, having
retired in 1980 from the Lincoln Telephone Company. From the collector-made drawers
to the carefully placed antennae, the time spent and the love shown in preparing this
collection is evident to the viewer.

J. F. GATES CLARKE, Department of Entomology, MRC NHB 127, National Museum
of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Journal of the Lepidopterists’ Society
43(2), 1989, 148-151

EVIDENCE FOR GENETIC DETERMINATION OF VARIATION IN
ADULT SIZE AND WING MELANISM OF PARNASSIUS PHOEBUS F.

Additional key words: phenotype, geographic variation, Papilionidae.

Parnassius phoebus F. (Papilionidae) ranges from Europe across Asia, and throughout
much of montane western North America. The species is highly variable in wing coloration
and size throughout its range, both within and between populations. C. D. Ferris (1975,
J. Res. Lepid. 15:1-22) described the taxonomic variation of non-Arctic North American
populations. I described some of the variation in wing color and size from an ecological
viewpoint (Guppy, C. S. 1986a, Can. J. Zool. 64:956-962; 1986b, Oecologia 70:205-213).

To arrive at an understanding of the systematic and ecological significance of phenotypic
variation of P. phoebus, it is necessary to know if the variation is due to genetic differences
between populations. Ferris (above) believed that variation in wing melanism is environ-
mentally controlled, and J. A. Scott (pers. comm.) believes it is genetically determined.
In this paper I provide evidence for the genetic basis of some geographic variation in
wing melanism and size (wing length) of P. phoebus.

Parnassius phoebus is a medium-sized to large butterfly with wings that are white with
various black markings and usually red ocelli. There is a predominantly black region at
the base of the hindwings which varies considerably in width, in proportion of black to
white scaling within the black region, and in density of scaling (transparency) of the
black region. This black region has a thermoregulatory significance (Guppy 1986b, above).
The forewing distal region has marginal and submarginal black markings which vary
greatly in development, especially in females. This region may be very transparent,
especially in females, but it apparently lacks thermoregulatory significance (Guppy 1986b,
above). Body size is highly variable, with a general trend of decreased size with increased
elevation (Guppy 1986a, above).

[ reared offspring concurrently from one or two arbitrarily selected females from each
of five P. phoebus populations (Table 1) under uncontrolled (outdoor) conditions in 1980.
Arbitrary samples from parent populations and all reared offspring were scored for six

VOLUME 43, NUMBER 2 149

TABLE 1. Parnassius phoebus sample origins and sizes. Eggs were obtained in 1979.

Sample size*

Locality eT Wer ee eee

no. Description Mw Mr Fw Fr

1 Montana, Missoula, elev. 1525 m 12 2 11 B

2 Alberta, Kananaskis Rd., Regal Ck., elev. 1525 m @. 6 2 8

3 British Columbia, Manning Park, Gibson Pass, iT 13 5 14
elev. 1370 m

4 British Columbia, Big Bar Creek, Poison Mt., elev. 9 5 g 4
2135-2195 m

5 British Columbia, Penticton, Mt. Apex, elev. 2190- 24 8 4 6
2247 im

* Number of males (M) and females (F) in wild (w) and reared (r) samples.

phenotypic characters by methods described and illustrated previously (Guppy 1986a,
above). Briefly, characters were defined as follows: “basal patch width’—proportion of
the centerline of the dorsal hindwing discal cell covered by the predominantly black
region. Basal blackness’ —proportion of scales in the basal black patch which were black
(the rest were white). “Basal transparency —proportion of the basal black patch which
was without scales (in the absence of scale erosion). “Distal blackness’—proportion of
100 quadrats in a microscope’s optical grid (oriented so outer corners at the points where
veins M, and Cu, met the forewing margin) in which >25% (males) or >50% (females)
of scales were black. “Distal transparency —proportion of distal forewing area not coy-
ered by scales (in the absence of scale erosion). Forewing length was measured with a
metric ruler from thoracic attachment point to wing apex. All phenotypic measurements
except forewing length were arcsine (square root) transformed before analysis to normalize
distributions.

Data were analyzed by nested analysis of variance (ANOVA), with PHENOTYPE a
function of LOCALITY, and ENVIRONMENT (reared vs. wild) nested within LOCAL-
ITY (Zar, J. H. 1974, Biostatistical analysis, Prentice-Hall, Englewood Cliffs, New Jersey,
620 pp.). In the ANOVA’s, if LOCALITY is significant, then there are significant differ-
ences between reared samples, and those differences are correlated with differences in
the wild populations. Therefore, such differences must be genetically controlled. The
ENVIRONMENT term could not be interpreted unambiguously because it included both
effect of developmental environments (six wild and one reared) and genetic effects due
to each reared sample having originated from eggs of 1-2 females instead of from a
random sample of eggs from all females in a population. However, if all reared samples
deviate in the same direction from corresponding wild sample phenotypes, it can be
concluded that developmental environment is important in determining phenotype. Ab-
sence of such consistent deviations does not necessarily mean that environmental effects
are absent, because the rearing environment may not have deviated in a consistent
direction relative to wild environments.

Basal patch width of males, distal transparency of both sexes, and wing length of males
(Fig. 1) differed significantly among reared samples, and those differences are correlated
with differences between wild populations (LOCALITY terms P < 0.05). Therefore,
genetic differences among populations cause at least some of the interpopulation variation
in these characters.

LOCALITY terms were nonsignificant (P > 0.05) for female basal patch width, female
basal blackness, and distal blackness for both sexes. ANOVA’s were not done for male
basal blackness, basal transparency for both sexes, and female forewing length because
of nonhomogeneous variances.

Basal blackness and basal transparency are apparently affected by developmental en-
vironment. Nine of the 10 reared samples (both sexes) were darker and less transparent

150 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

BASAL WING
PATCH WIDTH (%) LENGTH (mm)

52-1 Male Male

4 9 3 Sone

DISTAL DISTAL
TRANSPARENCY (%) TRANSPARENCY (%)
16

Female

50

Locality Locality

Fic. 1. Characters of Parnassius phoebus for which a genetic basis was detected.
Dark bars, reared samples; light bars, wild samples. Vertical lines represent 1 SE. Sample
origins and sizes are given in Table 1.

than the corresponding wild samples. Locality No. 4 males showed no difference between
reared and wild samples in basal darkness and transparency. In addition, 9 of the 10
reared samples have shorter forewing lengths than the corresponding wild samples (Lo-
cality No. 5 reared sample averaged 1 mm longer than the wild sample). Therefore, size
as indicated by forewing length (Miller, W. E. 1977, Ann. Entomol. Soc. Am. 70:253-
256) is also affected by developmental environment.

Significant ENVIRONMENT terms (P < 0.05) occurred for female basal patch width,
female basal blackness, male distal blackness, male distal transparency, and male forewing
length, but, as mentioned above, interpretation of ENVIRONMENT is ambiguous.

VOLUME 43, NUMBER 2 Abt

This study provides evidence of a genetic basis for interpopulation variation in three
of six phenotypic characters examined for P. phoebus. There is also evidence for devel-
opmental environment affecting phenotype for three characters. In light of the small
sample sizes, failure to detect either a genetic or an environmental component to variation
in a character does not mean that these components are unimportant, merely that they
were not detected.

This research was partly supported by a Natural Sciences and Engineering Research
Council of Canada (NSERC) operating grant to Judith H. Myers at the University of
British Columbia.

C. S. Guppy, Biological Collections Section, Royal British Columbia Museum, Victoria,
British Columbia, Canada V8V 1X4.

Received for publication 23 February 1988; accepted 15 November 1988.

Journal of the Lepidopterists’ Society
43(2), 1989, 152-153

EFFECT OF REFRIGERATION ON EGG INCUBATION PERIOD OF THE
TASAR SILK INSECT ANTHERAEA MYLITTA DRURY (SATURNIIDAE)

Additional key words: _ sericulture, India.

Antheraea mylitta Drury is a semidomesticated tasar silk insect reared thrice in a year
during July-August, September—October and November—December. January to middle
of June is the diapause period. In recent years, 40-60 percent of fertile egg production
from May to mid-June has gone unutilized for rearing due to lack of quality leaves in
tasar food plants and excessive outdoor temperatures (39 + 4°C). This situation causes
loss and scarcity of eggs for subsequent commercial tasar crops. It would be desirable to
prolong egg incubation by some suitable means to enable utilization of those eggs at the
onset of the favorable rearing period, and to synchronize hatching of all egg batches for
simultaneous rearing.

The egg of A. mylitta normally requires seven days of incubation at room temperature,
and hatches on the eighth day after oviposition. Refrigeration is a common means to
delay hatching of other silkworm eggs. Information is available on the effect of low
temperature on the incubation periods of the mulberry silkworm, Bombyx mori L. (Tay-
ade, D. S., M. D. Jawale & P. K. Unchegaonkar 1987, Sericologia 27:297—299) and the
Eri silkworm, Philosamia ricini H. (Choudhury, S. N. 1982, Eri silk industry, Directorate
of Sericulture and Weaving, Govt. of Assam, Gauhati, 177 pp.; Viswakarma, S. R. 1982-
83, Indian J. Seric. 21—22:36-39). Since no information was available on the effect of
refrigeration of eggs of A. mylitta, this investigation was made.

In the Mayurbhanj district of Orissa, India, 29,000 freshly oviposited eggs were collected
from 290 DFI’s (disease free layings from 290 healthy mated females) of the Sukinda
trivoltine race of A. mylitta on 22 May 1987 at 0900 h, and were kept at room temperature
(31 + 2°C) as a common stock. Every day at 0900 h from the first to seventh day after
oviposition, 4000 eggs (40 DFI’s) were taken from the common stock and divided into
four equal groups for 1, 2, 8, and 4 days of refrigeration treatment at 10 + 1°C, after
which they were again allowed to incubate at room temperature until hatching. The
remaining 1000 eggs (LODF!’s) served as the control. The incubation period of the treated
groups was then compared with the control. The experiment was repeated five times
during the same period and under the same conditions.

The incubation period of control eggs was seven days. One and two days of refrigeration
of Ist- (fresh or 0-day-old) and 2nd-day (1-day-old) eggs increased the incubation period
to 12 days (Table 1), 5 days more than the control. Three and four days of treatment to
such eggs increased the incubation period to 13 days (Table 1), 6 days more than the
control.

One and two days of refrigeration increased the incubation period by two days beyond
the control in 3rd- (2-day-old), 4th- (8-day-old), and 5th-day (4-day-old) eggs, and by

TABLE 1. Incubation period of Antheraea mylitta eggs refrigerated at different ages
for different periods.

Incubation period when refrigerated for:

Day after Age of eggs

oviposition (days) 1 day 2 days 8 days 4 days
Ist 0 12 2 13 13

(fresh eggs)

2nd I 12 12 13 13
3rd 2 9 9 Il 11
Ath 3 3) 9) 10 10
5th 4 9 9 11 11
6th iB) 8 8 10 10
7th 6 8 8 9 9

VOLUME 43, NUMBER 2 153

TABLE 2. Analysis of variance.

Source of variation df SS MS F
Between refrigeration periods 3 61.71 10.28 108.27*
Between ages 6 14.28 4.76 aO12"
Error 18 La7fdl 0.09 --
Total 2M Wiha! — —

Critical difference (P < 0.05) for refrigeration period = 0.35
* Significant (P < 0.05).

one day in 6th- (5-day-old) and 7th-day (6-day-old) eggs. Similarly three and four days
of refrigeration increased the incubation period by four days in 3rd- and 5th-day eggs,
by three days in 4th- and 6th-day eggs, and by two days in 7th-day eggs.

The data were analyzed as two-way classified. There is significant variation (P < 0.05)
among different refrigeration treatments as well as among different ages of eggs; further,
the critical difference indicates that the two-day refrigeration treatment differed signif-
icantly from the three-day (Table 2).

Thus the fresh and one-day-old eggs of A. mylitta refrigerated for three and four days
showed maximum increase of the incubation period amounting to six days. Viswakarma
(above) observed that the incubation period of P. ricini eggs when refrigerated at 7 +
2°C for five days increased by four to five days. Choudhury (above) reported prolongation
of the incubation period of P. ricini eggs by four days of refrigeration at 15°C. Tayade
et al. (above) observed one or two days extension of the incubation period in B. mori
eggs with increase of refrigeration at 5°C to 55 days. Studies on the effect of different
degrees of temperature on incubation and embryonic development of A. mylitta eggs
should be carried out.

B. K. NAYAK AND A. K. Dasu, State Sericultural Research Station, Orissa, Baripada-
757001, India.

Received for publication 22 January 1988; accepted 5 August 1988.

Journal of the Lepidopterists’ Society
43(2), 1989, 154

BOOK REVIEW

THE SATURNIIDAE OF AMERICA. CERATOCAMPINAE, by Claude Lemaire. 1988. 480 pp.,
64 pls., 379 text figs. Mus. Nac. Costa Rica. Soft cover. $80.00.

_ This is the third volume by Lemaire in his series of monographs of the New World
Saturniidae; its predecessors concerned Attacinae (=Saturniinae) (1978) and Arsenurinae
(1980). All three follow the same format; the two earlier volumes were illustrated in black
and white, but the present one shows the moths in color.

The use of Ceratocampinae may come as a surprise to some, rather than Citheroniinae,
which has been used by recent workers. Fourteen subfamily names are listed as being
available for this group of moths; the oldest, by 25 years, is Ceratocampinae. Two ad-
ditional available family-group names also have priority over Citheroniinae.

This subfamily is second only to Hemileucinae in numbers of species; 170 are covered
in this volume, and they are placed in 27 genera. The group is restricted to the New
World, with the taxa being distributed from southern Canada to southern South America.
The highest percentage of endemic species is found in Mexico and Central America, the
next largest group in the general area of southeastern Brazil.

The introductory section of the book gives morphological characters for the subfamily,
geographic distribution, a summary of knowledge of the early stages (with six color plates
showing caterpillars of 24 species), a discussion of taxonomy and name usage, and phy-
logeny, followed by a key to genera. Each genus has its bibliography, followed by most
of the subjects listed above. Following keys to included species, each taxon is treated in
a similar fashion. Drawings of male and female genitalia, plus distribution maps, are
always present; antennae, venation, and legs are illustrated for most genera. Each species
(and subspecies when present) is illustrated in color, usually with several examples being
shown.

While the text is in French, each taxon, from subfamily to subspecies, has a diagnostic
summary in English; in addition, there is a Spanish summary for the subfamily and for
each genus. This makes the book readily understandable to those who do not read French;
Lemaire is to be highly commended for including these extremely useful additions.

The taxonomic approach is based on study of specimens from the entire New World.
This method, rather than defining genera by use of species from a restricted geographic
area, has led to some name changes. On the generic level, the only change for the North
American fauna is that Syssphinx is used instead of Sphingicampa. Lemaire takes a
conservative approach to nomenclature; his treatment of some species and subspecies
differs from some recently published papers. It is a pleasure to see how he handles these
problems, utilizing his knowledge and perspective, and shedding new light on some areas
that need this type of analysis.

This volume, like the two before it, is handsomely done; the color plates are a great
improvement over the earlier black and white illustrations. In a work of this size it is not
surprising that a few errors have inadvertently been made; an included erratum sheet
covers most of these. Lemaire is to be congratulated; we look forward to each additional
volume in this series by the leading specialist of New World Saturniidae.

This and the two previous volumes will be the standard by which identifications and
curating will be followed for decades to come. They will be of interest to anyone curious
about this family of moths. Now that the basic taxonomy has been done, the invitation
is there for much needed work on ecology, life histories, food plants, and behavior of
these interesting moths, to mention a few possible fields of study.

Copies may be obtained by sending a check for $80.00 (U.S.), made to Fundacion
Neotropica, Museo Nacional de Costa Rica, Aptdo. 749-1000, San Jose, Costa Rica; for
airmail delivery, add $5.00 to the price. To obtain Vols. 1 (Attacinae) and 2 (Arsenurinae),
I suggest contacting the author directly, as he had these volumes privately printed. The
address of this Lepidopterists’ Society member is La Croix des Baux, F-84220 Gordes,
France.

FREDERICK H. RINDGE, Department of Entomology, American Museum of Natural
History, New York, New York 10024.

Journal of the Lepidopterists’ Society
43(2), 1989, 155

FEATURE PHOTO

Unusual moth pupa, Gonionota sp. (Oecophoridae), from Ecuador. It is covered with
blunt projections and two lateral flanges, and is shown in dorsal view. It measures 10 mm
high by 6 mm maximum width and was found upright and exposed on a leaf surface. It
probably mimics an inedible object. Other Gonionota spp. (J. A. Powell 1973, Smith.
Contr. Zool. 120, 302 pp.) and Hypertropha (I. F. B. Common 1980, Entomol. Scand.
11:17-31) share a similar mode of pupation. Photo taken with a Minolta X-570 and 80
PX ring flash on a 50 mm macrolens. Pupa collected in Ecuador, Pichincha Prov., Hotel
Tinalandia, during the period 5-15 May 1988 by S. Passoa, on an undetermined shrub;
adult emerged 19 May 1988.

STEVEN Passoa, Department of Entomology, University of Illinois, 320 Morrill Hall,
Urbana, Illinois 61801.

Journal of the Lepidopterists’ Society
43(2), 1989, 156

MANUSCRIPT REVIEWERS, 1988

The merit of a scientific journal depends on the quality of its reviewers as well as of
its authors, but the former are usually unknown to readers. The Journal acknowledges
with gratitude the services of the people listed below from whom manuscript reviews

were received in 1988.

P. R. Ackery, London, England
Peter H. Adler, Clemson, SC

Paul H. Arnaud, San Francisco, CA
Matthew P. Ayres, East Lansing, MI

R. Robin Baker, Manchester, England

Gregory R. Ballmer, Riverside, CA

Gary D. Bernard, New Haven, CT

Carlos R. Beutelspacher, México, D.F.,
Mexico

M. Deane Bowers, Cambridge, MA

John W. Brown, Washington, DC

Keith S. Brown Jr., Campinas, Sao Paulo,
Brazil

John M. Burns, Washington, DC

Karen M. Calloway, Long Beach, CA

Donald R. Davis, Washington, DC

John C. Downey, Sarasota, FL

Boyce A. Drummond, Woodland Park,
CO

J. N. Eliot, Taunton, Somerset, England
Thomas C. Emmel, Gainesville, FL

Douglas C. Ferguson, Washington, DC
Clifford D. Ferris, Laramie, WY :

Lawrence F. Gall, New Haven, CT
Patricia Gentili, Washington, DC
George L. Godfrey, Champaign, IL
Michael D. Greenfield, Los Angeles, CA

David F. Hardwick, Perth, Ontario,
Canada

John B. Heppner, Gainesville, FL

Ronald W. Hodges, Washington, DC

Dale W. Jenkins, Sarasota, FL

Roy O. Kendall, San Antonio, TX
Joel G. Kingsolver, Seattle, WA
Ian J. Kitching, London, England

Be

Jean-Francois Landry, Ottawa, Ontario,
Canada

Robert C. Lederhouse, East Lansing, MI

Sanford Leffler, Seattle, WA

C. Don MacNeill, Oakland, CA
Sterling O. Mattoon, Oroville, CA
Tim L. McCabe, Albany, NY
James S. Miller, New York, NY
Lee D. Miller, Sarasota, FL
Thomas A. Miller, Frederick, MD

H. H. Neunzig, Raleigh, NC
Paul A. Opler, Fort Collins, CO

Ken Pivnick, Saskatoon, Saskatchewan,
Canada
Jerry A. Powell, Berkeley, CA

Hector E. Quintero, San German, PR

Donald F. Ready, West Lafayette, IN
Thomas J. Riley, Baton Rouge, LA
Frederick H. Rindge, New York, NY
Robert K. Robbins, Washington, DC
Gaden Robinson, London, England

Theodore D. Sargent, Amherst, MA
Malcolm J. Scoble, London, England
James A. Scott, Lakewood, CO
Arthur M. Shapiro, Davis, CA
Oakley Shields, Mariposa, CA

John A. Shuey, Columbus, OH
Steven R. Sims, St. Louis, MO

M. Alma Solis, Washington, DC

J. Bolling Sullivan, Beaufort, NC

Michael E. Toliver, Eureka, IL
James P. Tuttle, Troy, MI

Ralph E. Wells, Jackson, CA
Diane C. Wiernasz, Seattle, WA
Ernest H. Williams, Clinton, NY

EDITORIAL STAFF OF THE JOURNAL

Boyce A. DRUMMOND, Editor WILLIAM E. MILLER, Retiring Editor
Natural Perspectives Dept. of Entomology
P.O. Box 9061 University of Minnesota
Woodland Park, Colorado 80866 U.S.A. St. Paul, Minnesota 55108 U.S.A.

Associate Editors:
M. DEANE BOWERS, DOUGLAS C. FERGUSON, LAWRENCE F. GALL,
ROBERT C. LEDERHOUSE, THOMAS A. MILLER, ROBERT K. ROBBINS,
THEODORE D. SARGENT

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories
are Articles, General Notes, Technical Comments, Book Reviews, Obituaries, Feature
Photographs, and Cover Illustrations. Reviews should treat books published within the
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contributors should prepare manuscripts according to the following instructions, and
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Abstract: An informative abstract should precede the text of Articles.

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Literature Cited: References in the text of Articles should be given as Sheppard (1959)
or (Sheppard 1959, 1961a, 1961b) and listed alphabetically under the heading LITERATURE
CITED, in the following format without underlining:

SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London.
209 pp.

196la. Some contributions to population genetics resulting from the study of

the Lepidoptera. Adv. Genet. 10:165-216.

In General Notes and- Technical Comments, references should be shortened and given
entirely in the text as P. M. Sheppard (1961, Adv. Genet. 10:165-216) or (Sheppard, P.
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CONTENTS

ELECTROPHORETIC COMPARISONS OF VICARIANT VANESSA: GE-
NETIC DIFFERENTIATION BETWEEN V. ANNABELLA AND V.
CARYE (NYMPHALIDAE) SINCE THE GREAT AMERICAN INTER-
CHANGE. Arthur M. Shapiro © Hansjurg Geiger

EVALUATION OF SPERMATOPHORE COUNTS IN STUDYING MATING
SYSTEMS OF LEPIDOPTERA. Robert C. Lederhouse, Matthew
P. Ayres & J. Mark Scriber 0 a

BEHAVIOR OF THE TERRITORIAL SPECIES LIMENITIS WEIDEMEYERII
(NYMPHALIDAE) WITHIN TEMPORARY FEEDING AREAS. Risa
H. Rosenberg us 3

A NEW NEOTROPICAL ANONCIA SPECIES (COSMOPTERIGIDAE).
David Adamski: 2.5 2 ee

A NEW SUBSPECIES OF NEONYMPHA MITCHELLII (FRENCH) (SA-
TYRIDAE) FROM NorTH CAROLINA. David K. Parshall &
Thomas W. Kral 0020 ee eee

REVISION OF CHLOROSTRYMON CLENCH AND DESCRIPTION OF
TWO NEW AUSTRAL NEOTROPICAL SPECIES (LYCAENIDAE).
Kurt Johnson 3.0

GENERAL NOTES
Papilio troilus L. on a new and rare larval food plant. M. Wayne Morris ....
Spencer collection given to Smithsonian. J. F. Gates Clarke 0c

Evidence for genetic determination of variation in adult size and wing mel-
anism of Parnassius phoebus F. C..S; Guppy 0 ee

Effect of refrigeration on egg incubation period of the tasar silk insect An-
theraea mylitta Drury (Saturniidae). B. K. Nayak & A. K. Dash ..........

Book REVIEW
The Saturniidae of America. Ceratocampinae. Frederick H. Rindge ...........
FEATURE PHOTOGRAPH

Steven Passoa

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

81

93

102

108

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Volume 43 1989 Number 3

ISSN 0024-0966

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS

7 September 1989

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

JACQUELINE Y. MILLER, President NIELS P. KRISTENSEN, Vice
JULIAN P. DONAHUE, Immediate Past President

President DONALD J. LAFONTAINE, Vice
RICHARD HOLLAND, Vice President President
WILLIAM D. WINTER, Secretary JAMES P. TUTTLE, Treasurer

Members at large:

Jo BREWER JOHN W. BROWN RICHARD A. ARNOLD
DALE W. JENKINS MOGENS C. NIELSEN SUSAN S. BORKIN
JOHN E. RAWLINS FLOYD W. PRESTON Davip L. WAGNER

EDITORIAL BOARD

PAUL A. OPLER (Chairman), FREDERICK W. STEHR (Member at large)
Boyce A. DRUMMOND (Journal), WILLIAM E. MILLER (Memoirs), JUNE PRESTON (News)

The object of the Lepidopterists’ Society, which was formed in May 1947 and for-
mally constituted in December 1950, is “to promote the science of lepidopterology in all
its branches, ... . to issue a periodical and other publications on Lepidoptera, to facilitate
the exchange of specimens and ideas by both the professional worker and the amateur
in the field; to secure cooperation in all measures” directed towards these aims.

Membership in the Society is open to all persons interested in the study of Lepi-
doptera. All members receive the Journal and the News of the Lepidopterists Society.
Institutions may subscribe to the Journal but may not become members. Prospective
members should send to the Treasurer full dues for the current year, together with their
full name, address, and special lepidopterological interests. In alternate years a list of
members of the Society is issued, with addresses and special interests. There are four
numbers in each volume of the Journal, scheduled for February, May, August and
November, and six numbers of the News each year.

Active members—annual dues $25.00
Student members—annual dues $15.00
Sustaining members—annual dues $35.00
Life members—single sum $500.00
Institutional subscriptions—annual $40.00

Send remittances, payable to The Lepidopterists’ Society, to: James P. Tuttle, Treasurer,
3838 Fernleigh Ave., Troy, Michigan 48083-5715, U.S.A.; and address changes to: Julian
P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, California
90007-4057 U.S.A. For information about the Society, contact: William D. Winter, Sec-
retary, 257 Common St., Dedham, Massachusetts 02026-4020, U.S.A.

For back issues of the Journal and News, write the Publications Coordinator at the
address below about availability and prices. Prices of The Lepidopterists Society com-
memorative volume 1945-1973 are $12.00 ($8.00 to members and subscribers); of A
catalogue/checklist of the butterflies of America north of Mexico, clothbound, $19.00
($12.00 to members and subscribers), paperbound, $10.50 ($7.00 to members and sub-
scribers). Order from the Publications Coordinator, Ronald Leuschner, 1900 John St.,
Manhattan Beach, California 90266-2608, U.S.A. Make remittance payable to “The Lep-
idopterists’ Society.”

Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for
$40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’
Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los
Angeles, California 90007-4057. Second-class postage paid at Los Angeles, California and
additional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’
Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, California 90007-
4057. ‘

Cover illustration: Female of Gastropacha populifolia (Esper) (Lasiocampidae) in nat-
ural resting position on a dead branch, Beijing, China. Submitted by Yu Xiangming, No.
31, Qian Men Wai Zhu Bao Shi, Beijing, China.

JOURNAL OF

Tue LeEerPipopTrerists’ SOCIETY

Volume 43 1989 Number 3

Journal of the Lepidopterists’ Society
43(3), 1989, 157-166

PRESIDENTIAL ADDRESS, 1988:
LEPIDOPTERISTS—COLLECTORS AND BIOLOGISTS?!

JERRY A. POWELL

Department of Entomological Sciences, University of California,
Berkeley, California 94720

Additional key words: food plants, diapause, behavior, longevity, seasonal abundance.

Traditionally this Society has invited the president to expose ideas
and opinions in an address, even though they may reflect little hard
data. Today is no exception. This discussion will try to encourage col-
lectors, especially amateurs, to devote part of their seemingly limitless
energy to the study of Lepidoptera biology.

For purposes of this discussion, an amateur is someone who has to
pay money to study Lepidoptera; a professional is someone who gets
paid to study Lepidoptera. We all know amateurs who do excellent
work and accomplish an astonishing amount, and some professionals
who don’t get much done. There may even be a few examples of the
reverse. Similarly, by this definition there are amateurs with Ph.D.-
level training in biology and professionals without it. Hence, there is
no inferior connotation in my use of the term amateur.

I thought it might be fun to begin by looking at a subject that is of
interest to spouses and other people who get dragged to these meetings
or into other embarrassing situations, that is: Why do we collect Lep-
idoptera?

The urge to accumulate collections is, of course, not restricted to
Lepidoptera—the affliction is widely expressed in non-biological arti-
facts, and it seems unrelated to genetic or environmental inheritance.

My earliest recollections of collecting, when I was 7 or 8 years old,
are of bottlecaps. (This was long before twist-top caps were invented,
and it was a challenge to get specimens in perfect condition, because

1 Delivered to the Annual Meeting of The Lepidopterists’ Society in Pittsburgh, Pennsylvania, on 16 July 1988

158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

people would bend the caps when popping them off with openers called
“church keys’; these were given out free with beer purchases and had
beverage company names imprinted, so some people collected church
keys, although I never found that particularly fascinating.) My parents,
who never had the slightest interest in collections (which is also true of
my brother, wives, and kids), watched, presumably bemused, as I pro-
gressed through matchbook covers, military implements, fossils, sea-
shells, and moths, assuming I suppose, that I would mature, get over
the penchant, and settle down to dentistry or some other respectable
career. They were, of course, wrong on all counts.

Recently I heard a M.D. who collects art and antiques, express it on
TV: “Collecting is an affliction that is intractable as any virus, one for
which there is no immunity nor cure.” We simply have it. I understand
that psychologists term it a “personality disorder’; but, quite frankly,
I find people who don’t have the addiction kind of deprived.

Gary Larsen’s cartoon depicting the guys returning triumphantly
from the hunt with a huge swallowtail tied to the hood of their car
probably gives a better perception of our feelings than most of us could
verbalize; nonetheless, I will try to analyze why collectors collect Lep-
idoptera. I divide the phenomenon into four components: 1) Lure of
Collectibles, 2) Hunter Instinct, 3) Acclaim from Peers, and 4) Satis-
faction in Discovery.

Probably any lepidopterist would think of other ways to dissect the
reasons why collecting is compelling, but most will recognize two or
three elements here that contribute to the urge to collect. Any one of
these might be the main source of pleasure for any given collector, but
probably most of us have never tried to explain it and don’t feel a need
to. It is only our incredulous friends and relatives that ask, or don’t ask
but just sigh and look the other way.

Lure of collectibles. It is unfathomable what constitutes collectibles.
Apparently, like a queue in England, it only takes two or three. A few
of anything that can be conceived of as constituting a set or series will
suffice to start a collection. I find it imponderable that someone recently
paid $650 for a copy of the high school yearbook of Don Mattingly’s
graduating class. In fact I don’t understand the urge to collect where
it depends mainly upon purchases, such as art, yet it must be incredibly
compelling. Every conceivable series of objects (or even non-series in
the case of Andy Warhol) can be and is collected. The urge has nothing
to do with biology, necessarily, as can be attested by the number of
lepidopterists who also collect postage stamps—one even collects mono-
grammed golf balls!

Pleasure from the collection itself is the primary goal in. some in-
stances, as epitomized by European collectors who buy specimens from

VOLUME 43, NUMBER 3 159

Tropical Regions-at auction for large sums. (Others of us buy them
discretely from Welling or Plaumann.) Many collectors seem to derive
a lot of their satisfaction and pleasure from the appearance of a neatly
curated collection. They actually like spreading and preparing speci-
mens, I guess. This aspect of the affliction provides continuing challenges
in time and effort of preparation of specimens, in attempting to obtain
perfect specimens to replace less aesthetically pleasing ones (if this is
the main goal), and in keeping up with costs of equipment and space
for storage (less a challenge with micros than with saturniids, of course).

Hunter instinct. Quite aside from the resultant collection, there is
considerable satisfaction derived from the challenge of the hunt—plan-
ning the quest, searching for the appropriate habitats, predicting the
timing of visits and so on. The anticipation is half the fun (often more).
Also satisfying is the skill required in stalking and catching the prey,
particularly for rare species and especially for those not seen before.
As we all know, those are the hardest ones to catch. This seems to be
the leading source of satisfaction for some collectors, to hear them boast.
It certainly must be more important than preparation and curating for
many collectors, to judge from the amount of papered material that
accumulates.

For many of us, I think, it is the lure of adventure that is a strong
factor. To see the open road ahead, leading to new and potentially
exciting areas (particularly if other lepidopterists have not visited them),
is the seduction, coupled with the anticipation that something new may
be discovered. The adventure: to collect in exotic areas is the need—
the specimens are secondary. Most lepidopterists, if given the choice,
obey Powell’s Law (Munroe, E. G. 1969, Proc. Entomol. Soc. Ontario
99:43), which can be paraphrased as, “No biologist willingly collects
within 1,000 miles of his home base.’ Thus, lepidopterists living in
California go to Mexico to collect, or to Costa Rica if we have a grant;
our host at the Carnegie leaves Pennsylvania to collect in Ecuador and
Taiwan; people in Kentucky and northward all go to Florida every
spring, while those in Florida are gone to Trinidad or Hispaniola (that
is not 1000 miles unless you are from Gainesville, but that’s OK because
it’s an island); people in Washington spend summers in Colorado and
Utah, except for Don Davis who collects everywhere else in the world;
everybody collects in southern Arizona except Arizonans, who go to
Mexico.

Doug Ferguson is the exception; they say he collects in his yard in
Maryland. Incidentally, Ferguson, our immediate past president, wrote
me and said he would not be able to attend the Executive Council
meeting here—he is collecting in British Columbia.

Simply the enjoyment of getting out to natural areas, away from

160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

phones, freeways, smog, commuting, demands of the job and respon-
sibilities at home has to be a big factor, for amateurs and professionals
alike. After all, collecting is a lot more fun than committee meetings,
preparing lectures or budget reports, etc.

Acclaim from peers. For some, there is pride in exhibiting accom-
plishments; presumably these often are the same people who get the
most satisfaction out of the collection itself. Competitiveness is a factor,
certainly more so for some people than others.

Most lepidopterists would not believe that fame is much of a factor
in why we collect (notoriety is a better descriptor), yet I wonder how
many of us would maintain enthusiasm if we thought absolutely nobody
else cared (as opposed to hardly anybody else)? Even though we go
collecting mainly for the enjoyment, challenge, and satisfaction in ob-
taining the specimens, can you really say that often you don’t think
“wait till so and so hears about this!’’?

I know one of the things I really enjoy is discovering things for other
researchers, and I think this is a prevalent feeling among many collec-
tors, amateur or professional (of course it is particularly enjoyable if it
is a species I think they have overlooked in areas they have or could
have worked).

The lure of patronyms should be mentioned. Some collectors are
unabashed in their admitted desire for this form of immortality; others
do not admit it, yet they look coyly away, suppressing a smile of delight,
if you mention it. Possibly some hardened professionals don’t care at
all, but you would be tempted to question their honesty. The indignant
condemnation of the increasing use of patronyms voiced by Dimock
(1984, J. Res. Lepid. 23:94-101) was misguided and pathetic—mis-
guided because he did not list the two most useful roles patronyms
fulfill, to acknowledge collectors’ efforts and to avoid secondary hom-
onymy, and pathetic because it will be ignored.

Satisfactory in discovery. Beyond the fun of collecting and the plea-
sure in curating the collection, for biologists there is the added feeling
of accomplishment in discovering new information, finding out things
that nobody has known before. I see this as a bonus to the lure of
collecting, one that you would not derive from collecting stamps or
baseball cards.

For sheer joy of accomplishment, I don’t think the discovery of facts
“new to science” is surpassed by any other aspect of collecting. Who
among us is not pleased by finding a new population or state record of
even a well-known species?

For specialists in microlepidoptera, finding a new species in a museum
collection is not very exciting; it means more dissections and descriptive

VOLUME 43, NUMBER 3 161

work—Ron Hodges has how many new Chionodes, 150? But finding
a new species that you recognize in the field—ah! that is another matter.
Then you feel you are the discoverer, not just a processor filling in
another space in the stamp album.

For me, there are two kinds of discoveries from which I derive the
most satisfaction. First, there is the finding of a “lost” species, one
collected and described long ago and perhaps known only from one or
a few specimens. For example, the rediscovery in Chihuahua of Apo-
demia phyciodoides a few years ago must have been a great thrill to
Richard Holland (although no doubt he showed no outward display of
excitement). Second, even more enjoyable to me, is the discovery of
the key to an insect’s biology, particularly a species that has been known
for a long time to lots of collectors.

It is this last, of course, that I wish to emphasize today—a satisfaction
that is available to everybody without obeying Powell’s Law, if you
spend some time watching the animals instead of taking the pinch-first-
and-ask-questions-later approach. I can share a couple of experiences
of these kinds of discoveries:

1) Rediscovery of Ethmia minuta. I began a study of Ethmia while
still a student. One day on a visit to the San Diego Natural History
Museum, I found specimens of this elegant little species—at the time
the smallest known member of the genus and the only one with marked
sexual dimorphism in wing color—the kind of thing that, as a specialist,
you say immediately: “that’s new.’ But these had been collected by
W.S. Wright in 1916 and labelled “San Diego.’ During the interim,
San Diego had grown from a village of several thousand people to a
city with a population of % million and huge urban sprawl, so there
seemed little hope of recovering the species. I will never forget the
thrill then, when a couple of years later in the foothills back of the city
I found adults of this “‘lost’’ (for 45 years) species at flowers of Crypt-
antha, which proved to be the key to its interesting biology, with the
female ovipositor greatly modified to penetrate the densely hirsute floral
buds.

2) The surprising biology of Ethmia scylla. I collected the first spec-
imen of this nondescript species at Mt. Diablo near my home in 1959.
John Burns and I went out the following spring and collected a nice
series, which was gratifying; but 10 years were to pass before I discov-
ered its biology. This involved repeated trips early each spring, mis-
guided in the belief that some borage or hydrophyll must be the host
plant because most ethmiids depend upon those plants. Finally I caged
females with unlikely (to me) plants from the habitat, and in one day
the females chose what they wanted for oviposition. The larvae feed

162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

in the flowers of Collinsia; Ethmia scylla is the only species in this
worldwide genus known to use Scrophulariaceae. It was a satisfying
find but also taught me a lesson about making assumptions.

WHAT KINDS OF BIOLOGICAL STUDIES ARE NEEDED?

‘In the remaining time, I will briefly summarize some examples of
biological studies of the kinds any of you can carry out with minimal
equipment in your local area.

Larval Foods and Habits

The most obvious biological characters to most lepidopterists are the
food plants. You might think that this aspect is pretty well documented,
but even for North American butterflies much remains to be discovered.
One of the most famous for his untiring efforts in this field is Roy
Kendall in Texas. In response to my inquiry he estimates that he has
reared more than 750 species of Lepidoptera, including about 330
species of butterflies. About 40% of these are thought to have been
previously unknown. He has more than 2000 vials of preserved larvae.
I would like to quote from a letter:

“I can’t recall anytime during the past 30+ years when there was
no livestock in my lab, and there is no end in sight [at age 76]. Although
many lepidopterists consider certain species ‘trash,’ I find them very
interesting and often rear these as well as ‘goodies’ numerous times
from different localities.’’ He also says, “Incidentally, I am an amateur
in every sense of the word. The only formal training received was a
3-hour high school course in zoology.’ Yet Kendall probably has con-
tributed more to our knowledge of larval biology of North American
Lepidoptera than any other single person. Publications by Kendall or
others with whom he readily shares unpublished data have recorded
host plants or other information on about 500 species.

While it often is a lot of work, compared to merely collecting and
killing adult Lepidoptera, I cannot overemphasize the need for this
kind of work: the repeated study of biologies of different populations
of the same species, in order to confirm existing records and to discover
and document geographical and seasonal variation in biological char-
acteristics. Just because a butterfly book states that a certain plant is
the host of a species does not mean that its biology is known. You should
question all such statements; errors are perpetuated by repeating from
such books, and, even if correct, the statement may be based on a single
record or apply only to a portion of the insect’s range. Moreover, when
one of the beautiful adults emerges, it is a lot more satisfying than
going to some locality listed in the Season Summary to recollect adults.

Important kinds of rearing studies that need to be carried out include

VOLUME 43, NUMBER 3 163

emphasizing diverse larval niches, not just external foliage feeding
caterpillars. Many species feed in leaf litter or as borers within roots or
stems, in seeds, galls, or leaf mines. Backyard studies, such as that
reported here yesterday by Bill Miller on sibling species of gall moths,
await the attention of lepidopterists in every part of North America.
Few places have been well surveyed for leaf mining species, yet the
various genera have highly characteristic forms of mines by which you
can learn to identify them, and they often live for leng periods in this
stage, so that the precise timing of search needed for the adults is not
so critical. Wagonloads of food plant and a pitchfork are not needed
as when you rear saturniids; just hold the leaf in a vial for a few days
and often a beautiful (and frequently undescribed) moth comes out.

Such studies are best carried out on a local basis, where you can
repeatedly visit a habitat. Any place in the Western Hemisphere will
have literally hundreds of species that have never been reared before,
or have only been studied in another region. John De Benedictis has
carried out a several-year survey at San Bruno Mountain near San
Francisco and to date has reared about 150 species of microlepidoptera;
still, each visit recovers larvae that he, and often anyone else, has never
seen before. Patience and painstaking search of the different ecological
horizons (roots, stems, flowers, fruit, mines, etc.) of all available potential
host plants are the requisites.

Before leaving this topic, I’ll make a pitch for preserving larvae. It
is easy to obtain good specimens by simply immersing in boiling water
for a few seconds or minutes and then preserving in drugstore rubbing
alcohol. Far more species have been reared than the number for which
we have material useful for larval studies, even in butterflies. Much of
the emphasis in the past has been to obtain perfect specimens of the
adults. Photographs of the larvae are not adequate for identification of
most moths, and our knowledge of larval taxonomy lags far behind that
of the adults for nearly all families.

Adult Behavior, Longevity

Mark-release-recapture studies of individuals, while time-consuming,
are fun to do. They yield information on dispersal, differential move-
ments of males and females, lifespan, feeding habits and so on, and
they have been carried out for rather few North American Lepidoptera.
These studies do not have to be very sophisticated to produce new
information. All you really need is a felt-tipped pen with permanent
ink, a notebook, and a net. For example, Smith (1982, J. Lepid. Soc.
35:172) marked and released common butterflies in his backyard in
Sacramento and learned from recaptures that individuals of Pieris rapae
and Papilio rutulus live up to 39 days, Battus philenor up to 44 days.

164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

We lack this kind of information for almost all Nearctic butterflies and
moths.

My backyard was the exotic locality where I studied mating behavior
of Incisalia iroides (1968, J. N.Y. Entomol. Soc. 76:47). The whole study,
which I think still records the most data on mating of any North
American thecline, took place at a small lemon tree that the males liked
to use as a perch. Mating occurred in late afternoon and extended into
evening, so I could easily handle the mated pairs, mark individuals,
and return them undisturbed to their perch. I suspect that mating habits
of theclines generally have been overlooked because the butterfly people
tend to keep bankers’ hours.

Waldbauer and Sternberg (1982, J. Lepid. Soc. 36:154-155) released
marked Hyalophora in Illinois and recorded recaptures of 18 males 6.8
miles away, using virgin females as bait; and, in a similar study, Toliver
and Jeffords (1981, J. Lepid. Soc. 85:76) recorded Callosamia move-
ments 14 and 36.5 miles from their release points. But for the vast
majority of Lepidoptera we have no data on dispersal capabilities.

Mark-release-recapture studies of skippers have been few and not
wholly successful. Handling most species evidently disturbs the indi-
viduals more so than is true of other butterflies. After releasing about
50 marked Paratrytone and never seeing one return, I developed a
method of marking them without capture. Using a brush made from
a feather, I found that males could be marked as they perched, with
a mixture of ink and paint. Residency and competition for perches
could then be monitored.

Studies of adult feeding also are needed. Paul Opler recorded floral
visitations of butterflies in Virginia and found their choices to be a
correlation of tongue length and corolla depth (Opler, P. A. & G. O.
Krizek, 1984, Butterflies east of the Great Plains, Johns Hopkins Univ.
Press, Baltimore, Maryland, 294 pp.), rather than just by color, or by
plant taxon, as butterfly enthusiasts often assume.

One of the most remarkable studies on feeding is that of Bill Miller,
who carried all his equipment to the stage when he reported the study
to us at Berkeley last year: a dixie cup, a water vial, and a wick. He
demonstrated increased fecundity in the spruce budworm when females
imbibe nutrients (1987, Environ. Entomol. 16:1291-1295). This may
not seem profound to you, but a recent bibliography recorded more
than 4000 references to this insect, easily the most intensively studied
species of Nearctic Lepidoptera, yet nobody had done this kind of study
previously.

“Mud puddling” has received some attention, but there are many
unanswered questions. Only one extensive study, that of Adler, has been
carried out (1982, J. Lepid. Soc. 36:161-173). He recorded 93 species

VOLUME 43, NUMBER 3 165

of moths at mud in New York; 99% were males. However, 80% of one
geomtrid visited flowers instead. Why don’t females do this, and why
is it so rare in California? Why do some species have this habit while
others do not?

Predation is another phase of biology that everybody seems to take
for granted but nobody does much about documenting. The observa-
tions by Paul and Anne Ehrlich on lizard predation of tropical butterflies
a few years ago is an example of how data can be recorded with a little
patience (1982, J. Lepid. Soc. 36:148-152).

Seasonal Abundance

This is another field wide open for investigation. The classic study
is Ehrlich’s team research on Euphydryas editha over a 25-year period
(1975, Science 188:221-228, et seq.). But such sophistication and fund-
ing are not necessary. The counts by Sidney Hessel of Catocala attracted
to mercury vapor lamps at one site in Connecticut during a 12-year
span, summarized in Sargent’s book (1975, Legion of night, Univ. Mas-
sachusetts Press, Amherst, 222 pp.), are almost without parallel. Indi-
cations of increase or decline seen during a five- or six-year period were
misleading when longer term fluctuations were observed. Smith (1984,
J. Lepid. Soc. 37:275-280) also did this by counting butterflies for two-
hour periods in his backyard for 12 years. There were large year-to-
year fluctuations but no general trends, such as are often alleged.

This is a reason that the annual counts of butterflies sponsored by
the Xerces Society are useful. We had 87 counts reported in 1987 [99
in 1988]; if we can obtain 150 or 200 that are reported on a continuing
basis, general trends in abundance, as well as migrations and other
comparative data, will be enhanced. A 15-mile diameter circle is se-
lected and all the butterflies seen in one day counted. The object is to
compare abundances from year to year at about the same date at each
site. Obviously a place like Berkeley is not going to have the species
richness of a site in southern Arizona or the Rocky Mountains, but after
14 years we have a good basis for predicting and explaining increases
and decreases in abundance from one year to another in our circle.

Diapause

For most species we have little information on diapause development.
The study by Sims of Papilio zelicaon (1988, J. Lepid. Soc. 37:29-37)
is a good example of what can be done. He showed that populations
on native umbells were univoltine, and modification of the diapause
pattern enabled adventive populations to colonize urban areas on sweet
fennel throughout the season. Incidentally, outdated terms such as
“breaking” and “triggering” should be dropped from your vocabulary;

166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

the process is a dynamic one that takes place over many weeks or
months. Treatments such as constant temperature chilling that results
in development in one instance may not do so for all populations of a
species or even all individuals of a population.

A special interest of mine has been prolonged diapause, the main-
tenance of dormancy for more than one year. I published a summary
of knowledge for Lepidoptera last year (1987, J. Res. Lepid. 25:83-
109). In yucca moths under optimum winter environments, all or nearly
all larvae complete development, while in adverse conditions, all or
nearly all maintain diapause. Adults emerge over several years, even
though neighbors in the same plant have completed development in a
prior season. I have emergences now up to 19 years [20 years in 1989]
after collection of the fully fed, prepupal larvae, so they are prepared
to wait out the adversity and the lepidopterists’ patience. One advantage
of such studies is that they are not very labor intensive.

In conclusion, the take-home message is that I think the anticipation
and realization of discovering something new is a major factor in the
attraction of collecting Lepidoptera. This part of the enjoyment and
satisfaction can be fulfilled in your local area if part of your effort is
devoted to study of biological or behavioral aspects of butterfly and
moth populations, rather than continuing an emphasis on subspecies
and county records.

Received for publication 22 May 1989; accepted 22 May 1989.

Journal of the Lepidopterists’ Society
43(3), 1989, 167-177

REPRODUCTIVE ENHANCEMENT BY ADULT FEEDING:
EFFECTS OF HONEYDEW IN IMBIBED
WATER ON SPRUCE BUDWORM

WILLIAM E. MILLER

Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108

ABSTRACT. Captive budworm adults obtained as wild pupae were divided into two
groups, and each group was offered liquids to imbibe once daily. One group received
plain water, the other water containing honeydew of hemispherical scale, Saissetia coffeae
(Walker), at concentrations averaging 6.5%. Two such experiments were conducted, a
preliminary one with 28 fertile pairs, and the main one with 34 fertile pairs. In both
experiments, honeydew prolonged female lifespan, egg maturation, and oviposition, the
latter two causing redistributions within apparently fixed oocyte complements. Honeydew
effects interacted with female body size, large females laying relatively more eggs than
small females. In the main experiment, imbibing did not begin until the third day of
adulthood. Thereafter, fertile females imbibed more often than infertile ones, the fre-
quency among the former peaking at 97% of their number during the seventh and eighth
days of adulthood. Amount imbibed per individual per day averaged 4.5 mg as determined
by weighing females before and after imbibing. In fertile females of average lifespan,
expected lifetime honeydew-water intake was 31.6 mg of liquid containing 2.05 mg dry
weight of honeydew, the latter corresponding to 3.5% of female average initial live weight.
Enhanced reproductive effects did not appear until late in adulthood.

Additional key words: Choristoneura fumiferana, Tortricidae, Tortricinae, fecun-
dity, Saissetia coffeae.

That tortricine adults are capable of imbibing has long been known
(Powell 1965:26), but only recently was reproduction in a tortricine
rigorously shown to be enhanced by imbibing. Thus, the spruce bud-
worm, Choristoneura fumiferana (Clemens) (Tortricidae), matured
more oocytes and laid more eggs when females received water con-
taining 15% bee honey than when they received plain water (Miller
1987). Imbibing habits of the adults are still obscure, however.

Honeydews might be available to spruce budworm adults in nature.
Precipitated water is often present, and such water is doubtless sweet-
ened at times by honeydews of co-occurring insects such as the balsam
twig aphid, Mindarus abietinus Koch (Aphididae). While bee honey
consists mostly of fructose and glucose (White 1975), honeydews may
contain these sugars as well as sucrose, other sugars, and nitrogenous
compounds (Auclair 1963).

Here, for captive budworm females given plain water or water con-
taining honeydew, I report several aspects of reproduction as well as
imbibing frequencies and amounts imbibed. Reproductive attributes
measured were lifespan, oviposition period, number of eggs laid daily,
egg viability, number of mature and immature oocytes in ovaries after
death, and attributes derived from these. The honeydew source was
hemispherical scale, Saissetia coffeae (Walker) (Coccidae).

168 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

MATERIALS AND METHODS

Two experiments were conducted, a preliminary one followed by
the main one. In both experiments, one group of moths received hon-
eydew solution for imbibing, and a second group received plain water.
The main experiment differed from the preliminary one chiefly in that
pupae and emergent adults were not cold-stored before use, female
imbibing was recorded, the different imbibing liquids were dispensed
in exactly the same way, and reproductive attributes were measured
in greater detail.

For both experiments, wild pupae reachable from the ground were
collected from 25 or more large trees of balsam fir, Abies balsamea
(L.) Mill. (Pinaceae), growing in one area of 0.1 ha or less. Pupal
collections were timed to coincide with incipient adult emergence. In
the preliminary experiment, pupae came from 12 km E of Meadow-
lands, St. Louis Co., Minnesota, and in the main experiment, from 8
km W of Grand Marais, Cook Co., Minnesota. In the preliminary
experiment, pupae and emergent moths were stored within 36 h of
collection at 8°C and held for two weeks during a start-up delay. In
the main experiment, pupae and emergent moths were in use within
36 h of collection.

The first step in both experiments was to sex pupae (Jennings &
Houseweart 1978) and emergent adults. Some females were freeze-
killed within 2 h of eciosion for ovarial study. Male-female pairs for
imbibing experiments were placed in 1-pint (0.48 |) cardboard ice cream
containers capped with Petri dish lids, one pair per container. A shoot
of balsam fir ca. 8 cm long was placed in each container as a substrate
for oviposition; in the preliminary experiment, the shoot also served as
an imbibing substrate for plain water. Containers were numbered and
assigned by equal and odd numbers to the two imbibing treatments.

Moth containers in both experiments were held in a temperature-
controlled room maintained at 23°C during the preliminary experiment,
and at 25°C during the main experiment. In the former, moths received
natural July light through a large N-facing window; in the latter, they
received fluorescent light on a 12L:12D schedule. Containers were
examined daily near mid-day, at which time imbibing liquids were
introduced and reproductive data were gathered.

The honeydew-providing colony of hemispherical scale infested a
2-m tall indoor-growing spineless yucca, Yucca elephantipes Regel (Lil-
iaceae). Upper surfaces of the plant’s leaves were nearly completely
coated with honeydew. In both experiments, segments ca. 6 cm? were
cut from the honeydew-laden leaves, misted with water just short of
runoff to form honeydew-water solution, and placed in moth containers

VOLUME 43, NUMBER 3 169

for imbibing. Segments were misted every day and replaced every
second or third day. Plain water for imbibing was provided in the
preliminary experiment by misting the balsam fir shoot, and in the
main experiment by misting yucca leaf segments also ca. 6 cm? from
which honeydew had been washed. All misting was done outside con-
tainers with a hand-powered household sprayer containing distilled
water.

Misted yucca leaf segments remained wet in moth containers for
1.2-1.5 h before drying naturally. Whether or not females imbibed was
determined by monitoring main-experiment individuals during this
interval on arbitrarily chosen days. Imbibing moths were spotted by
their characteristic preimbibing head movements, and by proboscises
extending to the wet yucca leaf segments.

Liquid intake was measured as the difference between pre- and
postimbibing weights of individual females preweighed just before they
were routinely offered liquids. Sample females were selected arbitrarily
for this purpose, and weights were recorded to the nearest 0.1 mg.
Imbibing occurred rapidly enough so that pre- and postimbibing weigh-
ings were seldom separated by more than 25 min. To verify that weight
differences truly represented intake, two females that walked and rested
on misted yucca leaf segments for 5 min without imbibing were weighed
in the same manner as imbibing females. These nonimbibers underwent
no weight gain, thus indicating that imbibing liquids were not absorbed
by body parts coming in contact with wet surfaces.

Eggs were spotted by examining shoots and container walls under a
2x reading magnifier. Shoots were removed from containers for this
examination. Deposited eggs were counted under stereomicroscope
magnifications up to 25x. In the preliminary experiment, female fer-
tility and egg viability were determined by observing 21-112 deposited
eggs per female for a week or until larval heads showed through cho-
rions. In the main experiment, fertility and viability were determined
by observing all deposited eggs until hatching or imminent hatching.

Oocytes in excised ovaries were identified as mature or immature
by size and stainability after 2-4 min exposure to ca. 0.2% aqueous
methylene blue. Chorionated (mature) eggs take up such stains less
readily than nonchorionated (immature) ones (Jennings 1974). Imma-
ture oocytes were counted at stereomicroscope magnifications up to
45x. Length of one forewing measured to the nearest 0.5 or 0.2 mm
in the preliminary and main experiments, respectively, was used as a
female body size index (Thomas et al. 1980, Results section of present
paper).

For chemical analyses of hemispherical scale honeydew, several hon-
eydew-laden yucca leaves were sprayed with distilled water, and 75

170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE l. Performance of spruce budworm females imbibing plain water and hon-
eydew-water in the preliminary experiment. SD preceded by +, range in parentheses.

Mean Departure
from plain-
Plain water Honeydew-water water group
Attribute [19 pairs] [9 pairs] (9)
Oviposition span, days! 8:8) == 1Si(6=12) 10:38 + 4.1 (5—17) 174
No. eggs and oocytes
Total 326 + 46 (243-395) 3839 + 75 (204-446) 4
Matured 200 + 46 (134-291) 245 + 71 (109-350) 22
Laid 197 + 46 (130-289) 235 + 76 (98-348) 19
Change in no. laid (L)
as function of fore-
wing length (W) 28.2 102.8 264¢
Unlaid immature 127 + 28 (88-176) 94 + 58 (0-184) = 202
Egg viability, % 75 + 20 (28-97) 81 + 15 (50-94) 8

1 Defined as time from first oviposition to death, usually one day longer than oviposition period.

aP < 0.01, 1-tailed Wilcoxon 2-sample test.

bP < 0.05, 1-tailed Student t-test.

¢P < 0.01, F-test. Values are slope coefficients from the regressions L = 28.2W — 126 (r = 0.40) and L = 102.8W —
930 (r = 0.75).

ml of runoff were collected. One aliquot was analyzed for sugars by
high-pressure liquid chromatography, one for total nitrogen by a high-
sensitivity Kjeldahl method, and one was oven-dried at 70°C to nomi-
nally constant weight for dry-weight conversions.

Honeydew concentration of imbibed honeydew-water was deter-
mined from weighings as follows. Honeydew-laden 25-50 cm? yucca
leaf segments were weighed (weight x), misted as for imbibing, re-
weighed (weight y), thoroughly washed, towel-dried, and again re-
weighed (weight z). These weightings were completed within 15 min,
and were to the nearest 0.1 mg. Honeydew concentration (c) was com-
olives! Ae == (Ge = 2) [ov = 6) SB Ge = 72)

In both experiments, attribute variances often differed significantly
between imbibing treatments (variance-ratio test). Treatment differ-
ences in such cases were analyzed nonparametrically by the Wilcoxon
two-sample test. Otherwise, treatment differences were analyzed para-
metrically by F- and Student t-tests. Because honeydew-water was
expected to have positive effects, most testing was one-tailed.

“Infertile’ here refers to females not producing viable eggs whether
mated or not. Standard deviation is abbreviated SD.

RESULTS

Preliminary experiment. Of 28 pupal and adult containers set up
for the honeydew-water imbibing treatment, and 29 set up for the
plain-water imbibing treatment, 9 and 19, respectively, produced viable
eggs and complete reproductive attribute records. Shortfalls were caused

VOLUME 43, NUMBER 3 171

No. eggs laid (L)

300

200

100

10.0 11.0 12.0 13.0
Forewing length (mm) (W)

Fic. 1. Role of spruce budworm female body size in determining number of eggs
laid in response to honeydew- and plain-water imbibing treatments. Each symbol rep-
resents one female. Line ® (closed squares) and line © (closed circles) depict honeydew-
water imbibers in the preliminary and main experiments, respectively; line © (open
squares) and line @ (open circles), plain-water imbibers.

by sexing errors, asynchronous eclosions, moth escapes, and unexplained
infertility.

Females given honeydew-water outperformed those given plain water
in four of seven reproductive attributes tabulated (Table 1). On average,
honeydew-water imbibers had a 1.5-day (17%) longer oviposition span,
matured 45 (22%) more oocytes, and contained 33 (26%) fewer im-
mature oocytes at death. Some tests were not robust enough to detect
biologically relevant attribute differences. The prime example concerns
number of eggs laid (Table 1, Fig. 1). In this and all attributes except
total oocyte number and egg viability, the effect of honeydew interacted
with body size. Thus an 11.0-mm female on honeydew-water laid 201

2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

eggs (11.0 x 102.8 — 930 = 200.8) while one on plain water laid 184
(11.0 x 28.2 — 126 = 184.2) (9% difference); but a 12.5-mm female
on honeydew-water laid 355 eggs (12.5 x 102.8 — 930 = 355.0) while
one on plain water laid 226 (12.5 x 28.2 — 126 = 226.5) (57% dif-
ference). Forewing length of females in the honeydew- and plain-water
treatments averaged 11.3 mm (SD = 0.6, range 10.5-12.0) and 11.2
mm (SD = 0.7, range 10.2-12.5), respectively. For attributes with vari-
ances shown in Table 1 (in SD, or square-root form), variances for the
honeydew-water treatment are usually greater numerically, two of them
greater statistically (P < 0.05, variance-ratio test), than for the plain-
water treatment.

In 2-h-old freeze-killed females, egg maturity averaged 7% (n = 8).
On average, these females contained 266 oocytes (SD = 40, range 228-
333), 60-73 fewer than the 326 and 339 totals shown in Table 1 for
imbibing females. The differences arose not because of inherently dif-
ferent oocyte numbers, but because freeze-killed females were smaller.
Their forewing lengths, averaging 10.5 mm (SD = 0.7, range 9.5-11.6
mm), were 0.7-0.8 mm less than for imbibing females. Regressions of
oocyte counts on forewing lengths among freeze-killed and imbibing
females did not differ significantly (P > 0.75, F-test of slope-coefficient
differences).

Dry-weight constituents of hemispherical scale honeydew (including
other water-soluble substances that also may have been on yucca leaf
surfaces) were fructose, 23.7%; glucose, 19.8%; sucrose, 12.8%; total
nitrogen, 0.6%; nonsugar and non-nitrogenous matter, 43.1%. These
amounts are similar to those previously reported for scale and aphid
honeydews (Auclair 1968).

Main experiment. Of 26 pupal and adult containers set up for the
honeydew-water imbibing treatment, and an equal number for the
plain-water imbibing treatment, 16 and 18, respectively, produced vi-
able eggs and complete reproductive attribute records. Shortfalls were
caused by the same factors as in the preliminary experiment.

Pairs given honeydew-water outperformed those given plain water
in 4 of 11 reproductive attributes tabulated (Table 2). On average,
honeydew-water imbibers had a 0.7-day (6%) longer female lifespan,
a 1.8-day (23%) longer oviposition period, and contained 41 (68%) fewer
immature oocytes at death. As in the preliminary experiment, some
tests were not robust enough to detect attribute differences, and number
of eggs laid is again the prime example (Table 2, Fig. 1). In this and
all attributes except total oocyte numbers and egg viability, the effect
of honeydew interacted with body size. Thus an 11.0-mm female on
honeydew-water laid 172 eggs (11.0 x 74.4 — 646 = 172.4) while one
on plain water laid 169 (11.0 x 25.0 — 106 = 169.0) (2% difference);

VOLUME 43, NUMBER 3 173

TABLE 2. Performance of spruce budworm pairs imbibing plain water and honeydew-
water in the main experiment. SD preceded by +, range in parentheses.

Mean or other value Departure
: from plain-
Plain water Honeydew-water water group
Attribute [18 pairs] [16 pairs] (%)
Preoviposition period,
days 22) 07 CL=3) 2.3 + 0.8 (1-4)
Lifespan, days
Female 10.7 + 2.0 (7-14) 11.4 + 3.4 (6-17) 64
Male 9.4 + 2.8 (5-14) 8.7 + 3.8 (4-17) sani
Oviposition period,
days Uc = P22 (GSM 9.5 + 3.6 (4-15) 23
No. eggs and oocytes
Total 238 + 60 (104-332) 283 + 80 (126—404) —2
Matured Nii 43 (104—288)) 234 78. 41-372) Zi
Laid 175 + 48 (102-288) 210 + 79 (104-370) 20
Change in no. laid (L)
as function of fore-
wing length (W) 25.0 74.4 198¢
Viable 156 + 36 (98-245) 181 + 78 (59-325) 16
Unlaid immature 60 + 41 (0-134) 19 + 29 (0-104) —684

2P < 0.01, 1-tailed Wilcoxon 2-sample test.
bP < 0.05, 1-tailed Student t-test.
ain < Oe Values are slope coefficients from the regressions L = 25.0W — 106 (r = 0.36) and L = 74.4W —
r = 0.81).
dP < (0.01, 1-tailed Student t-test.

but a 12.5-mm female on honeydew-water laid 284 eggs (12.5 x 74.4
— 646 = 284.0) while one on plain water laid 206 (12.5 x 25.0 — 106
= 206.5) (38% difference). For 9 of the 10 attributes with variances
shown in Table 2 (in SD, or square-root form), variances for the hon-
eydew-water treatment are greater numerically, four greater statisti-
cally (P < 0.05, variance-ratio test), than those for the plain-water
treatment.

In 2-h-old freeze-killed females, egg maturity averaged 10% (n =
13). On average, these females contained 259 oocytes (SD = 83, range
132-424), a number not inherently different from the 238 and 233
totals shown in Table 2 for imbibing females (P > 0.75, F-test of
differences in slope coefficients of oocyte number-forewing length
regressions). Forewing length of freeze-killed females averaged 10.7
mm (SD = 1.2, range 10.1-12.7).

Daily oviposition records in both imbibing treatments were similar
until the latter half of the oviposition period (Fig. 2). Beyond day 8 of
adulthood, live female-days/female averaged 3.5 in the honeydew-
water group and 1.7 in the plain-water group, and respective numbers
of eggs laid/female were 35 and 10.

Individual females were monitored for imbibing more than 150 times

174 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Mean no. eggs laid/female

200
_— ae ae
160 Aa
4
“a
a
7
honeydew-water group
— — — — plain-water group
Daily
|
|
ea leees
1 3 5 7 9 11 13 15

Day no. of adulthood

Fic. 2. Oviposition records for fertile spruce budworm females in the main experi-
ment. Sixteen females in the honeydew-water group, 18 in the plain-water group. Vertical
bars represent SD.

during the daily period of liquid availability. Imbibing started within
20 min after liquids were offered, usually much sooner, and individual
imbibing episodes lasted as long as 5 min. No monitored female imbibed
more than once daily, and none was seen excreting liquids anally. After
drying, honeydew presumably was not consumed until the next misting.

Females did not imbibe on the first or second days of adulthood (n
= 11 and 12, respectively). During days 3-12 of adulthood, fertile
females imbibed more often than infertile ones, the former using 78%
of their opportunities (62/79), the latter 50% (23/46). The difference
is significant (P < 0.01, 2 x 2 contingency table, G-test).

Among fertile females, imbibing frequency was unrelated to presence
or absence of honeydew. During days 3-12 of adulthood, sample fe-
males in the plain-water treatment imbibed essentially as often as those
in the honeydew-water treatment, the former using 77% of their op-
portunities (33/43), the latter 80% (29/36). Imbibing frequency was
affected by female age, however. Thus imbibing frequency rose from
60% on days 3-4 of adulthood to 97% on days 7-8, then fell to 54% on
days 11-12 (Table 3). Pooling by two-day intervals in Table 3 increased
the base and reliability of percentages.

VOLUME 43, NUMBER 3 175

TABLE 3. Imbibing frequencies by age of fertile spruce budworm females offered
honeydew-water or plain water in the main experiment.

Day no. of adulthood No. females observed Percent of females imbibing!
3-4 3) 60
5-6 15 80
7-8 30 97
9-10 16 69
1lj-12 13 o4

1 Underlying frequencies significantly dependent on female age (P < 0.01, 2 x 5 contingency table, G-test with
Williams correction).

For 10 fertile females monitored for imbibing on two successive days,
30% imbibed both days (3/10), and 70% imbibed one day only (7/10).
These females were in the 5th to 11th day of adulthood at the first
observation, averaging day 8.

Regressing weights of fertile females determined just before daily
imbibing episodes (M, range 13.8-86.0 mg) on forewing length (F,
range 9.8-13.0 mm) and day number of adulthood (D, range 1-12)
produced the equation M = 11.7F — 4.6D — 70.8 (R = 0.91, n = 87).
Based on this equation, live weights of honeydew-water imbibing fe-
males of average forewing length on days 1, 4, 7, and 10 of adulthood
were, respectively, 59.2, 45.4, 31.6, and 17.8 mg. Such weight decline
during adulthood reflects egg laying and depletion of stored reserves.
Forewing length of females in the honeydew- and plain-water treat-
ments averaged, respectively, 11.5 mm (SD = 0.9, range 9.38-13.0) and
11.25 mm (SD = 0.6, range 10.0-12.2).

Intake per imbibing episode did not differ significantly between
plain- and honeydew-water or fertile and infertile classifications in any
partition of data in Table 4 (P > 0.18, one-tailed Student t-tests); it
likewise varied independently of female size and age (r-range 0.03-
0.57, none approaching P = 0.05). Intake per episode averaged 4.5 mg

TABLE 4. Intake per investigated imbibing episode by individual spruce budworm
females offered liquids daily in the main experiment. SD preceded by +, range in
parentheses.

Fertility No.
status observed Mean age (days) Mean forewing length (mm) Amount imbibed (mg)
Plain-water imbibers
Fertile gl 8. 2246-12) 111 £08 405-11.7) 3.9\4 28 (0.9-81)
Infertile 9) 46 + 1.8 (8-7) 10.7 + 0.6 (9.8-11.2) 4.3 + 1.5 (2.7-6.5)
Honeydew-water imbibers

Fertile ri 8.0 = 1.0 (7-10) 11.7 = 0.8.(10.8-13.0) 5.2 + 2.5 (1.7-106.0)
Infertile i 5. + ks (4—7) 10.8 + 0.5 (9.8-11.2) 3.0 + 1.8 (1.0-6.5)

All imbibers
Mixed 28 6.7 + 2.3 (8-12) 11.1 + 0.7 (9.8-18.0) 4.5 + 2.4 (0.9-10.0)

176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

of liquid (Table 4), or 8, 10, 14, or 25% of the live female weights
computed above on days 1, 4, 7, and 10 of adulthood.

Honeydew concentration in honeydew-water on yucca leaf segments
averaged 6.5% by weight (SD = 2.1, range 3.5-9.4, n = 5).

During and immediately after adult eclosions observed under a bin-
ocular microscope (4 females, 2 males), the paired galeae were separate
for most of their length as reported for lepidopterans generally (Hep-
burn 1971). Within 15-30 min after pupal skin splitting, however,
galeae had engaged along their full lengths to form proboscises. During
much of the engagement process, which seemed assisted by the narrow
corridor formed by palpi, galeae twitched and intertwined. In two cases,
a droplet of clear fluid appeared near the base of galeae during the
engagement period.

DISCUSSION

Honeydew in imbibed water clearly had positive reproductive effects.
In both experiments, these effects were prolonged female lifespan,
oocyte maturation, and oviposition, the latter two causing redistribu-
tions within apparently fixed total oocyte complements (Tables 1 & 2).
Effects were strongest in the largest females (Fig. 1).

With only 7-10% of oocytes mature at female eclosion, a range
consistent with observations by Outram (1971), much oocyte maturation
necessarily takes place after eclosion. Thus an opportunity exists for
adult imbibing to influence oocyte maturation. However, females were
slow to begin imbibing under a regime that provided a once daily
opportunity. There was no evidence that they imbibed on the first or
second days of adulthood; even after four days, 40% of fertile females
still had not imbibed (Table 3). This explains why preoviposition period
was unaffected in this study, and perhaps also why positive reproductive
effects appeared late in adulthood. Preoviposition period was affected
in earlier work when the period lasted longer, and a stronger nutrient
solution (15% bee honey) was available constantly (Miller 1987). Ab-
stinence from imbibing in early adulthood did not appear to be mor-
phological in origin; proboscises were formed and presumably func-
tional within 0.5 h after eclosion.

In imbibing frequency and intake per imbibing episode, females
showed no clear preference for honeydew-water over plain water. This
result suggests that the moths merely respond to liquid.

Products (I) of age-specific imbibing frequencies (Table 3) and av-
erage intake per imbibing episode (4.5 mg, Table 4) plotted on female
age (A) resulted in a nearly straight line. This line is closely described
by the equation I = 3.45A — 7.7, and computed I-values differ by no
more than 7% from actual ones. Based on this equation, lifetime ex-
pected honeydew-water intake for fertile females of average lifespan

VOLUME 43, NUMBER 3 77

(11.4 days, Table 2) is 31.6 mg. Since honeydew concentration in hon-
eydew-water on yucca leaf segments averaged 6.5% by dry weight,
corresponding honeydew intake is 2.05 mg (31.6 x 0.065 = 2.05).
Attribute departures among honeydew-water imbibing females may
therefore be ascribed to average consumption of ca. 2 mg of dry hon-
eydew per individual, or ca. 3.5% of initial live weight of females of
average forewing length in the main experiment (2.05/59.2 = 0.085).

The greater attribute variability (variances) noted among honeydew-
water imbibers in both experiments probably results from the inter-
action between honeydew effect and female body size. However, in an
experiment in which bee-honey concentration was a constant 15%, and
honey-water was constantly available, no tendency to heterogeneous
variability appeared (Miller 1987).

How much the positive reproductive effects of imbibed honeydew
might influence the dynamics of natural populations would seem to
hinge on how long females survive in nature, as well as on how readily
available honeydew is to them.

ACKNOWLEDGMENTS

Ingman Laberatories Inc., Minneapolis, Minn., performed the chromatography, and
University of Minnesota Soil Science Analytical Services the Kjeldahl analysis. I thank
Jana Campbell for help in collecting pupae, J. M. Muggli for the honeydew source, A.
B. Hamon for expert identification of hemispherical scale, R. L. Jones for laboratory
space, D. A. Andow for statistical advice, G. T. Harvey, R. W. Hansen, D. W. Ragsdale
and three anonymous readers for useful manuscript reviews, and R. C. Lederhouse for
serving as special Journal editor for this manuscript.

LITERATURE CITED

AUCLAIR, J. L. 1963. Aphid feeding and nutrition. Ann. Rev. Entomol. 8:439-—490.

HEPBURN, H. R. 1971. Proboscis extension and recoil in Lepidoptera. J. Insect Physiol.
17:637-656.

JENNINGS, D. T. 1974. Potential fecundity of Rhyacionia neomexicana (Dyar) (Ole-
threutidae) related to pupal size. J. Lepid. Soc. 28:131-136.

JENNINGS, D. T. & M. W. HousEWeEarT. 1978. Sexing spruce budworm pupae. U.S.
Dept. Agr. For. Serv. Rep. Northeast. For. Exp. Sta. NE-255, 2 pp.

MILLER, W. E. 1987. Spruce budworm (Lepidoptera: Tortricidae): Role of adult im-
bibing in reproduction. Environ. Entomol. 16:1291-1295.

OuTRAM, I. 1971. Morphology and histology of the reproductive system of the female
spruce budworm, Choristoneura fumiferana (Lepidoptera: Tortricidae). Can. Ento-
mol. 103:32—44.

POWELL, J. A. 1965. Biological and taxonomic studies of tortricine moths, with reference
to the species in California. Univ. Calif. Publ. Entomol. 32, 317 pp.

THomas, A. W., S. A. BORLAND & D. O. GREENBANK. 1980. Field fecundity of the
spruce budworm (Lepidoptera: Tortricidae) as determined from regression relation-
ships between egg complement, fore wing length, and body weight. Can. J. Zool. 58:
1608-1611.

WuitE, J. W. 1975. Composition of honey, pp. 157-206. In E. Crane (ed.), Honey: A
comprehensive survey. Heinemann, London. 608 pp.

Received for publication 14 November 1987; accepted 8 May 1989.

Journal of the Lepidopterists’ Society
43(3), 1989, 178-183

WORLD NUMBERS OF BUTTERFLIES

OAKLEY SHIELDS
6506 Jerseydale Road, Mariposa, California 95338

ABSTRACT. World butterflies number about 17,280 species representing described
taxa that have not been synonymized, and are currently grouped into 1855 genera, 35
subfamilies, and 7 families. Butterflies constitute only 9-12% of all lepidopteran species.

Additional key words: faunal realms, genera, subfamilies, families, conservation.

Published estimates of the total number of butterfly species in the
world range from 7700 (Kirby 1872) to 20,000 (Fox & Fox 1964, Vane-
Wright 1978, Landing 1984), although most authors do not cite specific
sources used in forming their estimates. An exception is Robbins (1982),
whose world total of 15,900-18,225 was compiled by faunal realms,
although he largely estimated numbers for the Neotropics and Orient
and did not adjust for the percentage of species that extend into two
or more realms. Affinities between the west Palearctic and Ethiopian
butterfly faunas are indeed minimal (De Jong 1976), but a modest
amount of exchange (perhaps 5-10%) is to be expected across the Nearc-
tic-Neotropical, east Palearctic-Oriental, and Oriental-Australian fron-
tiers (cf. Schmidt 1954, Rapoport 1971). The total number of butterfly
species, coupled with the land area they occupy, measures butterfly
richness and density on a global scale, of interest to ecologists and
conservationists. Here I present a current estimate of butterfly species
numbers as part of the continuing effort to determine just how many
species of living organisms there are on earth (cf. Jones 1951, May 1988,
Wilson 1988).

METHODS

Species totals for most butterfly subfamilies (Table 1) were obtained
from the recently published catalogs of Bridges (1988a, 1988b, 1988c,
1988d), which counts species that extend into two or more realms only
once. Some of the Nymphalidae subfamilies, however, have not been
recently cataloged. The number of Nymphalinae species was estimated
by multiplying its number of genera (181) by 14 (the average number
of species per genus in Danainae s. str., its close relative) to yield ca.
2500 species. This figure compares well with the roughly 2700 species
I estimate for Nymphalinae using faunal lists. Numbers of species for
Satyrinae and Morphinae are also estimates (see footnotes 6 and 7, Table
1). For the Danainae and Brassolinae, species numbers are based on
figures provided by researchers currently revising these groups (see
footnotes 4 and 8, Table 1).

VOLUME 438, NUMBER 3 179

RESULTS AND DISCUSSION

The grand total of described butterfly species is about 17,280 (see
Table 1). This total is higher than the 13,000 estimate of Owen (1971),
the 14,750 estimate of Scott (1986), the 15,600 low estimate (when
Hesperiidae are included) of Ehrlich and Raven (1964), and the 15,900
low estimate of Robbins (1982) and is lower than the world maximum
estimates of 18,225 (Robbins 1982), 18,600 (Ehrlich & Raven 1964),
and 20,000 (Fox & Fox 1964, Vane-Wright 1978, Landing 1984). The
total is only 945-1320 species less than the maximum estimates of
Ehrlich and Raven (1964) and Robbins (1982). Bridges (1988d) lists
1855 world genera, though, of course, generic limits are largely sub-
jective. For example, Bridges’ (1988d) figure of 1326 genera (excluding
skippers) sharply contrasts with the estimates of 730-930 and 953 genera
given by Ehrlich and Raven (1964) and Scott (1986), respectively. The
number of described species of butterflies in the world constitutes 9-
12% of all described Lepidoptera species, whose total numbers are
estimated to be 150,000-200,000 (Kristensen 1984).

Subfamilies with the greatest numbers of species are (in descending
order) Nymphalinae, Satyrinae, Theclinae, and Hesperiinae (greater
than 2000), followed by Pyrginae, Polyommatinae, and Riodininae
(greater than 1000). Baroniinae, Curetinae, Styginae, Pseudopontiinae,
Libytheinae, and Calinaginae are morphologically archaic subfamilies
each containing only one or two genera and fewer than two dozen
species. Three families (Hesperiidae, Lycaenidae, and Nymphalidae)
comprise 82% of all butterfly species and are the only families to use
both dicots and monocots extensively as larval hostplants (cf. Ehrlich
& Raven 1964). About 30% of all butterfly species feed only on mono-
cots, especially the Trapezitinae, Hesperiinae, Megathyminae, Satyri-
nae, Morphinae (in part), and Brassolinae. The world species richness
of butterflies, 17,280, when divided by 128,811,340 km?, the total land
area of the earth excluding Antarctica and inland waters, yields an
average density of 0.000134 species per km?. Roughly two-thirds of the
species occur in the tropics.

The numbers of butterfly species presented here represent described
taxa that have not been synonymized. Only in the best-known family,
Papilionidae, is it possible to estimate closely the true number of species.
Subfamilies like Theclinae, Polyommatinae, Riodininae, Nymphalinae,
Calinaginae, and Satyrinae are so poorly known taxonomically that
their counts probably inflate their actual species totals by including
many unsynonymized names. Most other subfamilies fall somewhere
between these two extremes. Cladistic analysis should aid in identifying
monophyletic subfamilies in Nymphalidae but has not yet been per-

180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TABLE 1. Numbers of butterfly species by family and subfamily.
Family and subfamily No. species
Hesperiidae 3592
Pyrginae 1198 Bridges 1988a
Coeliadinae 80 Bridges 1988a
_Pyrrhopyginae 155 Bridges 1988a
Trapezitinae 67 Bridges 1988a
Hesperiinae 2048 Bridges 1988a
Megathyminae 49 Bridges 1988a
Papilionidae 566
Papilioninae 511 Bridges 1988b
Parnassiinae 54 Bridges 1988b
Baroniinae il Bridges 1988b
Lycaenidae! 4089
Lipteninae Spall Bridges 1988c
Poritiinae 52 Bridges 1988c
Liphyrinae 20 Bridges 1988c
Miletinae Ta Bridges 1988c
Curetinae 22 Bridges 1988c
Theclinae 2128 Bridges 1988c
Lycaeninae 97 Bridges 1988c
Polyommatinae 1132 Bridges 1988c
Riodinidae 1366
Hamearinae 97 Bridges 1988c, Robbins 1988a
Euselasiinae 154 Bridges 1988c, Robbins 1988a
Riodininae 1114 Bridges 1988c, Robbins 1988a
Styginae 1 Robbins 1988a, 1988b
Pieridae 1215
Pseudopontiinae 1 Bridges 1988b
Dismorphiinae 95 Bridges 1988b
Pierinae 905 Bridges 1988b
Coliadinae 214 Bridges 1988b
Libytheidae 12
Libytheinae 12 Shields 1985
Nymphalidae 6440
Nymphalinae? 2500 Bridges 1988d, see text
Argynninae® 155 Warren 1944, 1955; dos Passos & Grey 1945;
Grey in litt; Common & Waterhouse 1972;
Brown 1981
Acraeinae 240 Pierre 1987
Calinaginae 16 Oberthur 1919, 1922; Wu 1938
Danainae‘* 462 Ackery & Vane-Wright 1984; Drummond &
Brown 1987
Apaturinae® 431 Stichel 1938, 1939; Le Moult 1950; Comstock
1961; van Someren 1975
Satyrinae® 2400 Gaede 1931; L. D. Miller, pers. comm.
Morphinae’ 55 D’Abrera 1984; Parsons 1984
Brassolinae® 81 Stichel 1932
World total 17,280 present study

' Eliot (in litt.) estimates 44 species of Poritiinae and 15 species of Curetinae based upon his unpublished research,
and he notes that the Neotropical Theclinae species listed by Bridges have many synonyms but also that hundreds of
discovered but undescribed species of Theclinae exist.

2 Includes the tribes Ageroniini (~Hamadryini), Biblidini (=Didonini, Ergolini, Eurytelini), Coeini (=Aganisthini,

VOLUME 43, NUMBER 3 181

formed; preliminary analysis indicates that some conventional nym-
phalid subfamilies are polyphyletic (DeVries et al. 1985).

The world number of species or “species richness’’ obscures much of
the interesting ecological diversity of butterflies. Many polytypic species,
e.g. in Heliconiini and Ithomiinae, have populations that differ mark-
edly in behavior, food plant relationships, and color patterns.

It is instructive to compare butterflies with birds, taxonomically the
best-known invertebrate and vertebrate groups. At the level of species,
“The taxonomy of no other group of animals is as mature as that of
birds” (Mayr 1982:292). Kirby (1872) estimated that the number of
species of butterflies and birds was about the same. By the early twen-
tieth century, Sharpe recognized 18,937 bird species (including fossil
species and all subspecies as full species), a figure that dropped to
10,000-16,000 by the early 1980’s (Mayr 1988). By 1935 careful zoo-
geographic and systematic research had reduced the estimated number
of birds to 8500 (Mayr 1946). Today’s best estimate of the number of
bird species is ca. 7000 + 200 (Mayr 1988), which is only 40% of the
number of butterfly species reported in the present study. Since 1935
only about 140 valid new bird species have been described, with the
reduction in the number of species names (by more than 60% over the
past 80 years) primarily coming about by revisional downgrading of
geographical isolates of polytypic species from the rank of species to
subspecies (Mayr 1988). This same process is now occurring in butter-
flies, but so far it has progressed to a far lesser extent than it has in
birds. Unlike birds, however, many new species of butterflies are still
being discovered and described each year, particularly from the tropics.

-—

Coloburini, Gynaeciini), Cyrestini (=Marpesiini), Epicaliini (=Callicorini, Catagrammini, Catonephelini, Dynamini,
Epiphilini, Eunicini), Limenitidini (=Abrotini, Adelphini, Bebeariini, Chalingini, Euthaliini, Neptini, Neurosigmatini,
Parthenini, Pseudacraeini), Nymphalini (=Araschniini, Cynthini, Diademini, Doleschallini, Euphydryini, Hypolimnini,
Junoniini, Kallimini, Melitaeini, Phyciodini, Vanessini), Pseudergolini.

8 Includes Heliconiinae, Cethosiini.

4 Includes Ithomiinae (=Ithomiini, Tellervini). Drummond and Brown (1987) cite 305 species of Ithomiinae and include
only one species of Tellervo, although Ackery (1987) claims there are 6. Other species estimates for Ithomiinae are
extremely variable: ca. 300 (Drummond 1986), 318 (Mielke & Brown 1979), 349 (Fox 1953), ca. 400 (D’Abrera 1984),
and 518 (Bryk 1937). Haensch (1909) in Seitz’s The Macrolepidoptera of the World listed 883 named forms, most of
which he treated as species. The best figure is 305 species, based largely on Brown’s ongoing study of ithomiine phylogeny
(fide Drummond). The Ithomiinae sarane show the gradual yeduetion in nee of species as a group becomes
better known (dropping by nearly two-thirds over the past 80 years), largely as a result of a growing recognition of the
ay widespread polytypic species in this group. There are 157 species of Danainae (s. an (Ackery & Vane-Wright
1984).

5 Includes Charaxinae.

6 Miller (1968) estimated between 2500-3000 Satyrinae species (including Brassolinae), but he now feels this is too
high. His new estimate (pers. comm.) is about half again as many as in Gaede (1931). As Gaede listed 1605 species, the
new estimate is ca. 2400 species.

7 Includes Amathusiinae. Parsons (1984) estimates there are ca. 100 species of Amathusiinae. Morphinae now includes
the Neotropical genera Morpho, Antirrhea, and Caerois (DeVries et al. 1985). D’Abrera (1984) lists 31 species of Morpho,
21 a and 3 Caerois. Smart (1977) estimates about 80 species of Morpho, probably too many (cf. D’Abrera
1984).

’ Smart (1977) lists 75 species. Preliminarily there are 86 species according to Casagrande (in litt.) who is currently
revising the subfamily.

182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ACKNOWLEDGMENTS

I thank Mirna M. Casagrande, Boyce A. Drummond III, John Eliot, Dale W. Jenkins,
Lee D. Miller, William E. Miller, Bruce A. Wilcox, and two anonymous reviewers for
their comments and suggestions; Keith S. Brown Jr., Boyce A. Drummond III, Glenn A.
Gorelick, L. Paul Grey, James R. Mori, and Robert K. Robbins for some references; and
Charles A. Bridges for loaning his catalogs.

LITERATURE CITED

ACKERY, P. R. 1987. The danaid genus Tellervo—A cladistic approach. Zool. J. Linn.
Soc. 89:203-274.

ACKERY, P. R. & R. I. VANE-WRIGHT. 1984. Milkweed butterflies: Their cladistics and
biology. British Museum (Natural History), London. 425 pp.

BRIDGES, C. A. 1988a. Catalogue of Hesperiidae. Privately published, Urbana, Illinois.

1988b. Catalogue of Papilionidae and Pieridae. Privately published, Urbana,

Illinois.

1988c. Catalogue of Lycaenidae and Riodinidae. Private published, Urbana,

Illinois.

1988d. Catalogue of family-group and species-group names (Rhopalocera).
Privately published, Urbana, Illinois.

Brown, K. S., JR. 1981. The biology of Heliconius and related genera. Ann. Rev.
Entomol. 26:427-456.

BrYK, F. 1937. Danaidae II: Subfam.: Ituninae, Tellervinae, Ithomiinae. Lepidopter-
orum Catalogus 80.

Common, I. F. B. & D. F. WATERHOUSE. 1972. Butterflies of Australia. Angus &
Robertson, Sydney. 498 pp.

Comstock, W. P. 1961. Butterflies of the American tropics: The genus Anaea. The
American Museum of Natural History, New York. 214 pp., 30 pls.

D’ABRERA, B. 1984. Butterflies of the Neotropical region. Part II. Danaidae, Ithomiidae,
Heliconidae and Morphidae. Hill House, Ferny Creek, Victoria. 384 pp.

DE JONG, R. 1976. Affinities between the west Palearctic and Ethiopian butterfly faunas.
Tijdschr. Entomol. 119:165-215.

DEVRIES, P. J., I. J. KITCHING & R. I. VANE-WRIGHT. 1985. The systematic position of
Antirrhea and Caerois, with comments on the classification of the Nymphalidae.
Syst. Entomol. 10:11-82.

Dos Passos, C. F. & L. P. Grey. 1945. A genitalic survey of Argynninae. Amer. Mus.
Novit. 1296, 29 pp.

DRUMMOND, B. A., III. 1986. Coevolution of ithomiine butterflies and solanaceous plants,
pp. 307-327. In D’Arcy, W. G. (ed.), Solanaceae: Biology and systematics. Columbia
Univ. Press, New York.

DRUMMOND, B. A., III, & K. S. BROWN JR. 1987. Ithomiinae: Summary of known larval
food plants. Ann. Missouri Bot. Gard. 74:341-358.

EHRLICH, P. R. & P. H. RAVEN. 1964. Butterflies and plants: A study in coevolution.
Evolution 18:586—-608.

Fox, R. M. 1953. The taxonomic value of the male genitalia in the Ithomiidae. Entomol.
News 64:141-143.

Fox, R. M. & J. W. Fox. 1964. Introduction to comparative entomology. Reinhold,
New York. 450 pp.

GAEDE, M. 1931. Satyridae. Lepidopterorum Catalogus 43, 46, 48.

HAENSCH, R. 1909. Familie: Danaidae, pp. 113-171, pls. 31-41. In Seitz, A. (eds.), Die
Gross-Schmetterlinge der Erde, Vol. 5. Kernen, Stuttgart.

JONES, G. N. 1951. On the number of species of plants. The Scientific Monthly 72:289-
294,

KirBy, W. F. 1872. On the geographical distribution of the diurnal Lepidoptera as
compared with that of the birds. J. Linn. Soc. Lond., Zool. 11:431-489.

KRISTENSEN, N. P. 1984. Studies on the morphology and systematics of primitive Lep-
idoptera. Steenstrupia 10:141-191.

VOLUME 43, NUMBER 3 183

LANDING, B. H. 1984. Factors in the distribution of butterfly color and behavior pat-
terns—Selected aspects. Privately published, Los Angeles. 200 pp.

Le Mouw.t, E. 1950. Revision de la classification des Apaturinae de l’ancien monde.
Miscellanea Entomologica, Suppl. 1, 68 pp., 19 pls.

May, R. M. 1988. How many species are there on Earth? Science 241:1441-1449.

Mayr, E. 1946. The number of species of birds. The Auk 63:64-69.

1982. The growth of biological thought. The Belknap Press of Harvard Univ.

Press, Cambridge, Massachusetts. 974 pp.

1988. Toward a new philosophy of biology. The Belknap Press of Harvard Univ.
Press, Cambridge, Massachusetts. 564 pp.

MIELKE, O. H. H. & K. S. BROWN Jr. 1979. Suplemento ao “Catalogo dos Ithomiidae
Americanos’ de Romualdo Ferreira D’Almeida. Univ. Federal do Parana, Curitiba.
216 pp.

MILLER, L. D. 1968. The higher classification, phylogeny and zoogeography of the
Satyridae. Mem. Am. Entomol. Soc. 24:1-174.

OBERTHUR, C. 1919. Description de deux espéces ou formes nouvelles de Lépidoptéres
provenant du Yunnan. Bull. Soc. Entomol. France 24:173-175.

1922. Note rectificative. Bull. Soc. Entomol. France 27:250-251.

OWEN, D. F. 1971. Tropical butterflies. Clarendon Press, Oxford. 214 pp.

PARSONS, M. 1984. Life histories of Taenaris (Nymphalidae) from Papua New Guinea.
J. Lepid. Soc. 38:69-84.

PIERRE, J. 1987. Systématique cladistique chez les Acraea. Ann. Soc. Entomol. France
(n.s.) 23:11-27.

RAPOPORT, E. H. 1971. The Nearctic-Neotropical frontiers. Proc. 13th Intern. Congr.
Entomol., Moscow 1968, 1:190-191.

ROBBINS, R. K. 1982. How many butterfly species? News Lepid. Soc. (1982) 3:40-41.

1988a. Comparative morphology of the butterfly foreleg coxa and trochanter

and its systematic implications. Proc. Entomol. Soc. Wash. 90:133-154.

1988b. Male foretarsal variation in Lycaenidae and Riodinidae, and the sys-
tematic placement of Styx infernalis. Proc. Entomol. Soc. Wash. 90:356-368.

SCHMIDT, K. P. 1954. Faunal realms, regions, and provinces. Quart. Rev. Biol. 29:322-
3o3l.

ScoTT, J. A. 1986. The butterflies of North America. Stanford Univ. Press, Stanford.
583 pp.

SHIELDS, O. 1985. Zoogeography of the Libytheidae (snouts or beaks). Tokurana 9:1-58.

SMART, P. 1977. The illustrated encyclopedia of the butterfly world. Hamlyn, London.
275 pp.

STICHEL, H. 1932. Brassolidae. Lepidopterorum Catalogus 51.

1938. Nymphalinae I. subfam.: Dioninae, Anetinae, Apaturinae. Lepidoptero-

rum Catalogus 86.

1939. Charaxidinae. Lepidopterorum Catalogus 91, 93.

VANE-WRIGHT, R. I. 1978. Ecological and behavioural origins of diversity in butterflies.
Symp. R. Entomol. Soc. Lond. 9:56-70.

VAN SOMEREN, V.G.L. 1975. Revisional notes on African Charaxes, Palla and Euxanthe,
part X. Bull. Brit. Mus. (Nat. Hist.), Entomol. 32:65-136.

WARREN, B.C. S. 1944. Review of the classification of the Argynnidi: With a systematic
revision of the genus Boloria. Trans. R. Entomol. Soc. Lond. 94:1-101.

1955. A review of the classification of the subfamily Argynninae. Part 2. Def-
inition of the Asiatic genera. Trans. R. Entomol. Soc. Lond. 107:381-391.

WILSON, E. O. 1988. The current state of biological diversity, pp. 3-18. In Wilson, E.
O. & F. M. Peter (eds.), Biodiversity. National Academy Press, Washington, D.C.

Wu, C.F. 1938. Catalogus Insectorum Sinensium, Vol. 4. Fan Mem. Inst. Biol., Peiping.
1007 pp.

Received for publication 20 September 1988; accepted 24 May 1989.

Journal of the Lepidopterists’ Society
43(3), 1989, 184-209

THE SPHINGIDAE (LEPIDOPTERA) OF
BAJA CALIFORNIA, MEXICO

JOHN W. BRowNn!

Department of Entomological Sciences, University of California,
Berkeley, California 94720

AND

JULIAN P. DONAHUE

Natural History Museum of Los Angeles County,
900 Exposition Boulevard, Los Angeles, CA 90007-4057

ABSTRACT. Baja California is a rugged, mostly xeric peninsula situated along the
northwestern coast of Mexico. With the exception of butterflies, the lepidopterous fauna
of this region is poorly studied. Twenty-six species of Sphingidae are known from the
peninsula, including one endemic species, Sphinx xantus Cary, and three endemic sub-
species, Manduca rustica cortesi (Cary), Pachysphinx occidentalis peninsularis Cary
[revised identification], and Callionima falcifera guaycura (Cary) [revised identification].
For each of the 26 species, information is presented on peninsular distribution, flight
period, and possible larval host plants.

Additional key words: peninsular effect, host plants, distributions.

The peninsula of Baja California (or Lower California; here termed
simply “Baja California’’) is situated along the northwestern coast of
Mexico, extending southeasterly approximately 1300 km from the in-
ternational border to its tip at Cabo San Lucas. It is bordered by the
state of California (Alta California) on the north, the Pacific Ocean on
the west, and the Sea of Cortés (Gulf of California) on the east (Fig.
1). The mainland Mexican states of Sonora and Sinaloa lie to the east
of the gulf. Much of the peninsula is a low lying desert. However, in
the north are two major mountain ranges, the Sierra Juarez and the
Sierra San Pedro Martir, which represent extensions of the Peninsular
Ranges of southern California. Two significant ranges occur in the
southern third of the peninsula: the Sierra de la Giganta, running par-
allel to the eastern coast, and the Sierra de la Laguna in the center of
the southern tip. Politically the peninsula is divided at the 28th parallel
into a northern state, Estado de Baja California (here termed Baja
California Norte to avoid confusion), and a southern state, Estado de
Baja California Sur (Baja California Sur).

Although comparatively depauperate, the lepidopterous fauna of
Baja California is nonetheless unique and diverse, primarily as a con-
sequence of the nearly 10° range in latitude the peninsula embraces,
and its relative isolation from mainland México. Except for butterflies,

' Research Associate, Entomology Department, San Diego Natural History Museum.

VOLUME 43, NUMBER 3 185

CALIFORNIA
Sierra ARIZONA

Juarez

Sierra
San
Pedro
Martir

SONORA

Baja
California
Norte

114°

Fic. 1. The peninsula of Baja California and adjacent areas.

however, few lepidopterous families have been studied intensively.
Prior to our study, a total of 20 species of Sphingidae had been credited
to the Baja California fauna. Mooser (1940) treated 145 species of
Sphingidae from the Republic of México, and specifically cited records
of 5 species from Baja California, mostly in the collection of Carlos C.
Hoffmann. Hoffmann (1942) cited 8 species, primarily Californian ele-
ments, as occurring specifically in Baja California, but provided no
locality data. Cary (1963) presented records of 244 specimens repre-

186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

senting 14 species taken in the Cape Region at the southern tip of the
peninsula. Only two of these had been listed previously by Hoffmann
(1942), the majority being widely ranging neotropical species cited by
Hoffmann as occurring throughout México without specific reference
to Baja California. In addition, Cary (1963) described four endemic
taxa. Her paper laid the groundwork for biogeographic comparisons
between the mainland and peninsular sphingid faunas. Schreiber (1978)
cited Baja California as within the range of 7 species in his work on
dispersal centers of neotropical Sphingidae; all of these had been listed
previously either by Hoffmann (1942) or by Cary (1963). Recent col-
lecting efforts have raised the number of sphingids known to occur in
Baja California to 26.

SPHINGID DISTRIBUTIONS

Although large and highly vagile insects, Schreiber (1978) has shown
that sphingids exhibit biogeographic patterns comparable to animals of
much lesser mobility. The species present in Baja California conform
reasonably well to the biotic provinces or phytogeographic regions pre-
sented by Shreve (1951) and Wiggins (1980). Four general patterns of
distribution are exhibited. 1) The Californian Province in the north-
western portion of the peninsula has a distinctive fauna composed
primarily of temperate species including several montane and oak
woodland associated elements (Figs. 8 and 15). 2) The Cape Region at
the southern end of the peninsula supports a limited fauna of tropical
elements similar to that of adjacent mainland Mexico, but with many
fewer species (Figs. 17 and 18). 3) A number of common, widespread
species range the length of the peninsula (Figs. 4, 5, and 20). 4) Two
species exhibit disjunct patterns shown by several butterfly species; they
are restricted primarily to the Californian Province but are represented
by isolated populations in the Cape Region (Figs. 9 and 10). Apparent
conformity to the phytogeographic regions probably has been enhanced
by lack of intensive collecting and by biases toward specific areas (e.g.,
the Cape Region has received far more attention than any other area).

Simpson (1964) first recognized that North American peninsulas had
fewer species present at their distal tips than at their mainland bases.
He suggested that this pattern was neither coincidental nor transient,
but was the result of an equilibrium between species colonization and
extinction. Subsequent studies on various North American vertebrates
(Cook 1969, Kiester 1971, Taylor & Regal 1978) corroborated Simpson’s
observation, and this pattern became known as the “peninsular effect.’’
Simply defined, the peninsular effect states that species density or rich-
ness decreases as a function of distance from the mainland base of a
peninsula.

VOLUME 43, NUMBER 3 187

number of
species
20

base tip

distance from base
(degrees of latitude)

Fic. 2. Species density histogram for Baja California Sphingidae (base = north end
of peninsula; tip = southern extremity).

Seib (1980) first questioned the general applicability of the peninsular
effect when he demonstrated that the reptiles of Baja California did
not conform to this pattern. Recently it has been shown that the but-
terflies of Baja California also do not conform to this biogeographic
principle (Brown 1987). Likewise, sphingids illustrate a pattern of species
richness contrary to that predicted by the peninsular effect (Fig. 2).
High species density occurs not only at the northern base but at the
distal tip as well. Species density appears to be a function of floral
diversity (community complexity) and proximity to mainland species
pools rather than distance from the mainland base of the peninsula.

SPECIES ACCOUNTS

Unless indicated otherwise, nomenclature follows that of Hodges
(1971). All capture records are listed for species represented by fewer
than 25 specimens. Locality data were transcribed from specimen labels,
thus there is a mixture of metric and English systems for distance and
elevation. The distribution maps include localities for all specimens
personally examined by us as well as all additional records cited (i.e.,

188 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

personal communication, etc.). Disposition of examined specimens is
indicated by abbreviations (e.g., AMNH) listed in the Acknowledg-
ments. Original descriptions are cited in full in the text; additional
references are in abbreviated style but are cited in full in the Literature
Cited section.

1. Agrius cingulatus (Fabricius) (Figs. 3, 21)
Sphinx cingulata Fabricius, 1775, Systema Entomologiae, 545.
Agrius cingulatus; Hodges, 1971:22.

This widespread Neotropical species occurs throughout much of Mexico, ranging into
the southern parts of the United States. Strays are known from as far north as Nova Scotia
(Hodges 1971). Although most Baja California records are from the Cape Region, A.
cingulatus probably occurs over most of the peninsula. It has been taken in coastal as
well as montane areas, and there is a single record from the northern desert. A. cingulatus
is probably multiple-brooded; adults fly from July through April in the southern portion
of the peninsula. Possible host plants in Baja California include Datura (Solanaceae) and
several species of Ipomoea (Convolvulaceae).

Specimens examined: BAJA CALIFORNIA NORTE: Rancho Santa Inez, 540 m, 1M,
27-VII-82, W. Clark and P. Blom (CI); SW end Isla San Esteban, 1M, 28-VII-86, D.
Faulkner (SDNHM). BAJA CALIFORNIA SUR: Ramal de Naranjas, 6 mi W Highway
near Santa Anita, 1M, 11-X-83, F. Andrews and D. Faulkner (SDNHM); 36.3 mi SE
Todos Santos, 1M, 10-X-83, F. Andrews and D. Faulkner (SDNHM); El Salto, 8 mi E
Todos Santos, 1F, 9-X-83, F. Andrews and D. Faulkner (SDNHM); Ricardo’s RV Park,
2 mi N Cabo San Lucas, 1M, 6-I-84, R. Wells (RW); 2 mi E El Triunfo, 1F, 12-VIII-66,
J. Chemsak, J. Doyen and J. Powell (UCB); Sierra de la Laguna, Rancho La Burrera, 1.9
rd mi S and 12.6 mi E Todos Santos, 1600’, 1F, 15-IX-85, J. and K. Donahue (LACM).

Additional records: BAJA CALIFORNIA SUR: San Bartolo microwave tower, 2000’,
24-VII-81, R. Holland (pers. comm.); Playa San Cristobal, 15 mi N Cabo San Lucas, 100’,
18-IV-84, J. Brown (sight record).

2. Manduca sexta sexta (Linnaeus) (Figs. 4, 22)
Sphinx sexta Linnaeus, 1763, Centuria Insectorum Rariorum, 27.
Phlegethontius sextus sextus; Cary, 1963:193.
Manduca sexta; Hodges, 1971:29.

M. sexta is abundant throughout the peninsula, ranging north well into California; it
is absent only at the highest elevations. Cary (1963) also records it from Sonora and
Sinaloa. In Baja California adults have been taken from March through December. It is
most common in the late summer and fall. Larvae feed on a wide variety of Solanaceae
including potato, tomato, and tobacco.

3. Manduca quinquemaculata (Haworth) (Figs. 5, 23)
Sphinx quinquemaculata Haworth, 1803, Lepidoptera Britannica 1:59.
Phlegethontius quinquemaculatus; Cary, 1963:194.
Manduca quinquemaculata; Hodges, 1971:31.

This species is very similar to M. sexta, from which it can be distinguished by the
presence of an almost continuous, straight, black subterminal line on the forewing. The
two are broadly sympatric throughout Baja California, however, M. quinquemaculata is
considerably less abundant. It is equally common in both the northern and southern halves
of the peninsula. Captures extend from March through October. Larval hosts are the
same as those given above for M. sexta.

4. Manduca rustica cortesi (Cary) (Figs. 6, 24)
Phlegethontius rusticus cortesi Cary, 1963, Ann. Carnegie Mus. 36:194.
Manduca rustica cortesi; Schreiber, 1978:41, 62.
Although capture records do not illustrate an extensive peninsular distribution, M.
rustica probably occurs throughout Baja California. It is frequent to the north in adjacent

VOLUME 43, NUMBER 3 189

southern California where it is probably resident. The subspecies cortesi was described
from specimens taken in the Cape Region, and the name is applicable to southern
California material as well. M. rustica cortesi averages smaller than the nominate sub-
species; its white over-scaling and black and white maculation are in strong contrast to
the typical brown and white markings of M. rustica rustica (Fabricius). Adults fly in the
summer and fall, with captures ranging from July to November. Hosts available in Baja
California include Bignonia (Bignoniaceae) and Verbena (Verbeneaceae), and probably
other members of these two families, such as the widespread Chilopsis linearis (Cav.)
Sweet and the more southern Tecoma stans (L.) Juss. (both Bignoniaceae).

5. Sphinx xantus Cary (Figs. 7, 25)
Sphinx xantus Cary, 1963, Ann. Carnegie Mus. 36:196; Schreiber 1978:45.

This endemic species has been found over a wide range of elevations in the Cape
Region, and ranges north along the eastern side of the peninsula to Bahia de las Animas
in Baja California Norte. Captures extend from August to November. S. xantus is the
peninsular counterpart of the mainland S. istar (Rothschild and Jordan). Cary (1963)
presents several morphological characters distinguishing the two: xantus is smaller in
forewing length and more somber brown than the lighter and variegated S. istar; the
male genitalia are also distinct. The differences between S. xantus and S. istar are not
as great as those between many species in the genus. However, until biological data to
the contrary become available, we will treat them as separate species. The larval host
plant is unknown.

Specimens examined: BAJA CALIFORNIA NORTE: Bahia de las Animas, sea level,
1M, 1F, 5-IX-85, J. and K. Donahue (#96,431, LACM). BAJA CALIFORNIA SUR: 7 mi
S San Pedro, 3M, 10-VIII-66, Chemsak, Doyen and Powell (UCB); Highway 19, 14.5 rd
mi NW Cabo San Lucas, 250’, 5M, 11-IX-83, J. and K. Donahue (LACM); Sierra de la
Giganta, Ligui microwave tower, 32 mi S Loreto, 1500’, 1M, 4-IX-84, J. and K. Donahue
(#88,114, LACM); Sierra de la Laguna, Rancho San Antonio de la Sierra, 11.6 rd mi SE
KP 147.6, 3000’, 2M, 11/12-IX-85, J. and K. Donahue (#97,169, LACM); San José del
Cabo, 1M (holotype), 26-XI-61, 1M (paratype), 27-XI-61, Cary-Carnegie Expedition
(CMNH).

6. Sphinx chersis (Hubner) (Figs. 7, 28)
Lethia chersis Htibner, 1823, Sammlung exotischer Schmetterlinge, 2.
Sphinx chersis; Hodges, 1971:58.

This widespread western U.S. species ranges into the northern portion of the peninsula.
Adults have been taken from May to September in adjacent southern California. Potential
larval hosts available in Baja California include Fraxinus (Oleaceae), Prunus (Rosaceae),
and Populus (Salicaceae) (Essig 1926, Hodges 1971).

Specimens examined: BAJA CALIFORNIA NORTE: 4 mi N Santo Tomas, 800’, 2M,
28-V-70, R. Holland (AMNH); Sierra San Pedro Martir, Meling Ranch [2200'], 1M, 13-
V-66 (LACM); 4 mi SW La Zapopita, Valle de Trinidad, 1M, 1F, 16-IV-61, F. Truxal
(LACM).

7. Sphinx libocedrus Edwards (Figs. 8, 27)
Sphinx libocedrus Henry Edwards, 1881, Papilio 1:115; Hodges, 1971:61.

There are two records of S. libocedrus from near the southern tip of the peninsula. It
is uncertain whether the specimens represent a resident population or stray individuals.
Hodges (1971) indicates that S. libocedrus flies from July through September in Texas
and Arizona; both Baja California captures are from September. Forestiera neomexicana
A. Gray (Oleaceae), the only documented larval host (Hodges 1971), occurs in the Cape
Region (Wiggins 1980).

Specimens examined: BAJA CALIFORNIA SUR: 10 mi SW San José del Cabo, 1M,
1-[X-59, Radford and Werner (CAS); 6 mi E El Aguaje, summit Canon Santo Tomas Rd,
3500’, 1M, 1-IX-87, R. Wells (RW).

8. Sphinx perelegans Edwards (Figs. 8, 26)
Sphinx perelegans Henry Edwards, 1874, Proc. Calif. Acad. Sci. 5:109; Hoffmann,
1942:221; Hodges, 1971:61.

190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

S. perelegans is common in the Californian Province, particularly in montane and oak
woodland areas, and ranges south along the west coast to San Quintin. Capture records
extend from April to September, probably representing two broods. Hodges (1971) suggests
Symphoricarpos (Caprifoliaceae) as the larval host; Essig (1926) mentions Prunus (Ro-
saceae) and Arctostaphylos (Ericaceae).

9. Sphinx sequoiae engelhardti Clark (Figs. 9, 29)
Sphinx dolli engelhardti Clark, 1919, Proc. New England Zool. Club 6:104.
Sphinx sequoiae engelhardti; Clark, 1920:66.

In Baja California, S. sequoiae Boisduval is known only from the pinyon-juniper wood-
land areas at the northern end of the Sierra Juarez and near Valle de la Trinidad. The
subspecies S. sequoiae engelhardti is primarily a desert inhabitant occurring in the south-
ern portion of the range of S. sequoiae; it is phenotypically very similar to S. dollii
Neumoegen. In adjacent southern California S. sequoiae has been collected from April
to August. The larval host is Juniperus californica Carr. (Cupressaceae).

Specimens examined: BAJA CALIFORNIA NORTE: 16 km S La Rumorosa, 2M, 27-
V-78, E. Sleeper (CSULB); 4 mi SW La Zapopita, Valle de la Trinidad, 1M, 16-IV-61,
F. Truxal (LACM); near Zapopita, Valle de Trinidad, 1M, 78-IV-61, F. Truxal (LACM).

10. Smerinthus cerisyi Kirby (Figs. 9, 30)
Smerinthus cerisyi Kirby, 1837, Fauna Boreali-Americana 4:301.
Smerinthus cerisyi cerisyi; Mooser, 1940:435.
Smerinthus cerisyi saliceti Boisduval, 1875, Histoire Naturelle des Insectes, Species
Général des Lepidoptéres Heterocéres 1:35; Hoffmann, 1942:222.
Smerinthus cerisyi ophthalmica Boisduval, 1855, Bull. Soc. Entomol. France 332;
Cary, 1963:197.

S. cerisyi occurs commonly throughout much of the Californian Province, particularly
at middle elevations and in riparian habitats, but it is also represented in the Cape Region
by an isolated population. The Cape Region of Baja California marks the southern limit
of this characteristically temperate species (Cary 1963); Mooser (1940) noted its presence
in Baja California Norte, without further detail. S. cerisyi has been recorded from March
through September in the north, and in November and January in the Cape Region. Salix
and Populus (Salicaceae) serve as larval hosts elsewhere (Comstock and Dammers 1943,
Hodges 1971).

Specimens examined: BAJA CALIFORNIA NORTE: Meling Ranch (San Jose), 1M,
30-VI-68, 1M, 2-VII-68, 1M, 1F, 29-VI-68, all D. Patterson (CAS), 1M, 5-IV-71, H. Real
(CAS); 1 mi N Meling Ranch, 1M, 17-III-72, J. Doyen and J. Powell (UCB); trail Las
Encinas to La Sanja, Sierra San Pedro Martir, 1M, 27-V-58, D. Patterson (CAS); 4 mi S
Las Encinas, 1M, 2-VI-58, D. Patterson (CAS); “Mexicali, Rubirosa,” 1M, 11-IX-61, D.
Patterson (CAS); Agua Caliente (San Carlos), 18.5 km E Maneadero, 1M, 1F, 6-VII-78,
P. Arnaud (CAS); 8 mi E Tecate, 1M, 6-VII-84, J. Brown and P. Tocco (SDNHM); Arroyo
Santo Domingo, 5.7 mi E Hamilton Ranch, 1M, 1F, 22-IV-63, H. Leech and P. Arnaud
(CAS); 3 mi S San José del Castillo, 1F, 16-VI-63, E. Sleeper (CSULB). BAJA CALIFOR-
NIA SUR: 4 mi W summit, El Aguaje-Miraflores, Sierra de la Laguna, 1F, 23-I-87, R.
Wells (RW); Arroyo San Bartolo, 4F, 1-XI-61, 2F, 15-XI-61, Cary-Carnegie Expedition
(CMNH); Arroyo San Bernardo, 3F, 17-XI-61, Cary-Carnegie Expedition (CMNH); Puer-
to Chileno, 1F, 26-XI-61, Cary-Carnegie Expedition (CMNH).

Additional records: BAJA CALIFORNIA NORTE: Mike’s Sky Ranch, Sierra San Pedro
Martir, 3600’, 18-VI-70, R. Holland (pers. comm. ).

lla. Pachysphinx occidentalis occidentalis (Edwards) (Figs. 10, 31)

Smerinthus modestus var. occidentalis Henry Edwards, 1875, Proc. Calif. Acad.
Sci. 6:92.

Smerinthus imperator Strecker, 1878, Lepidoptera, Rhopaloceres and Hetero-
ceres, Indigenous and Exotic, 125.

Pachysphinx modesta imperator form kunzei Rothschild and Jordan, 1903, Novit.
Zool. 9(suppl.):3438.

Pachysphinx modesta occidentalis; Mooser, 1940:436.

VOLUME 43, NUMBER 3 191

Pachysphinx modesta imperator; Hoffmann, 1942:223.
Pachysphinx occidentalis; Hodges, 1971:91.

The nominate subspecies is found sporadically throughout the northern portion of the
peninsula. It is most common at middle elevations and in riparian areas where Populus
and Salix (Salicaceae), its larval hosts, grow.

Specimens examined: BAJA CALIFORNIA NORTE: Meling Ranch (San José), 1M,
30-VI-68, 1M, 1-VII-68, D. Patterson (CAS); Agua de Chale, 22 mi S San Felipe, 1M, 18-
VI-68, D. Patterson (CAS); Low. Corona, Sierra San Pedro Martir, 1M, 14-VI-61, E.
Sleeper (CSULB); San José del Castillo, 4M, 3-IX-61, 1M, 1F, 15-VI-61, E. Sleeper (CSULB);
3 mi S San José del Castillo, 1M, 1F, 15-VI-63, E. Sleeper (CSULB); 10 mi S San Matias
Peak, Sierra San Pedro Martir, 1F, 28-VIII-60, E. Sleeper (CSULB).

Additional records: Rothschild and Jordan (1903) cite a pair of specimens from Lower
California (in the Paris Museum) in their original description of P. modesta imperator
form kunzei, hesitating to recognize the taxon as a distinct subspecies for lack of enough
material. We have not examined these specimens, but they may refer to P. occidentalis
peninsularis (see below).

1lb. Pachysphinx occidentalis peninsularis Cary (Fig. 10)
Pachysphinx modesta peninsularis Cary, 1963, Ann. Carnegie Mus. 36:198; Schrei-
ber 1978:48.

A unique population of P. occidentalis was discovered in the Cape Region by the Cary-
Carnegie Expedition (Cary 1963). No additional specimens have been collected. The type
series is from San José del Cabo near the coast, but the insect also may inhabit the Sierra
de la Laguna where its probable larval hosts, Populus and Salix (Salicaceae), occur.

Specimens examined: BAJA CALIFORNIA SUR: San José del Cabo, 2F (holotype and
paratype), 25-X-61, Cary-Carnegie Expedition (CMNH).

12. Erinnyis ello (Linnaeus) (Figs. 11, 32, 33)
Sphinx ello Linnaeus, 1758, Systema Naturae (10th ed.) 1:491.
Erinnyis ello; Cary, 1963:198; Hodges, 1971:99.

This widespread species of the American tropics ranges throughout the peninsula,
uncommon only in the mountains. E. ello is most abundant in the Cape Region. In the
northern portion of the peninsula E. ello flies from July through September; in the southern
portion captures range from July through January. Larval hosts include a variety of plants
in the Euphorbiaceae.

13. Erinnyis crameri (Schaus) (Figs. 12, 34)
Dilophonota crameri Schaus, 1898, Entomol. News 9:136.
Erinnyis crameri; Hodges, 1971:100.

E. crameri occurs throughout most of mainland Mexico (Hoffmann 1942). It has been
collected only once in Baja California, and may not be a breeding resident. Hodges (1971)
indicates that all documented larval host plants are in the Apocynaceae.

Specimens examined: BAJA CALIFORNIA NORTE: Hiway 1, ca. 10 mi NNW Ca-
tavina, 2400’, 1M, 1/2-IX-83, J. and K. Donahue (LACM).

14. Erinnyis obscura obscura (Fabricius) (Figs. 12, 35)
Sphinx obscura Fabricius, 1775, Systema Entomologiae, 538.
Erinnyis obscura; Cary, 1963:200; Hodges, 1971:101.

This little sphingid occurs throughout the lowlands, but is common only in the southern
third of the peninsula, where it has been taken from sea level to 3000 feet. It is widespread
on the mainland, ranging north well into the southern United States. It is occasional in
southern California where it may be resident. Captures range from August through March
in the Cape Region. It is both sexually and seasonally polymorphic, and there is some
confusion whether E. obscura and E. domingonis (Butler) represent separate species or
merely color forms of the same species (Hodges 1971). The two are genitalically indis-
tinguishable, and domingonis-like individuals may be taken sympatrically with E. ob-
scura. Comstock and Dammers (1935) report Philibertia (Asclepidaceae) as the larval
host.

192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

15. Pachylia syces syces (Hiibner) (Figs. 13, 36)
Enyo syces Htibner, 1822, Verzeichniss bekannter Schmettlinge, 132.
Pachylia syces; Cary, 1963:200.

This widespread Neotropical species ranges north at least to the state of Sinaloa on the
Mexican mainland (Hoffmann 1942). It was first reported from Baja California by Cary
(1963). On the peninsula, P. syces is uncommon, and is confined to the Cape Region.
D’ Almeida (1944) reports Ficus (Moraceae) as the larval host; several species are available
in the Cape Region.

Specimens examined: BAJA CALIFORNIA SUR: Los Barriles, 1F, XI-67, V. Stuart
(RW); Hotel Hacienda, Cabo San Lucas, 1F, 16-II-80, J. McBurney (LACM); Los Cabos
airport, 30 mi NE Cabo San Lucas, 1F, 5-IX-83, E. Hawks (LACM); 6 mi W Los Barriles,
El Coro Rd, 1F, 25-I-87, R. Wells (RW); Bahia de las Palmas, 1F, 12-XI-61, Cary-Carnegie
Expedition (CMNH).

16. Callionima falcifera guaycura (Cary) (Figs. 14, 37)
Hemeroplanes parce guaycura Cary, 1963, Ann. Carnegie Mus. 36:200.
Callionima parce guaycura; Schreiber, 1978:51.

Although long treated as C. parce (Fabricius) (Hoffmann 1942, Cary 1963, Hodges
1971, Schreiber 1978), J. Cadiou (pers. comm.) has examined the type specimens of this
and related species, and advises that C. falcifera (Gehlen) is the correct name for this
western Mexican taxon. The weakly distinguished, endemic subspecies C. falcifera guay-
cura is widely distributed throughout nearly the entire southern third of the peninsula,
including the southern portion of the Vizcaino Desert. It is most common in the lowlands
of the Cape Region, at times abundant at beach localities, but has been collected up to
about 1000 m in the Sierra de la Laguna near San Antonio and Miraflores. Captures range
throughout the year with peaks in September and October and again in April and May.
During peak flight periods, C. falcifera may be taken in abundance. Although the early
stages are unknown, the larvae of other members of the genus are known to feed on
plants in the Apocynaceae.

17. Aellopos clavipes (Rothschild and Jordan) (Figs. 15, 38)
Sesia tantalus clavipes Rothschild and Jordan, 1903, Novit. Zool. 9(suppl.):436.
Aellopos clavipes; Hodges, 1971:111.

A. clavipes is abundant in the Cape Region; there is a single record from the northern
portion of the peninsula. It occurs from the immediate coast to about 1300 m in the Sierra
de la Laguna. Adults are diurnal and avidly visit flowers. Captures range from August
to February. Other members of the genus utilize Rubiaceae as larval hosts.

Specimens examined: BAJA CALIFORNIA NORTE: San Quintin, 1F, 12-VIII-54,
alfalfa, Rohlf (SDHNM). BAJA CALIFORNIA SUR: 2 mi S La Paz, 2M, 6-VIII-66, J.
Chemsak (UCB), 1F, 11-VIII-66, J. Powell (UCB); San José del Cabo, 1M, 1116-IX-67,
J. Chemsak and A. Michelbacher (UCB); 26 mi W La Paz, 1M, 10-VIII-66, J. Powell
(UCB); 7 mi S San Pedro, 1M, 10-VIII-66, J. Doyen (UCB); 9 mi SW La Paz, 1M, IF,
14-VIII-66, J. Powell (UCB); Hotel Finisterra, Cabo San Lucas, 1M, 28-XI-80, J. and P.
Brown (SDNHM); 4.2 mi W Miraflores, 1F, 30-IX-80, F. Andrews and D. Faulkner
(SDNHM); 27 km NE Todos Santos, 900’, on flowers of Antigonon leptopus, 1F, 8/9-X-
75, R. Snelling (LACM); Cafion Santo Tomas Rd., 6 mi E El Aguaje, 3500’, 3M, 1-II-87,
R. Wells (RW).

18. Hemaris diffinis (Boisduval) (Figs. 15, 39)
Macroglossa diffinis Boisduval, 1836, Histoire Naturelle des Insectes, Species Gene-
ral des Lépidoptéres, 1:pl. 15.
Hemaris diffinis; Hodges, 1971:117.

This Nearctic species is restricted to the middle elevations (1300-1700 m) of the Sierra
Juarez and the Sierra San Pedro Martir. Captures range from June through September;
it is most common in July. Adults are diurnally active and strongly attracted to the flowers
of low growing annuals, especially the purple flowers of Monardella (Lamiaceae). Al-
though H. senta (Strecker) is reported as occurring in Baja California Norte by Mooser
(1940), Hoffmann (1942), and Schreiber (1978), based on specimens in the Hoffmann

VOLUME 43, NUMBER 3 193

collection, their citations probably refer to H. diffinis. Reported larval hosts include
Symphoricarpos and Lonicera (Caprifoliaceae) (Hodges 1971).

Specimens examined: BAJA CALIFORNIA NORTE: 15 mi E Meling Ranch, Sierra
San Pedro Martir, 4M, 1F, 20-VI-79, J. Brown and D. Faulkner (SDNHM); Las Encinas,
Sierra San Pedro Martir, 4M, 14-VII-80, J. Brown and D. Faulkner (SDNHM); 6 mi N
Laguna Hanson, Sierra Juarez, 1F, 21-VII-80, J. Brown and D. Faulkner (SDNHM); 2-
5 km S El Condor, Sierra Juarez, 1F, 5-IX-83, D. Faulkner (SDNHM); 19 mi E Ojos
Negros, 1M, 21-VII-1980, J. Brown and D. Faulkner (SDNHM).

19. Eumorpha satellitia (Linnaeus) (Figs. 16, 40)
Sphinx satellitia Linnaeus, 1771, Mantissa Plantarum Altera, 539.
Eumorpha satellitia; Hodges, 1971:123.

This widespread Neotropical species is uncommon in Baja California. The few speci-
mens are from the Cape Region, and were collected from July through October. Moss
(1920) reports the larval host as Cissus (Vitaceae).

Specimens examined: BAJA CALIFORNIA SUR: 2.3 mi SW San Bartolo, 1F, 1-X-81,
F. Andrews and D. Faulkner (SDNHM); Punta Lobos, 1M, 20-VII-71, H. Real and R.
Main (CAS); Miraflores, 1M, 1-VIII-71, H. Real and R. Main (CAS); Los Cabos airport,
30 mi NE Cabo San Lucas, 1F, 5-IX-83, E. Hawks (LACM); Sierra de la Laguna, Rancho
La Burrera, 1.9 rd mi S and 12.6 mi E Todos Santos, 1600’, 1M, 15-IX-85, J. and K.
Donahue (#97,345, LACM); Sierra de la Laguna, Rancho San Antonio de la Sierra, 11.6
rd mi SE KP 147.6, 3000’, 5M, 3F, 11/12-IX-85, J. and K. Donahue (#97,169, LACM).

20. Eumorpha achemon (Drury) (Figs. 16, 41)
Sphinx achemon Drury, 1778, Illustrations of Natural History 2:51.
Pholus achemon; Hoffmann, 1942:229.
Eumorpha achemon; Hodges, 1971:124.

E. achemon occurs throughout much of the eastern United States ranging west to
Arizona and California (Hodges 1971), and south into the Mexican states of Sonora,
Chihuahua, and Durango (Hoffmann 1942). In Baja California, E. achemon has been
collected rarely. Commonly associated with Vitis species (Vitaceae) elsewhere, E. ache-
mon may eventually be encountered in the northwestern portion of the peninsula where
grapes are cultivated.

Specimens examined: BAJA CALIFORNIA SUR: La Presa de San Ysidro, near La
Purisima, 1M, 2129-V-84, N. Bloomfield (SDNHM); Highway 1, KP 20, 12 rd mi NE
Villa Insurgentes, 250’, 1M, 7-IX-83, J. and K. Donahue (LACM).

Record: “Baja California’ (no further data) (Hoffmann 1942:229).

21. Eumorpha vitis (Linnaeus) (Figs. 17, 42)
Sphinx vitis Linnaeus, 1758, Systema Naturae (10th ed.) 1:491.
Pholus vitis; Cary, 1963:202.
Eumorpha vitis; Hodges, 1971:126.

Although confined to the Cape Region, this tropical species is the most common sphingid
in the southern portion of the peninsula. It may be encountered in almost every habitat
from the coasts to the mountains. Captures range from July through November. Hodges
(1971) lists Vitis (Vitaceae) as the larval host in the southern United States and mainland
Mexico. Two males from Sierra de la Laguna (LACM) have orchid pollinia attached to
the eyes, suggesting the use of orchids as a nectar source.

22. Eumorpha fasciata (Sulzer) (Figs. 17, 43)
Sphinx fasciatus Sulzer, 1776, Abgekurtze Gesch. der Insecten 1:151.
Pholus fasciatus; Cary, 1963:202.
Eumorpha fasciata; Hodges, 1971:126.

Although widely distributed from northern Argentina to Nova Scotia (Hodges 1971),
E. fasciata is extremely rare in Baja California. The single specimen listed by Cary (1963)
is the only Baja California record to date. Moss (1912) reports the food plant as a member
of the Onagraceae.

Specimen examined: BAJA CALIFORNIA SUR: San José del Cabo, 1F, 25-X-61, Cary-
Carnegie Expedition (CMNH).

194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

23. Euproserpinus phaeton Grote and Robinson (Figs. 18, 44)
Euproserpinus phaeton Grote and Robinson, 1865, Proc. Entomol. Soc. Philadelphia
5:178; Hoffmann, 1942:231; Hodges, 1971:143.

This diminutive, diurnal sphingid is exceptionally inconspicuous as it flies rapidly within
a few inches of the ground. It is most frequently found in dry washes and flat areas in
the desert regions of southern California. It ranges south into Baja California at least as
far as Valle de la Trinidad, a desert intrusion between the Sierra Juarez and the Sierra
San Pedro Martir. The single spring brood flies from late February to April in southern
California. Comstock and Dammers (1935) report Oenothera (Onagraceae) as the larval
host.

Specimens examined: BAJA CALIFORNIA NORTE: Aguajito Spring, Valle de la
Trinidad, 1F, 20-III-36, C. Harbison (SDNHM).

Additional records: BAJA CALIFORNIA NORTE: Hiway 8, 4.7 mi N Valle de las
Palmas, 2M, 3F, 2023-II-72, J.-M. Cadiou (JC).

24. Xylophanes tersa (Linnaeus) (Figs. 18, 45)
Sphinx tersa Linnaeus, 1771, Mantissa Plantarum Altera, 538.
Xylophanes tersa; Cary, 1963:202; Hodges, 1971:150.

Although seldom encountered in numbers, X. tersa may be locally and seasonally
common in Baja California, where it is confined to the southern tip of the peninsula. It
occurs from the coastal lowlands to about 900 m in the Sierra de la Laguna. It is apparently
single-brooded with adults having been taken from September to early December. Else-
where the larvae feed on Rubiaceae; larval hosts in Baja California are unknown.

25. Xylophanes pluto (Fabricius) (Figs. 19, 46)
Sphinx pluto Fabricius, 1777, Genera Insectorum, 274.
Xylophanes pluto; Hodges, 1971:149.

Hoffmann (1942) indicated that X. pluto occurred throughout Mexico with the excep-
tion of the northwestern region. We have examined single specimens from both Sinaloa
(UCB) and Baja California. The probable host in Baja California is Chiococca (Rubiaceae).

Specimen examined: BAJA CALIFORNIA SUR: Hwy 19, 14.5 rd mi NW Cabo San
Lucas, 250’, 1M, 11-IX-83, J. and K. Donahue (LACM).

26. Hyles lineata (Fabricius) (Figs. 20, 47)
Sphinx lineata Fabricius, 1775, Systema Entomologiae, 541.
Celerio lineata; Cary, 1963:1538.
Hyles lineata; Hodges, 1971:158.

H. lineata is by far the most common and widespread sphingid in Baja California. It
ranges from the coastal lowlands to the mountains; it is common in the desert areas; it is
also known from several islands both in the Sea of Cortés (e.g., Isla Angel de la Guarda
and Isla Mejia) and along the Pacific coast (e.g., Isla de Cedros and Isla Guadalupe). In
the north it is on the wing from March through October; in the south captures range
throughout the year with a peak in September through November. Larval hosts encompass
many genera in several families including Rosaceae, Solanaceae, Onagraceae, Portula-
caceae, and Nyctaginaceae.

Possible Species

The Californian Province in the northwestern portion of the peninsula
represents a significant southern intrusion of Nearctic elements, both
floral and faunal. Physiographically, this region is an extension of the
area to the immediate north. Most of the sphingids present in southern
California occur here as well. Two Californian species for which recent
Baja California records are conspicuously absent are Proserpinus clark-
iae (Boisduval) and Arctonotus lucidus Boisduval. Mooser (1940) cites
an unspecified number of each from Baja California Norte in the col-

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204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 21-26. 21, Agrius cingulatus. 22, Manduca sexta. 23, Manduca quinque-
maculata. 24, Manduca rustica. 25, Sphinx xantus. 26, Sphinx perelegans.

VOLUME 438, NUMBER 3 205

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Smerinthus cerisyi. 31, Pachysphinx occidentalis. 32, Erinnyis ello (male). 33, Erinnyis
ello (female). 34, Erinnyis crameri.

lection of Carlos C. Hoffmann, a reference which subsequently has
been repeated, without examination of the material, by Hoffmann
(1942), Hodges (1971), and Schreiber (1978), for the former species,
and by Hoffmann (1942) for the latter. Hodges (pers. comm.) indicates
that he has not examined specimens from this area. Both Hodges’ and

206 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 35-41. 35, Erinnyis obscura. 36, Pachylia syces. 37, Callionima falcifera
(female). 38, Aellopos clavipes. 39, Hemaris diffinis. 40, Eumorpha satellitia. 41, Eu-
morpha achemon.

VOLUME 438, NUMBER 3 207

Fics. 42-47. 42, Eumorpha vitis. 43, Eumorpha fasciata. 44, Euproserpinus phae-
ton. 45, Xylophanes tersa. 46, Xylophanes pluto. 47, Hyles lineata.

208 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Schreiber’s citations presumably are based on Mooser’s and/or Hoff-
mann’s records. There is no apparent reason why either of these species
should be absent from the northern chaparral or montane regions; both
have been taken to the immediate north in San Diego County, Cali-
fornia.

The Neotropical sphingid fauna of the Cape Region at the southern
end of the peninsula is exceedingly depauperate compared with that
of the Mexican mainland. Species recorded from Sinaloa and Sonora
that eventually may be discovered in this region include Sphinx merops
Boisduval, Erinnyis yucatana (Druce), Pachylia ficus (Linnaeus), Cau-
tethia spuria (Boisduval), Eumorpha labruscae (Linnaeus), and Xylo-
phanes falco (Walker). In addition, J. Cadiou (pers. comm.) has sug-
gested that the following widely ranging Neotropical species may
eventually be documented from the peninsula: Erinnyis alope (Drury),
E. domingonis, Enyo lugubris (Linnaeus), and Pseudosphinx tetrio
(Linnaeus).

ACKNOWLEDGMENTS

We acknowledge with thanks the assistance of the following individuals whose coop-
eration permitted the use of specimens in their care (the following abbreviations are used
in the text to indicate disposition of specimens examined): F. Rindge, American Museum
of Natural History (AMNH), New York; P. Arnaud, Jr., California Academy of Sciences
(CAS), San Francisco; W. Clark, College of Idaho (CI), Caldwell; J. Rawlins, Carnegie
Museum of Natural History (CMNH), Pittsburgh, Pennsylvania; E. Sleeper, California
State University, Long Beach (CSULB); C. Hogue, Los Angeles County Museum of Natural
History (LACM); E. and D. Hawks, San Diego, California (now LACM); J.-M. Cadiou
(JC) personal collection, Saint Cloud, France; J. McBurney (JM) personal collection (now
LACM), Anaheim, California; R. Wells (RW) personal collection, Jackson, California; D.
Faulkner, San Diego Natural History Museum (SDNHM), San Diego, California; J. Powell,
Essig Museum of Entomology (UCB), University of California, Berkeley; R. Schuster,
Bohart Museum of Entomology (UCD), University of California, Davis. Duplicate spec-
imens from the LACM collection are being deposited in the Instituto de Biologia, Uni-
versidad Nacional Autonoma de Mexico, D. F., Mexico.

We thank Ron Hodges for comments on the text, and Jerry Powell for reviewing an
early draft of the manuscript. Jean-Marie Cadiou provided helpful comments on a pre-
liminary checklist of species. We are particularly grateful to the Entomology Department
of the San Diego Natural History Museum, which provided one of us (JWB) with funds
for travel and a base of operations for investigations into Baja California. We also thank
the National Science Foundation for grant number BSR84-10742 to the Los Angeles
County Museum of Natural History, which funded a technician who prepared some of
the material studied.

LITERATURE CITED

BROWN, J. W. 1987. The peninsular effect in Baja California: An entomological assess-
ment. J. Biogeogr. 14:359-365.

Cary, M. M. 1963. Reports on the Margaret M. Cary and Carnegie Museum Expedition
to Baja California, 1961. 2. The family Sphingidae. Ann. Carnegie Mus. 36:193-204.

CLARK, B. P. 1920. Sixteen new Sphingidae. Proc. New England Zool. Club 7:65-78.

Comstock, J. A. & C. M. DAMMERS. 1935. Notes on the early stages of three butterflies
and six moths from California. Bull. So. Calif. Acad. Sci. 34:120-142.

VOLUME 43, NUMBER 3 209

1943. California species of Smerinthus with notes on the early stages of S.
cerisyi ophthalmicus. Bull. So. Calif. Acad. Sci. 42:42—45.

Cook, R. E. 1969. Variation in species density in North American birds. Syst. Zool. 18:
63-84.

D’ALMEIDA, R. F. 1944. Estudios biologicos sobre algunas lepidopteros do Brasil. Ar-
quivos Zool. Est. SAo Paulo 4:33-72.

Essic, E. O. 1926. Insects of western North America. Macmillan Co., New York.
1035 pp.

HopcEs, R. 1971. Sphingoidea. In The moths of America north of Mexico, fasc. 21. E.
W. Classey Ltd. and R.B.D. Publ. Inc., London. 158 pp. + plates.

HOFFMANN, C. C. 1942. Catalogo sistematico y zoogeografico de los lepidopteros Mex-
icanos, tercera parte: Sphingoidea y Saturnioidea. Anales Inst. Biol. Univ. Auton. Mex.
13:213-256.

KIESTER, A. R. 1971. Species density of North American amphibians and reptiles. Syst.
Zool. 20:127-137.

Mooser, O. 1940. Fauna Mexicana. III (Insecta, Lepidoptera, familia Sphingidae).
Enumeracion de los Esfingidos Mexicanos (Insecta, Lepiddéptera), con notas sobre su
morfologia y su distribucion en la Republica. An. Escuela Nac. Ciencias Biol. 1(3 +
4):407-495 (incl. pls. 57-75).

Moss, A. M. 1912. On the Sphingidae of Peru. Trans. Zool. Soc. London 20:73-134.

1920. Sphingidae of Para, Brazil. Novit. Zool. 27:333—424.

ROTHSCHILD, W. & K. JORDAN. 1908. A revision of the lepidopterous family Sphingidae.
1:CXXXV + 456. Novit. Zool. 9(suppl).

SCHREIBER, H. 1978. Dispersal centres of Sphingidae (Lepidoptera) in the Neotropical
region. Biogeografica 10. Dr. M. Junk B.V., Publ., The Hague and Boston. 195 pp.

SEIB, R. L. 1980. Baja California: A peninsula for rodents but not for reptiles. Am. Nat.
115:613-620.

SHREVE, F. 1951. Vegetation of the Sonora Desert, pt. 1. Carnegie Inst. Washington
Pub> 591.

SIMPSON, G. G. 1964. Species densities of North American recent mammals. Syst. Zool.
13:57-78.

TAYLOR, R. J. & P. J. REGAL. 1978. The peninsular effect on species diversity and the
biogeography of Baja California. Am. Nat. 112:583-598.

WiccIns, I. 1980. Flora of Baja California. Stanford Univ. Press, California. 1025 pp.

Received for publication 20 November 1988; accepted 15 May 1989.

Journal of the Lepidopterists’ Society
43(3), 1989, 210-216

BIOLOGY AND IMMATURE STAGES OF
SCHINIA MASONI (NOCTUIDAE)

BRUCE A. BYERS

Natural Science Program, Campus Box 331,
University of Colorado, Boulder, Colorado 80309-0331

ABSTRACT. Schinia masoni (Smith) was studied using field observations, laboratory
rearing, and data from museum collections. Its larval host plant, not previously reported,
is Gaillardia aristata. Adults also take nectar from this plant and are well camouflaged
on its blossoms. The flight period of S. masoni is synchronized with the blooming of
Gaillardia aristata, both peaking in late June. Eggs are deposited between disk-flowers
of the host, there are five larval stages, and pupation occurs in the soil. Museum records
suggest that this species occurs only in Colorado. Schinia masoni and forms with coloration
almost identical to closely-related S. volupia occur sympatrically on Gaillardia aristata
in east-central Colorado, and individuals with intermediate coloration are common in
this area, raising a question about the systematic relationship of these two species.

Additional key words: Schinia volupia, Gaillardia aristata, Heliothinae.

Schinia masoni (Smith 1896), a heliothidine flower moth, feeds in
the larval stage on flowers and developing seeds of blanketflower, Gail-
lardia aristata Pursh (Asteraceae). The burgundy wings and yellow
head and thorax of adults make them extremely well camouflaged when
feeding or resting on G. aristata blossoms (Cockerell 1910, Ferner &
Rosenthal 1981, Owen 1980). Cockerell (1927) wrote that this species
“was discovered by Mr. J. Mason, formerly of Denver, through the
picking of a Gaillardia flower on which a moth happened to be resting.”’
In his description of the species, Smith (1896) stated that it was collected
on flowers of Rudbeckia. Several facts suggest that this may be an error:
the flowers of Gaillardia and Rudbeckia are somewhat similar and are
commonly confused by non-botanists; the colors of Schinia masoni do
not match those of Rudbeckia; and it was never observed on Rudbeckia
(n = 400 blossoms) during this study even when these were interspersed
with Gaillardia aristata on which S. masoni was observed. Cockerell
(1910) observed it only on Gaillardia aristata.

Three photographs of Schinia masoni resting on Gaillardia aristata
have been published (Brower & Brower 1956, Ferner 1980, Owen 1980),
as well as one paper and two notes on how its behavior relates to
camouflage (Brower & Brower 1956, Cockerell 1910, Ferner & Rosen-
thal 1981). The species was illustrated in Holland (1903) as Rhododipsa
masoni; this generic name was later synonymized with Schinia (Hard-
wick 1958).

A combination of field observation, laboratory rearing, and data from
museum collections was used in this study, which reports for the first
time the larval host plant and immature stages of Schinia masoni, and

VOLUME 43, NUMBER 3 Has WA

considers the systematic relationship between Schinia masoni and
closely-related S. volupia.

Biology

All but 6 of 90 specimens in several museums (American Museum
of Natural History, Canadian National Collection, Denver Museum of
Natural History, Los Angeles County Museum of Natural History, Uni-
versity of Colorado Museum, and the U.S. National Museum) come
from Colorado, specifically the foothills of the Rocky Mountains be-
tween Denver and Ft. Collins. Six specimens from the U.S. National
Museum are labelled from “Utah” or “U.T.”’ These specimens have no
dates of collection, and their locality data are incomplete and uncertain
(R. W. Poole pers. comm.). Schinia masoni has been collected from
only a very small part of the range of Gaillardia aristata, which occurs
northward from Colorado into Canada and westward to Washington,
Oregon, and British Columbia (Biddulph 1944).

Dates of collection of the 90 museum specimens of Schinia masoni
range from 10 June-15 July, with a peak during the last week of June.
Adults were observed in Boulder Co., Colorado, from 14 June-6 July,
1988, at elevations from 1820 to 2730 m. Adults appeared at the lowest
elevations first.

During the day adults were usually observed resting on the disk of
a flower (n = 7) or under the ray-flowers (n = 2). Gaillardia aristata
has brick-red disk-flowers and yellow ray-flowers. Moths resting on the
tops of flowers were often oriented with their yellow heads and thoraces
outward over the bases of the yellow ray-flowers and their burgundy
wings over the brick-red disk, the most advantageous position for cam-
ouflage, as noted by Cockerell (1910), Brower and Brower (1956), and
Ferner and Rosenthal (1981). Adults were observed actively flying and
seeking nectar at dusk (n = 5).

Captive adults (n = 4) were kept in the laboratory in 4-| glass jars
with four or five blossoms of G. aristata in a small container of water.
Blossoms were replaced daily; no other water or food was added. The
laboratory room was open to free circulation of outdoor air, and tem-
peratures were essentially the same as outdoors, ranging from 21° to
29°C. Captive moths were most active during the late afternoon and
early evening, and rested on blossoms during the day. Captive adults
lived up to 6 days.

Eggs oviposited by captive females were always laid between disk-
flowers (n = 27), although in the field a few eggs were found on the
surfaces of buds in which the disk-flowers were still very tightly packed.
Three captive females oviposited in the laboratory. Oviposition was
observed five times between 1200 and 2115 h, but most often (3/5

2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

cases) at dusk. Captive females commonly laid more than one egg in
each blossom; one laid seven eggs in one blossom. In the field it was
not uncommon to find two or three eggs or small larvae in a flower.
Unhatched eggs were found in the field on blossoms from the bud stage
to those that were almost finished blooming (14% weeks past the bud
stage).

~ Newly-hatched larvae tunnelled into an adjacent disk-flower. Blos-
soms containing small larvae showed patches of brown and shrunken
disk-flowers; larger larvae pushed up patches or ridges of disk-flowers
in feeding on the developing seeds underneath.

Larvae burrowed into the soil to pupate. Pupae removed from sandy
soil in the laboratory had bound soil particles together with silk to a
distance of approximately 5 mm in all directions to form a weak pu-
pation chamber (n = 16). This species appears to be univoltine.

Description of Stages

Descriptions of immature stages are based on both field-collected
and laboratory-reared larvae. Head widths and body lengths were mea-
sured on larvae collected in the field. Larvae were reared in the lab-
oratory using techniques described by Hardwick (1958). They were
examined every second day, and changes in body length, color pattern,
and evidence of molting were noted. Duration of each larval stage was
estimated from dated evidence of molts and body length measurements
of laboratory-reared larvae, supplemented with head-width and body-
length measurements from field-collected larvae. Unless otherwise not-
ed, numerical data are means and standard deviations.

Adult. (Figs. 1, 2) (n = 19) Smith’s (1896) original description seems generally accurate
although the following differences or additions should be noted: Abdomen: dark grayish
brown, rather than “blackish” as stated by Smith, often with row of yellow scales at
posterior margin of each segment; some dark purplish pink scales ventrally and laterally.
Forewing: burgundy or crimson due to mixture of dark purplish red and dark grayish
brown scales (not “black” as stated by Smith). Lines and spots of very pale yellow;
distinctness of lines variable. Antemedial (am) line with even outcurve and often with
white teeth pointing basally. Postmedial (pm) line bisinuate, narrowing median space
toward inner margin. Orbicular and claviform spots very pale yellow; claviform spot
often appearing to connect am and pm lines. Subterminal line very pale yellow, often
indistinct. Fringe grayish pink to pinkish fawn. Underside of forewing dark grayish brown
(not “black” as stated by Smith) with carmine or burgundy around costal and outer
margins. Fringe grayish pink to pinkish fawn. Hindwing: medium to dark grayish brown
(not “black” as stated by Smith); outer margin and anal angle sometimes with burgundy
tinge. Fringe grayish pink to pinkish fawn. Underside of hindwing mostly carmine or
burgundy, often with dark grayish brown around inner margin and humeral and anal
angles. Fringe pale pinkish yellow to pinkish fawn. Sexes same coloration. Forewing
length: 10.8 + 0.7 mm (n = 11).

Egg. (Fig. 3) (n = 10) White, sometimes with very pale yellow tint, iridescent. Elongate,
0.8-0.9 mm long and 0.3-0.4 mm wide, blunter and more rounded at micropylar end.
Often deformed by compression between tightly-packed disk-flowers. Remains same color

VOLUME 43, NUMBER 3 213

Fics. 1-4. Schinia masoni (Smith); adults and immature stages. 1, Paralectotype S.
masoni, male, Denver Museum of Natural History; 2, adult feeding on its food plant
Gaillardia aristata (not in head-out resting position most advantageous for camouflage);
3, two eggs deposited between disk-flowers of G. aristata; 4, fifth instar larvae, dorsal
and lateral views.

until black head and thoracic shield become visible through chorion shortly before hatch-
ing. Incubation period: 3-4 days (n = 10).

First instar. (n = 16) Head, prothoracic, and suranal shields black. Body white. Spiracles
with dark rims. Head width: 0.26 mm (n = 1). Duration: 4.5 days (n = 7).

Second instar. (n = 10) Head very dark brown. Prothoracic and suranal shields dark
brown to black; sometimes solid color, sometimes with three longitudinal lines of cream
or very pale yellow, lines usually less distinct than in later instars. Mid-dorsal line varying
from medium brown, pale reddish brown and pale tan, to pale pink or yellowish pink.
Subdorsal area cream or very pale yellow, usually with two subdorsal lines of same color
as, or slightly paler than, mid-dorsal line. Supraspiracular line same color as mid-dorsal
and subdorsal lines. Subdorsal and supraspiracular lines sometimes not well developed,
pigmented only in middle of segments. Spiracular line and suprapodal area cream or
very pale yellow. Spiracles with black rims. Thoracic legs varying from cream or very
pale yellow to caramel. Head width: 0.56 + 0.06 mm (n = 5). Duration: 4 days (n = 10).

Third instar. (n = 9) Head light to dark caramel, sometimes mottled with darker brown
or black. Prothoracic and suranal shields black or very dark brown with three longitudinal
stripes of ivory or very pale yellow. Mid-dorsal line varying from dark brown or dark
purplish brown, through reddish brown and brick red, to pale pink or yellowish pink.
Subdorsal area cream or very pale yellow with two subdorsal lines of same color as, or
slightly paler than, mid-dorsal line. Supraspiracular line same color as mid-dorsal and
two subdorsal lines. Spiracular line cream or very pale yellow. Spiracles with black rims.
Suprapodal area usually cream or very pale yellow, sometimes with pale rose or brown
color. Thoracic legs varying from cream or very pale yellow to caramel. Head width:
0.92 + 0.06 mm (n = 19). Duration: 2.5 days (n = 9).

214 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fourth instar. (n = 9) Generally same as third instar. Pigmentation of suprapodal area
often darkening; concolorous with, but paler than, mid-dorsal, subdorsal and supraspi-
racular lines. Head width: 1.45 + 0.10 mm (n = 15). Duration: 3-3.5 days (n = 9).

Fifth instar. (Fig. 4) (n = 9) Head light to dark caramel, sometimes mottled with
darker brown or black. Prothoracic and suranal shields black or very dark brown, divided
into four bars by three longitudinal stripes of ivory or very pale yellow. Mid-dorsal line
varying from dark brown or dark purplish brown, through reddish brown and brick red,
to pale pink or yellowish pink. Subdorsal area ivory or very pale yellow with two subdorsal
lines of same color as, or slightly paler than, mid-dorsal line. Supraspiracular line same
color as mid-dorsal and two subdorsal lines. Spiracular line ivory or very pale yellow.
Spiracles with black rims. Suprapodal area concolorous with, but usually paler than, mid-
dorsal, subdorsal and supraspiracular lines, with a broken longitudinal line of ivory or
very pale yellow. Thoracic legs varying from ivory or very pale yellow to caramel. Setal
arrangement same as that of members of the elliptoid-eyed group of the genus (Hardwick
1958, fig. 87). On the first eight abdominal segments (Al to A8), SD2 is minute and
variably absent; on the first and second thoracic segments (T1 and T2), SD1 and SD2 are
sometimes absent (Stehr 1987). Head width: 2.36 + 0.06 mm (n = 11). Duration: 5.5-6
days (n = 9).

Total duration of larval life, laboratory rearing at 21-29°C: 19.9 + 1.3 d (n = 9). At
elevations between 2560 m and 2730 m collections of larvae made 18 days apart suggest
that larval development may require up to twice as long as above, undoubtedly because
of colder temperatures.

Pupa. (n = 16) Orange-brown. Spiracles on segments 2 and 3 borne on weak projections
of cuticle; on segment 4 on a level with general surface of cuticle; on segments 5-7 in
shallow depressions. Anterior margins of segments 5-7 with band of conspicuous pits.
Proboscis terminating at apices of wings. Cremaster usually consisting of four setae borne
on prolongation of 10th abdominal segment. Apical abdominal segments similar to those
of S. pallicinta (Hardwick 1972a) or S. jaegeri (Hardwick 1972b) except for number of
setae. Setae often slightly curved ventrally; inner pair (approx. 0.3-0.4 mm long) slightly
longer than outer pair (approx. 0.2 mm long), which is directly lateral to inner pair. One
or both outer setae occasionally much reduced or absent. Length from anterior end of
pupa to posterior margin of fourth abdominal segment: 7.9 + 0.5 mm (n = 16).

Larval Diagnosis

In the elliptoid-eyed members of the genus Schinia, Hardwick (1958)
found that “chaetotaxy ... has no significance on the specific level ... .
There is rather wide latitude in the setal arrangement of individual
larvae but no interspecific variation is evident.” In fact, throughout the
genus Schinia setal patterns are of very little diagnostic use, whereas
larval color patterns are very often diagnostic (D. F. Hardwick pers.
comm.).

The color pattern of the fifth instar distinguishes Schinia masoni
from other described species of Schinia. The four black or very dark
brown bars on the prothoracic shield distinguish it from all but S.
pallicinta (Hardwick 1972a), which was formerly placed in the genus
Rhododipsa along with S. masoni. The prothoracic shield of S. jaegeri
is similar, but the dark bars are not as well defined (Hardwick 1972b).
The presence of a mid-dorsal band, two subdorsal bands, and a su-
praspiracular band, and the reddish pigmentation of these bands, dis-

VOLUME 43, NUMBER 3 Dal ts,

tinguishes S$. masoni from both S. pallicinta and S. jaegeri. These
diagnostic pigmentation patterns may be seen in Fig. 4.

Larval Feeding Ecology

To compare the larval development of S. masoni with the rate of
development of Gaillardia aristata blossoms, 10 blossoms were marked
at the bud stage and photographed twice a week for three weeks. This
record showed that it took 2% weeks for a flower to go from the bud
stage (ray-flowers absent or just emerging) to the early seed-head stage
(ray-flowers dried and shriveled, or dropped; seeds beginning to dry
and harden).

Nearly three weeks were required for S. masoni to complete its
development from egg to pupa in the laboratory. In the field, it appeared
that some larvae had completed development on a single blossom.
However, captive fifth instar larvae ate the developing seeds of an
entire blossom approximately every two days for about the last four
days before pupation. Such a feeding rate makes it seem unlikely that
a larva could complete development in the flower on which its egg was
laid. Movement of larvae from flower to flower was observed in the
field: several late third or early fourth instar larvae were seen crawling
on uneaten blossoms near ones that had been eaten but that contained
no larvae. On the other hand, many larvae were found on isolated
blossoms many meters from any other, making it seem unlikely that
they could locate and move to another blossom to complete their de-
velopment. Clarification of this aspect of the larval feeding ecology of
S. masoni will require further research.

Systematic Status

The original description of Schinia masoni (Smith 1896) recognized
its close resemblance to Schinia volupia (Fitch), and these species are
still considered to be closely related (D. F. Hardwick pers. comm.). The
larval and adult food plant of S. volupia has not previously been re-
ported; during this study it was found to be Gaillardia pulchella Fou-
geroux, at least in eastern Colorado. Specimens of S. volupia in the
museums listed above were collected in Colorado, Kansas, Oklahoma,
Texas, New Mexico, and Louisiana; this area overlaps most of the range
of Gaillardia pulchella (Biddulph 1944).

During this study an area was found on the Palmer (Platte-Arkansas)
Divide between Denver and Colorado Springs where typical S. masoni
and individuals with coloration almost identical to S. volupia occur
together on Gaillardia aristata. Schinia volupia has light pink to car-
mine-pink forewings and hindwings, and none of the specimens from

216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

eastern Colorado or New Mexico I examined (n = 12) had any grayish
brown scales on either forewings or hindwings. The pale volupia-like
forms from the Palmer Divide all had some grayish brown scales on
the upper hindwings, giving them a visible brownish tinge not seen in
typical volupia. Individuals with intermediate coloration were common
in this area. If these intermediate forms are hybrids, it is possible that
S. masoni and S. volupia are subspecies rather than full species.

ACKNOWLEDGMENTS

My warmest thanks to A. Armstrong who first called my attention to this moth; R. S.
Peigler of the Denver Museum of Natural History for his invaluable help and encour-
agement; D. F. Hardwick for advice on larval taxonomy and rearing; U. N. Lanham and
M. J. Weissmann of the University of Colorado Museum for advice and use of equipment;
J. P. Donahue of the Los Angeles County Museum of Natural History, J. D. Lafontaine
of the Biosystematics Research Center, Agriculture Canada, R. W. Poole of the U‘S.
National Museum, and F. H. Rindge of the American Museum of Natural History for
data on the specimens in those collections; and R. Holland for specimens of S. volupia
collected in New Mexico.

LITERATURE CITED

BIDDULPH, S. F. 1944. A revision of the genus Gaillardia. Res. Stud. State Coll. Wash-
ington 12:195-256.

BROWER, L. P. & J. V. BROWER. 1956. Cryptic coloration in the anthophilous moth
Rhododipsa masoni. Am. Natur. 90:177-182.

COCKERELL, T. D. A. 1910. Notes on some heliothid moths. Entomol. News 21:343-
344.

1927. Zoology of Colorado. Univ. Colorado, Boulder, Colorado. 262 pp.

FERNER, J. W. 1980. [Schinia masoni photo.] Bioscience 30 (Feb.):cover.

& M. ROSENTHAL. 1981. A cryptic moth Schinia masoni (Noctuidae) on Gail-
lardia aristata (Compositae) in Colorado. Southw. Nat. 26:88—90.

HARDWICK, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed species
of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae. Can. Entomol.
Suppl. 6, 116 pp.

1972a. The life history of Schinia pallicincta (Noctuidae). J. Lepid. Soc. 26:

29-33. ;

1972b. The life history of Schinia jaegeri (Noctuidae). J. Lepid. Soc. 26:89-93.

HOLLAND, W. J. 1903. The moth book. Doubleday Page & Company, New York.
479 pp.

OwEN, D. 1980. Camouflage and mimicry. Univ. Chicago Press, Chicago. 158 pp.

SMITH, J. B. 1896. A new species of Rhododipsa. Entomol. News 7:284—285.

STEHR, F. W. 1987. Immature insects. Kendall/Hunt Publishing Co., Dubuque, Iowa.
768 pp.

Received for publication 18 October 1988; accepted 15 May 1989.

Journal of the Lepidopterists’ Society
43(3), 1989, 217-228

GENETIC DIFFERENTIATION AMONG CALIFORNIA
POPULATIONS OF THE ANISE SWALLOWTAIL
BUTTERFLY, PAPILIO ZELICAON LUCAS

MARK L. TONG’ AND ARTHUR M. SHAPIRO?
Department of Zoology, University of California, Davis, California 95616

ABSTRACT. The anise swallowtail butterfly, Papilio zelicaon Lucas, is widely dis-
tributed in California. California zelicaon are composed of low- and high-elevation eco-
types defined by host-plant preference and diapause physiology. Electrophoretic-genetic
surveys of 14 loci over 10 populations (157 samples total) demonstrate great similarity
among these ecotypes, suggesting that their adaptive differences may be defined by a
small number of loci rather than broad genomic differentiation.

Additional key words: ecotypes, electrophoresis, Papilionidae.

The anise swallowtail butterfly, Papilio zelicaon Lucas, is native to
western North America, where it is widely distributed (Tyler 1975). In
central California, zelicaon is found in a wide variety of habitats from
sea level to tree line (Table 1, Fig. 1). Populations at the same latitude
exhibit diapause phenologies from univoltine (one generation/yr) to
multivoltine (up to four/yr) as a function of habitat elevation, length
of growing season, and larval host plant (Sims 1979).

Populations in the Coast Range and the Sierra Nevada above 400 m
primarily utilize native Umbelliferae including Lomatium, Angelica,
and Cymopterus. These native plants are available to zelicaon larvae
from spring to midsummer when the onset of hot, dry weather renders
the leaves too hard and dry for the larvae to ingest. These populations
are univoltine, though in the montane Sierra a second generation oc-
casionally occurs (Sims 1979, Shapiro unpubl.).

Lowland populations (below 400 m) today feed almost exclusively
on sweet fennel (Foeniculum vulgare Miller, Umbelliferae), which is
common throughout coastal and interior lowland California (Munz
1970), and also on orange (Citrus sinensis Osbeck, Rutaceae) which
has been grown commercially in California since 1841 (Opitz & Platt
1969). Both plants are available to zelicaon 8-12 months per year,
allowing these populations to breed continuously (Sims 1983). Fennel
and orange were introduced to California by Spanish missionaries in
the 18th Century (Hutchinson 1969). Both produce natural compounds
similar to those in native Umbelliferae (Dethier 1941). Before this
zelicaon was presumably univoltine, being limited by ephemeral host
plants at low elevations and the short growing season in the mountains
(Sims 1983). The introduction of these perennial host plants probably

‘Current address: 109 Berwick Drive, Pittsburgh, Pennsylvania 15215.
* To whom reprint requests should be sent.

218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

enabled zelicaon to “‘switch’’ its ovipositional preference to the intro-
duced plants or to disperse to areas where only the introduced species
were available, or both. This, in turn, allowed multivoltinism to evolve
in the lowland areas where these plants are abundant (Shapiro & Masuda
1980, Sims 1983).

_ Papilio zelicaon populations are consistent with the ecotype concept
first proposed by Turesson (1922) for hawkweed (Hieracium, Astera-
ceae): plants from different habitats were shown to be phenotypically
distinct even when grown under identical conditions, thus demonstrat-
ing a genetic basis for the differences.

Using wild zelicaon and a laboratory strain selected for nondiapause,
Sims (1979) demonstrated that univoltine zelicaon populations have
significantly higher diapause incidence (photophase required to induce
diapause) and intensity (duration of chilling needed to terminate dia-
pause) than multivoltines, and that this phenomenon is genetically based.
Sims (1983) showed that incidence and intensity are polygenically in-
herited, with intensity being affected by maternal phenotype. These
characteristics are maintained under varied environmental regimes (Sims
1979, Shapiro unpubl.).

The present study began as an attempt to use electrophoretic analysis
to determine whether orange-feeding zelicaon in northern California
evolved independently of orange-feeding zelicaon in southern Califor-
nia, or had been introduced inadvertently from the south. Papilio zeli-
caon was reported as an orange pest as early as 1909 near Visalia, Tulare
Co. (Coolidge 1910), and in the 1960’s near Chico, Butte Co. (Shapiro
unpubl.). This study also investigates the degree to which differentiation
of zelicaon into low- and high-elevation diapause ecotypes is reflected
by electrophoretically detectable genetic variation.

MATERIALS AND METHODS

Electrophoresis, a commonly used method in biochemical system-
atics, is based on the movement of charged particles under the influence
of an electrical field (Ferguson 1980). Proteins carry a net electrical
charge depending on amino acid structures and environmental pH. The
rate at which proteins migrate through a support medium is related to
their size and shape and is proportional to net charge. Different proteins
with different electrophoretic properties migrate at different rates un-
der identical test conditions.

Differential migration of homologous proteins is detectable and of
special interest in biochemical systematics. Such differentiation is pre-
sumed to reflect differences in nucleic acid sequences that encode pro-
teins. The degree of electrophoretically detected differentiation is
thought to reflect the extent of evolutionary divergence between sam-

VOLUME 43, NUMBER 3 219

1 Gazelle

2 Washington

3 Hemet

4 Orland

5 Butts Canyon

6 Suisun

7 ~~ Blue Ridge

8 Rancho Cordova
9 Castle Peak

Auburn

Fic. 1. Location of populations studied.

pled taxonomic groups, although the structural proteins studied rep-
resent only one segment of the overall genome.

Adult zelicaon were collected during 1984 and 1985 flight seasons
at the sites in Table 1. Captured specimens were frozen live and stored
at —70°C to prevent protein denaturation.

In preparation for analysis, thoraces were excised and homogenized
in 600 ul of glass-distilled water with a Teflon-coated tissue grinder.
Homogenate was absorbed onto 2 x 9 mm wicks of #3 Whatman paper
and applied to the gels. Horizontal slab gels were made with Sigma
starch and were prepared and run for 5 h using methods described in
Ayala et al. (1972, 1974a).

After running, gels were cut into four 2-mm-thick slices so that each

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

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VOLUME 43, NUMBER 3 Repel

TABLE 2. Enzymes assayed.

Enzyme Abbreviation Buffer*
Phospho-glucose isomerase PGI REG
Aldolase ALDO REG
a-Glycerophosphate dehydrogenase aGPD REG
Glutamate-oxaloacetate transaminase GOT-1 REG
Hexokinase HK-1 REG
Phospho-gluco mutase PGM REG
Fumarase FUM-2 REG
Mannose phosphate isomerase MPI REG
Malic enzyme ME-1 JRP
Glucose-3-phosphate dehydrogenase G38PD DH
Glucose-6-phosphate dehydrogenase G6PD DH
Hydroxybutyrate dehydrogenase HBDH DH
Esterase EST-1 DH

EST-2 DH

* REG: Gel buffer—9 mM Tris, 3 mM citric acid, pH 7.0. Electrode buffer—135 mM Tris, 45 mM citric acid. JRP:
Gel buffer—76 mM Tris, 5 mM citric acid, pH 8.65. Electrode buffer—300 mM boric acid, 60 mM NaOH. DH: Gel
and electrode buffer—8.7 mM Tris, 8.7 boric acid, 1 mM EDTA, 1 mM 6-NAD*, pH 9.0.

sample was tested for four enzymes. Table 2 lists the enzymes assayed.
Specific staining systems and gel fixation techniques are described in
Ayala et al. (1972, 1974a).

Fixed gels were scored after each run using a light box. Loci were
characterized and interpreted as for Pieridae, for which the genetic
basis of the electrophoretic banding patterns has been demonstrated in
an extensive breeding program (Geiger 1981, Burns & Johnson 1971).
Electromorphs were recorded as distance (mm) migrated from the
origin.

Electromorph frequencies (considered as allelic frequencies) were
used to calculate I, a statistic of genetic identity between taxa (Nei
1972), for all pairwise comparisons of populations. I-values were ana-
lyzed using the UPGMA method of cluster analysis (Ferguson 1980).

G-tests (Sokal & Rohlf 1981) were performed on genotype frequen-
cies in the populations represented by large samples (= 14 individuals)
to determine whether observed frequencies for each population were
consistent with Hardy-Weinberg equilibrium, and whether all popu-
lations can be considered to represent a single panmictic population.

RESULTS

Table 3 shows the electromorph frequencies for each population. Of
the 14 loci assayed, three are polymorphic: PGI, PGM and MPI.

Results of G-tests are displayed in Table 4. Most loci exhibit Hardy-
Weinberg equilibrium. However, weighted-average results show that
among the populations examined, zelicaon does not exhibit Hardy-
Weinberg equilibrium and cannot be considered a single, panmictic
population. Genotype frequencies are shown in Table 7.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

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VOLUME 43, NUMBER 3 Doe

TABLE 4. G-test of genotype frequencies in populations =14.

G-statistics for populations

Butts Rancho Castle
Locus (df) Hemet Orland Canyon Suisun Cordova Peak Average
BEC) 0 4.194* 6.605* 0 1.010 2.966 12. 776*
PGM (6) 24.349* 20.526* 4.502 L2oS2* 21.838* 6.846 48.90*
MPI (10) 3.224 3.918 1.582 1.976 10.606 10.334 30.926*

BS
© ; df = 4(n? — n) (Ferguson 1980).

*P < 0.05. G = 25 OBS |
P < 0.0. O EXP

Table 5 shows the I-values calculated from electromorph frequencies
for each pairwise comparison of populations. The average I-value is
0.980 + 0.139, indicating a very high level of genetic similarity.

To highlight differentiation without altering phenetic clustering of
populations, I-values were recalculated using only data for the three
polymorphic loci (Table 6). By excluding the background of mono-
morphic loci, a more useful graphic analysis can be generated. The
UPGMA dendrogram derived from these data is shown in Fig. 2.

DISCUSSION

The high identity values (Table 5) are consistent with similar studies
on other insects. Ayala et al. (1974b) reported I of 0.970 + 0.006 for
Drosophila willistoni populations sampled throughout Central and South
America, substantiating previous conclusions about their relatedness
based on reproductive relations. Brussard et al. (1985) surveyed genetic
identity findings comparing 14 insect taxa (including 3 Lepidoptera),
and reported I of 0.97 for local populations. It is likely that these values
are conservative. Mutants of crucial glycolytic or catabolic enzymes are
likely to be eliminated by selection (Bell 1976; Turner 1974; Zera et
al. 1985).

Reproductive relations in zelicaon are not clear. Breeding trials among

TABLE 5. Genetic identity I-value matrix using 14 loci (I = 0.980 + 0.139).

Wash- Butts Blue Rancho Castle

ington Hemet Orland Canyon Suisun Ridge Cordova Peak Auburn
Gazelle 0:97.18) 0:96) © (O:96%, -0:055:0:96" + 0:96. '0:98."—- "0:97: 0:98
Washington — O93), LOO £09800 «O98 201934" 0-98 10:99
Hemet — 0:99" ~ 0:99" *0:99" 0:99." 0.99" 0.98 °° 0:97
Orland — O99 0:99" “0:99: 0:99). | O98 O97
Butts Canyon — O99 7, 5 0:99.2 hf O99) + O:99) GOT
Suisun — O95" O99" O98.) 0:96
Blue Ridge _ 0.99 0.99 0.97
Rancho Cordova — 0.99 0.98
Castle Peak a 0.98

Auburn wth

224 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 6. JI-value matrix using only the three polymorphic (I = 0.867 + 0.100) loci.

Wash- Butts Blue Rancho Castle

ington Hemet Orland Canyon Suisun Ridge Cordova Peak Auburn
Gazelle 165.. .7138 ..707 -.656 .7238 676 (S07 2 oOmeeae
Washington — 890 .8380 .842 .795 .812 (849°) “S50 Neeo5o
Hemet — .993- .962 .960 (982 —/953) (985 aeTe4
Orland — 930 .974 .953 (9579 [9iGaase
Butts Canyon — 900 .984 901 ‘979 7808
Suisun — 952 .966 .964 .719
Blue Ridge — 949) Ganummers4
Rancho Cordova — iO Eee
Castle Peak — .808

Auburn —

multivoltines from northern and southern California and the Central
Valley show these populations to be intercompatible (Shapiro unpubl.).
Alternatively, Sims (1983) suggests that univoltines and multivoltines
are not fully intercompatible because of male-biased hybrid broods.
However, control (within-population) data are not available in adequate
numbers to validate this conclusion.

While I-values suggest that all zelicaon populations are conspecific,
the weighted-average G-tests show that zelicaon is, of course, not pan-
mictic over its entire range. Figure 2 suggests that populations can be
clustered on the basis of geographic proximity.

Gazelle (Shasta Valley) is most genetically dissimilar, and is probably
more geographically isolated as well. Washington and Auburn are Sier-
ran west slope univoltines. Orland, Suisun and Rancho Cordova are
Central Valley multivoltines. Castle Peak, Butts Canyon, Blue Ridge,
and Hemet represent univoltine and multivoltine populations in a mix-
ture of very diverse ecological contexts.

The germinal issue is to what degree populations are reproductively
isolated by host-plant selection and physical distance. Certainly, the
breeding trials and high I-values suggest that all zelicaon populations
are potentially intercompatible. However, voltinism may be a geneti-
cally heritable trait that divides zelicaon into low- and high-elevation
ecotypes (Clarke & Sheppard 1970).

Our data support Sims’s (1983) contention that zelicaon diapause
physiology and host-plant selection are highly plastic. The clustering
of the orange-feeding Orland population with other Central Valley
populations rather than with Hemet implies that the northern and
southern orange-feeders evolved separately. While fennel is abundant
in lowland areas, and is heavily used (Shapiro 1974a, 1974b), the use
of orange may allow zelicaon to increase its range despite the inferiority
of orange as a host plant (Masuda 1981).

VOLUME 43, NUMBER 3 225

TABLE7. Observed genotype frequencies for the three polymorphic loci in populations
where n = 14.

Populations
(Electromorph- Butts Rancho Castle
Genotype electromorph) Hemet Orland Canyon Suisun Cordova Peak
PGI 13-13 0) 0) 0.14 1) 0 0.13
13-7 0 0.05 0.07 0 0.13 0.11
7-7 1.00 0.95 0.80 1.00 0.89 0.86
PGM 32-32 0.21 0.11 0) 0.08 0) 0.06
32-26 0 0) 0 0.08 0.31 0.11
32-23 0.05 0.05 OWT. 0.08 0.19 0.11
32-20 0 8) 0.17 0.08 0 0.09
26-26 Oulgh 0.21 0 0.21 0.06 0.09
26-23 0.26 0.53 0.33 0.25 0.19 0.23
26-20 0) 0) 0) 0.42 0.13 0.09
23-23 0.21 0.05 0.17 0.42 0) 0.14
23-20 0.05 0 0.17 0 0.13 0.03
20-20 0.11 0.05 0) 0.13 0 0.06
MPI 44-44 1) 0) 0) 0 0 0
44-40 0) 0) 0) 0) 0) 0
44-37 0 0) 0 0 0 0
44-33 0) 0.10 0) 0.14 0) 0.03
44-28 0 0 0 0) 0) 0
40-40 0 0 0 0 0) 0)
40-37 1) 0) 0 0) 0) 0
40-33 0) 0.10 0 0.14 0 0.10
40-28 0 0 0 0 0) 0)
37-37 Omit 0.10 0.13 0) 0) 0.13
37-33 0.32 0.20 0.50 0.29 0.10 0.29
37-28 0.05 0 0 0 0 0.03
33-33 0.42 0.40 0.25 0.43 0.70 0.32
33-28 0.05 0.10 0.13 0) 0 0.67
28-28 0.05 0) 0) 0) 0.20 0.03

Butts Canyon (North Coast Range serpentine) and Castle Peak (Sier-
ran volcanic alpine) probably represent relict zelicaon populating rocky,
unforested environments with endemic host plants. Other Lepidoptera
are known to be similarly disjointly distributed between the Coast Range
serpentines and the alpine Sierra; Papilio indra, Pieris sisymbrii, and
Euchloe hyantis all occur obligately in these areas with few or no
intervening populations (Shapiro unpubl.). Clustering of Blue Ridge
(east of the Vaca Hills, the easternmost part of the Inner North Coast
Ranges in Yolo and Solano cos.) with these postulated relict populations
rather than with other multivoltines in the Central Valley is especially
interesting. Papilio zelicaon was not seen in the Vaca Hills during
summer in field studies initiated by Shapiro (unpubl.) in 1972. Males
were seen on the ridge-tops, but only in spring coinciding with such
behavior on Coast Range serpentines to the north. At this time, the site
had one patch of 10 fennel plants. In 1975, females were observed

226 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Blue Ridge
990

Pi Peak
Hemet

972
Butts Canyon

932 Orland

974

814 ene Suisun

Rancho Cordova
Washington
Oe 950

Auburn

Gazelle

0.800 0.900 1.000

ha V ANE E
Fic. 2. Phenogram of P. zelicaon populations (UPGMA; Ferguson 1980).

ovipositing on fennel. By 1978, fennel was spreading rapidly in dis-
turbed areas and zelicaon showed evidence of four generations in one
year. Presently, there are over 500 fennel plants along three miles of
road in this area, and it continues to spread. It has been presumed that
the multivoltine Vaca Hills zelicaon are upslope colonists from multi-
voltine Central Valley populations. Our study suggests, rather, that they
are at least partially downslope colonists from univoltine ridge-top (Coast
Range) populations. If this is the case, they have very rapidly evolved
multivoltinism, apparently as an adaptation to the spread of fennel.
This supports the plasticity of host plant- and diapause-“‘switching”’
proposed by Sims (1983) to explain the evolution of multivoltinism.
Certainly, zelicaon is physically capable of having colonized these can-
yons from the Coast Range. Shields (1967) demonstrated that zelicaon
is a hilltopping species; adult males and receptive females congregate
on summits to mate, thereby promoting gene flow among neighboring
populations. Shields determined that adults are capable of traveling
several km per day.

Studies by Ehrlich and Raven (1969) and Endler (1973) suggest that
populations undergoing sufficiently strong divergent selection will dif-
ferentiate despite the counter-effects of continuous gene flow. This has
been observed in wild Lepidoptera with populations showing differ-

VOLUME 43, NUMBER 3 DOT:

entiation in metrical traits as a result of differential selection, despite
close proximity and gene flow (Creed et al. 1959, Clarke & Sheppard
1962). If gene flow along the Coast Range ridge-tops has been contin-
uous, the Vaca Hills population has not only become multivoltine within
three years time, but has done so with constant influx of univoltines
from the Coast Range. Multivoltinism may be evolving through hy-
bridization, or through selection. Multivoltinism shortens generation
time and should, all other factors being equal, be selectively advanta-
geous.

Wright (1948a, 1943b) theorized that a continuously distributed species
exposed to different conditions of selection would differentiate if sub-
divided into partially isolated “islands’’ separated through inbreeding
or limited dispersal ability. Papilio zelicaon is certainly distributed
throughout habitats with different selection conditions and appears to
be sufficiently vagile to be essentially continuous in distribution through-
out major portions of its range. More finely focussed studies of nongly-
colytic enzymes and mark-release-recapture studies on movement would
help to determine the size and location of hilltopping regions and the
appropriateness of Wright’s “island’’ models to zelicaon.

Papilio zelicaon is composed of low- and high-elevation ecotypes
defined by host-plant preferences and diapause physiology. These traits
may be determined by a relatively small number of loci that are under
strong selection pressure and whose distribution is not reflected by
electrophoretically accessible glycolytic enzyme loci, which show great
genetic similarity among populations.

ACKNOWLEDGMENTS

We thank F. J. Ayala for use of laboratory facilities, Hansjiirg Geiger and P. Ward for
helping us interpret electrophoretic data, and John Emmel for the Hemet specimens; also
Douglas Engfer for developing software to compute I-values. This study forms part of
California Agricultural Experiment Station Project CA-D*-AZO-3593, “Host Switching
by the Anise Swallowtail,’ A. M. Shapiro, Principal Investigator.

LITERATURE CITED

AYALA, F. J., J. R. POWELL, M. L. TRACEy, C. A. MOURAO & S. PEREZ-SALAS. 1972.
Enzyme variability in the Drosophila willistoni group. IV. Genic variation in natural
populations of D. willistoni. Genetics 70:113-139.

1974a. Genetic variation in natural populations of five Drosophila species and

the hypothesis of the selective neutrality of protein polymorphisms. Genetics 77:343-

384.

1974b. Genetic differentiation during the speciation process in Drosophila.
Evolution 28:576-592.

BELL, M. A. 1976. Evolution of phenotypic diversity in Gasterosteus aculeatus super-
species on the Pacific Coast of North America. Syst. Zool. 25:211-227.

BRUSSARD, P. F., P. R. EHRLICH, D. D. MurpHy, B. A. WILCOX & J. WRIGHT. 1985.
Genetic distances and the taxonomy of checkerspot butterflies (Nymphalidae: Nym-
phalinae). J. Kansas Entomol. Soc. 58:403-412.

228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

BURNS, J. M. & F. M. JOHNSON. 1971. Esterase polymorphism in the butterfly Hemiargus
isola: Stability in a variable environment. Proc. Nat. Acad. Sci. 68:34-37.

CLARKE, C. A. & P. M. SHEPPARD. 1962. Disruptive selection and its effect on a metrical
character in the butterfly Papilio dardanus. Evolution 16:214—226.

1970. Is Papilio gothica a good species. J. Lepid. Soc. 24:230-233.

CoOOLIDGE, K. R. 1910. A California orange dog. Pomona College J. Entomol. 2:533-

084.

CREED, E. R., W. H. DOWDESWELL, E. B. FORD & K. G. MCWHIRTER. 1959. Evolu-
tionary studies on Maniola jurtina: The English mainland, 1956-57. Heredity 13:
363-391.

DETHIER, V.G. 1941. Chemical factors determining the choice of food plants by Papilio
larvae. Am. Nat. 75:61-78.

EHRLICH, P. R. & P. H. RAVEN. 1969. Differentiation of populations. Science 165:1228-
1232.

ENDLER, J. A. 1973. Gene flow and population differentiation. Science 179:243-250.

FERGUSON, A. 1980. Biochemical systematics and evolution. Blackie, Glasgow. 194 pp.

GEIGER, H. J. 1981. Enzyme electrophoretic studies on the genetic relationships of
pierid butterflies (Lepidoptera: Pieridae) I. European taxa. J. Res. Lepid. 19:181-
195.

HUTCHINSON, W. H. 1969. California: Two centuries of man, land and growth in the
Golden State. American West Publishing Co., Palo Alto, California. 351 pp.

Masupa, K. K. 1981. The effects of two host plants, Foeniculum vulgare Mill. and
Citrus sinensis Osbeck, on adult oviposition, behavior and larval and pupal biology
of the anise swallowtail butterfly (Papilio zelicaon, Lucas) (Lepidoptera: Papilioni-
dae). M.S. Thesis, University of California, Davis. 82 pp.

Munz, P. A. 1970. A California flora. Univ. of California Press, Berkeley. 1681 pp.

NEI, M. 1972. Genetic distance between populations. Am. Nat. 106:283-292.

Opitz, K. W. & R. G. PLATT. 1969. Citrus growing in California. Calif. Agric. Exper.
Station Service. 59 pp.

SHAPIRO, A. M. 1974a. The butterfly fauna of the Sacramento Valley, California. J. Res.
Lepid. 13:73—-82, 115-122, 137-148.

1974b. Butterflies of the Suisun Marsh, California. J. Res. Lepid. 13:191—206.

& K. K. MasupDa. 1980. An opportunistic new citrus pest. Cal. Agric. 34:4—5.

SHIELDS, O. 1967. Hilltopping: An ecological study of summit congregation behavior
of butterflies on a southern California hill. J. Res. Lepid. 6:69-178.

Sims, S. R. 1979. The reproductive biology and diapause dynamics of Papilio zelicaon
(Lepidoptera: Papilionidae). Ph.D. Dissertation, University of California, Davis. 87
pp. Diss. Abstr. Order No. DA7815518.

1983. Inheritance of diapause induction and intensity in Papilio zelicaon. He-
redity 51:495-500.

SOKAL, R. R. & F. J. ROHLF. 1981. Biometry: The principles and practice of statistics
in biological research. 2nd ed. Freeman, San Francisco. 859 pp.

TURNER, B. J. 1974. Genetic divergence of Death Valley pupfish species: Biochemical
versus morphological evidence. Evolution 28:281—294.

TURESSON, G. 1922. The genetic response of the plant species to the habitat. Hereditas
3:211-350.

TyLer, H. A. 1975. The swallowtail butterflies of North America. Naturegraph Pub.,
Healdsburg, California. 192 pp.

WRIGHT, S. 1943a. Isolation by distance. Genetics 28:114-138.

1943b. An analysis of local variability of flower color in Linanthus parryae.
Genetics 28:139-156.

ZERA, A. J., R. K. KOEHN & J. G. HALL. 1985. Allozymes and biochemical adaptation,
pp. 633-674. In Kerkut, G. A. & L. I. Gilbert (eds.), Comprehensive insect physiology,
biochemistry and pharmacology, Vol. 10. Pergamon Press, Elmsford, New York.

Received for publication 5 July 1988; accepted 26 May 1989.

Journal of the Lepidopterists’ Society
43(3), 1989, 229-238

A NEW SUBSPECIES OF COENONYMPHA TULLIA (MULLER)
(NYMPHALIDAE: SATYRINAE) CONFINED TO THE
COASTAL DUNES OF NORTHERN CALIFORNIA

ADAM H. PORTER
Department of Zoology, University of California, Davis, California 95616

AND

STERLING O. MATTOON
2109 Holly Ave., Chico, California 95926

ABSTRACT. Coenonympha tullia yontocket is described from a single known pop-
ulation confined to the coastal sand dunes north of Crescent City, Del Norte County,
California. It is most similar in phenotype to C. tullia eunomia Dornfield, but may be
distinguished by wing characters. A population of C. tullia eryngii Hy. Edwards occurs
ten kilometers away; these two populations show no clear signs of reciprocal introgression
in wing characters. Electrophoretic analysis indicates that yontocket retains the high
genetic variability characteristic of other tullia-group taxa, but no diagnostic alleles were
found. The high genetic variability is most likely maintained by gene flow from eryngii.
Coenonympha tullia subspecies yontocket, eunomia, eryngii, and california Westwood
are genetically very similar (Nei’s unbiased genetic distance <0.035). The data justify
the placement of yontocket as a subspecies rather than a species. This subspecies is a
likely candidate for listing as threatened in California; collectors and developers are urged
to protect this population from extinction.

Additional key words: Coenonympha tullia yontocket, taxonomy, electrophoresis, ring-
lets, threatened species.

Investigation of the coastal dunes of northern California has turned
up a unique population of the widespread Ringlet butterfly, Coeno-
nympha tullia (Miller). This population occurs in the vicinity of Cres-
cent City, Del Norte Co., and has an ochre ground color; it is wholly
contained within the range of C. tullia eryngii Hy. Edwards, a wide-
spread subspecies with a whitish ground color. It is quite similar phe-
notypically to C. tullia eunomia described by Dornfield (1967), whose
nearest known population is 250 km away in the Umpqua River drain-
age in southeastern Oregon (Porter & Geiger 1988). This new population
flies in the fog belt, and shows the heavy melanization of the wings
and body characteristic of butterflies from this type of environment
(Hovanitz 1941, McCorkle & Hammond 1988).

Herein, we provide a description of this population as a new sub-
species, and justify our taxonomic placement with genetic evidence
from electrophoretic analysis. We did not examine genitalic or larval
characters: Davenport (1941) indicated that all tullia-group taxa were
indistinguishable genitalically despite high levels of intrataxon vari-
ability, and description of the immature stages would be of little taxo-
nomic use given our small series and the lack of comparative material.

230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

VOLUME 43, NUMBER 3 231

Coenonympha tullia yontocket, new subspecies

(Figs. 1-3)

Description. Holotype (Fig. 1): male; dorsal ground color dull ochraceous; medium to
light gray scaling along the costal and distal forewing margins, extending proximally
along the veins. Dorsal hindwing with gray scaling along distal margins, stronger in anal
area. Eyespots absent; ventral whitish markings barely visible from above. Both dorsal
surfaces strongly melanized subbasally. Ventral forewing ground color deep ochraceous,
almost orange; medium band whitish, extending from R veins to Cu,; ground color fades
to whitish, then greenish gray in costal and apical regions, becoming strongly suffused
with melanized scales; eyespots absent. Ventral hindwing ground color brownish ochre
in discal region, fading to greenish gray beyond the median markings. Median band
whitish, well marked; absent only between Cu, and Cu,. Whitish basal patch present at
radial vein. Eyespots absent. Darkened, single marginal line on all wing surfaces, well
expressed ventrally. Head, thorax, and ventral hindwing bases covered with long hairs
matching ventral hindwing ground color.

Morphological variation (Figs. 2, 3). Forewing length, males: 14-18 mm (n = 65);
females: 15-19 mm (n = 10). Spring brood averages slightly larger (males: x = 16.3 mm;
n = 52) than fall brood (males: x = 14.7 mm; n = 17). Spring brood: gray scaling dorsally
along the distal margins of both wings may be almost absent, but may extend proximally
in extreme individuals (n = 2) so that the outer third of the wing is pale gray. Ventral
forewing median band whitish; extends from R veins to Cu, or Cu,. Single ventral forewing
eyespot absent in most individuals, but may be up to 1 mm diameter, unpupilled ochra-
ceous or yellow, or yellow pupilled with black. Ventral hindwing: ground color sometimes
obliterated by melanized scaling in discal area; wholly brownish or wholly greenish in
some individuals. Median band sometimes weakly expressed between Cu, and Cu,, rarely
absent below M, (Fig. 3b). Whitish basal patches often present, connecting to median
markings via the costa in extreme individuals (n = 2) (Fig. 3a). Eyespots absent in almost
all individuals; rarely up to three, yellow or yellow with black pupils, most likely between
Cu, and Cu,. Marginal line often double. Females (Fig. 2) tend towards less melanization,
broader wings. Fall brood: markings similar to spring brood, more animals with brownish
rather than greenish ventral ground color, and more likely expression of ventral basal
patches.

Diagnosis. Separable immediately from nearby populations of eryngii by the ochra-
ceous ground color. Separable from eunomia ventrally by stronger expression of the
medial markings; from ewnomia and ampelos ventrally by the frequent occurrence of
basal patches, and dorsally by gray scaling along the veins and outer wing edges.

Distribution. Known only from Del Norte Co., California, among the coastal sand
dunes north of Crescent City, beginning at the north shore of Lake Earl and extending
north 7.5 km to the south bank of the Smith River (Fig. 4). This area is hereby designated
as the type locality. Seemingly suitable habitat between Lake Earl and Point St. George
may also be populated by yontocket. Not present in abutting disturbed habitats to the
east (mostly cow pastures), or at the dunes north of Arcata Bay in Humboldt County,
California. Replaced by eryngii 10 km to the east on exposed serpentine hilltops.

Material examined. Holotype: Male, California, Del Norte Co., 4 km W Fort Dick, 8-
IX-1979, leg. S. O. & E. Mattoon. Deposited in the Bohart Museum at the University of
California, Davis. Paratypes: California, Del Norte Co., 4 km W Fort Dick, end of Kellogg
Rd. S to north shore of Lake Earl, 2-VI-1979 (15 males), 30-VI-1979 (4 males) & 8-IX-

—_—

Fics. 1-3. 1, Coenonympha tullia yontocket holotype male; (a) dorsal and (b) ventral
surfaces. 2, Coenonympha tullia yontocket paratype female; (a) dorsal and (b) ventral
surfaces. Unlike this specimen, many females do show basal ventral hindwing patches.
3, Coenonympha tullia yontocket ventral surfaces, showing the extremes of expression
of maculation. The reduced pattern of (b) is characteristic of C. tullia eunomia popu-
lations.

232 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fic. 4. Map showing localities sampled in northwestern California and southwestern
Oregon. Note the close proximity of the eryngii population to the type locality of yontocket
(inset). Neither subspecies occurs in the intervening cow pastures.

1979 (16 males, 2 females), S. O. & E. Mattoon, leg. These will be deposited in the Bohart
Museum, the California Department of Food and Agriculture, the California Academy
of Sciences, the Los Angeles County Museum, the Allyn Museum, and the National
Museum of Natural History, Smithsonian Institution. Additional material: wing vouchers
from specimens used for electrophoretic analysis (29 males, 8 females), collected from
the Yontocket Archeological Site at the north end of the population range.

Biology. Flight periods May—July and September-October. Habitat: elev. 2 m; in grassy
areas among dunes with coniferous lee slopes and grassy exposed slopes, and among dunes
on slightly elevated ground around seasonally marshy sphagnum bogs which fill during
the rainy season. 2 females oviposited (5 observations) on dry grass stems (mixed species
composition) approx. 2-5 cm above soil in areas free from flooding. Larvae and pupa (n
= 2 larvae; 1 pupated) are apparently not different from those of eryngii (n = 8 larvae;
2 pupated). Larval host(s) presently unknown.

Etymology. Coenonympha tullia subspecies are often given American Indian names.
This population is dedicated to the memory of the Yontocket tribe, which once had
seasonal settlements in these dunes.

In deciding to name this population, we considered two points: (i) is
it sufficiently distinct from C. t. eunomia to warrant taxonomic rec-
ognition?, and (ii) given that an apparently permanent population of

VOLUME 43, NUMBER 3 230

C. t. eryngii occurs in serpentine grassland habitat on a hilltop 10 km
to the east and within sight of the yontocket population (Fig. 4), should
yontocket be given species status? To address these questions, we per-
formed starch gel electrophoresis to provide insights into the genetic
relationships among the yontocket population, the nearby eryngii pop-
ulation, a previously studied eunomia population from Riddle, Oregon,
and a C. t. california Westwood population from near Myers Flat,
Humboldt Co., California. Each of these populations comes from rel-
atively isolated areas of grassland habitat, providing a control on the
potential for genetic differentiation resulting solely from variation in
local population structure. Previous work has established that eryngii,
california, and eunomia are members of a single polytypic species
(Porter & Geiger 1988).

MATERIALS AND METHODS

Butterflies were netted and temporarily stored on wet ice, hand-
carried or mailed back to Davis, then frozen alive at —80°C for storage
until analysis. Electrophoretic analysis followed the protocol of Ayala
et al. (1972) and Geiger and Shapiro (1986), with one modification:
rather than using sponge wicks to complete the circuit between the
electrode buffer solutions and the gels, gel molds were used which
allowed the ends of the gels to contact the electrode buffer directly.
We scored 13 loci: adenylate kinase (AK-1), aldolase (ALDO), fumarase
(FUM), glutamic-oxaloacetic transaminase (GOT-1, GOT-2), glycer-
aldehyde-3-phosphate dehydrogenase (GAPDH), a-glycerophosphate
dehydrogenase (a-GPDH), isocitrate dehydrogenase (IDH-1), malate
dehydrogenase (MDH-1, MDH-2), phosphoglucomutase (PGM), phos-
phoglucose isomerase (PGI), and superoxide dismutase (SOD-1). Zymo-
grams were scored as described in Porter and Geiger (1988), and data
were analyzed using the computer program BIOSYS-1 (Swofford &
Selander 1981).

RESULTS

Allelic frequencies for the yontocket, eryngii, and california popu-
lations are given in Table 1; allelic frequencies at these loci were pre-
viously given for the eunomia population in Porter and Geiger (1988).
All populations show high levels of genetic variability. characteristic of
Coenonympha tullia populations elsewhere (Table 2; Porter & Geiger
1988); this is an indication that the yontocket population has not been
through a significant genetic bottleneck in its recent past. Table 3 shows
genetic relationships among these populations using Nei’s unbiased min-
imum distance and identity measures (Nei 1978). The phenogram con-
structed based on these values using UPGMA (Fig. 5; see Sneath &

234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Nei's (1978) unbiased genetic identity
097 pga.» “gles 1.00

california

eryngii
yontocket

eunomia

0.03 0.02 0.01 0.00

Nei's (1978) unbiased genetic distance

Fic. 5. Phenogram of genetic relationships constructed using the UPGMA algorithm
on data from Table 3. Yontocket does not cluster with eunomia, but all populations are
genetically very similar.

Sokal 1973 for methodological details) does not group yontocket with
eunomia, despite their general similarity in wing characteristics. These
distance-identity values indicate a very low level of genetic differen-
tiation overall, corresponding to subspecies-level differentiation in most
taxa (Thorpe 1983), including butterflies (AHP unpubl. data; H. J.
Geiger, pers. comm.).

DISCUSSION

Neither the yontocket nor eryngii population in Fig. 4 has colonized
intervening, non-native grassland presently used for grazing. There is
also no clear evidence of introgression in wing pattern traits in the
animals we sampled. The pale gray along the veins and wing edges
dorsally in yontocket may well be evidence of such introgression, but
populations from the Pit River drainage in eastern California, where

VOLUME 43, NUMBER 3 Zoe

TABLE 1. Allelic frequencies of Coenonympha tullia-group taxa. Population localities
and locus abbreviations given in the text.

Taxon Taxon
Locus and EE Ee eee TEU feara a eek red ca EL a
allele california} eryngii? _ yontocket® allele california! _eryngii? yontocket®
AK-1 MDH-1
76 0.056 91 0.028 0.014
86 0.023 93 0.028
90 0.361 0.275 0.500 100 0.861 0.986 0.932
100 0.556 0.675 0.432 105 0.028
102 0.028 110 0.056 0.068
110 0.050 0.045 MDH-2
ALDO 96 0.028
100 1.000 1.000 1.000 100 0.972 0.917 0.932
FUM 105 0.069 0.054
100 1.000 1.000 1.000 110 0.014 0.014
GAPDH PGI
100 1.000 1.000 1.000 81 0.014
GOT-1 88 0.028 0.014
89 0.028 0.014 94 0.028
91 0.194 OxIgeh 0.135 97 0.083
94 0.041 100 0.306 0.333 0.284
100 0.778 0.806 0.730 103 0.056 0.028
102 0.056 107 0.306 0.389 0.662
108 0.014 0.054 105 0.042
110 0.041 Hila 0.028 0.069
GOT-2 114 0.194 0.056 0.027
100 0.944 0.944 0.973 Tey 0.014
1, 0.056 0.056 0.027 AL 0.042
a-GPDH PGM
15 0.028 90 0.014
90 0.042 94 0.028 0.042
96 0.014 SG 0.028 0.027
100 0.972 0.931 1.000 100 0.361 0.486 0.662
110 0.014 106 0.528 0.389 0.311
IDH-] 110 0.083 0.028
90 0.028 0.014 Jy 0.014
a2 0.028 0.028 SOD-1
100 0.417 0.583 0.730 89 0.028
103 0.194 0.208 0.108 100 0.944 1.000 1.000
106 0.306 0.125 0.162 120 0.028
Ed 0.028 0.042

In = 18.
2n = 36, except at AK-1, where n = 20.
3n = 87, except at AK-1, where n = 22.

california and eryngii (white ground color), and ampelos (ochre ground
color) intergrade, produce many specimens of wholly intermediate
background coloration (Porter & Geiger 1988). These observations sug-
gest that differentiation is maintained by behaviors related to habitat
and(or) host-plant selection—but not necessarily by reproductive bar-
riers.

Phenograms based on genetic distance-identity indices are often used

236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 2. Genetic variability statistics for the three populations given in Table 1.
Mean number of alleles per locus = x,y... Percent of loci polymorphic = P. Observed
heterozygosity = H,,,. Heterozygosity calculated from Hardy-Weinberg proportions =
H.,,. Standard errors in parentheses.

Population Tee P H.. HS
california 8.2 (0.5) 84.6 0.303 (0.090) 0.283 (0.080)
eryngii 8.5 (0.7) 69.2 0.240 (0.078) 0.246 (0.076)
yontocket 2.5 (0.4) 61.5 0.199 (0.070) 0.210 (0.064)

to approximate phylogenetic relationships between species, but these
measures can only reflect overall genetic differentiation within a species.
Within a species, the degree of differentiation expressed among pop-
ulations reflects a balance between the forces of natural selection, ge-
netic drift, mutation, and gene flow acting at each locus. The fact that
yontocket is more similar to california and eryngii than to eunomia
implies that gene flow between yontocket and eunomia is interrupted:
it seems unreasonable to consider them consubspecific. This interpre-
tation is also in agreement with the disjunct distribution of these taxa.

The high level of variability in yontocket enzyme characters also
requires explanation. The yontocket population probably has an effec-
tive breeding population of moderate size, and is likely to be affected
somewhat strongly by genetic drift. If yontocket is fully reproductively
isolated from eryngii, then exceedingly strong selection on these en-
zymes is required to maintain such high numbers of alleles; on the
other hand, infrequent influxes of eryngii phenotypes could easily main-
tain this variability. Given that there is evidence of some gene flow
between ewnomia and eryngii in southeastern Oregon (Porter & Geiger
1988) (the subspecies separated by the greatest geographic distances in
the phenogram of Fig. 5); and that yontocket is of intermediate simi-
larity, we think it is wise to place yontocket as a subspecies of tullia
unless subsequent studies on reproductive biology demonstrate intrinsic
barriers to gene flow. The level of current gene flow between these two
adjacent tullia-group populations, based on their present constellations
of allelic frequencies, indicates that these populations exchange between
four and five breeding individuals every generation on average (Porter,
in prep.), further supporting the taxonomic placement proposed here.

The evolutionary origins of the diagnostic yontocket traits are ex-
plainable by a number of plausible scenarios (many non-diagnostic traits
may be attributable to gene flow from eryngii). The most likely scenario
is that these traits arose from eunomia or even columbiana Mc-
Dunnough, which may have had more southerly distributions during
the last glacial stages. A population of Polites mardon (Edwards) (Hes-
periidae) also occurs in Del Norte Co., California, disjunct from nearest

VOLUME 438, NUMBER 3 231

TABLE 8. Nei’s (1978) unbiased genetic identity (above diagonal) and distance (below
diagonal) values between population pairs. Populations given in the text.

Taxon

Taxon california eryngii eunomia yontocket
california 0.997 0.976 0.982
eryngii 0.003 0.974 0.990
eunomia 0.030 0.035 0.979
yontocket 0.018 0.010 0.026

known populations in southwestern Washington State (Scott 1986; T.
C. Emmel, J. F. Emmel & S. O. Mattoon, in prep.). However, this alone
does not explain the high incidence of the basal ventral hindwing
patches, a characteristic of populations in the Great Basin and Rocky
Mountains. Whether the basal ventral patch is adaptive, ancestral in
North America, or exists in yontocket as a result of past gene flow from
the east, is unknown. Functionally unrelated traits can clearly have
independent geographic ranges within a species. Thus, the conclusions
we draw concerning the historical biogeography of tullia-group traits
depend largely on whether or not biological species boundaries exist
within the complex (and if they do exist, where they are).

We think it is particularly important to recognize the threat of ex-
tinction to C. t. yontocket caused by habitat destruction. The southern
end of the population distribution occurs in habitat patches within an
abandoned gridwork of streets originally intended as a housing devel-
opment. With the spread of development by the tourist industry around
the recently formed Redwood National Park, the yontocket habitat is
likely to become attractive to developers of beachfront property—both
for private and public use. Given the failure of yontocket to invade
adjacent cow pastures, a habitat used by tullia subspecies elsewhere in
western North America, it is likely that such development will have
severe impact on this population. We urge lepidopterists to refrain from
collecting in this fragile ecosystem, and to provide support for groups
dedicated to the preservation of this and other threatened taxa along
the Pacific Coast.

ACKNOWLEDGMENTS

AHP was supported during this study by a grant entitled Cross-disciplinary Studies
of Population Structure to the Institute of Ecology at UC Davis from the Alfred P. Sloan
Foundation. A Graduate Research Award from the University of California—Davis to
AHP partially covered travel to the study area. Rick Harris took the photographs; Cali-
fornia Agricultural Experiment Station project CA-D*-AZO-3994-H, Climatic Range
Limitations of Phytophagous Insects, to Art Shapiro, funded the electrophoresis; and
Brad Shaffer kindly provided laboratory facilities. Chuck Hageman, Ken Hansen, and
Eileen Mattoon accompanied SOM when the first series of yontocket was collected.

238 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

LITERATURE CITED

AYALA, F. J., J. R. POWELL, M. L. TRACEy, C. A. MouRAO & S. PEREZ-SALAS. 1972.
Enzyme variability in the Drosophila willistoni group. IV. Genetic variation in natural
populations of Drosophila willistoni. Genetics 70:113-139.

DAVENPORT, D. 1941. The butterflies of the Satyrid genus Coenonympha. Bull. Mus.
Comp. Zool. 87:215-348.

DORNFIELD, E. J. 1967. On the yellow forms of Coenonympha tullia (Satyridae) in
Oregon. J. Lepid. Soc. 21:1-7.

GEIGER, H. J. & A. M. SHAPIRO. 1986. Electrophoretic evidence for speciation within
the nominal species Anthocharis sara Lucas (Pieridae). J. Res. Lepid 25:15-24.
Hovanitz, W. 1941. Parallel ecogenotypical color variation in butterflies. Ecology 22:

259-284.

McCorRKLE, D. V. & P. C. HAMMOND. 1988. Biology of Speyeria zerene hippolyta
(Nymphalidae) in a marine-modified environment. J. Lepid. Soc. 42:184-195.

NEI, M. 1978. Estimation of average heterozygosity and genetic distance from a small
number of individuals. Genetics 89:583-590.

PorTER, A. H. & H. J. GEIGER. 1988. Genetic and phenotypic population structure of
the Coenonympha tullia complex (Lepidoptera: Nymphalidae: Satyrinae) in Cali-
fornia: No evidence for species boundaries. Can. J. Zool. 66:2751-2765.

ScoTT, J. A. 1986. The butterflies of North America. Stanford University Press, Stanford,
California. 583 pp.

SNEATH, P. H. A. & R. R. SOKAL. 1973. Numerical taxonomy. W. B. Freeman, San
Francisco. 573 pp.

SWOFFORD, D. L. & R. B. SELANDER. 1981. A computer program for the analysis of
allelic variation in genetics. J. Hered. 72:281-283.

THORPE, J. P. 1983. Enzyme variation, genetic distance and evolutionary divergence
in relation to levels of taxonomic separation, pp. 131-152. In Oxford, G. S. & D.
Rollinson (eds.), Protein polymorphisms: Adaptive and taxonomic significance. Ac-
ademic Press, New York. 405 pp.

Received for publication 17 December 1988; accepted 25 April 1989.

Journal of the Lepidopterists’ Society
43(3), 1989, 239-243

A PROCEDURE FOR EXAMINING THE GENITALIC
MUSCULATURE OF LEPIDOPTERA

JOHN A. DE BENEDICTIS' AND JERRY A. POWELL

Department of Entomological Sciences, University of California,
Berkeley, California 94720

ABSTRACT. The functional morphology of the genitalia (characteristics of the scler-
otized parts and the presence and position of the associated musculature) has been the
basis of recent phylogenies of Tortricidae and several other groups of Lepidoptera. Ex-
amination of this musculature can be difficult. Procedures for fixing muscles and preserving
specimens for future preparation, for dissecting, cleaning and staining the genitalia, and
for treating the preparation for viewing are presented. Using these methods, one can
stain muscles selectively, minimize handling during cleaning to reduce the potential of
physical damage, and view the musculature through transparent sclerotized parts.

Additional key words: functional anatomy, dissection, staining, male genitalia.

The sclerotized parts of the genitalia often are the best or only means
of identifying species of Lepidoptera and can provide important char-
acters for determining taxonomic relationships. The associated func-
tional musculature has been used as another source of data on phylo-
genetic relationships, particularly of higher taxa in Lepidoptera in recent
decades.

Forbes (1939) described differences in the male genitalic musculature
among six species in five families of Lepidoptera that he examined and
two other species illustrated by Snodgrass (1985:fig. 308). Utilization of
these characters in taxonomic studies of Lepidoptera was proposed by
Stekol’nikov (1965), who examined males of five and females of two
species of Noctuidae. Stekol’nikov (1967a, 1967b) constructed a phy-
logeny of butterflies based on the functional morphology of the genitalia
and discussed evolutionary trends in the genitalia of primitive Lepi-
doptera. An ensuing series of papers by Kuznetsov and Stekol’nikov
(1981, 1984, 1985) proposed the higher classifications of various groups
of Lepidoptera based largely upon genitalic musculature of the males,
and others (e.g., Razowski 1981) have employed these characters in
studies of Tortricidae and other Lepidoptera.

Workers wishing to investigate these conclusions further and those
intending to use their system to assign troublesome genera to tribe or
subfamily may have difficulty preparing specimens for examining mus-
culature, as there is little published information on the methodology.
The following procedure, derived by trial and error and from the
suggestions of colleagues, may greatly simplify this task. We worked
with Tortricidae, and the techniques should be applicable to all Lep-

' Present address: Department of Entomology, University of California, Davis, California 95616.

240 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

idoptera although we did not attempt dissection of smaller moths such
as leaf miners. With this procedure, much of the fatty tissue is dissolved,
so handling time during cleaning is minimized. The sclerotized parts
turn transparent which enables one to view the internal muscles and
their connections.

Preparation of Lepidopterous Genitalic Musculature

The best preparations of genitalic musculature result from proper
preservation of specimens, from careful dissection, cleaning, and stain-
ing, and from treating the stained, dissected genitalia for viewing.
Sectioning with a microtome may be necessary for minute Lepidoptera,
but the following procedure was effective on a species of Diactenis,
one of the smallest tortricid moths. Although it was not tried, the
procedure should be satisfactory for examination of female musculature
as well.

Some internal muscles and their attachment points are seen more
easily when the valvae are spread. The valvae of some specimens will
spread automatically when killed or immersed live in the preservative,
or they can be forced open by squeezing the pregenital segments prior
to preservation in fluid. However, using our procedure, the sclerotized
parts of the genitalia become transparent, and attachment points can
be determined. Consequently, most preparations were of specimens
with the valvae closed, because it was easier to position them for dorsal,
ventral, or lateral viewing.

Preservation of the specimen: Musculature of dried, pinned speci-
mens can be observed, but better preparations are obtained from spec-
imens preserved in fluid soon after capture. Specimens placed directly
into 70% alcohol (we used either isopropanol or ethanol) are usually
satisfactory, but the muscle tissue may deteriorate slowly, and older
specimens may be unusable.

We obtained better results by first immersing the specimen in Kahle’s
fluid for 12-24 hours to fix the muscles. Peterson (1964:67-68) and
Borror et al. (1976:736) give different, but equally effective recipes for
Kahle’s fluid.

The moth can be immersed directly into Kahle’s fluid; then a few
drops of 70% alcohol should be added to enhance wetting. Alternatively,
the specimen can be dipped into 70% alcohol or cellusolve (ethylene
glycol monoethy] ether) for a few seconds until soaked, then transferred
to Kahle’s fluid. When it is impractical to preserve specimens imme-
diately, muscles of those killed in dry cyanide vials can be similarly
fixed if treated soon after capture.

After fixing the muscles, the specimen can be dissected immediately
or retained in 70% alcohol. Specimens transferred from Kahle’s fluid

VOLUME 43, NUMBER 3 24)

to 70% alcohol were in excellent condition more than five years after
preservation. Prolonged immersion in Kahle’s fluid or preservation in
95% or absolute alcohol makes the muscles brittle and gives them a
greater tendency to detach from the sclerotized structures.

A few preparations were made from dried, pinned specimens by
removing the abdomen and soaking it in warm water until the viscera
softened. The muscles of previously dried specimens are inelastic, some-
times shrunken and contorted, and easily detached, and they do not
stain as well as those of specimens preserved in fluid, so results vary.

Dissection and staining: Remove the abdomen from the specimen
and place it in 70% ethanol. Shallowly insert the tip of each of two pair
of No. 5 jeweler’s forceps, one on each side of the pleuron, into the
intersegmental membrane anterior to the last or next to last visible
pregenital abdominal segment, and carefully peel away the integument.
Continue to remove the integument of the pregenital segments from
the genitalia until none can be removed without risking damage to the
genitalic muscles.

Large agglomerations of fat will inhibit staining, but at this time
attempt to remove only the larger looser globules atop the base of the
aedeagus to avoid damaging the muscles.

Place the excised genitalia in a drop or two undiluted van Giesen’s
muscle stain (1 part of 2-3% acid fuchsin and 9 parts of saturated picric
acid) on a stain plate for 3-10 minutes or until stain begins to penetrate
but does not completely stain the internal muscles of the tegumen. Next,
soak the preparation in 70% alcohol for several hours to allow stain to
penetrate internal muscles while washing away excess.

After soaking in alcohol, muscles of a properly stained preparation
will be red throughout, fat will be paler, and the sclerotized parts should
be mostly unstained. Some staining of the sclerotized parts is unavoid-
able, especially the aedeagus, some of the more membranous parts, and
around the margins of other structures.

If too much stain was removed, the exposed muscles on the aedeagus
will have lost much of their color while the internal muscles remain
red. Place such preparations briefly into a drop of stain then wash off
the excess in 70% alcohol.

If understained, stain will not have penetrated the internal muscles,
and the staining-soaking step should be repeated. Sometimes, partic-
ularly in previously dried specimens, some muscles will not stain well,
and additional attempts will only stain the sclerotized structures.

Subsequent steps will remove some additional stain, so slight over-
staining is acceptable. If the sclerotized portions are stained excessively,
additional soaking in clean 70% alcohol may eventually remove the
excess. If still overstained, immerse the preparation in hydrogen per-

242 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

oxide solution, then restain if necessary. Final cleaning follows the next
step.

Transfer the preparation to cellusolve and soak for 10 hours or longer.
The solvent will dehydrate the preparation and dissolve much of the
fat and cause much of the remainder to agglomerate into easily re-
movable globules. The dehydrated remnants of pregenital segments
and unwanted extrinsic muscles can be abraded from the genitalia easily
with forceps or fine probes, and most of the remaining fat globules can
be teased free of the preparation with fine probes. Smaller traces of fat
will dissolve or turn transparent in the next step, so it is not necessary
to risk damage by trying to remove small bits trapped between muscles.

Preparation for viewing: Place the cleaned preparation in methyl
salicylate (oil of wintergreen) for viewing. From about 30 minutes to
about five days following immersion, the sclerotized structures will be
sufficiently transparent to view the internal muscles, yet will retain
enough pigmentation to determine the attachment points of the muscles.

With prolonged immersion in methy] salicylate, the preparation be-
comes increasingly brittle and more easily damaged during handling,
and the sclerotized parts slowly darken, presumably due to infusion of
the stain. Partial darkening may help identify sclerotized structures,
but after about five days, some musculature may be difficult to see
through the sclerotized parts. A darkened preparation can be bleached
in hydrogen peroxide after being washed of methyl] salicylate in cel-
lusolve then 70% alcohol. It can be restained and treated for viewing
as before, but it will be more brittle and somewhat inferior overall.

We positioned preparations for viewing with glass chips in the depres-
sion of a culture slide filled with methyl] salicylate. A camera lucida
attached to a binocular dissecting microscope facilitated sketching the
preparation. When necessary, a compound microscope was used to
determine musculature attachment points accurately.

After examination, the preparation was washed free of methy] salic-
ylate in cellusolve, then bathed in 70% alcohol. It was preserved in 70%
alcohol with the remainder of the specimen. In the alcohol, the stain
slowly leaches from the preparation. It can be restained and cleared
for re-examination, but will be of lesser quality than after the first
treatment.

ACKNOWLEDGMENTS

Major problems in staining the muscles and clearing the sclerotized parts were overcome
by using techniques developed by Dr. Thomas F. Hlavik, San Francisco, California, which
were suggested to us by Dr. John T. Doyen, Department of Entomological Sciences,
University of California, Berkeley. Our procedure was refined during a study of the
musculature of tortricid moths funded by National Science Foundation grant BSR-8411459.

VOLUME 438, NUMBER 3 243

LITERATURE CITED

Borror, D. J., D. M. DELONG & C. A. TRIPLEHORN. 1976. An introduction to the
study of insects. 4th ed. Holt, Rinehart and Winston, New York. x + 852 pp.

FORBES, W. T. M. 1939. The muscles of the lepidopterous male genitalia. Ann. Entomol.
Soc. Am. 32:1-10.

KUZNETSOV, V. I. & A. A. STEKOL’NIKOV. 1981. Functional morphology of the genitalia
of some Asiatic moths (Lepidoptera, Papilionomorpha: Epiplemidae, Uraniidae, Dre-
panidae, Callidulidae) and their systematic position. Proc. Zool. Inst. Acad. Sci. USSR
103:19-48.

1984. Classification and phylogenetic relations of the families and superfamilies

of the gelechioid moths (Lepidoptera, Papilionomorpha: Copromorphoidea, Elach-

istoidea, Coleophoroidea, Gelechioidea) with regard of functional morphology of the

male genitalia. Proc. Zool. Inst. Acad. Sci. USSR 122:3-68

1985. Comparative and functional morphology of the male genitalia of the
bombycoid moths (Lepidoptera, Papilionomorpha: Lasiocampoidea, Sphingoidea,
Bombycoidea) and their systematic position. Proc. Zool. Inst. Acad. Sci. USSR 134:
3-48.

PETERSON, A. 1964. Entomological techniques. How to work with insects. 10th ed.
Edwards Brothers, Inc., Ann Arbor, Mich. v + 485 pp.

RAZOWSKI, J. 1981. Musculature of the male genitalia in Tortricinae (Lepidoptera,
Tortricidae). Polskie Pismo Entomol. 51:3-12.

SNoDGRASS, R. E. 1935. Principles of insect morphology. McGraw-Hill, New York. x
+ 667 pp.

STEKOL NIKOV, A. A. 1965. Functional morphology of the copulatory apparatus in some
Lepidoptera. Entomol. Obozr. 44:258-271.

1967a. Phylogenetic relations within the Rhopalocera (Lepidoptera) on the basis

of the functional morphology of the genital apparatus. Entomol. Obozr. 46:3-24.

1967b. Functional morphology of the copulatory apparatus in the primitive

Lepidoptera and general evolutionary trends in the genitalia of the Lepidoptera.

Entomol. Rev. 46:400-409.

Received for publication 27 December 1988; accepted 1 March 1989.

GENERAL NOTES

Journal of the Lepidopterists’ Society
43(3), 1989, 244

ON THE LOCATION OF SOME H. A. FREEMAN SKIPPER
HOLOTYPES (HESPERIIDAE)

Additional key words: American Museum of Natural History, Mexico.

In “Records, New Species, and a new Genus of Hesperiidae from Mexico,” Journal of
the Lepidopterists’ Society, Vol. 23, Supplement 2, 1969, I stated that the holotypes of
most of the species described were to be placed in the United States National Museum,
Washington, D.C. Actually, these holotypes were deposited in the American Museum of
Natural History (AMNH), New York, in 1981 along with my entire collection of Mexican
Hesperiidae. Thus, holotypes of the following species can be found in the AMNH: Pyr-
rhopyge tzotzili, Mysoria wilsoni, Epargyreus windi, Epargyreus brodkorbi (designated
in 1969 paper for Museum of Zoology, Univ. of Michigan), Astraptes louiseae, Astraptes
gilberti, Polythrix mexicanus, Aethilla chiapa, Mimia chiapaensis, Windia windi, Sta-
phylus veytius, Staphylus zuritus, Quadrus francesius, Enosis matheri, Dalla ramirezi,
Vettius argentus, Niconiades comitana, Anthoptus macalpinei, Cynea nigricola, Pher-
aeus covadonga, Carystoides escalantei, Carystoides abrahami, Carystoides floresi, Car-
ystoides mexicana, Atrytone mazai, Atrytone potosiensis, Mellana montezuma, Euphyes
chamuli, and Tirynthia huasteca.

HuGH A. FREEMAN, 1605 Lewis Drive, Garland, Texas 75041.

Received for publication 01 April 1989; accepted 18 April 1989.

Journal of the Lepidopterists’ Society
43(3), 1989, 244-247

EFFECTS OF HANDLING ON EUPHYDRYAS EDITHIA (NYMPHALIDAE)

Additional key words: Mark-release-recapture, wing wear, aging.

A central component of most studies of insect population dynamics is mark-release-
recapture (MRR). It is generally assumed that handling insects during MRR does not
affect either their survival or behavior, but rarely have these assumptions been tested.
Several previous studies have looked at possible effects of handling on recapture proba-
bilities. R. H. T. Mattoni and M. S. B. Seiger (1963, J. Res. Lepid. 1:237-244) compared
observed with expected values of multiple recaptures of Philotes sonorensis and found
no decrease in observed recaptures, as would be expected if repeated handling had a
negative effect on recapture probability. Other studies, however, found reduced proba-
bilities of recapturing handled butterflies in the area of first capture (Singer, M. C. & P.
Wedlake 1981, Ecol. Entomol. 6:215-216; Morton, A. C. 1982, Oecologia 53:105-110;
Gall, L. F. 1984a, Biol. Conserv. 28:139-154).

Studies attempting to determine the age-structure of butterfly populations commonly
use wing-wear as an indicator of age (Watt, W. B., F. S. Chew, L. R. G. Snyder, A. G.
Watt & D. E. Rothschild 1977, Oecologia 27:1-22; Ehrlich, P. R., A. E. Launer & D. D.
Murphy 1984, Am. Nat. 124:525-539; Gall, L. F. 1984b, Biol. Conserv. 28:111-138).
Butterflies captured with undamaged (fresh) wings are considered young, while butterflies
with worn wings are scored as old. In such studies, it is important to determine whether
the MRR technique itself measurably wears the insects; such an effect would increase
age estimates of repeatedly handled butterflies and possibly decrease survival. In this

VOLUME 438, NUMBER 3 245

TABLE 1. Linear regression of change in condition on number of handling events, by
days in residence (males).

Days in 95% confidence limits Correlation Power*
residence Sample size Slope (m) or the slope coefficient (R) (1 — B)
3 24 0.08 —0.30 0.47 0.10 0.12

4 30 0.00 =) PA) 0.30 0.01 0.13

3) 32 —0.18 —0.39 0.04 0.29 0.52

6 28 All 4 1. 0.00 0.37 0.71

7 LS 0.12 —0.13 0.37 0.20 0.25

8 Pall 0.07 Se OEE 0.26 0.17 0.27

9 25 —0.03 —0.19 0.13 0.08 0.12

10 19 0.09 —0.10 0.28 0.23 0.21

* Power values for correlation coefficients from Cohen, J. 1977, Statistical power analysis for the behavioral sciences,
rev. ed., Academic Press, New York, 474 pp.

study, we attempt to determine if handling during MRR studies causes an increased rate
of wing-wear.

At Stanford University’s Jasper Ridge Biological Preserve, Euphydryas editha bayensis
(Sternitzky) populations have been under experimental observation since 1960. In the
past twenty-eight years, extensive data from MRR studies have been collected (Ehrlich,
P. R. 1965, Evolution 19:327-836; Ehrlich, P. R., R. R. White, M. C. Singer, S. W.
McKechnie & L. E. Gilbert 1975, Science 188:221-228; and Baughman, J. F., D. D.
Murphy & P. R. Ehrlich 1988, Oecologia 75:593-600).

In 1981, an intensive MRR study was carried out at the Jasper Ridge Area H demo-
graphic unit from 23 March to 1 May (Ehrlich et al. 1984, above). Butterflies were handled
on all of the days that they flew; a total of 478 individuals were handled at least once
during the season (310 males and 168 females). Males are more likely to be caught than
females because of differences in flight behavior. Three experienced field workers at-
tempted to capture all of the butterflies present on each day of the flight season. The
MRR protocol followed that of P. R. Ehrlich and S. E. Davidson (1960, J. Lepid. Soc. 14:
227-229), with each individual given a characteristic mark with a felt-tipped, permanent-
ink pen. Between capture and release, individuals were kept in glassine envelopes with
their wings together to keep them from moving; these envelopes were then placed in
slotted boxes appropriately marked by sex and area of capture. After collecting was
completed, butterflies were removed from the envelopes with forceps, marked (on initial
capture), examined, and released.

At capture and at each subsequent recapture, the individual’s age, as estimated by
wing-wear, was recorded on a scale of 0.5 to 3.5, in increments of 0.5, with 0.5 indicating
a newly emerged individual and 3.5 a very worn one (for an alternate technique, see
Watt et al., above). In this study, both loss of scales and nicks were used as indicators of
wear. When making age estimates, an effort was made to ignore obvious handling damage
(such as fingerprints) and to score only naturally induced wear. For consistency, the same
three people performed all of the sampling and two checked each rating.

TABLE 2. Linear regression of change in condition on number of handling events by
days in residence (females).

Days in 95% confidence limits Correlation Power

residence Sample size Slope (m) for the slope coefficient (R) G3)
3 18 0.22 —0.07 0.52 0.37 0.54
4 20 0.02 —0.21 0.24 0.04 0.11
5 14 0.20 —0.19 0.59 0.30 0.28
6 13 —0.12 —0.54 0.30 0.19 0.16
7 11 0.02 —(0.41 0.44 0.03 0.09

246 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Field records indicate how many times each individual was captured, the day each
capture or recapture occurred, and the estimated condition at the time of each handling.
From these data, the length of time between first and last capture (days in residence),
the number of handling events that occurred (initial capture plus total number of recap-
tures), and how much the butterfly aged (change in condition), were determined for each
individual.

To determine if handling the butterflies influenced the rate at which they aged (as
indicated by wing-wear), a linear regression of change in condition on number of handling
events was performed (Model I linear regression with >1 value of Y for each value of
X; for details, see Sokal, R. R. & F. J. Rohlf 1981, Biometry, 2nd ed., W. H. Freeman
and Co., New York, 859 pp.). Previous studies (Ehrlich et al. 1984, above) have shown
that male and female Euphydryas wear at significantly different rates; therefore, the data
were pooled by sex. For each sex, individuals were pooled by number of days in residence
in order to separate natural wear from wear induced by handling. Individuals captured
only once were not included in the analysis. Only males in residence between 3 and 10
days (210 individuals), and females in residence 3 to 7 days (76 individuals), were con-
sidered. Too few were in residence for longer and shorter periods to make analysis reliable.

The results of the regressions are summarized in Tables 1 and 2. In all cases, regression
line slopes are not significantly different from zero. Although with small sample sizes it
is not possible to affirm the null hypothesis at a satisfactory power (only males 6 days in
residence had a test power >0.70; for most of the other regressions, the probability of
rejecting a false null hypothesis (1 — 6) was <0.30), the results suggest that there is no
significant relationship between amount of handling and change in condition. In addition,
a linear regression of change in condition on days in residence was performed for each
sex, pooling across number of handling events. In both cases, slopes were significantly
different from zero (males, m = 0.12, P < 0.001; females, m = 0.13, P < 0.001), indicating
a significant relationship between time and change in condition, as would be expected.
The majority of males (157 of 210) and females (56 of 76) had an initial condition of 0.5
and over half of the remaining individuals in each case had initial conditions of 1.0;
consequently, further subdividing the butterflies into wing-wear cohorts (grouping by
initial condition) did not change the significance of any of the results.

It is doubtful, however, that handling never causes wear. Different investigators, because
of varying amounts of practice or ability, probably cause different amounts of wear to
the butterflies they handle. The same person may occasionally cause a great deal of wear
to a single butterfly (due to difficulty in disintangling the butterfly from the net, for
example) while normally causing very little wear. It is probable that the greatest handling-
induced change in condition occurs during the initial capture and marking. Subsequent
recaptures may not greatly affect overall condition. Singer and Wedlake (above) found
that Graphium sarpedon (L.) handled while being marked were much less likely to be
recaptured than those not handled while marked, which they interpreted as a change in
dispersal behavior due to the initial capture. A marking effect limited to the date of
capture was found in Boloria acrocnema (Gall & Sperling) by Gall (1984b, above).
Capturing and marking the butterflies disrupted their flight activity immediately following
release, but this effect did not appear to last beyond the marking date. Wear induced by
initial capture and marking would not cause an increased rate of wing-wear, but possibly
could affect survival.

The conclusion that increased handling does not significantly change the amount of
wear observable on Euphydryas editha has two important implications for MRR studies.
First, it indicates that handling may not significantly “age” Euphydryas editha individuals.
Secondly, it suggests that, when done carefully, it is possible to estimate age reliably using
wing-wear as an indicator.

We thank Alan Launer and Dennis Murphy for assistance in the field, and Paul Ehrlich
for advice and encouragement. Lawrence F. Gall and Oakley Shields reviewed the
manuscript and provided helpful suggestions. This work was supported by a series of
grants to P. R. Ehrlich from the National Science Foundation, most recently BSR8206961.

VOLUME 43, NUMBER 3 QA47

MarIA E. ORIVE* and JOHN F. BAUGHMAN, Department of Biological Sciences, Stan-
ford University, Stanford, California 94301.

* Current address: Department of Zoology, University of California, Berkeley, California 94720.

Received for publication 20 July 1988; accepted 3 April 1989.

Journal of the Lepidopterists’ Society
43(3), 1989, 247-249

PERFORATED CUPOLA ORGANS ON LARVAE OF
EUSELASIINAE (RIODINIDAE)

Additional key words: Euselasia aurantiaca, E. mystica, Hades noctula, ultrastruc-
ture.

Perforated cupola organs (PCO’s) are minute, epidermal secretory organs, homologous
to setae, found on larvae of many Lycaenidae (Malicky, H. 1970, J. Lepid. Soc. 24:190-
202). They also occur on larvae of Riodinidae. These organs are known to secrete amino
acids in some species (Pierce, N. E. 1983, Ph.D. Thesis, Harvard University, Cambridge,
Massachusetts, 286 pp., Diss. Abs. Int. 44:1708B), and are thought to be involved in
maintenance of ant associations in myrmecophilous species even though they are also
found on amyrmecophilous larvae (Malicky, above; Kitching, R. L. & B. Luke 1985, J.
Nat. Hist. 19:259-276). These organs have been relatively well-studied in Lycaenidae
(DeVries, P. J., D. J. Harvey & I. J. Kitching 1986, J. Nat. Hist. 20:621-633 and included
references; Kitching, R. L. 1987, J. Nat. Hist. 21:535-544), but there is little information
on their occurrence in Riodinidae (sometimes considered a subfamily of Lycaenidae).
They have been illustrated using scanning electron microscopy in one amyrmecophilous
species of Old World Hemearinae, Hamearis lucina (L.) (Kitching & Luke, above), and
one myrmecophilous species of New World Riodininae, Pandemos palaeste Hewitson
(Harvey, D. J. & L. E. Gilbert, J. Nat. Hist. in press). They have not been illustrated,
however, for larvae of a third subfamily, Euselasiinae, although their presence in this
group has been alluded to (Harvey, D.J. unpubl., cited in DeVries et al., above). Larvae
of the remaining subfamilies, the monotypic Styginae and Corrachiinae, are unknown
(Harvey, D. J. 1987, pp. 446-447 in Stehr, F. (ed.), Immature insects, Vol. 1, Kendall/
Hunt, Dubuque, Iowa, 754 pp.).

Euselasiinae consists of three genera: Euselasia with over 130 species, Hades with 2
species, and the monotypic Methone (Harvey, D. J. 1987, Ph.D. Thesis, University of
Texas, Austin, Texas, 216 pp., Diss. Abs. Int. 49:625B). Distribution and morphology of
PCO’s on mature larvae of three euselasiines, E. mystica (Schaus), E. aurantiaca (Godman
& Salvin) and H. noctula Westwood, are described here.

Larvae were examined with a Wild stereomicroscope. Material for scanning electron
microscopy was coated with gold-palladium in a Hummer V sputter coater, and micro-
graphs taken with an ISI Super IIIA.

All three species have the same distribution pattern of PCO’s. Some are scattered along
lateral and posterior margins of the prothoracic shield (Fig. 1). All remaining PCO’s on
larvae are restricted to clusters around abdominal (A) spiracles (Fig. 2). Long, tactile
setae, present elsewhere on the larvae are absent from these clusters, though they may
be immediately adjacent. The PCO’s are set in fields of microtrichia (Figs. 2-4). The
numbers of abdominal PCO’s on larvae of the three species are as follows (A segment

248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 1-5. Scanning electron micrographs of perforated cupola organs (PCO’s) on
larvae of Euselasiinae (Riodinidae). 1, Euselasia mystica, PCO on prothoracic shield
(scale bar = 10 um); 2, Hades noctula, cluster of PCO’s around spiracle on A4, left side
(scale bar = 200 um); 3, H. noctula, individual PCO (scale bar = 10 ym); 4, E. aurantiaca,
PCO in field of microtrichia, near spiracle on A4, left side (scale bar = 10 ym); 5, E.
aurantiaca, sieve plate of PCO showing pores (scale bar = 4 um).

number: number on right and left side; “?” denoting larva damaged and PCO’s present
but uncountable):

E. mystica: A1:15,17; A2:33,31; A3:24,27; A4:18,15; A5:14,10; A6:17,14; A7:22,26; A8:
26,21.

E. mystica: A1:?,49; A2:55,52; A8:?,45; A4:?,?; A5:?,385; A6:39,?; A7:53,?; A8:70,?.

H. noctula: A1:36,38; A2:86,91; A3:47,56; A4:34,36; A5:33,28; A6:29,27; A7:42,51; A8:
45,47.

In addition to identical placement of PCO’s, all three species also show a similar pattern
in relative numbers of PCO’s on different segments: maximum numbers on the anterior
segments are found on A2, usually followed by A3; and on the posterior segments, on A7
and A8.

Diameters of the PCO’s average 21 um for E. mystica (on prothoracic shield), 27 wm
for E. aurantiaca (on A4), and 28 um for H. noctula (on A4). Minute pores are visible
on the central “sieve plate” of the PCO’s (Figs. 1, 5). No pores are evident on sieve plates
of H. noctula, which have minute crenulations (Fig. 3), of unknown function.

Larvae of Euselasia and Hades are amyrmecophilous; ant mutualisms are restricted
to the subfamily Riodininae (Harvey, above; Harvey & Gilbert, above).

PCO’s of Riodininae differ from those on larvae of the amyrmecophilous H. lucina,
which lack pores, are not clustered, and are more sparse (Kitching & Luke, above). PCO’s
are also present on first instars of both myrmecophilous (Eurybia, Calospila among others)

VOLUME 43, NUMBER 3 249

and amyrmecophilous (Apodemia, Calephelis among others) riodinines, where they form
a regular component of chaetotaxy (Harvey unpubl.). Despite broad taxonomic occurrence
of PCO’s, their function (if any) in euselasiines and other amyrmecophilous riodinids is
obscure.

Pattern of PCO distribution in Euselasiinae examined in this study is consistent, and
may be taxonomically significant. It resembles that described for the curetine lycaenid
Curetis regula Evans (DeVries et al., above), where PCO’s are also clustered near ab-
dominal spiracles. However, Curetis differs in having PCO’s near the prothoracic spiracle
(rather than on the shield), and in their closer spacing (without intervening microtrichia).
In addition, their form is more elevated, and waxy exudates are present on sieve plates.
On the other hand, Euselasiinae differ from observed Riodininae, which usually have
PCO’s (when present) in several pairs of clusters per segment, or if single clusters are
present (as in Euselasiinae), they are more dorsal on segments (Harvey & Gilbert, above;
Harvey unpubl.). Restriction of PCO’s to clusters around spiracles and prothoracic shield
may be the primitive configuration in Riodinidae, perhaps also in Lycaenidae. In light
of this possibility, description of PCO’s from additional members of Hamearinae, from
Styx infernalis Staudinger and Corrachia leucoplaga Schaus, and from the lycaenid
subfamilies Lipteninae and Poritiinae, would be of interest.

I thank P. J. DeVries, N. Greig and J. Mallet for providing fluid-preserved larvae, J.
C. Downey, R. L. Kitching, and P. J. DeVries for comments on the manuscript, C. Drake
and the late S. Meier for assistance with scanning electron microscopy.

DONALD J. HARVEY, Department of Zoology, University of Texas, Austin, Texas 78712.
Present address: Department of Entomology, National Museum of Natural History,
NHB Stop 127, Smithsonian Institution, Washington, D.C. 20560.

Received for publication 1 July 1987; accepted 22 September 1988.

BOOK REVIEWS

Journal of the Lepidopterists’ Society
43(3), 1989, 250-251

CATALOGUE OF LYCAENIDAE & RIODINIDAE (LEPIDOPTERA: RHOPALOCERA), by Charles
A: Bridges. 1988. Printed by the author. 816 pp.: vi, ii + 377 pp., ii + 115 pp., ii + 140
pp., ii + 100 pp., ii + 37 pp., ii + 1 p., ii + 10 pp. 21 x 28 cm, hardcover. $95.00 in
North America, $97.50 elsewhere.

Bridge's Catalogue of Lycaenidae and Riodinidae is an extremely useful publication
for any lepidopterist interested in the systematics of these two families. It provides
information on original descriptions and other literature, authors, periodicals, and the
current systematic placement of taxa. I have found the catalogue easy to use and a great
time-saver for tracking down names and references.

The catalogue is divided into a brief introduction, six parts and two appendices, each
with separate pagination. Part I consists of an alphabetic list of species-group names. Each
entry includes the author, date of publication, abbreviated literature citation (cross-
referenced to Part IV), and the original genus for each species-group name. The status
of each name (i.e., available and valid species, subspecies, or synonym; available invalid;
or unavailable) is indicated by a letter code. For species subsequently transferred to other
genera, the current genus is given along with a reference(s) (cited in full in Part IV) for
the transfer. In addition, many entries include information on the type locality, location
of type specimen, its sex, and/or references to life history. Part IJ is an index to genera
that includes a list of species-group names under each genus. Part III is the index to the
bibliography. Publications are listed by author, date, and journal (cross-referenced to the
full citation in Part IV). Under each publication, species group names are listed, along
with reference to volume and page number, and to plate and figure numbers if the taxon
is illustrated. Part IV, the bibliography, lists complete citations for 4258 publications.
Each publication is given a unique number by which it is cross-referenced in other parts.
This part also provides information on when and where some authors were born and
died, and the disposition of their collections. Part V, the index to journals and serials, lists
them by abbreviation and includes their full titles. Under each is a list of included papers
(cross-referenced to Part IV) with the author, date, volume and page numbers. Part VI,
the index to the bibliography by year, lists the unique numbers of each publication in
Part IV under its year of publication. Appendix I is a synonymic list of family-group
names. Appendix II is a synonymic list of genus group names arranged according to
family, subfamily, tribe, subtribe (if any) or section (if any). Although not explicitly stated,
the higher classifications follow Eliot for the Lycaenidae and Stichel for the Riodinidae.

There are some problems with the first 40 copies that will be corrected in later editions.
In Part I, a block of 290 names is missing “between” pages 275-276, and a block of names
is duplicated on pages 287-293. In addition, some changes in format also will be made:
in Part IJ each genus will have an indication of its place in the higher classification that
will facilitate finding it in Appendix IJ, and Part VI will include the names of authors.
A series of annotations is being issued which lists additions and corrections to the catalogue.

The catalogue appears to contain relatively few errors for a work of its size. I found
two typographical errors in the Introduction (misspellings of Julian P. Donahue’s and
Jacqueline Y. Miller’s names). The authorships of riodinid taxa described by Le Cerf and
by Lathy in a paper by Rebillard are incorrectly attributed to Rebillard in the catalogue.
The generic name Balocna Moore is listed as a synonym of Zemeros Boisduval in Appendix
II but it is actually a synonym of Dodona Hewitson. Although the author states in the
introduction that no new names are introduced in the catalogue, the subtribe Sarotiti is
apparently proposed in Appendix I as a replacement name for Charitini Stichel. These
problems are minor, and the author is to be commended for providing a mechanism for
correcting such errors.

The catalogue holds at least one nomenclatorial surprise. The name Orimba Herrich-
Schaeffer (1858), used by Stichel and all subsequent authors for a genus of neotropical
riodinids, is actually a synonym of Setabis Westwood (1851).

VOLUME 438, NUMBER 3 Zon

The amount of time and effort required to produce this catalogue must have been
immense, and such enterprises are often thankless tasks. Bridges has done us a great favor
by providing a careful, well-planned and exhaustive work. This catalogue is an indis-
pensible reference that belongs in the library of all who work on the systematics of
lycaenids and riodinids.

DONALD J. HARVEY, Department of Entomology, NHB Stop 127, Smithsonian In-
stitution, Washington, D.C. 20560.

Journal of the Lepidopterists’ Society
43(3), 1989, 251-252

PAPILLONS ET CHENILLES DU QUEBEC ET DE L’EST DU CANADA [Lepidoptera and Larvae
of Quebec and of Eastern Canada], by Jean-Paul Laplante, 1985. 280 pp., 65 color plates,
with many other color figures in text. Editions France-Amerique, 170 Benjamin Hudon,
Montreal, Quebec H4N 1H8, Canada. Hardcover. About $25.00.

This beautiful book makes an important contribution to our knowledge of the butterflies
and moths of Quebec and of eastern Canada in general. Covering more than 300 species
and subspecies of Lepidoptera with illustrations of the adults, eggs, larvae, pupae, and
habitats (over 1000 separate color figures), this book would be of value to anyone with
an interest in the Canadian fauna. It is currently available only in a French edition, but
Latin insect and plant names, locality names that can be identified on any map, etc.,
make it readily usable even if one should not know French.

The author has worked for more than 30 years on the lepidopteran fauna of eastern
Canada, especially Quebec, and has collaborated with many workers in Canada to as-
semble the knowledge and photographs displayed in this volume. He begins with a general
introduction to the evolution and biogeography of butterflies and moths, their ecology,
and life history. Excellent text drawings and scanning electron micrographs, as well as
color photographs, illustrate scale structure and other features. Concise but well-done
summaries of geographic and genetic variation of butterflies and moths are presented,
along with a fascinating discussion of mimicry that includes unusual illustrations not
appearing in any other book. Likewise, a short section is devoted to the enemies and
diseases of Lepidoptera, and there is a valuable descriptive section on the characteristics
of each family of butterflies and some of the major moth groups.

The author then presents a quite usable key to the species of diurnal Lepidoptera in
Quebec as well as to species of certain genera in major moth families (Sphingidae,
Lasiocampidae, Saturniidae, Arctiidae, Agaristidae, Notodontidae, and Lymantriidae).
The last key, interestingly enough, is solely to the larvae of the species in the genus
Dasychira in Quebec, because they offer the best distinguishing characters for the genus.

The outstanding and immaculately reproduced color plates, however, carry the major
load of identification of specimens. The male, female, and underside of each species is
shown, along with seasonal and geographic variation across eastern Canada. The plates
are among the very best ever produced for a book on a North American faunal region.
After the 34 color plates of adult specimens, photographed crisply on a blue background,
the author includes a series of 30 plates of eggs, larvae, and pupae of the illustrated
butterflies and moths, as well as a plate of eight habitat photographs. Technically, it
would be hard to suggest any improvement that could be made in the beautiful photog-
raphy that illustrates this book.

The author discusses in detail the vegetation zones of Quebec and northeastern Canada,
north to Hudson Bay and west into Ontario as well as east into Labrador and Newfound-
land. One of the most interesting features is a complex yet highly readable table presenting
a summary of biological notes on 282 species and 11 subspecies of Lepidoptera, including
134 butterflies and 159 moth species. This table neatly shows the distribution, flight period,
abundance in habitat, cross-references to illustrations in the text, number of annual
generations, the hibernation or aestivation stage, and the larval characteristics, including
host plants, period of activity, living habits (solitary, gregarious, etc.), and the average

202 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

body length, as well as cross-references again to the illustrations in the main text. The
book closes with a brief but very adequate discussion on how to collect, prepare and
preserve butterflies and moths. An excellent glossary and selected bibliography, as well
as a comprehensive index, close the book.

Jean-Paul Laplante has produced an excellent book on the butterflies and many of the
interesting larger moths found in Quebec and the other areas of eastern Canada. The
wonderful color illustrations of the larvae of virtually all the species of butterflies and
major moth groups in Quebec would make this book a sound investment on that basis
alone. The extraordinarily low cost of this beautiful book and its ready intelligibility even
to readers lacking a good reading knowledge of French should prompt many lepidopterists
to purchase it for their personal libraries.

THOMAS C. EMMEL, Division of Lepidoptera Research, Department of Zoology, Uni-
versity of Florida, Gainesville, Florida 32611.

Journal of the Lepidopterists’ Society
43(3), 1989, 252-253

SLUG AND NETTLE CATERPILLARS: THE BIOLOGY, TAXONOMY AND CONTROL OF THE
LIMACODIDAE OF ECONOMIC IMPORTANCE ON PALMS IN SOUTH-EAST ASIA, edited by M.
J. W. Cock, H. C. J. Godfray, and J. D. Holloway. 1987. 270 pp., 18 color plates. CAB
International, Wallingford, Oxon, UK. Hardcover. $99.00.

This book is an invaluable tool for tropical biologists in the coconut and oil palm
industries of South-east Asia. It also is important in a broader geographic and economic
sense because the larvae of Limacodidae, which are highly polyphagous, are pests of
palms and other tropical plantation crops worldwide. Although less than comprehensive,
the work presents a review of recent literature on natural enemies of New World limacodid
palm pests along with a wealth of information on Limacodidae in general.

The organization of the book is as follows: chapters 1 and 2 present introductory
information on Limacodidae; chapter 3 provides systematic accounts of palm pests of
South-east Asia; and chapter 4 is a short, preliminary account of tropical Australasia pests.
Chapters 5-17 deal with aspects of pest management and include systematic accounts of
parasitoids and predators, and fungal, viral, and chemical control. Most of these final 13
chapters are brief, encompassing about half of the total text.

The book includes 36 plates comprising genitalic preparations, color photographs of
spread specimens (with useful, identified black and white duplicates on facing pages),
and striking photographs of live larvae, cocoons, natural enemies, and adults in natural
postures.

As one who has reared limacodids for several years, I can appreciate the amount of
intensive labor that the book represents. I found the information on rearing methods
(chapter 2) particularly enlightening. A minor shortcoming, however, is a sense that larvae
were reared by someone other than the authors. The statement that larvae appear “re-
markably stupid” because they must be manually transferred to new host material is
absurd, as this is a manifestation of rearing these specialized larvae in captivity. Slug
caterpillars, especially in early instars, have a difficult time moving from one leaf to
another because of the small thoracic legs and absence of abdominal prolegs. Difficulties
in rearing slug caterpillars that the authors fail to mention include: 1) their tendency to
become immobilized with frass due to the sticky nature of their ventral surface; and 2)
their movement off the host material and onto the container, perhaps a preference for
the smoothest available substrate (most limacodids are found on hosts with smooth leaves).

Chapter 3 on systematics of the South-east Asian pest species considerably expands our
knowledge of the region’s fauna, with 35 new species and four new genera described by
Holloway, and 28 new synonyms, new combinations, and other nomenclatural changes.
Relationships among genera are proposed on the basis of the signum type of the female
genitalia as in Holloway (1986, The moths of Borneo: Key to families; Cossidae, Metar-

VOLUME 43, NUMBER 3 253

belidae, Ratardidae, Dudgeoneidae, Epipyropidae and Limacodidae, Malay. Nat. J. 40:
1-166). While the family represents a well defined monophyletic group, there is no widely
accepted supergeneric classification for the world fauna. Use of the signum in relating
genera may have merit, but caution should be exercised since the congruence of this
character with other morphological and/or behavioral characters has not been examined
in a phylogenetic (cladistic) context.

Chapters 5-9 deal with parasitic Hymenoptera associated with South-east Asian Li-
macodidae. The chapters on Ichneumonidae, Braconidae, and Chalcidoidea, constituting
nearly a fourth of the book, have keys to the parasitoids, with scanning electron micro-
graphs and line drawings of the former two families. I found chapter 5 very informative
in its division of the life styles of ichneumonid wasps by taxonomic groups. Chapters 10
and 11 summarize dipteran parasitoids in the Tachinidae, Sarcophagidae, and Bomby-
liidae, and chapter 12 reviews hemipteran predators. Chapter 18, a half page description
of a pyralid cocoon predator, Ectomyelois ceratoniae (Zeller), would have been better
summarized in the introductory matter or mentioned with other predators. Reviews of
classical biological control, fungal pathogens, viruses, and chemical control of limacodids
are presented in the final four chapters.

Although the book is full of important life history information and literature citations,
several important references on limacodid life histories are not included. In the 60th
anniversary since the passing of Harrison G. Dyar (1866-1929), it seems appropriate to
mention his contribution to this subject. Even though Dyar described the early stages of
primarily Nearctic species, many of these taxa have obvious phylogenetic connections
with the Asian fauna. Reference to Dyar’s work would have added support to statements
on the origin of non-stinging, smooth (‘gelatine’) types of caterpillars from those with
stinging scoli. Dyar recognized the ancestral plan of Limacodidae as possessing two rows
of scoli, and hypothesized two independently derived lineages of smooth larvae (Dyar,
H. G. 1899, The life-histories of the New York slug caterpillars, J. N.Y. Entomol. Soc. 7:
234-253, pls. 6-8). Each of these lineages possess rows or rudiments of scoli in the first
instar that are lost in later instars.

While I applaud the information on rearing methods, there is no mention of obtaining
life-history data by capturing adult females and inducing oviposition. This is a viable
alternative, particularly since most if not all limacodids can be reared on palm without
previous knowledge of the host owing to their polyphagous nature. This procedure, used
by Dyar, is practical for associating adults with larvae and obtaining good series of both.

Appendix 1 is a list of host plants of South-east Asian limacodids mentioned in the text.
Unfortunately, this information is not indexed, making it difficult to find the species of
limacodids associated with each plant.

The price of this book may be prohibitive to those with only a casual interest in
Limacodidae or without an economic stake in the subject. However, since the work
represents a significant contribution to our knowledge of the early stages, behavior, and
systematics of Limacodidae, it is indispensible for the serious student.

MakC E. EPSTEIN, Department of Entomology, National Museum of Natural History,
NHB 127, Smithsonian Institution, Washington, D.C. 20560.

Journal of the Lepidopterists’ Society
43(3), 1989, 253-255

THE GUILD HANDBOOK OF SCIENTIFIC ILLUSTRATION, edited by Elaine R. S. Hodges,
1989. xv + 575 pp. Van Nostrand Reinhold, 115 Fifth Avenue, New York, New York
10003. 22 x 29 cm, hardcover. $79.95.

In 1968 U.S. natural science illustrators organized themselves professionally; since then,
the Guild has brought to a formerly disconnected occupation a unity of strength and
purpose. The Guild Handbook of Scientific Illustration is one very tangible result of this
union, its high-tone pages bringing together in one volume the wide variety of principles

254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

and procedures that define the field of natural science illustration (practiced by scientitic
illustrators dedicated to biological and medical subjects).

In five major parts, 30 chapters introduce the reader first to the “Basics: steps in the
process of illustration, outfitting the studio, materials, and the play of light on subjects.
“Rendering Techniques’ follow: an in-depth survey of the use of media from simple line
and ink to complex color applications. The majority of the text consists of 11 chapters
covering the special problems and approaches to the principal “Subject Matter:” plants,
fossils and extinct vertebrates, invertebrates (including insects), fishes, amphibians and
reptiles, birds, mammals, animals in their habitats, humans and their artifacts, and medical
subjects. Several advanced topics follow in “Beyond Basics:” using the microscope, charts
and diagrams, cartography, copy photography, and the printing process. Finally, the
“Business of Scientific Illustration” is given attention, where copyright law, making con-
tracts and operating a free-lance business are discussed. Included are a Bibliography and
Appendix with names of sources of supplies and other compact information.

The voluminous text is generally well written by 45 motivated and capable authors
and is fully illustrated with over 600 clean, instructive figures in black and white and
color. The matter discussed is immense in depth and diversity, requiring and receiving
a masterful editing job by Hodges.

It is hard to find fault in a work obviously so lovingly and meticulously produced. I
would, perforce, comment on a few minor imperfections, mostly trivial, which mar only
slightly an otherwise superior technical publication:

1. While the somewhat varied approaches taken by authors to accommodate the quirks
of their disciplines are highly appropriate, they allowed for some redundancy (e.g., where
techniques overlap) and at least one contradiction (sans serif lettering advocated by Allen
on p. 500, the contrary by Lynch on p. 459).

2. I find some bothersome pedantries and truisms, the most common being the oft
repeated statement that taking classes in this or that discipline of biology makes one a
better illustrator. I think that many items in the lists of instruments and materials in Part
3 are part of any studio; mention of such items as magnifiers, rulers, a camera, French
curves, etc., could give way to more discussion of things peculiar to the subjects at hand.
It does not seem necessary to say, on p. 394, right column: “The illustrator may be called
upon to work on a wide range of subjects in a variety of settings.’’ Or on p. 376, middle
column: “There is considerable variety in the field of illustrating mammals.”

3. Some confusing statements have crept into the text: p. 23, left column: “To this end,
adaptations of some traditional drawing media have been developed in some techniques.”
Also, on p. 264, first sentence: “Invertebrates that are not arthropods do not have jointed
legs.”

4. I miss discussion on some important issues: Nowhere is perspective or the problem
of parallax discussed, yet lighting is given a detailed treatment. Needed is a more unified
review of the types of symmetry, axes, planes, and regions in organisms, and terms
pertaining thereto. A glossary would be a welcome addition to the next edition.

5. Some authors fail to include mention of those involved in the historical development
of their field. While this may be excused by the primarily prosaic purposes of the book
and the availability of other works on the history of biological illustration, the omission
is definitely to the detriment of the reader, especially when classic examples are not
shown.

6. Much of the flavor of the book is towards drawing for taxonomy. I would like to
have seen more shift given to the branches of anatomy and behavior (especially regarding
insects).

7. The list of suppliers in the index leans heavily toward the eastern U.S.; we out west
have many fine outlets too.

8. Some errors bear noting: p. 197, right column: “Plants having roots, stems, leaves,
and a vascular system are called gymnosperms.” should read “. . . are called thallophytes.”
The sentence is repeated on p. 199, left column but ends in “angiosperms.” also erro-
neously. In the list of types of specimens given on p. 4, left column, mounted specimens
are illogically omitted (see #6 below).

As an entomological illustrator myself, I have a bit more to say of the pages dealing

VOLUME 438, NUMBER 3 255

with insects. These are small issues, as those above, the treatment on the whole being
excellent.

1. I notice again the lack of acknowledgment of achievements of historical workers.
Superb examples date back as far as Hooke’s louse and Lyonnet’s goat moth larva of the
early 18th century. In my opinion the finest entomological illustrator was Hermann Weber.
His rendering of homopteran mouthparts are masterpieces of analytical anatomical graph-
ics. Others important in establishing the field were E. O. Detmold, A. J. E. Terzi, and G.
Ferris. Some useful and important technical publications might have been cited [Edy, R.
1968, Some illustrations of microsculpture in the Hymenoptera, Proc. Entomol. Soc.
London, ser. A 43:66f.; King, R. & H. Akai (eds.) 1982, 1984, Insect ultrastructure. Vols.
1-2; Catts, E. & J. Young 1959, A chalkboard technique for making illustrations, Pan-
Pac. Ent. 35:168f.]. A nice little book that teaches much on the art of posturing for live
insect illustrations is N. Weaver's How to draw insects (Studio Pub., London. 1958).

2. Some additional techniques are: individual sand grains are suggested for propping
specimens; a bed of fine silica sand gives an even more versatile matrix for holding
specimens in any position. Specimens may also be embedded temporarily in clear gelatin
to hold them for drawing. Insect membrane is commonly indicated by light stippling
while sclerites are left clear in anatomical works. Precautions for putting away microscope
slides are given on p. 261; I would add that the box or tray should be stored so that the
slides are flat, with specimen on top, to prevent gravity from tugging at the medium.

3. There are a few mistakes: p. 290, first paragraph of left column: “... a dorsal
segment is a tergum or tergite’;. . . , should read, “a dorsal sclerite is a tergum or tergite;”
p. 289, top of middle column: Myriapoda is a category that contains millipedes (Diplop-
oda) and is not synonymous with them; p. 297, center column: carbolic acid or phenol
crystals, not naphthalene, are usually added to relaxers to inhibit mold growth. Instructions
for calibrating microscope micrometers in the “Eyepiece Scale Value Method” (p. 31),
transpose “stage micrometer’ for “ocular micrometer (reticle) ”’.

4. Terms that need more explanation are “sclerotized’’ (used to infer hardness and/or
pigmentation); “spines’’ (as distinct anatomically from setae); “minutens’ (unfamiliar to
the non-entomologist).

5. Some indefensible or inane statements appear: p. 293, center column: “Tarsal struc-
ture is second in importance only to antennal form in many insects . . . for identification
to family.” And on p. 3801, last paragraph: “Because these animals vary so widely in size,
appearance, anatomy, and requirements for preservation, the techniques for handling
and drawing them also vary.”’

6. There seems to be a confusion of what is meant by “mounting” and “propping” (p.
260). In entomology a “mounted” specimen is one that has been prepared in some way
(on pin, wings spread, etc.). These may need propping as much as an unmounted specimen.

In summary, this fine work is encyclopaedic and copies will no doubt be put on the
reference shelf by many librarians. But more than that, it is also a voluptuous handbook,
so full of practical data and sound conceptual advice, and beauty as well, that most copies
sold will surely never be found far from the illustrator’s hand.

CHARLES L. HOGUE, Curator of Entomology, Natural History Museum of Los Angeles
County, Los Angeles, California 90007.

Journal of the Lepidopterists’ Society
43(3), 1989, 255-257

BUTTERFLIES OF NEPAL (CENTRAL HIMALAYA), by Colin Smith. 1989. 352 pp., 355 color
figures, 3 maps. Tec Press Service L.P., 487/42 Soi Wattanasilp, Pratunam, Bangkok,
Thailand. 15 x 23 cm, hardcover. $50.00 U.S., plus $5.00 airmail postage.

The Central Himalaya Mountains have always held a fascination for lepidopterists
interested in both temperate and tropical butterflies. At last, we have a field guide to the
butterfly fauna of Nepal—and one to match the demands of its incredible diversity.

256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The great number of butterfly species known to occur in Nepal and the rugged
mountains culminating in Mt. Everest, along with adjacent rich tropical lowlands, have
brought many biologists to this landlocked country to explore and enjoy its natural history
and scenic beauty. The country is partitioned lengthwise into Palearctic and Oriental sets
of floral and faunal provinces. It ranges in elevation from some 500 feet above sea level
on the Ganges plain to the highest peaks in the world in the Central Himalaya (to over
29,000 feet on Mt. Everest in northeastern Nepal). It is little wonder that altitude seems
to be the critical factor for butterfly distribution here. In fact, a significant dividing line
(shown in Smith’s map on p. 31) appears at about 3000 meters elevation (between 9000
and 10,000 feet); some 90% of the species above this line show Palearctic affinities, while
below it, about 90% of the species are of Oriental origin. Although many references to
this diverse fauna have been made in books on high-altitude entomology and in scattered
publications on the characteristics of different groups of butterflies from Nepal, Smith’s
book is the first to cover this extremely interesting fauna in a comprehensive format.

In this new publication Colin Smith has set forth an outstanding introduction to the
614 species known to occur in Nepal. The illustrations cover more than 70% of the species
(90% of the 266 genera), using more than 200 photographs of butterflies in their natural
living state and another 100 photographs of mounted specimens to show uppersides and
undersides. In addition to the extensive taxonomic section, Smith includes a general
discussion of the biology of butterflies, including particular examples of migration, mim-
icry, etc. from Nepal. He also presents a fascinating introduction to the country of Nepal
and to the natural geographic divisions of the Central Himalaya region, including climate,
seasonality, ecology, habitats, and other attributes. The book thus provides an indispensable
introductory guide to the natural history of Nepal, as well as the sole popular introduction
to the butterfly fauna of that country.

For the lepidopterist, the innovative features of this book include a classification chart
at the beginning of each family that shows nicely in columnar format a series of facts
(subfamilies, tribes, genera, new genera, subgenera, region of origin, total number of
species worldwide, and number of species in Nepal) about the classification and status of
that family. Additional tables are offered to help in the identification of certain complex
groups, such as the Lycaenidae. A general summary for the identification of the Nym-
phalidae is presented in a running tabular form. Each genus is numbered within a family,
and author’s name, date, type species, and general diversity worldwide are given. For
each individually numbered species account, Smith gives the complete species and sub-
species name, author, date of publication, common name if available, range of wingspan,
comments on distribution (usually to district within Nepal), seasonality, elevational range,
distribution outside Nepal, and the species’ relative abundance.

Following the taxonomic section, the author traces the history of butterfly collecting
in Nepal and includes a summary of species and subspecies endemic to the country, a
record of principal collectors who have taken Nepalese butterflies and the species and
subspecies they have taken, and a list of the butterflies recorded from Nepal (based on
all authenticated Nepal records known to the author) with their habitats, altitude, sea-
sonality, and common name. The book concludes with a selective bibliography of 48
publications on the butterflies of Nepal, indices to scientific and common names, and a
brief biography of the author.

Overall, this book represents a remarkable individual accomplishment in its presentation
of the first general faunal coverage of the butterflies of Nepal and the Central Himalaya.
Exploration of the Palearctic high country of the Central Himalayan and Trans-Hima-
layan regions has lagged far behind that of the Oriental region in Nepal. Some Himalayan
areas such as border districts and national parks are often closed to collecting, and limited
access by road or air-strip often necessitates carefully-planned, multi-man expeditions by
backpack or porters. The incredibly diverse fauna and terrain will undoubtedly produce
many more species with further lepidopterological exploration of the country, as the
author is the first to admit (he added two Nepalese species new to science in 1986 alone).
Overall, the book is well designed and printed. Although some of the color photographs
are not reproduced as clearly or as crisply as would be desirable, the overall impression
of the book is quite favorable and the illustrations can be used to identify specimens.

VOLUME 43, NUMBER 3 DoT

Perhaps most importantly, the beautiful photographs of habitats throughout Nepal,
coupled with the author’s enthusiasm for photographing living butterflies and studying
their natural history, will help to engender greater worldwide interest in the wilderness
conservation programs and butterflies of this fascinating country of the Central Himalaya.
This book belongs on the library shelf of any lepidopterist interested in temperate and
tropical Old World faunas, or in beautiful butterfly books in general. The book also will
be of interest to biogeographers, ecologists, and conservationists worldwide.

THOMAS C. EMMEL, Department of Zoology, University of Florida, Gainesville, Florida
32611, and OAKLEY SHIELDS, 6506 Jerseydale Road, Mariposa, California 95338.

Journal of the Lepidopterists’ Society
43(3), 1989, 258

FEATURE PHOTOGRAPH

Split-level Dining: Three female Godyris zavaleta amaretta (Haensch, 1903) (Nym-
phalidae: Ithomiinae) share a piece of fruit in the interior of primary rainforest in eastern
Ecuador. The leaf on which these three are perched is 1.5 m above the forest floor. In
the canopy 60 m overhead is a flock of yellow-headed parrots (Amazona ochrocephala)
gorging themselves on fruit, fragments of which fall to the forest understory below,
providing a food source for deep forest Lepidoptera and other insects. Male Godyris visit
tree-fall gaps and stream margins to seek flowers for nectar, while females remain in the
forest interior and feed on detritus. Photograph taken at Limoncocha, Napo Province,
Ecuador (0°24’S, 76°38’W; 280 m elev.) on 26 July 1974 with a Pentax SP-1000 with a
50 mm macrolens (Kodak Plus-X, natural light: %, sec £4.0).

Boyce A. DRUMMOND III, Natural Perspectives. P.O. Box 9061, Woodland Park,
Colorado 80866.

Journal of the Lepidopterists’ Society
43(3), 1989, 259

ANNOUNCEMENT

COLOR ILLUSTRATIONS IN THE JOURNAL

Many Lepidoptera are colorful animals, both as juveniles and adults, and black and
white illustrations rarely do them justice. Color illustrations accompanying some articles
in the Journal should enhance both the information content and the esthetic quality of
our publication. Fortunately, the cost of color printing has declined in recent years, now
making the use of color in the Journal financially feasible. Although author page charges
for color illustrations will always be significantly higher than regular page charges, the
Executive Council of the Society recently approved a policy of subsidizing part of the
cost of appropriate color illustrations in the Journal. As a result, authors are encouraged
to submit color illustrations for publication in the Journal.

The cost of color printing varies with the size and format of the color illustration, so
authors who wish to use color should contact the Editor before submission to discuss the
nature of the illustration, the special submission requirements, and the cost. Please note
that the newly established FEATURE PHOTOGRAPH category will also accept color sub-
missions, although no Society financial subsidy is available for color illustrations in this
category.

Society support for the appearance of color in the Journal will come from the Society’s
Color Illustration Fund, which gladly accepts donations both private and corporate.
Contributions to the Color Illustration Fund may be sent to the Treasurer of the Society.

Boyce A. DRUMMOND, Editor

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EDITORIAL STAFF OF THE JOURNAL

Boyce A. DRUMMOND, Editor
Natural Perspectives
P.O. Box 9061
Woodland Park, Colorado 80866 U.S.A.

Associate Editors:
M. DEANE Bowers (USA ), LAWRENCE F. GALL (USA),
ROBERT C. LEDERHOUSE (USA ), ROBERT K. RoBBINs (USA),
CHRISTER WIKLUND (Sweden)

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories
are Articles, General Notes, Technical Comments, Book Reviews, Obituaries, Feature
Photographs, and Cover Illustrations. Reviews should treat books published within the
past two years. Obituaries must be authorized by the President of the Society. Require-
ments for Feature Photographs and Cover Illustrations are stated on page 203 in Volume
42(3). Journal submissions should be sent to the editor at the above address. Short manu-
scripts concerning new state records, current events, and notices should be sent to the
News, June Preston, Editor, 832 Sunset Drive, Lawrence, Kansas 66044 U.S.A. Journal
contributors should prepare manuscripts according to the following instructions, and
submit them flat, not folded.

Abstract: An informative abstract should precede the text of Articles.

Key Words: Up to five key words or terms not in the title should accompany Articles,
General Notes, and Technical Comments.

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en-
tirely double-spaced, with wide margins, on one side only of white, letter-sized paper.
Titles should be explicit, descriptive, and as short as possible. The first mention of a plant
or animal in the text should include the full scientific name with author, and family.
Measurements should be given in metric units; times in terms of the 24-hour clock (0930 h,
not 9:30 AM). Underline only where italics are intended.

Literature Cited: References in the text of Articles should be given as Sheppard (1959)
or (Sheppard 1959, 1961a, 1961b) and listed alphabetically under the heading LITERATURE
CITED, in the following format without underlining:

SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London.
209 pp.

196la. Some contributions to population genetics resulting from the study of

the Lepidoptera. Adv. Genet. 10:165-216.

In General Notes and Technical Comments, references should be shortened and given
entirely in the text as P. M. Sheppard (1961, Adv. Genet. 10:165-216) or (Sheppard, P.
M., 1961, Sym. R. Entomol. Soc. London 1:23-30) without underlining.

Illustrations: Only half of symmetrical objects such as adults with wings spread should
be illustrated, unless whole illustration is crucial. Photographs and drawings should be
mounted on stiff, white backing, arranged in the desired format, allowing (with particular
regard to lettering) for reduction to fit a Journal page. Illustrations larger than letter-
size are not acceptable and should be reduced photographically to that size or smaller.
The author’s name and figure numbers as cited in the text should be printed on the back
of each illustration. Figures, both line drawings and photographs, should be numbered
consecutively in Arabic numerals; “plate” should not be employed. Figure legends must
be typewritten, double-spaced, on a separate sheet (not attached to illustrations), headed
EXPLANATION OF FIGURES, with a separate paragraph devoted to each page of illustrations.
Color illustrations are encouraged; contact editor for submission requirements and cost.

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for
tables should not be capitalized. Tabular material must be typed on separate sheets, and
placed following the main text, with the approximate desired position indicated in the
text. Vertical lines as weil as vertical writing should be avoided.

Voucher specimens: When appropriate, manuscripts must name a public repository
where specimens documenting identity of organisms can be found. Kinds of reports that
require vouchering include life histories, host associations, immature morphology, and
experimental enquiries.

Proofs: The edited manuscript and galley proofs will be mailed to the author for
correction of printer's errors. Excessive author's changes at this time will be charged to
authors at the rate of $2 per line. A purchase order for reprints will accompany proofs.

Page charges: For authors affiliated with institutions, page charges are $20 per Jour-
nal page. For unaffiliated authors, page charges are $10 per Journal page with a $50
maximum. Authors of Book Reviews and Obituaries are exempt from page charges.
Authors unable to pay page charges for reasons such as difficulties with foreign exchange
should apply to the editor for free publication. Four free publication slots are available
annually in the Journal.

Correspondence: Address all matters relating to the Journal to the editor.

PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A.

CONTENTS

PRESIDENTIAL ADDRESS, 1988: LEPIDOPTERISTS—-COLLECTERS AND
BIOLOGISTS?) “Jerry A.’ Powell... 0 Ui)

REPRODUCTIVE ENHANCEMENT BY ADULT FEEDING: EFFECTS OF
HONEYDEW IN IMBIBED WATER ON SPRUCE
BUDWORM. \ William E. Miller 00 a a

WORLD NUMBERS OF BUTTERFLIES. Oakley Shields

THE SPHINGIDAE (LEPIDOPTERA) OF BAJA CALIFORNIA,
Mexico. John W. Brown & Julian P. Donahue .W..

BIOLOGY AND IMMATURE STAGES OF SCHINIA MASONI (NOCTUI-
DAE). Bruce A. Byers 0005

GENETIC DIFFERENTIATION AMONG CALIFORNIA POPULATIONS OF
THE ANISE SWALLOWTAIL BUTTERFLY, PAPILIO ZELICAON
Lucas: Mark L. Tong < Arthur M. Shapiro ae

A NEW SUBSPECIES OF COENONYMPHA TULLIA (MULLER)
(NYMPHALIDAE: SATYRINAE) CONFINED TO THE COASTAL
DUNES OF NORTHERN CALIFORNIA. Adam H. Porter ¢ Ster-
ling O. Mattoon 00

A PROCEDURE FOR EXAMINING THE GENITALIC MUSCULATURE OF
LEPIDOPTERA. John A. De Benedictis & Jerry A. Powell .

GENERAL NOTES

On the location of some H. A. Freeman skipper holotypes. (Hesperi-
idae), Hugh:A. Freeman 00080) 2

Effects of handling on Euphydryas editha (Nymphalidae). Maria E. Orive
& John F. Baughman 00 Nes

Perforated cupola organs on larvae of Euselasiinae (Riodinidae) Donald J.
Flare gy 3 Oe EI sl NNT

BooK REVIEW

Catalogue of Lycaenidae & Riodinidae (Lepidoptera: Rhopalo-
cera). Donald, J Harvey soy jol a

Papillons et chenilles du Quebec et de lest du Canada [Lepidoptera and
larvae of Quebec and of eastern Canada]. Thomas C. Emmel ...........

Slug and nettle caterpillars: The biology, taxonomy and control of the Li-
macodidae of economic importance on palms in southeast Asia. Mare

Fe ESCO eee eto ee ce

The Guild handbook of scientific illustration. Charles L. Hogue .......

Butterflies of Nepal (Central Himalaya). Thomas C. Emmel & Oakley

Shields sisi a i UL OO NC
FEATURE PHOTOGRAPH

SPLIT-LEVEL DINING. Boyce A. Drummond LID wcccccccecccecenn

ANNOUNCEMENT
Color illustrations inthe) Journal Viigo

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

239

244

244

247

250

251

Volume 22 1968 Number 4

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN

In This Issue

EVOLUTION AND SIGNIFICANCE OF MULTIPLE PAIRING
FACTORS IN ECLOSION OF PIERIS RAPAE
BUTTERFLIES FROM COAHUILA
CONTINUOUS VARIATION IN CATOCALA
LIFE HISTORY OF CHLOSYNE FULVIA

(Complete contents on back cover)

15 November 1968

THE LEPIDOPTERISTS’ SOCIETY

EDITORIAL COMMITTEE

Jerry A. PoweEL., Editor of the Journal
Paut A. Oper, Assistant Editor
E. J. Newcomer, Editor of the News
| S. A. Hesse, Manager of the Memoirs
P. F. BELLINGER E. G. MUNROE C. L. Remincton’ F. T. THORNE

EXECUTIVE COUNCIL

F. Martin Brown (Colorado Springs, Colo.), President
E. B. Forno (Oxford, England), Ist Vice President

J. Kumescu (Linz, Austria), Vice President

H. StemMprrer (Paris, France), Vice President

Roy O. KenpAautut (San Antonio, Texas), Treasurer
Joun C. Downey (Carbondale, Illinois), Secretary

Members at large (three year terms): P. R. Euruicn (Stanford, Calif.), 1968
C. D. MacNemut (Oakland, Calif.), 1968 P.D. Syme (Sault St. Marie, Can.), 1968
D. R. Davis (Washington, D.C.), 1969 C. L. Hocue (Los Angeles, Calif.), 1969
F. T. TuHorne (El Cajon, Calif.), 1969 J. F. G. Crarxe (Wash., D.C.), 1970
H. K. Cxencu (Pittsburgh, Pa.), 1970 B. Wricur (Halifax, Nova Scotia), 1970

The object of the Lepidopterists’ Society, which was formed in May, 1947 and
formally constituted in December, 1950, is “to promote the science of lepidopterology
in all its branches, . . . to.issue a periodical and other publications on Lepidoptera,
to facilitate the exchange of specimens and ideas by both the professional worker
and the amateur in the field; to secure cooperation in all measures” directed towards
these aims.

Membership in the Society is open to all persons interested in the study of
Lepidoptera. All members receive the Journal and the News of the Lepidopterists’
Society. Institutions may subscribe to the Journal but may not become members.
Prospective members should send to the Treasurer full dues for the current year,
together with their full name, address, and special lepidopterological interests.
In alternate years a list of members of the Society is issued, with addresses and
special interests. There are four numbers in each volume of the Journal, scheduled
for February, May, August and November, and eight numbers of the News each year.

Active members—annual dues $6.00
Sustaining members—annual dues $15.00
Life members—single sum $125.00
Institutional subscriptions—annual $7.00

Send remittances, payable to The Lepidopterists’ Society, and address changes
to: Roy O. Kendall, 135 Vaughan Place, San Antonio, Texas, 78201, U.S. A.

The Lepidopterists’ Society is a non-profit, scientific organization. The office of
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520.
Second class postage paid at Lawrence, Kansas, U.S.A. 66044.

JOURNAL OF

Tue LEerpiporprTeERISTS’ SOCIETY

Volume 22 1968 Number 4

THE EVOLUTIONARY AND BIOLOGICAL SIGNIFICANCE
OF MULTIPLE PAIRING IN LEPIDOPTERA

Rocer W. PEASE, Jr.
Department of Biology, College of Wooster, Wooster, Ohio!

The frequency of pairing in Lepidoptera may be related to population
structure (Labine, 1964), to courtship behavior (Marshall, 1901) or
to the balance of a polymorphism in a mimetic population (Burns, 1966).
In this paper a sample of female Utetheisa ornatrix bella (1.) (Arctiidae )
from the polymorphic Florida population is analyzed by phentotpye and
pairing frequency for evidence of sexual selection; data taken from the
literature are summarized; and the importance of mating frequency for
the evolution of behavior and of population structure is discussed.

Studies of copulation have shown that in many Lepidoptera the male
constructs a sperm containing bag (spermatophore) in the genitalia
of the female (Callahan and Cascio, 1963: Callahan and Chapin, 1960;
Khalifa, 1950; Norris, 1932; Weidner, 1934). Mating refers to copula-
tion; successful mating requires transfer of at least one spermatophore.
Since spermatophores remain in the bursa copulatrix, the number of
successful pairings can be ascertained by dissecting the bursa and
counting spermatophores. The spermatophore is reported to disintegrate
as soon as it is formed in certain Microlepidoptera (Callahan and
Casico, 1964: 554). While spermatophore remnants are difficult to
count, the hardened duct (collum) through which sperm pass from
the spermatophore to the ductus seminalis is retained more or less
complete in Utetheisa ornatrix, and the number of colli is an index
of pairing frequency. At most one spermatophore is produced during
a single copulation of this species. Of twelve virgin females mating
once in controlled breeding experiments in 1961, all had one and only
one spermatophore in the bursa copulatrix.

In the pink bollworm, Pectinophora gossypiella (Saunders) (Gele-
chiidae), spermatophores were formed in 199 of 219 laboratory matings
of virgin females (Ouye, et al., 1965, experiment 1). Seventeen of the

1 Present address: 36 Lincoln Street, New Britain, Connecticut,

198 PEeAsE: Multiple pairing Vol. 22, no. 4

20 pairs which did not form spermatophores remained in copulo no
longer than 30 minutes. Thus, a certain minimum time is required for
spermatophore construction and no more than one spermatophore is
formed during a single copulation.

The distribution of spermatophore frequencies in a sample of 89
females of U. o. bella collected in Florida is tabulated in Table 1. One
bursa was lost in the process of dissection. Fifty-three males were
collected with the females. The mean number of spermatophores was
3.489 with standard deviation 2.656. One individual had copulated at
least 11 times, and eight had not mated successfully.

Before dissection, the condition (fresh vs. worn) of each specimen
was estimated on the basis of external appearance and graded A, B,
C, or D. Not unexpectedly, the mean number of spermatophores in-
creases from 1.455 for very fresh (A) individuals to 6.250 for very
worn (D) individuals (Table 2). Moths reared under constant con-
ditions (80°F, 12-16 hours light per day) usually mated at night al-
though a few males collected in the field mated in the afternoon by
artificial light. Diurnal mating may be an adaptation to the Florida
winter season with its cold nights and warm days. A refractory period
may follow copulation during which time the male will not mate suc-
cessfully (cf. Khalifa, 1950: 39, Galleria mellonella (1.) (Pyralidae);
Ouye, et al., 1965 experiment 2, Pectinophora gossypiella).

The sample was tabulated separately for five wing pattern characters
and tested for differences in mean spermatophore number between
categories (Table 3). The characters were subdivided into categories
as follows: (1) forewing ground color—yellow or orange, red orange and
orange red, or red; (2) the distribution of forewing ground color—colored
or streaked, intermediate and white; (3) black spotting on forewing—
spotted or unspotted; (4) hindwing black markings—wide, semi or
narrow; (5) hindwing ground color—red, pink and flush versus white
(Pease, 1968).

When the t-test was applied independently to each of the five char-
acters in Table 3, only the difference between the mean number of
individuals with spotted versus unspotted forewings was statistically
significant with probability less than .02 of a difference between the
means as large or larger. However, the probability is about % that
one or more of five independent tests is significant at the .05 level,
(1 — .95°). Since the statistical hypothesis was formulated after looking
at the data, the result does not favor a hypothesis that wing pattern is
‘a factor determining mating frequency. This is consistent with the
observation that this species mates at night and that the bright pigmenta-
tion serves as a warning stimulus (aposematic coloration) to potential

1968 Journal of the Lepidopterists’ Society 199

diurnal predators. Toxic substances have been found in the haemolymph
by M. Rothschild. The species exudes a frothy bubble at the tegulae
when seized suddenly (reflex bleeding).

The average number of spermatophores in the tiger swallowtail
(Papilio glaucus L.), a species with dimorphic females, was greater
for yellow females than for dark females in samples from Mountain
Lake, Virginia and Baltimore, Maryland (Burns, 1966). The resem-
blance of dark females to the unpalatable blue swallowtail (Battus
philenor (L.)) is believed to confer protection from predators whose
vision enables them to distinguish the two forms. The population fre-
quencies of dark and yellow females are affected by two antagonistic
forces of natural selection (an example of disruputive selection). An
extra-specific environmental factor, mimicry, favors the mimetic dark
female; an intra-specific factor, sexual selection, favors the yellow female.

The preference of the male tiger swallowtail, which is always yellow,
for the yellow female is relatively independent of the frequencies of
dark and yellow females. At Baltimore the frequency of dark females is
4483 and the mean number of spermatophores was .3366 greater in
yellow than in dark females. At Mountain Lake, Virginia where the
frequency of dark females is almost doubled (.8571) the difference in
mean spermatophore number between yellow and dark females is nearly
the same (.3889) (Burns, 1966 Tables 1 and 2).

The difference in means for spermatophores in dark and yellow forms
is significant at the 0.2 level for the sample from Mountain Lake, Vir-
ginia and is significant between the 0.2 and 0.3 level for the sample
from Baltimore County, Maryland (Appendix 1). The “true” difference
between the means is important to the theory of polymorphic populations.
The experimental biologist can determine the correct sample size to
prove or disprove the theory by using the data given below.

Population parameters for the frequency of dark and yellow forms
of the tiger swallowtail and the mean number of spermatophores for
each follow (Burns, 1966):

Mountain Lake, Virginia Baltimore County, Maryland
Mean number of Mean number of
Frequency of Spermatophores Standard Frequency of Spermatophores Standard
Female Type Per Female Deviation-(s) Female Type Per Female Deviation-(s)
Dark oT 1.694 781 448 1.538 .776
Yellow 143 2.083 1.379 002 1.875 .619
Combined - 1.750 .890 - 1.724 .702

Data

For an assessment of how many specimens should be collected from
each population so that the estimated difference in the average number

200 PEAsE: Multiple pairing Vol. 22)enom-

TABLE 1. DISTRIBUTION OF SPERMATOPHORES IN FIELD SAMPLES OF
VARIOUS LEPIDOPTERA AND EXPERIMENTS ON MATING BEHAVIOR

N—sample size; X—mean or average; s2—sample variance, mean square or square
_ of the sample standard deviation.

Number of spermatophores) No 07 01) 2 3) 455 6a 7S Sel Opie s2

Utetheisa ornatrix bella (L.) 88 8 12 15 201154423 3 1 3.4886 7.0573
( Arctiidae )—Archbold Biol.

Station, Lake Placid, Florida

(net collection) —

Pseudaletia
unipuncta (Haw. ) 417 182 107 76 36 14 2 - —-—-— —
( Noctuidae )—Louisiana
Callahan and Chapin (1960)
Table 1—p. 779
(light trap )

Peridroma saucia Hbn. 939 203 16 5 8 61-—----—- —-
(= margaritosa Haw. )

( Noctuidae )—Louisiana

Callahan and Chapin (1960)

Table 2—p. 780

(light trap? )

Heliothis zea ( Boddie ) 1295 519.455 227 77 16 1 —--—--—-— -— — .9336 .9338
(Noctuidae )—Louisiana

Callahan (1958 )

Table 6—p. 427

1.0384 1.3447

33805 .8525

(light trap )
N less than 2 2 or more
Euphydryas editha (Bdv.) 23 14 9
(Nymphalidae )—California
Labine (1964)
Neh Oa taal Dw ta es x s2
Battus philenor (L.) SOMO Line Gla prosumer eile oll 7/273 9545

(Papilionidae )—Mtn. Lake
Biol. Sta., Virginia
Burns, 1966—table 2.

Papilio glaucus L.
( Papilionidae )
Burns, 1966—table 2.

Mtn. Lake Biol. Station, Virginia
dark females TOP Ones Oy mola wile by ILGY4/4 .6095

yellow females 12 27,0833) egos
Total SECO Com ee OP Pe LOO 7922,

S
(ep)
XS)
bo
—
_

Baltimore County, Maryland
dark females 13) COR 8) WISE G2) = JS SSS Ss .6026
yellow females 16 1.8750 3833
Total 29 oO D2! Swine as > ie Gala 4926

S
pS
_
>)
NS)
|
|

1968 Journal of the Lepidopterists’ Society 201

TABLE 1 Continued

Pectinophora gossypiella—( Saunders )

1. Number of copulations (spermatophores not counted) during the lifetime of
individual pairs maintained in laboratory population cages.

val

No. of copulations No. of moths N (O) stir et ciety 2 aarti 1 ante
in Population
Cage: 6 @
Lukefahr and Griffin IL OO 22a Gao ad il 3500 5934
(1957, 1967)

Ouye, et al. (1964) Wee i OAS (OAs SO a Att a pte .9894 6343
Ouye, (in litt. 1967 )

s2

2. Number of spermatophores produced during lifetime exposure to moths of
opposite sex under laboratory conditions (Ouye, et al., 1965, experiment 4—
table 3; experiment 5—table 4, and Ouye, in litt. 1967).

No. of No. of

spermatophores Moths

formed in

during Pop. Cage

lifetime Ae TOMNG 20 geste 12 1 ene Gn SOW LO eat 52
Males S206 10 19) 1208 251 29 2OmiG 9 Oras) 4.952475 SiG
Females

Cw mC@rowdedus (axl L963) 49°74, 45029. = = 9 99A> 1186
GE mOncrowded 1 6 2668) 64) 63 69.346 (f= 1) 2:3995 1.4334
Combined (A+B) 496 1111315711456 8 2 —1 — — 2.2792 1.2993

* Amends N in experiment 4 and table, Ouye, et al., 1965.

3. Spermatophore formation during the first 24 hours after eclosion of females
(Ouye, et al., 1965, experiment 3 table 2, Ouye, in litt. 1967).
No. of spermatophores N 0 Hf 2 3 4 XxX s2
Time exposed to males
ir population cages

15-21 hours B13 = 98972224) 44> 16 2 9812 0176
9-15 hours Mey Meisy = 2) i + - 0763 TL
3-9 hours 16 14 i - J) - .2500 .6000

4. Distribution of spermatophore number in a sample of females collected at light
traps in the vicinity of Brownsville, Texas (Graham, et al., 1965; and Ouye,
in litt. 1967).
Namopispermatopnores N @O) 91 92 "3 4°15 6G xX s?
No. of moths STD) Prove US TOMER Son Bie: Bo veal 1.0911 .4067

of spermatophores lies within .05 of the “true” value 95% of the time,
see Appendix 2.

Disruptive selection in the tiger swallowtail may be an example of
evolutionary homeostasis at the population level; that is, constant intra-
specific factors of selection counterbalance variable extra-specific en-
vironmental factors and tend to restore primitive population conditions.

202 Pease: Multiple pairing Vol. 22, no. 4

TABLE 2. Utetheisa ornatrix bella In EACH OF FOUR GRADES OF
CONDITION WITH MEAN NUMBER OF SPERMATOPHORES PER FEMALE

Condition # Individuals Mean Number of Spermatophores
A pap 1.455
B 32 2.781
C 26 Ooi
D 8 6.250

If this is true, when the environment ceases to favor the evolutionary
novelty (the dark female), the force of sexual selection will restore a
uniformly yellow population. This hypothesis is consistent with the
observed correlation between the distribution of the blue swallowtail
and a high frequency of the dark female form of the tiger swallowtail.

Data on spermatophore frequency in Lepidoptera are summarized
in Table 1. The maximum number of spermatophores counted in a
female was 11 (Utetheisa ornatrix). The maximum number of sper-
matophores formed by a male was 11 (Pectinophora gossypiella—Ouye,
in litt. 1967).

The mating habits of the pink bollworm moth (Pectinophora gossy-
piella) have been thoroughly studied (Lukefahr and Griffin, 1957; Ouye,
et al., 1964; Ouye, et al., 1965; Graham, et al., 1965; Ouye, in litt. 1967).
Data are summarized in Table 1. Pairs of moths copulated an average of
.300 times during their lifetime in the experiments of Lukefahr and
Griffin and .989 times in the experiments of Ouye and his workers
(spermatophores were not counted). The two means are different
(Appendix 3).

Females mate successfully as many as four times during the first
24 hours after eclosion; males produce no more than one spermatophore
in a 24 hour period. The average number of spermatophores formed
during the life of a male is 4.252 under laboratory conditions. This is
almost double the lifetime average of 2.279 for the combined data of
females in crowded (A) and uncrowded (B) population cages. (Table
1, experiments 2 and 3) (Appendix 4).

Thus, while the female pink bollworm moth mates successfully more
often in one day, the male can mate successfully almost twice as many
times as the female in the moths’ lifetimes.

The greater number of spermatophores formed by the male is con-
sistent with a hypothesis that natural selection acts more strongly on
the male than on the female. Two factors reduce the male’s average
under the competitive conditions in the field. Males may compete more
actively for females than females compete for males (intra-specific
sexual selection). Extra-specific factors of natural selection sometimes

1968 Journal of the Lepidopterists Society 203

TABLE 3. NUMBER OF INDIVIDUALS AND MEAN NUMBER OF
SPERMATOPHORES IN EACH CATEGORY OF THE FIVE CHARACTERS
FOR WING PATTERN AND PIGMENTATION IN
Utetheisa ornatrix bella rrom FLORIDA

Mean Number of

Character Phenotype Number of Individuals Spermatophores
IL Colored/streaked 36 Sula)
Intermediate 22, 3.136
White 30 3.433
py) Yellow/orange 76 3.526
Red/orange red/red orange 10 3.100
(“Redless” Aberration)' 2} 4.000
3 Spotted 69 3.145
Unspotted 19 A731
4 Wide 55 3.618
Semi 30 3.367
Narrow 3 DROOS
5 Red/pink/flush 82 3.304
White 4 2.000
(“Redless” Aberration)* 9) 4.000

+ Specimens in which the red pigment is missing on both the upper and under surface of
the wings.

favor survival of the female at the expense of the male, as for example,
when the female is protectively colored. Thus, at the population level,
the male’s greater reproductive potential compensates for individual
competition among males and for the greater risk involved in being a
male. |

It seems intuitive that the competition for mates affects the distribu-
tion of spermatophores formed by the two sexes, perhaps, by increasing
the variation in the number of spermatophores formed by the males.
However, the design of a practical experiment to collect data, and a
method of analysis are a challenge to the ingenuity of the experimental
biologist.

In samples of Lepidoptera collected at light traps, the average number
of spermatophores varies from 1.0911 (Pectinophora gossypiella) to .3305
(Peridroma saucia Hbn. (Noctuidae) ). The average number of sper-
matophores in a collection made with a net was 3.4886 for Utetheisa
ornatrix. These data are not comparable to laboratory data because of
the mixed age distribution in feral populations.

Some calculations (by Graham, ef al., 1965) suggest that the first
mating of the pink bollworm is density dependent, but that multiple
paring is density independent. This hypothesis is based on a correlation
(or absence of) between the log of the number of moths collected in
light traps and (a) the proportion of moths which had mated, (b) the

204 Pease: Multiple pairing Vol. 22) aime

TABLE 4. SPERMATOPHORE NUMBER IN FALL-WINTER VERSUS THE
OVERWINTERING SPRING POPULATIONS OF D. plexippus IN
CALIFORNIA (DATA FROM TABLE 16, WiLLiAMs, et al., 1942) —

’ Date Location N 0 1 2 3. 2S aes

Oct. 1938—Feb. 1939. Pacific Grove 38 17 19 2 0 0 0 “605 (2353
San Diego
El Carrito

April 1939 San Francisco 5 0 O 2 (0) Po e2escote7300

proportion of once mated or multiply mated moths, (c) the mean
number of spermatophores for mated and unmated moths combined,
and (d) the mean number of spermatophores for mated moths.

Observations on the sex ratio, migration, spermatophore number,
adult activity and mating behavior of the winter population of the
monarch butterfly (Danaus plexippus (L.) (Nymphalidae) ) in Cali-
fornia are reported by J. A. Downes (Williams et al., 1942: 160-165).
No more than two spermatophores were found in any female until
after February (Table 4). Although both sexes migrate, the estimated
male: female ratio in the population remaining at Pacific Grove in
winter quarters was 1000 to 1 by the second week in May. Females
apparently migrate first. Some mating occurs in the hibernating popula-
tion at Pacific Grove (no reports from November to January, however)
even though egg follicles remain unripe until the last week in March.

Species in which both sexes pair several times contrast with those
in which the female is prevented from multiple insemination by a
sphragis or plug which the male constructs in the genitalia (Acraea :
Marshall, 1901, 1902, and Eltringham, 1912; Parnassius : Eltringham,
1925; Bombyx mori L. : Omura, 1938). Marshall hypothesized that
differences in courtship behavior separate sphragis building genera
(e.g., Acraea and Parnassius) from many other butterflies, “marriage
by capture” (Poulton, 1911) versus “marriage by courtship.” In other
words, sphragis construction complements a behavior pattern in which
the male “grapples” with the female after a rudimentary display; in
species with more complex behavior patterns, courtship may be term-
inated at any of several stages (e.g., Brower, et al., 1965).

Eltringham (1912) suggests that the sphragis may block the release
of a chemical attractant which serves to assemble males. If this hypo-
thesis is correct, the “capture” system may represent only the final
stages of courtship.

In terms of evolutionary potential, no matter how effective “no” signals

1968 Journal of the Lepidopterists’ Society 205

are, the rare male which succeeds in breaking through the defense
mechanisms of an already mated female will leave more offspring than
the male which does not mate under such circumstances. Unless there
is a selective advantage to multiple insemination such as a short life span
of sperm or a prolonged oviposition period, the balance between multiple
versus single copulation should occur when the effort required to copulate
with an already mated female will reduce, first, the probability of
insemination of unmated females and, second, the males contribution
to the gene pool of the next generation. For multiple insemination to
persist under such circumstances, the number of fertile offspring left
by the more versatile male should outnumber those left by the virgin
inseminator.

If this were not so, the evolution of more effective mechanisms against
multiple pairing would be expected.

A relationship between mating frequency and interpopulation gene
flow has been suggested (Labine, 1964—Euphydryas editha (Bdv.)
(Nymphalidae) ). Populations in which the female mates before mi-
gration and only once can be contrasted with those in which the female
mates several times and especially with endemic males after migration.
In the first instance, a migrating female produces offspring with a gene
complement derived exclusively from the parent population while all
offspring produced by migrant males mated to endemic females receive
half their genes from the parent population and half from the other.
In contrast, if both sexes mate after migration, essentially, all offspring
are interpopulation hybrids. Thus, multiple pairing may increase the
proportion of hybrid offspring among progeny of migrants and hasten
the breakdown of introduced gene complexes. On the one hand multiple
pairing may increase variation through recombination between the two
gene pools and thereby influence the speed of adaptive change. On
the other hand, if the crossing of the two gene pools proves deleterious
and non-adaptive, F; offspring of migrants will be at a competitive
disadvantage even though offspring homozygous for genes from the
parent population compete successfully in the new habitat. Thus,
hybridization is an effective strategy on the part of the endemic popula-
tion for increasing the probability of beneficial combinations, and at
the same time, serves to reduce the competitive advantage of a closely
related invading species provided that the reproductive potential of the
endemic population is great enough to prevent swamping.

ACKNOWLEDGMENTS

The author wishes to thank Professor L. P. Brower of Amherst College
whose stimulating discussion suggested the paper. The sample of

206 PEASE: Multiple pairing Vol. 22) neie4:

Utetheisa ornatrix bella was collected at the Archbold Biological Station
of the American Museum of Natural History during the tenure of an
NSF pre-doctoral Fellowship at Yale University.

LITERATURE CITED

ANDERSON, R. L. & T. A. BANcRoFT. 1952. Statistical Theory in Research. Mc-
Graw-Hill Book Company, Inc.

Brower, L. P., J. V. Z. BRowrEr, & F. P. Cranston. 1965. Courtship Behavior
of the Queen Butterfly, Danaus gilippus berenice (Cramer). Zoologica, 50:
1-39, 7 plates.

Bryk, F.. 1918. Griindzuge der Sphragidologie. Archiv fiir Zoologie, 11, No. 18: 7.

Burns, J. M. 1966. Preferential mating versus mimicry: disruptive selection and
sex-limited dimorphism in Papilio glaucus. Science, 153: 551-553.

CALLAHAN, P. S. 1958. Serial morphology as a technique for determination of
reproductive patterns in the corn earworm, Heliothis zea (Boddie). Ann. Ent.
Soc. Amer., 51: 413-428.

CaLLaHAN, P. S., & T. Cascio. 1963. Histology of the reproductive tracts and
transmission of sperm in the Corn Earworm, Heliothis zea. Ann. Ent. Soc.
Amer., 56: 535-556.

CALLAHAN, P. S., & J. B. Cuapin. 1960. Morphology of the reproductive system
and mating in two representative members of the family Noctuidae, Pseudaletia
unipuncta and Peridroma margaritosa, with comparison to Heliothis zea. Ann.
Ent. Soc. Amer., 53: 763-782.

CiLarKE, C. A., & P. M. SHEPPARD. 1962. Offspring from double matings in
swallowtail butterflies. Entomologist, 95: 199-203.

ELTRINGHAM, H. 1912. A monograph of the African species of the genus Acraea,
Fab., with a supplement on those of the Oriental Region. Trans. Roy. Ent. Soc.
London, 1912: 1-374 plus 16 plates.

1925. On the source of the sphragidial fluid in Parnassius apollo (Lep.) Trans.
Roy. Ent. Soc. London, 1925: 11-15.

Fraser, D. A. S. 1958. Statistics: an Introduction. J. Wiley & Sons, Inc., New
York.

GraHaM, H. M., P. A. Gricx, M. T. Ouye, & D. F. Martin. 1965. Mating
frequency of female Pink Bollworms collected from light traps. Ann. Ent. Soc.
Amer., 58: 595-596.

Kuauira, A. 1950. Spermatophore production in Galleria mellonella L. (Lepi-
doptera). Proc. Roy. Ent. Soc. London (A), 25: 33-42.

LaBINE, P. A. 1964. Population biology of the butterfly, Euphydryas editha. I.
Barriers to multiple insemination. Evolution, 18: 335-336.

LuKEFAHR, M., & J. GrirFin. 1957. Mating and oviposition habits of the pink
bollworm moth. J. Econ. Ent., 50: 487-490.

MarsHaL., G. A. K. 1901. On the female pouch in Acraea. The Entomologist,
34: 73-75.

1902. Five years observations and experiments (1896-1901) on the bionomics
of South African insects, chiefly directed to the investigation of mimicry and
warning colors. Trans. Roy. Ent. Soc. London, 50: 287-584 (pp. 539-540).

MicuaEL, H. 1923. Uber den Bau der Geschlechtsapparate und die Kopulation
von Bombyx mori. Arch. Naturg., 89(A): 25-52.

Norris, M. J. 1932. Contributions toward the study of insect fertility. I. The
structure and operation of the reproductive organs of the genera Ephestia and
Plodia (Lepidoptera, Phycitidae). Proc. Zool. Soc. London, 1932: 595-611,
Biapls:

Omura, S. 1938. Studies on the reproductive system of the male of Bombyx mori.
II. Post testicular organs and post-testicular behavior of the spermatozoa. J.
Faculty Agric. Hokkaido Imperial University, XL: 111-128.

1968 Journal of the Lepidopterists’ Society 207

OuyEe, M. T. 1967. Letter, February 1967 giving data on mating in Pectinophora
gossypiella.

Ouye, M. T., R. S. Garcia, H. M. Granam, & D. F. Martin. 1965. Mating
studies on the Pink Bollworm, Pectinophora gossypiella (Lepidoptera: Gele-
chiidae), based on the presence of spermatophores. Ann. Ent. Soc. Amer.,
58: 880-882.

OuyeE, M. T., H. M. Granam, C. A. RicoMonp, & D. F. Martin. 1964. Mating
studies in the Pink Bollworm. J. Econ. Ent., 57: 222-225.

Pease, R. W., Jr. 1968. Evolution and hybridization in the Utetheisa ornatrix
complex (Lepidoptera: Arctiidae). I. Inter- and intra-population variation and
its relation to hybridization. Evolution, 22 (in press).

Peterson, W. 1907. Uber die Spermatophoren der Schmetterlinge. Z. wiss. Zool.,
88: 117-130.

1928. Uber die Sphragis und das Spermatophragma der Tagfaltergattung Parnas-
sius. (Lep.) Deutsch. Ent. Zeitschr. (Berlin), 1928: 407-413.

Poutton, E. B. 1911. Observations on the courtship of Planemia alcinoe, Feld.
Proc. Ent. Soc. London, 59: xcev.

Travassos, L. 1946. Contribuicao ao conhecimento dos “Arctiidae” XI. Genero
“Utetheisa’ Hubner, 1819. Verificacao de “U. Pulchella” (L., 1958) Kirby
1892, no Nordeste do Brasil. Rev. Brasil. Biol., 6: 343-354.

Werner, H. 1934. Beitrage zur Morphologie und Physiologie des Genital-
apparates der weiblichen Lepidopteren. Zeitschr. fiir angew. Ent., 21: 239-290.

Wixurams, J. L. 1938. The mating of Ephestia kuehniella Zeller and its results
Ent. News, 49: 104-107, 121-126, 2 pls.

1939. The occurrence of spermatophores and their measurement in some British
Lepidoptera. Trans. Soc. Brit. Ent., 6: 137-148.

1941. The relation of the spermatophore to the female reproductive ducts in
Lepidoptera. Ent. News, 52: 61-65.

WituiaMs, C. B., G. F. Cockpit, M. E. Gisss, & J. A. Downes. 1942. Studies
in the migration of Lepidoptera. Trans. Roy. Ent. Soc. London, 92: 101-282,
2 pls.

APPENDIX
Statistical Notes (Anderson and Bancroft, 1952: Fraser, 1958)

1. Tests for equality of the average number of spermatophores in
samples of dark and yellow females of Papilio glaucus L. (Burns, 1966).

Hypothesis Test Degrees of Value of Value of Tabulated Significance Conclusion
Freedom Experimental Statistic Level
Statistic

Mountain Lake Biological Station, Virginia
Dark Yellow

Ppo=py Ut 71 11 .13496 1980<t’<2.201 .05 means equal
(Cockran and Cox)
(Note: a value of the experimental statistic as large or
larger is expected on the basis of chance alone about .8 of
the time).

Baltimore County, Maryland
Dark Yellow

Poy t 12; 15 1.080 2131<t < 2179 05 means equal
(Note: a value of the experimental statistic as large or

208 PeAsE: Multiple pairing Vol, 22aores

larger is expected on the basis of chance alone between
.7 and .8 of the time).

2. An approximate solution can be found using the following ex-
- pression for the confidence limits for the difference of two means
(Fraser, 1958: 281):

to; —is the t statistic with N-—2 degrees of freedom but evaluated
here for an infinite number of degrees of freedom since the
sample size is large but unknown.

—Sample size.

—frequency of dark females in sample.

—frequency of yellow females in sample.

—square root of the variance. Since the variance for spermato-
phore numbers may differ in the two types of females, an
approximate solution is obtained by using the population
variance calculated from the combined data.

Ia 1S) ay

The solution is obtained by choosing N such that

lt 1 i
sal ate YN = 05

Or

i) 3 pe al IL )
N = 400s?( 1.96) aos
The answer depends on the relative frequencies of dark and yellow
females in the sample. The smallest adequate sample is one in which
the frequencies of dark and yellow females are equal (i.e., D = Y = .5).
When equal numbers of both types of female are collected, 3028 females
comprise an adequate sample in Maryland and 4870 in Virginia.

3. Means differ at the .02 level of significance, but variances are
equal for the two sets of data.

Lukefahr and Griffin, 1957 N, = 100 X= 7300 Sep == be}

Ouye, et al., 1964 Ne 94 X = .989 $75 = 4005
Hypothesis Test Degrees of Value of Value of Significance Conclusion
Freedom Experimental Tabulated Level
Statistic Statistic

O17 =O 6~ F-test 93,99 1.070 1.59<?9.<1.87 .02 variances equal
ee fl, t-test 192 7.256 2:33 << %o.< 2:36 (02) meansmotscameall

1968 Journal of the Lepidopterists’ Society 209

4, Data for females in crowded (A) versus uncrowded (B) popula-
tion cages were tested for equality of means and variances. Combined
data for females under both conditions were compared with the data
- from males for equality of means and variances.

Crowded females (A) ING = 196 N= 2204 SIA
Uncrowded females (B) N; = 266 Xe = 2320 So Asa
Combined data for females No = 462 Ke 2.279 s?¢ = 1,299

Males Nar == 206 Xx == 4952, SoM =O
Hypothesis Test Degrees of Value of Value of Significance Conclusion
Freedom Experimental Tabulated Level
Statistic Statistic

we—onek vest 269,195 1.251 “128 <F 95 < 1.59 02 variances equal

app t-test 460 A10 2.33 < too<2:36 02) means equal

oc’=oy F-test 205,461 4.091 1.00<F 92<1.48 .02 variances unequal

oH = 205,461 11.664 2.33<t' o92.<2.35 .02 means unequal
(Cockran and Cox)

LEONARD STEVENS PHILLIPS (1908-1968 )

Leonard Stevens Phillips was born December 4, 1908 at Le Claire,
Iowa. He died suddenly in Chicago, Illinois, February 13, 1968. He was
the son of Clyde and Winifred Phillips. His marriage to Merle Olive
Garton took place May 26, 1937. She survives him; there were no chil-
dren.

He attended the public schools of Le Claire and received his B.A. de-
gree from the State University of Iowa in 1932. He did graduate work
there and at the Iowa State University of Agriculture, and received his
teacher's certificate from the Iowa State Teachers’ College in 1938.

He engaged in private business from 1936 to 1946, then became a
laboratory assistant in the Stritch School of Medicine, Loyola University,
Chicago, a position which he held until 1950. Following a period as
laboratory technician with Swift and Company in Chicago, he became
Assistant Biologist at the Illinois Institute of Technology Research In-
stitute. Here he was in charge of the animal room and worked on many
projects involving the use of small animals in behavioral studies and bio-
chemical research. In 1965 he joined the Loop City College of Chicago
as a laboratory assistant, and in 1967 returned to private business, in
which he was engaged at the time of his death.

Leonard was an active and enthusiastic collector of Lepidoptera. He
collected personally in every state of the continental United States and

210 Irwin: Phillips obituary Vol. 22, no. 4

maintained an active correspondence and exchange with fellow lepidop-
terists throughout the world. He contributed several papers to the pages
of this Journal; a bibliography is given below. His collection of some
5,600 specimens of worldwide Lepidoptera is being retained by his
- widow for the present, but will be presented to Buena Vista College,
Storm Lake, Iowa.

Among his other interests were collections of pressed plants, minerals,
and stamps; woodworking, and amateur art. He was active in Boy Scout,
boys’ club, and church work. He was a member of the Chicago En-
tomological Society, and had been a member of the Lepidopterists’
Society since 1948.

Leonard’s many friends and correspondents will miss his friendly, out-
going personality, his enthusiasm for his avocation, and his willingness
to be of service to others.

BIBLIOGRAPHY OF LEONARD S. PHILLIPS

1936. A red-bellied woodpecker. Iowa Bird Life, March 1936.

1955. A Papilio flight pattern. Lepid. News, 9: 143.

1956. Monarchs numerous in an Iowa ravine. Lepid. News, 10: 44.

1960. Fluorescence in the colors of certain Lepidoptera observed under ultraviolet
light. J. Wepid: Soe, 13: 73—17.

1961. Nymphalis j-album captured at fluorescent light in Chicago. J. Lepid. Soc.,
15: 101.

1965. Flight habits of Boloria toddi. J. Lepid. Soc., 19: 104.

1966. Nymphalis californica in Illinois and Iowa. J. Lepid. Soc., 20: 124.

Roperick R. Irwin, 24 East 99th Place, Chicago, Illinois.

1968 Journal of the Lepidopterists’ Society DALI

THE EFFECT OF BAROMETRIC PRESSURE AND OTHER
FACTORS ON ECLOSION OF THE CABBAGE BUTTERFLY
PIERIS RAPAE (PIERIDAE)

JouN M. KoLyEerR AND HaRoLp B. PALMER
50 Chimney Ridge Drive, Convent, New Jersey, U.S.A.

The literature describes various effects of barometric pressure on
insects. For example, there is evidence that slightly reduced pressure
increases the rate of development of insects while slightly increased
pressure has no positive influence (Wellington, 1946). Pieris rapae (L.)
is said to lay more eggs when the barometric pressure is low (Stephens
and Bird, 1949), while low pressure appears to be disadvantageous to
at least one insect activity in that a slight depression is reported to
prevent the silkworm from secreting silk (Markovic-Giaja, 1957).

Of particular interest is the assertion that lowering of the barometric
pressure is necessary for successful eclosion of butterflies (Pictet, 1933),
including Pieris rapae and Pieris brassicae (L.); it was recorded that
91.3% of 1758 pupae eclosed during atmospheric depression. Also,
certain Lepidoptera are supposed to be so sensitive to the effect of
barometric pressure that they eclose when the pressure falls only 1.7—-3.4
mm of mercury below the daily maximum (Mell, 1939).

The present work is in large part an evaluation of the effect of pressure
on eclosion of the subject species by means of observations at ambient
conditions and also by controlled experiments.

ECCLOSION UNDER AMBIENT CONDITIONS

The pressure in the central area of a typical “high” is about 765-773
mm, while a “low” is normally 743-750 mm (Anonymous, 1960). In
the New York City area the mean pressure in summer generally is about
762 mm.

The data of Table 1 are for successive broods in a culture started
with eggs laid by Pieris rapae taken at Flemington, New Jersey on
May 1, 1965. It is apparent that the barometric pressure was relatively
high, and certainly not considerably depressed, during eclosion. The
impression is that the time of eclosion was controlled simply by the
time required for maturation of the pupa (7-10 days, approximately ).
The possibility would seem to remain that eclosion might be delayed
by an unusually high pressure, e.g. 788 mm as recorded in New York
City for record highs in 1927 and 1949 (Hansen, 1961), and the effect
of extremely high pressures was studied m the experiments discussed
below.

Dale, KOLYER AND PALMER: Eclosion of Pieris Vol. 22) nem

TABLE 1. BAROMETRIC PRESSURE DURING ECLOSION UNDER
AMBIENT CONDITIONS

Conditions during eclosion?

Num- Pupa- Relative Barometric
ber tion Eclosion Date eclosion humidity Temperature pressure

eclosed (day) (day) started (%) (AR) (mm )
38 0-6 9-15 June 6, 1965 38-55 75-90 754-766"
51 0-5 14-18? July 11, 1965 49-62 77-87 758-766
37 0-3 7-10 Aug. 11, 1965 48-60 78-89 761-765
16 0-5 9-13 Sept. 16, 1965 60-65 72-84 760-770

1 The barometric pressure readings are corrected to O°C and sea level.
2The pressure was in the 759-766 mm range except for the last day.
3'The pupae had been refrigerated from day = 6 to day = 12.

ECLOSION UNDER CONTROLLED CONDITIONS—EXPERIMENTAL

Rearing of larvae.—Larvae were reared in cardboard boxes with gauze
windows and fed cabbage leaves from refrigerated heads as in previous
work (Kolyer, 1966).

The pupae used for experiment 1 derived from eggs laid on August
6 and 7, 1966 by females taken at Berkshire Valley and Morristown,
New Jersey. Pupation took place 17-21 days after the inception of
hatching on August 9. The larvae were reared in a room at 73-91°F
and 40-70% relative humidity.

The pupae for the remaining experiments were reared from eggs
obtained from N. R. Spencer of the U. S. Department of Agriculture
(see Acknowledgment). A minor portion of the final-instar larvae
evidenced black spots on the integument, but fortunately there was no
effect on pupation or eclosion. Pupation took place 26-29 days after
the inception of hatching on April 7, 1967. The room was at 67-79°F
and 25-34% relative humidity during the larval period. Incidentally,
two male adults from this brood were of the canary-yellow form (one
eclosed in experiment 2 and one in experiment 5).

TABLE 2. SUMMARY OF EXPERIMENTAL CONDITIONS

Experiment Relative
Number Pressure (mm) Photoperiod* Humidity (%)
1 735-740 and 765-770? no AS vat oe
2 735-740 and 765-770? yes Ar wate es
5 690-700 and 790-800" no AS vation
4 825-830 yes AB at Wb
5 ambient yes Si satamonls
6 ambient yes 43 at 72°

1 Diffuse sunlight (from windows with southern exposure) from 7:27 AM to 5:03 PM.
2Cycled from one range to the other every 4 hours and 48 minutes (five times per day).

1968 Journal of the Lepidopterists’ Society

Fig. 1. Apparatus used for experiments 1-4.

Final-instar larvae were sorted quite reliably into males and females
by means of the testes visible in the male as done in previous work
(Kolyer, 1966), and the sexes were allowed to pupate in separate boxes.

Apparatus and procedure—The apparatus shown in Figure 1 was
used for experiments 14 (summarized in Table 2). This consisted
simply of two one-liter flasks connected through their side-arms and
fitted with a stopcock (and pinch clamp, not shown, for perfect seal)
and mercury-containing manometer open to the atmosphere. Gauze
was provided in the flasks so that the emerging butterflies could climb
up and expand their wings.

Vials (approximately one inch inside diameter) containing saturated
potassium carbonate solution in contact with solid potassium carbonate
hydrate were suspended inside the flasks to regulate the relative hu-
midity. A value of 42.8% relative humidity at 77°F is given by Stokes
and Robinson (1949), and 43% at 72°F was found experimentally by
confining a calibrated hygrometer with the potassium carbonate system.
The capacity of the system to absorb water was demonstrated by intro-
ducing one milliliter of water into a one-gallon jar containing a hygrom-
eter and the regulating system in a 2.5 inch diameter dish. In about five

214 KOLYER AND PALMER: Eclosion of Pieris Vol. 22) mom

OD 85
30 B ee
a “1
=<
e =
Py)
——>> >
—_—
© 80 ¢
25 —O-O a
oO me
e
75
ia
Ww 20 0
(@)
—
O
Lu
oO
=a
¥ @
== 45 O
&
= D
= G
=)
O

8 9 10 11
TIME, DAYS FROM START OF PUPATION
EXPLANATION OF GRAPH |

Cumulative number eclosed vs. day from start of pupation for experiment 1.
A record of temperature is included.

hours the relative humidity had risen to a maximum of 81% and at 50
hours it had fallen back to 50%; in the absence of the system 100%
relative humidity was attained about 10 hours after adding the water.
Pupae, detached by clipping the silken girths and pulling free from
the silk button at the caudal end, were dropped into the flasks (males
in one flask, females in the other) three days before eclosion began,
and maintenance of conditions as defined in Table 2 was initiated.

1968 Journal of the Lepidopterists’ Society PAS

25

20

ECLOSED

15

CUMULATIVE NO.

| J

TAL 12 13 14
TIME, DAYS FROM START OF PUPATION

EXPLANATION OF GRAPH 2
Cumulative number eclosed vs. day from start of pupation for experiment 2

In the graphs and tables each day is arbitrarily taken to begin at
2:39 AM and is divided into five equal periods (beginning at 2:39 AM,
7:27 AM, 12:15 PM, 5:03 PM, and 9:51 PM). At the start of each period
the barometric pressure and the room temperature were noted, and the
pressure was cycled in the case of experiments 1-3. The object of cycling
was to give the pupae a choice of high or low pressure every five hours.

216 KOLYER AND PALMER: Eclosion of Pieris Vol. 22, mom

25

20

ECLOSED

—
ol

10

CUMULATIVE NO.

11 12 13 14 5

TIME, DAYS FROM START OF PUPATION

EXPLANATION OF GRAPH 3

Cumulative number eclosed vs. day from start of pupation for experiment 3.

For reference, the first period of the 8th day on Graph 1 was a high cycle,
as was the first period of the 11th day for Graphs 2 and 3.

The data are presented in Graphs 1-7. The temperature record for
experiment | is included in Graph 1, while the temperature record for
experiments 2-6 (all done at the same time) and the barometric pressure
record applicable to experiments 5 and 6 are shown in Graph 7.

1968 Journal of the Lepidopterists’ Society ZN.

jo)

5 Le{GIL(O}S E10)

CUMULATIVE NO

i 12 13 14 15

TIME, DAYS FROM START OF PUPATION

EXPLANATION OF GRAPH 4
Cumulative number eclosed vs. day from start of pupation for experiment 4.

The barometric pressure readings in Graph 7 were used to adjust
the pressure in the flasks. For example, if the atmospheric pressure
was 760 mm and a pressure of 740 mm was desired in the flask, a
differential of 20 mm on the manometer was produced by drawing out
air by lung power and closing off the stopcock and the pinch clamp.
A rubber bulb was used to pressure the flasks in the high cycles. To
correct the pressure readings, which were taken at Convent, New Jersey
at an elevation of 290 feet, to 0°C and sea level, approximately 4 mm
must be added (Perry, 1950). The flask pressures listed in Table 2 are
ranges because some fluctuations necessarily accompanied temperature
variations.

In experiments 5 and 6, wide-mouth jars (approximately 3.5 inches
inside diameter by 5 inches deep) contained the pupae and humidity-
regulating system. The jars were closed tightly enough to maintain
regulated humidity but not to hold a pressure differential relative to
the atmosphere. The potassium carbonate system was used in experi-
ment 6, while in experiment 5 a saturated sodium carbonate solution in
contact with solid hydrated sodium carbonate was included. The
sodium carbonate system gave 87% relative humidity at 75°F experi-
mentally; Lange (1946) lists 92% at 65°F.

ECLOSION UNDER CONTROLLED CONDITIONS—RESULTS AND DISCUSSION

The five factors considered were barometric pressure, light, tempera-

218 KOLYER AND PALMER: Eclosion of Pieris Vol. 22, no. 4

TABLE 3. ECLOSION DURING ALTERNATING CYCLES OF HIGH AND LOW
PRESSURES

Number Eclosed

-Experiment Theoretical
Number Pressure Male Female Total Random Distribution
il low 7 16 23, 25
high 12 15 Dil 5)
2 low 18) 13 26 24
high itil ll YD) 24
3 low 7 is: 20 24
high y/ iLL 28 24

ture, relative humidity, and sex. The results are discussed in terms of
each of these.

Barometric pressure—The data summarized in Table 3 show no
significant trend with respect to barometric pressure. By the chi-square
method of testing goodness of fit (Sinnott and Dunn, 1939) it is found
that the differences observed in the total numbers eclosed at high vs.
low pressure can very possibly be explained by chance alone (Table 4).

In experiment 3 the low pressure was below one of the lowest on
record (721 mm) for New York City (Hansen, 1961), and the high
pressure was above a record high (788 mm). In experiment 4, in which
the pressure was held constant at an abnormally high level, the butter-
flies had no notable difficulty in eclosing (one unexpanded and one
with shriveled forewing vs. two imperfect specimens in experiment 1,
three in experiment 2, and one in experiment 5). Also, eclosion in
experiment 4 was not delayed and proceeded over about the same time
interval as in experiments 2, 3, 5, and 6.

In experiment 1 the larvae were diseased, and only 11 specimens (4
males and 7 females) expanded normally. It is interesting that 7 of
these (all but 4 of the females) eclosed during the high pressure cycle,
showing the lack of advantage of low pressure even when expansion
ability was marginal.

TABLE 4. SIGNIFICANCE OF DATA (TOTAL ECLOSED) OF TABLE 3

Experiment Approximate Probability of
Number x? Observed Deviation by Chance Alone’
1 320 58
2 333 57
) 138 25

1For one degree of freedom.

1968 Journal of the Lepidopterists’ Society 219

TABLE 5. ECLOSION VS. TIME OF DAY

Experiment 1] Experiment 2 Experiment 3
Period' Male Female Total Male Female Total Male Female Total
1 0 4 4 1 D, 6: ik 4 5
Dy i 2) 9 21 18 39 7 i 14
3 6 14 20 2 2 4 7 9g 16
4 5 6 ILI 0 2 2 4A 3 7
5 iL 5 6 0 0 0 5 1 6
Experiment 4 Experiment 5 Experiment 6

Period’ Male Female Total Male Female Total Male Female Total

if 0 0 0 0 1 1 2 0 2
2 7 10 U7 9 6 15 4 7 ee
3 2 2 4 il + 5 3 2 bY
4 0 0 0 0 0 0 1 0 1
5 0 0 0 0 0 0 0 2 2

1The day was divided into five equal periods beginning at 2:39 AM.

Pupae which are prepared to eclose appear to be able to wait several
hours for the stimulus of light and so might have been expected to take
advantage of the occurrence of low pressure cycles every four hours
and 48 minutes if reduced pressure also is a stimulus.

As expected, there was no real correlation of eclosion with ambient
barometric pressure in experiments 5 and 6. For example, eclosion in
experiment 5 was concluded at the high point of the pressure record.
In view of the data of Table 1 and the results of experiments 1-4, the
fortuitous drop in pressure seen in Figure 7 probably had no bearing
on eclosion.

Light—In experiments 1 and 3 the pupae were kept in darkness
except for brief intervals of light when the pressure was adjusted
between periods. The data (Table 5) indicate a preference for periods
2-4 (7:27 AM—9:51 PM) in experiments 1 and 3, and the chi-square
test (four degrees of freedom) shows less than a 5% probability that
the results are due to chance alone. It is possible that temperature
variation and/or brief admission of diffuse sunlight at the beginning
of the favored periods was responsible. However, David and Gardiner
(1962) report a rhythm of eclosion for Pieris brassicae in darkness.

In experiments 2, 4, 5, and 6, in which diffuse sunlight was available
during periods 2 and 3, there resulted a very marked preference for
eclosion during the photoperiod. Only 40% of the pupae should have
eclosed during the photoperiod by chance, while the result was 90%.
The chi-square test shows that in all cases the deviation from chance

220 KOLYER AND PALMER: Eclosion of Pieris Vol. 22. mone

Q
Lij
Tp)
ro)
wi
oO
LJ
10
oO
z
bij
>
-_
<
ahi
=
= 5
oO

14 is
TIME, DAYS FROM START OF PUPATION

EXPLANATION OF GRAPH 5
Cumulative number eclosed vs. day from start of pupation for experiment 5.

distribution is highly significant. Graphically, the stepwise nature of
the curves for experiments 2, 4, 5, and 6 is conspicuous.

The impression is that the mature pupa can wait several hours for the
arrival of the photoperiod before eclosing, and it is reported (Mell,
1939) that butterflies generally eclose in the early morning and that
the coming of light seems to be the stimulus. In a study including
Pieris rapae, 85.9% of 1758 pupae eclosed from 8 AM to 6 PM and
the remaining 14.1% at night (Pictet, 1933).

Temperature.—In experiments 2-6 the temperature varied over a

1968 Journal of the Lepidopterists’ Society OOK.

ECLOSED

CUMULATIVE NC.

12 13 14 NS

TIME, DAYS FROM START OF PUPATION

EXPLANATION OF GRAPH 6
Cumulative number eclosed vs. day from start of pupation for experiment 6.

range of 9°F (Graph 7). Low points occurred at the beginning of
period 1 (2:39 AM) in all cases, and high points occurred at the
beginning of periods 3 or 4. The fluctuations were similar in magnitude,
though less regular, in experiment 1 (Graph 1).

It would seem that the effect of light as a factor is predominant in
the experiments having a photoperiod (2, 4, 5, and 6), but some tempera-
ture fluctuation may be required for photoperiod to promote eclosion;
David and Gardiner (1962) report that for Pieris brassicae eclosion
takes place in the dark period with photoperiod 6 AM—10 PM when
the temperature is constant but that eclosion is delayed until moming
or afternoon when the temperature fluctuates.

Relative humidity —Very high humidity may delay eclosion of certain
moths (Mell, 1939). However, in experiment 5 eclosion at 87% relative
humidity was certainly not hindered, nor was it judged significantly
accelerated or delayed vs. eclosion at 43% relative humidity. As in the
other experiments, light appeared to be the dominant factor (though
possibly through interaction with temperature fluctuation patter).
Eclosion of the males was completed during the highest phase (763-766
mm, corrected to 0°C and sea level) of the ambient pressure record,

222 KOLYER AND PALMER: Eclosion of Pieris Vol. 22, smone4

760

Ss
ol
ol

BAROMETRIC PRESSURE, MM
~S
Oo
fo

745

75

TEMPERATURE, CF

70

11 12 13 14 15
TIME, DAYS FROM START OF PUPATION

EXPLANATION OF GRAPH 7
Record of barometric pressure and temperature during experiments 2-6.

and it is suspected, though it was not experimentally demonstrated, that
controlled variations of the pressure would have had no more effect
at 87% relative humidity than they were found to have in experiments
1-3 at 43%.

At a lower relative humidity under ambient conditions (23-34%
relative humidity, 72-76°F), five males eclosed at 764, 763, 761, 766,
and 759 mm (corrected to 0°C and sea level). Again, reduced pressure
was not required.

1968 Journal of the Lepidopterists Society 223

Sex.—In Pieris napi (L.) and P. bryoniae (Ochs.) the males are said
to tend to emerge before the females (Bowden, 1953). However, in
experiments 1-3 the sexes eclosed over almost exactly the same time
interval and at about the same rate; if anything, the females tended to
eclose a little earlier in experiments 2 and 3. In experiments 4 and 6
the females tended to eclose considerably earlier, though the significance
of this is dubious because of the limited numbers involved. Certainly,
there was no trend for the males to emerge before the females in any
of the experiments. This conclusion is, of course, applicable only to the
specific conditions of the tests.

CONCLUSION

Care must be taken in drawing generalized conclusions from the data
because of the possibility of the interaction of factors, the possibility
of discontinuities in cause-effect relationships, and the possible effect
of rate of change of variables. For example, at some critical values for
the three variables the relative humidity, temperature, and barometric
pressure might interact so that the pressure does influence eclosion. Or
the effect of pressure might be nil at slight or major depressions but
unexpectedly apparent at medium depressions. Or eclosion might be
promoted not simply by low barometric pressure but by the dynamic
factor of falling pressure.

Therefore, the present work does not prove that barometric pressure
cannot influence eclosion of Pieris rapae but only that it does not
influence eclosion under specific ambient conditions or in certain
controlled environments. In fact, even the strain of a species conceivably
could have an effect. Still, it seems a reasonable conclusion that light
is a principal factor (as appears in the literature) and that barometric
pressure is not a significant factor under certain typical summer ambient
conditions or when cycled between extreme values, or held at a constant
high value, in experiments at constant relative humidity, slightly fluctuat-
ing temperature (9°F maximum variation), and controlled photoperiod.

The indication is that the time of eclosion was controlled to the nearest
day or so simply by the rate of development of the pupa (in turn con-
trolled by the temperature history; David and Gardiner (1962), for
example, list a pupal period for Pieris brassicae of 40 days at 54.4°F
and only 7.5 days at 86°F). Then, under the particular conditions of
the tests, which involved some temperature fluctuation, the mature
pupa showed a strong tendency to await the coming of light as stimulus
for eclosion. No obvious effect of sex, relative humidity, or barometric
pressure upon this process was seen.

224 KOLYER AND PALMER: Eclosion of Pieris Vol. 22, now

SUMMARY

Under typical summer conditions, pupae of Pieris rapae (L.) were
observed to eclose at up to 770 mm barometric pressure with no apparent
preference for atmospheric depression, although reduced pressure has
been described in the literature as a requirement for eclosion of certain
Lepidoptera.

Under controlled conditions, male and female pupae were studied
separately. The relative humidity was held constant, the temperature
fluctuated over a maximum of 9°F with minima at night, and a photo-
period (diffuse sunlight) of either 0 or 9.6 hours was provided. The
barometric pressure was held at a constant high level (approx. 830 mm)
or cycled from approx. 735 to 770 mm or from approx. 690 to 800 mm
five times per day to give eclosing pupae a choice of high or low pressure.
No significant dependence of eclosion on barometric pressure or sex
was found, but light stimulated eclosion under the test conditions. There
was no notable difference in eclosion at 87% vs. 43% relative humidity.

The indication is that the time of eclosion was controlled within a
day or so simply by the rate of development of the pupa (dependent
on temperature history), and that light, if available, then was the im-
mediate stimulus for eclosion in a very significant proportion of cases.
It is emphasized that conclusions must be confined to the particular
experimental conditions.

ACKNOWLEDGMENT

We wish to acknowledge the aid and advice of Mr. N. R. Spencer,
U. S. Department of Agriculture, Entom. Res. Div., Columbia, Missouri,
who kindly supplied us with eggs from his Pieris rapae culture.

LITERATURE CITED

Anonymous. 1960. McGraw-Hill Encyclopedia of Science and Technology, Vol. 1.
McGraw-Hill Book Co., Inc., New York City. (p. 640).

BowbeNn, S. R. 1953. Timing of imaginal development in male and female hybrid
Pieridae (Lep.). Entomologist, 86(11): 257-264.

Davin, W. A. L., & B. O. C. Garpiner. 1962. Observations on the larvae and
pupae of Pieris brassicae L. in a laboratory culture. Bull. Ent. Res., 53(2):
417-436.

Hansen, H. (Editor). 1961. The World Almanac. New York World-Telegram
and The Sun, New York City. (p. 445).

Kotyer, J. M. 1966. The effect of certain environmental factors and chemicals
on the markings of Pieris rapae (Pieridae). J. Lepid. Soc., 20(1): 13-27.
Lance, N. A. 1946. Handbook of Chemistry (6th Ed.). Handbook Publishers,

Inc., Sandusky, Ohio. (p. 1397).

Markovic-Graja, L. 1957. Contribution to the physiology of the silkworm. Acta
Physiol. et Pharmacol. Neerland, 6: 339-345.

Mei, R. 1939. Beitrige zur Fauna Sinica. XVIII. Der Schliipfmoment siid-
chinesischer Lepidopteren. Zeitschr. Morph. u. Okol. Tiere, 35(1): 139-168.

1968 Journal of the Lepidopterists’ Society pep,

Perry, J. H. 1950. Chemical Engineers’ Handbook (3rd Ed.). McGraw-Hill
Book Co., Inc., New York City. (p. 365).

PicteT, A. 1933. Les éclosions de papillons et la météorologie. Lambillionea,
33(4): 89-97; (7): 158-164.

SINNOTT, E. W., & L. C. Dunn. 1939. Principles of Genetics (3rd Ed.). McGraw-
Hill Book Co., Inc., New York City. (p. 77).

STEPHEN, W. P., & R. D. Birp. 1949. The effect of barometric pressure upon
oviposition of the imported cabbageworm, Pieris rapae L. Canad. Ent., 81(5):
132.

Stokes, R. H., & R. A. Ropinson. 1949. Standard solutions for humidity control
at 25°C. Ind. Eng. Chem., 41(9): 2013.

WELLINGTON, W. G. 1946. The effects of variations in atmospheric pressure upon
insects. Can. J. Res., Sect D Zool. Sci., 24(3): 51-70.

FOODPLANTS OF CALLOPHRYS (INCISALIA) IROIDES

Jerry A. POWELL
University of California, Berkeley

In contrast with other members of the subgenus, which are restricted
in host selection, I. iroides (Boisduval) is polyphagous. The diverse host
plants credited to this West Coast butterfly are summarized by Clench
(1961), who indicates some of the early records are doubtful.

Recorded foodplants for iroides include “young apples” (Malus, Rosa-
ceae ) in British Columbia (Bethune, 1904) and both Ceanothus (Rham-
naceae) and Cuscuta (Polemoniaceae ) in southern California (Comstock
and Dammers, 1933). Field oviposition was observed and larvae reared
on Cuscuta, a leafless, parasitic plant which lacks chlorophyll. Clench
also lists Gaultheria and Arbutus (Ericaceae) as hosts but does not cite
the original source of these records.

Recent investigations during California Insect Survey activities con-
firm use of two of these foodplants in central and southern coastal Cali-
fornia and have disclosed the use by I. iroides of Chlorogalum pomeri-
dianum, a monocotyledenous plant in the foothills of the Sierra Nevada.

A nearly mature larva was collected on Arbutus menziesii at China
Camp, Marin County, June 3, 1964, from which an adult iroides was
reared, emerging on April 19, 1965. Ceanothus probably is commonly
used by iroides over much of its range. One larva was swept from C.

226 Powe: Callophrys iroides foodplants Vol. 22, no. 4

cuneatus near Middletown, Lake County, on May 14, 1966. It fed on the
green fruit of this plant in the laboratory prior to pupation in mid-June.
Emergence did not occur, but a fully developed adult was dissected
from the pupal shell in May, 1967. Additional larvae were taken on an
unidentified species of Ceanothus at the north end of Casitas Reservoir,
Ventura County, on March 15, 1967, by P. A. Opler. One iroides emerged
April 16, 1967, suggesting, as did Comstock and Dammers’ observations,
that populations in southern California develop two spring generations.

Six larvae of varying ages were found on Chlorogalum pomeridianum
(Liliaceae) about two miles south of Grass Valley, Nevada County, Cali-
fornia, on July 3, 1967. They were located on lateral branches in the
spreading inflorescences, feeding on the flowers and buds. Pupation oc-
curred by late July and adults emerged April 21 and May 15, the follow-
ing year.

Incisalia iroides was early reported to feed on Sedum (Crassulaceae )
in California. Comstock (1927) states that the larva and pupa were
described by Henry Edwards (1878) from this plant. Possibly this record
refers to I. fotis (Strecker), which is represented by a recently redis-
covered Sedum-feeding race near San Francisco, where Edwards did
much of his work. Adults of the two butterflies are similar in appear-
ance.

It is curious that I. augustinus (Westwood) of the eastern United States
is restricted to Ericaceae (Cook & Cook, 1904, 1906; Clench, 1961), yet
is considered by Clench to comprise with iroides a single widespread,
polytypic species. Other workers have treated augustinus and iroides as
closely related species, each with its own subspecific diversity (e.g., dos
Passos, 1964). Perhaps further data on host selection by the various
races of this complex will help clarify taxonomic relationships.

LITERATURE CITED

BETHUNE, C. J. S., 1904. Editor's footnote. Canad. Ent., 36: 136.

Ciencu, H. K., 1961. Family Lycaenidae. Blues and metal marks. 175-251, in:
Ehrlich, P. R. and A. H. Ehrlich, How to know the butterflies. Wm. C. Brown
Co., Dubuque, Iowa.

Comstock, J. A., 1927. Butterflies of California. Publ. by author, Los Angeles,
Calif., 334 pp.

Comstock, J. A. AND C. M. Dammers, 1933. Notes on the life histories of four
Californian lepidopterous insects. Bull. So. Calif. Acad. Sci., 32(2): 77-83.

Cook, J. H. anp H. Coox, 1904. Notes on Incisalia augustinus. Canad. Ent., 36: -

136.
1906. Studies in the genus Incisalia Il. Incisalia augustinus. Canad. Ent., 38:
214-217.
pos Passos, C. F., 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. Soc.,
Mem. 1, 145 pp.

Epwarps, H., 1878. Pacific Coast Lepidoptera [27: 2]. Proc. Calif. Acad. Sci.
[ Not seen].

i)
i)
~I

1968 Journal of the Lepidopterists’ Society

BUTTERFLIES FROM COAHUILA, MEXICO

Harry K. CLENCH
Camegie Museum, Pittsburgh, Pennsylvania

Dr. C. J. McCoy is Assistant Curator of Amphibians and Reptiles at
Carnegie Museum. In June, 1966, accompanied by Mr. Arthur Bianculli,
he made a trip! to central Coahuila to collect and study these animals.
Dr. McCoy also maintains a considerable interest in butterflies and,
as his regular work permitted, made a collection of them in the area.
Because almost nothing is known about Coahuilan butterflies, a list of
his captures should be useful.

The collection was made in three different localities, all in the vicinity
of Cuatro Ciénegas de Carranza (about 70 airline km west of Monclova),
central Coahuila, Mexico. A sketch map of the area, additional de-
scription and some landscape photographs may be found in Taylor
(1966). Muller (1947) gives a general description and map of the
regional vegetation, also accompanied by photographs.

The three localities are as follows, bracketed portions not being
repeated in the species list below:

1. Rio Canon [ca. 1000 m., 3 mi N Cuatro Ciénegas]

Collecting was done along the river, a spring-fed permanent stream
in a deep, narrow canyon with precipitous walls up to 500 feet high.
The narrow part is about five miles long and opens at either end onto
broad desert basins. Vegetation in the canyon consists of bunch grasses
with scattered willows and narrow-leaf cottonwoods along the river,
and thickets of mesquite and acacia on higher ground. In side canyons
there are hackberry trees and some oaks. Most of the smaller butterflies
were taken at the flowers of a low Verbenaceous plant (Phyla lance-
olata). Lower Sonoran zone: Chihuahuan Desert Scrub (Muller, 1947).

The butterflies here (31 species taken, three others seen) are typical
of the Lower Sonoran facies of the Chihuahuan Desert fauna, which
extends with little variation over much of the lower elevations on the
northern Mexican Plateau and into western Texas, southern New Mexico
and southeastern Arizona. With progressive depletion it also occurs
in arid and semi-arid Upper Sonoran regions of the southern Plateau
and also well into the prairie regions of central United States, and in
very dilute form even into eastern United States.

A striking feature of this fauna is the large proportion of known or
probable wanderers (species that migrate regularly, whether or not they
do so en masse): K. lyside and castalia, E. nicippe and lisa, P. protodice,

1 Supported by funds from Carmegie Museum and a research grant-in-aid from the Society
of the Sigma Xi.

228 CxLENcH: Coahuila butterflies Vol. 22, no. 4

D. gilippus, A. vanillae, E. claudia, V. atalanta, Libytheana sp., H. isola,
L. marina, B. exilis, H. phyleus. These total 14 species, or about 41%
of all species taken or seen. In addition to their regular and often long
migrations, these wanderers share (a) broad environmental tolerance,
both to temperatures and to vegetation type; (b) generally high levels
of abundance, some of them inclined to frequent eruptions; (c) broad
choice of larval food plant species; and (d) frequent occurrence in
disturbed environments. The combination is conspicuously one of op-
portunistic species, capable of rapidly exploiting a region where condi-
tions are stringent and suitable environments few, widely scattered,
and often transient. |

It is worth noting that there is no trace of regional endemism in this
fauna, such as occurs in some degree in the reptiles, and to a truly
remarkable degree in the fresh water fish and especially the fresh water
mollusks (Taylor, 1966).

2. Rio Salado [de los Nadadores, 7.3 mi W Sacramento, 650 m.|

The area is extensively farmed, part of a large desert basin. Collecting
was done along the man-made channel of the river and along the edges
of fallow, weed-grown fields. Hedgerows of fig and pomegranate were
present, and scattered cottonwoods along the river. Most of the butter-
flies were taken from the yellow-and-pink flowers of Cryptantha. The
locality appears to be in the Subtropical zone: Tamaulipan Thorn
Scrub (Muller, 1947).

Twelve species were taken here, and two others seen. Despite more
intensive and prolonged collecting, five of these 14 were not found at
Rio Cafon: P. sennae, P. tharos, B. hyperia, M. amymone, S. columella.
In this latitude, all of these are regional residents of the Subtropical
zone. P. tharos ranges far into cooler zones in eastern United States,
but not locally. P. sennae and M. amymone are wanderers, but do not
appear to reside anywhere in cooler zones, though they may sometimes
occur as transients. B. hyperia and S. columella are both regionally
confined to the Subtropical zone. S. columella especially is a good zonal
indicator regionally: it is fairly common, multiple brooded, tolerates
arid and semi-arid conditions, and is not known to wander at all. On
the basis of these species, particularly columella, I conclude that this
locality lies in the Subtropical zone, but probably near its boundary
with the Lower Sonoran.

3. El Caprifio [2.4 mi E Sacramento, 550 m.]

A few butterflies were collected at weeds along the roadside, in
mesquite grassland. The land is open, rocky, hilly, grazed by goats
but not farmed. Probably Subtropical zone.

1968 Journal of the Lepidopterists’ Society 229

The Rio Cafion is only about 16 airline miles from the Rio Salado
locality. Nonetheless, Muller (1947) shows that they are in different
vegetation zones and the butterflies, as described above, indicate dif-
ferent life zones. Dr. McCoy tells me that there is also a striking
difference in the herpetofaunas of the two localities. This difference
in the localities is explained in part by elevation (Rio Cafion is about
300 meters—1,100 feet—higher than Rio Salado), and in part by the
westward decrease in precipitation. The Rio Cafion is itself a well-
watered locality, so the latter effect may be less important to the butter-
flies than the former.

SpEcIES LIST

Papilio polyxenes asterius Cramer
Rio Canon 9-12.VI (34 29)
Nathalis iole Boisduval
Rio Canon 10-19.VI (7¢ 11¢); Rio Salado 30.VI (2¢ )
Kricogonia lyside (Godart)
Rio Canon 9-19.VI (5¢@ )
This is a known migrant and probably is non-resident.
Kricogonia castalia (Fabricius )
Rio Canon 9-26.VI (8¢: 2, no apical hind wing black bar;
1, very thin bar; 5, normal bar); Rio Salado 30.VI (16, no bar).

I am not convinced of the distinctness of this and the preceding species, but
follow Comstock (1944: 515) in discriminating them. Like the preceding, castalia
is a renowned migrant, probably nonresident at the Rio Canon at least.

Eurema mexicana (Boisduval )
Rio Canon 18.VI (12)
Eurema nicippe (Cramer )
Rio Canon 10-22.VI (44 29)
Eurema lisa lisa Boisduval & LeConte
Rio Canon 26.VI (1¢4 )
Eurema nise (Cramer )
Rio Canon 18.VI (1¢4 )
This may be a stray from the Subtropical zone.
Phoebis sennae (Linnaeus )
Rio Salado (seen, not taken)
Pieris protodice (Linnaeus )
Rio Cafion 8-18.VI (34 92); Rio Salado 30.VI (23 29)
Danaus gilippus strigosus (Bates )
Rio Canon 9-19.VI (2¢ 29)
Agraulis vanillae incarnata (Riley )
Rio Canton 9.VI (1¢)
Euptoieta claudia (Cramer )
Rio Canon 9.VI (1¢9 ?)

A pair in copula (10:30 A.M., ¢ flying) was also taken in Nuevo Leon: 6 mi
S Villa de Garcia (25° 49’ N, 100° 35’ W), 770 m., 2.VII.
Chlosyne lacinia adjutrix Scudder

Rio Cafon 10-19.VI (1¢ 59); Rio Salado 30.VI (1é¢ 192); El Caprio
DBAVA (C2) a O”)

To judge by the condition of the specimens, a brood was just coming to

230 Cxiencu: Coahuila butterflies Vol. 22, no. 4

an end in late June, represented almost entirely by badly worn females. At the
same time a new brood was beginning to appear, represented chiefly by fresh males.

Phyciodes vesta (Edwards)
Rio Canon 16-18.VI (2¢); Rio Salado 30.VI (36 19)
Phyciodes tharos (Drury )
Rio Salado 30.VI (1@ )
Phyciodes phaon (Edwards )
Rio Cafion 9-26.VI (106 792); Rio Salado 30.VI (44 );
El] Capriho 23.VI (2¢ 19)
Phyciodes (Tritanassa) texana texana (Edwards )
Rio Canon 10.VI (1¢ )
Nymphalis antiopa (Linnaeus )
Rio Canon (seen, not taken)
Vanessa atalanta (Linnaeus )
Rio Canon (seen, not taken)
Mestra amymone (Ménétriés )
Rio Salado 30.VI (192 ?)
Biblis hyperia (Cramer)
Rio Salado (seen, not taken)
Asterocampa leila (Edwards), subspecies
Rio Canon 9-19.VI (9¢ 5¢@)
Anaea aidea (Guérin-Méneville )
Rio Canon 18.VI (19 )
A female was also taken in Nuevo Leon: 6 mi S Villa de Garcia (25°
49’ N, 100° 35’ W), 770 m., 2.VII.
Libytheana carinenta mexicana Michener
Rio Canon (a Libytheana, probably this, seen but not taken);
Rio Salado 30.VI (1)

I am not certain that this entity is really distinct from L. bachmanii larvata
(Strecker). It is a well known migrant and perhaps not resident.

Calephelis species
Rio Canon 15 specimens

These will be determined by Mr. W. S. McAlpine.

Strymon melinus franki Field
Rio Canon 10-26.VI (2¢ 199); Rio Salado 30.VI (146 49)

Dr. McCoy tells me that when he arrived in the area (Rio Canon) in
early June there were no melinus at all, but that they became common towards
the end of the month. Curiously, however, the few early specimens are all rather
fresh, the late ones much worn. Perhaps these latter are immigrated, rather than
newly emerged, specimens.

Strymon columella istapa (Reakirt )
Rio Salado 30.VI (16 192)
Hemiargus (Echinargus) isola alce (Edwards)
Rio Canon 9-25.VI (126 62)
Leptotes marina (Reakirt )
Rio Cafion 9-18.VI (26); El Caprifo 23.VI (14 )
Brephidium exilis exilis (Boisduval )
Rio Canon 9-18.VI (66 4¢)
Systasea evansi (Bell)
Rio Canon 9-19.VI (2)
Pyrgus oileus philetas Edwards
Rio Canon 9-10.VI (1¢é 29)

1968 Journal of the Lepidopterists’ Society 231

Pyrgus sp. (group of P. communis (Grote) )

Rio Canon 12-19.VI (4¢ 292); Rio Salado 30.VI (29 )
Pholisora catullus (Fabricius )

Rio Canon 10.VI (1)
Ancyloxypha arene (Edwards)

Rio) Canon 16-18: VL (lS 19)
Copaeodes aurantiaca (Hewitson )

Rio Canon 18—-26.VI (6)
Hylephila phyleus (Drury )

Rio Canon 10.VI (1¢ 1¢@)
Amblyscirtes nysa Edwards

Rio Canon 26.VI (19 )

LITERATURE CITED

Comstock, W. P., 1944. Insects of Porto Rico and the Virgin Islands, Rhopalocera
or Butterflies. Sci. Survey Porto Rico and Virgin Is. (New York Acad. Sci.),
12: 421-622, 12 pls., 29 text figs.

Mu.uer, C. H., 1947. Vegetation and climate of Coahuila, Mexico. Madrono,
Sooo pls., | text fig.

Taytor, D. W., 1966. A remarkable snail fauna from Coahuila, Mexico. The
Veliger, 9: 152-228, pls. 8-19, 25 text figs.

CONTINUOUS VARIATION IN RELATED SPECIES OF THE
GENUS CATOCALA (NOCTUIDAE)

Morton S. ADAMS AND MARK S. BERTONI
Newark Road, Sodus, New York

The genus Catocala has been extensively studied for more than a cen-
tury. In fact, at the turn of the century, American journals dealing with
the Lepidoptera sometimes devoted the bulk of their coverage to this
genus. Even with all this attention many taxonomic problems remain.
These problems have defied classical morphological techniques, perhaps
because they centered around characters differing in kind rather than
amount. This study is, in the main, descriptive of the variation existing
in several frequently used diagnostic characters. However, the species
used in the examples were selected to suggest the utility of these statis-
tical descriptions in taxonomic studies. They may supplement a know]-
edge of classical morphology and ecology.

METHODS
An unselected sample of over 1500 Catocala of 30 species was taken
during the summer of 1961 at a Mercury vapor light operated on the
edge of a deciduous wood at the University of Michigan, Edwin S. George

232 ADAMS AND BERTONI: Variation in Catocala Vol. 22, no. 4

Fig. 1. Schematic diagram of the right clasper of the genus Catocala, showing
the measurements used.

Reserve, Pinkney, Michigan. From this sample the following series were
studied:

78 male C. ilia Cramer

25 male C. palaeogama Guenee

25 male C. retecta Grote

17 male C. sordida Grote?
7 male C. gracilis Edwards!

The following measurements were made on each of the 152 specimens:
wing span (WS), total right valva (clasper) length (C a + b), length
of distal clasper segment (Ca) and length of clasper projection (Cp)
(Fig. 1). The measurement WS was considered a reflection of the
overall size of the moth. Cp and Ca, being heavily sclerotized, are more
reliable measures than C a + b.

RESULTS

Table 1 presents the mean and standard deviation of the four variables
for the five species considered.

Table 2 presents the coefficients of correlation of each of the variables
on all other variables. The overall size of the moth as measured by WS
is not significantly correlated to the size of the genitalia. However, the
various genitalia measurements are not independent. Cp was chosen for
further analysis.

1 Determined by A. E. Brower, Augusta, Maine, to whom we are grateful for many helpful com-
ments.

1968 Journal of the Lepidopterists’ Society 233

24

20

16

TZ

Mane 19 20 2\22 23 24 25:26 27
UNITS

Fig. 2. Frequency distribution of the clasper projection (Cp) in Catocala ilia.

Figure 2 presents the frequency distribution of Cp for Catocala ilia.
It approaches the normal distribution.

Figure 3 presents the frequency distribution of Cp for C. palaeogama
superimposed on C. retecta. These two species are closely related but
distinct species. The measurement of Cp definitely indicates two pop-
ulations with some overlap. The “t” test of difference of the means is
significant at a P < 0.001.

Figure 4 presents the frequency distribution of Cp for C. gracilis
superimposed on C. sordida. These species are closely related and in-
dividual specimens are often impossible to determine with certainty.

ADAMS AND BERTONI: Variation in Catocala

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1968 Journal of the Lepidopterists’ Society 935

TABLE 1: MEAN AND STANDARD DEVIATION OF FOUR VARIABLES FOR
FIVE SPECIES OF CATOCALA!

Mean Standard deviation
ilia WS 77.14103 2.97926
Ca+b 78.20513 4.95003
Ca 40.02564 2.61849
C p 22.64103 1.51164
palaeogama. WS 67.96000 2.09126
ca+b 59.20000 4,46281
Gia 32.88000 3.07300
Crp 17.24000 2.14632
retecta WS 71.48000 2.90287
(Ge se |p 71.64000 4.51738
Gra 40.68000 3.36304
Cp 20.68000 1.46401
gracilis WS ' 41.71429 LAST
CG Ages |s Ba OAS IBN 2.63674
Cia 20.14286 1.57359
Cep 10.85714 0.89974
sordida WS 41.70588 1.21268
@a-=b 38.64706 2.64436
Ca 20.23529 1.25147
Gop 11.00000 1.00000
1 (WS measured in mm.; Ca and Cp measured in units, 50 units = 7 mm.)

Here the measurement of Cp does not indicate two populations and the
“t? test is not significant, P—0.50-0.80.

DiIscussION

Variation in the clasper of these five species of the genus Catocala is
clearly continuous. The frequency distributions are nearly normal, im-
plying that the character is controlled by small additive contributions
from many genetic factors, no one of which is individually measurable
(i.e., multifactorial inheritance ).

It is not surprising that closely related but distinct species (C. retecta
and C. palaeoguma) showed some overlapping values. The bulk of their
genetic contribution is probably of identical origin. It is even less sur-

<

Fig. 3. Frequency distribution of the clasper projection (Cp) in Catocala palaeo-
gama and C. retecta.

Fig. 4. Frequency distribution of the clasper projection (Cp) in Catocala gracilis
and C. sordida.

236 ADAMS AND BERTONI: Variation in Catocala Vol. 22) nom
TABLE 2: CORRELATION COEFFICIENTS FOR FOUR VARIABLES IN
FIvE SPECIES OF CATOCALA!
WS Ca+b (Gz C p
ilia WS 1.0000 0.24635 0.19764 0.16711
Ca+b 1.00000 0.76008 0.31197
Ca 1.00000 0.34030
Cp 1.00000
palaeogama WS 1.00000 —0.15536 0.44011 0.14147
Ca-+b 1.00000 0.26007 0.16008
Ga 1.00000 0.50362
Cp 1.00000
retecta WS 1.00000 0.37913 0.54990 0.53767
C a--b 1.00000 0.45561 0.00076
Cra 1.00000 0.57073
Cp 1.00000
gracilis WS 1.00000 —0.10805 0.70007 0.25332
Ca-b 1.00000 0.38448 0.45163
Ca 1.00000 0.48769
Cp 1.00000
sordida WS 1.00000 0.55031 0.25436 0.15462
Ca+b 1.00000 0.61213 —0.04727
Ca 1.00000 —0.04994
Cp 1.00000
1 (WS measured in mm.; Ca and Cp measured in units, 50 units = 7 mm.)

prising that C. gracilis and C. sordida completely overlap, since they are
of similar size, shape and coloration. Their eggs and larvae are nearly
identical. They feed on the same food plant (Vaccinium). Several pos-
sibilities exist to explain this degree of overlap. The sample may be too
small to demonstrate a difference. However the frequency distributions
give no evidence that these samples are abnormal. There may be hy-
bridization in Michigan. This is very possible if the two species are
isolated mainly by weak ecological factors which may be ineffective in
this area where C. gracilis is on the very edge of its range. Finally it is
recognized that parallel varietal forms occur (é.g., some specimens of
both species may have a dark shade along the inner third of the fore-
wing ). It is possible that these two species are in fact a single breeding
population which has been artificially separated on the basis of mono-
meric traits having diverse gene frequencies in various geographic areas.

The study of continuously varying characters, such as those considered
in this report, is unlikely to give definitive results. However it is likely
that most adaptive radiation is on the basis of quantitative rather than
monomeric traits. Thus such characters are appropriate material for the
study of racial and geographic variation.

NS)
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1968 Journal of the Lepidopterists’ Society

THE LIFE HISTORY AND HABITS OF
CHLOSYNE FULVIA (NYMPHALIDAE)

JAMeEs A. SCOTT
60 Estes Street, Lakewood, Colorado

In the summer of 1961 two larvae were found on paintbrush, Castilleja
integra A. Gray, west of Pueblo, Pueblo County, Colorado. They were
reared and found to represent Chlosyne fulvia (Edwards). In order to
obtain a more complete description of the life history, the author confined
several females with the foodplant on May 16, 1964. Four females laid
approximately 100 eggs on May 16 and 17. Descriptions of the egg and
larvae from 1964 specimens and a description of the pupae from 1961
specimens follow. In addition, notes on the foodplant and field habits
of the species have been included.

FieLp HABITS

C. fulvia flies in juniper woodland in the Upper Sonoran Zone wherever
its foodplant abounds, usually on low hills formed from gypsum-rich
shale. Adults fly slowly and alight often on the ground, and are thus
easy to capture. Males enjoy the few flowers available. Males are not
hilltoppers. There are three broods at Pueblo, May 5 to June 10, a second
flight occurs in July, and the third from August 23 to September 2.
Adults are most abundant in late May and late August.

FOODPLANT

Castilleja integra has crimson bracts and slender leaves one inch in
length. It is the only species of paintbrush at the localities near Pueblo
where C. fulvia flies. Two other undetermined species of Castilleja
from the Wet Mountains in Pueblo County were offered to the larvae
but were refused.

Larvae consume only the fleshy bracts; when the fleshy parts of
transplanted plants dried, larvae ate the leaves. One larva devoured
part of the ovary and some of the premature seeds.

Eggs are laid in clusters of ten to 30 on stem, leaves, or bracts. Eggs
may be laid singly in the field, however. Most of the eggs laid May
16 and 17 hatched May 21.

DESCRIPTION OF EARLY STAGES

Ecc: Pale yellow. Spherical, with slightly flattened base, diameter 0.5 mm; upper
half with approximately 18 vertical ridges, lower half pitted with many small, roughly
pentagonal cavities.

First Instar: Length 1.5 mm. Cylindrical, pale grayish green, first two thoracic

938 Scorr: Chlosyne life history Vol. 22. mom

8A TA

EXPLANATION OF FIGURES

Figs. 1-4. Mature larva of Chlosyne fulvia Edwards. 1, setal map; 2, posterior
view of larva; 3, ocelli and ocellar setae; 4, frontal view of head. In figs. 1-2 tiny
circles represent unbranched setae, and larger circles represent branching spines;
dotted lines delineate borders of sclerotized areas.

1968 Journal of the Lepidopterists’ Society 239

EXPLANATION OF FIGURES

Figs. 5-7. Pupa of Chlosyne fulvia Edwards. 5, ventral view; 6, lateral view;
7, dorsal view.

segments darker. Head black. Body covered with dark setae, arranged on I-VIII
abdominal segments, as follows: one long (1 mm) dorsolateral seta, one short
(0.5 mm) lateral seta just below and slightly posterior to dorsolateral seta, one long
seta below the short seta, in line with dorsolateral seta; below and slightly behind the
spiracle two short setae, one below the other, on thoracic and IX abdominal segments
the supraspiracular setae consist of two long setae, forming four equally-spaced setae
on the top half of larva in dorsal view. An additional short dorsolateral seta between
VIII and IX segments. Larvae molted mostly on May 23.

SECOND Instar: Length 2.5 mm. Anterior half pale green, posterior half pale
yellow. Pinaculi black, the largest around the longest setae. Prothoracic shield with
six long setae and two shorter setae posteriorly. Setae arranged as in first instar.
White internal structures appear around each setae in late stages lending a slightly
more mottled appearance. Most larvae molted May 26.

Tuirp INstar: Length 4.5 mm. Similar to mature larva, dull green fading to
greenish yellow at end of abdomen, becoming yellow prior to molting. Setae replaced
by branched spines (scoli). Base of spines reddish brown, distal portion black.
Larvae appear very black and spiny, spines almost as large as those of mature larvae.
Each spine with approximately six setae. Dark patches surrounding spines almost
touch, forming thin dorsal line along length of larva. Spines arranged as in mature
larva. Larvae molted mostly on June 3.

FourtH Instar: Length 8 mm. Ground color ochre yellow, vented surface darker.
Spines arranged as in mature larva. Thin, dark dorsal line as in mature larva. Dark
patches surrounding spines; in abdominal segments I—VIII and thoracic segments
2-3 both lateral spines above spiracles surrounded by a common dark pinaculum.
Most larvae molted June 12.

FirtH INstar: Length 14 mm. Similar to mature larva. Ground color ochre-
yellow. Spines arranged as in mature larva. Thin brown line connecting subdorsal
as well as dorsal brown pinaculi. Most larvae molted on June 18.

Mature Larva: Length about 25 mm. Ground color ochre-yellow; spines black,
slightly brownish at base. Body tapering anterior to thoracic segment 3 and posterior
to abdominal segment VII. Dark pinaculi surrounding dorsal spines on abdominal

240 Scott: Chlosyne life history Vol, 22, aime:

segments I—VIII, and much larger dark brown pinaculi surround both dorsolateral
spines on these segments. A narrow dorsal line from thoracic segment 2 to abdominal
segment VIII. Thoracic segments 2-3 with dark brown pinaculi around upper
dorsolateral spine. A heavy line connecting subdorsal dark pinaculi from thoracic
segment 2 to abdominal segment VIII. Ventral surface light brown; boundary
~ between brown and ochre yellow occurring between the upper and lower rows of
subspiracular spines. Ventral surface, especially prolegs, covered with small reddish
brown setae and hundreds of smaller transparent setae. Arrangement of spines
(scoli) as in Figure 1. Setae of the most ventral spine in abdominal segments I, II,
and VII unpigmented. Each large spine on dorsal half of body covered with about
20 minute setae, the longest (about 1.3 mm) at base and shortest at distal end of
spine. Shorter spines with fewer setae. Leg with black trochanter and tarsal claw,
other segments reddish brown. Ventral surface of legs covered with setae. Crochets
biordinal, forming a lateral penellipse. Anal plate shown in Figure 2. Anterior
lobe of anal plate dark brown, remainder reddish brown. Head reddish brown.
Adfrontal sutures darker, separated from rest of head by pale sutures (Fig. 4).
Ocelli and ocellar setae shown in Figure 3. Head with many dorsal and lateral
setae; only those which have a constant position shown in Figure 4. Larvae began
wandering on June 24; most pupated the following day.

Pura: Length 15 mm. White, mottled with black stripes and spots as in Figures
5-7. Degree of melanism variable; in one individual many black areas were broken
into separate spots, presenting a lighter appearance. Light brown showing faintly
on dorsal surface: between black spots that are close together; in grooves between
segments of abdomen (especially the grooves posterior to wing cases and one groove
anterior to these grooves); and outlining wing cases. Light brown not showing on
dorsum in a median one mm-wide strip except a few days before eclosure, when
the segments posterior to the wing cases turn reddish brown. Ventral surface with
light brown in the small spaces between the black in the space between the wing
cases. Pupal stage lasts about eight days.

ADDENDUM

In the article “Study of fluorescent pigments in Lepidoptera by means of paper
partition chromatography” by George W. Rawson (J. Lepid. Soc., 22 (1): 27-40,
1968), the following additions and corrections should be made.

On page 31, the author of Melanargia galathea is Linnaeus, not Seitz.

On page 36, the names of the 14 Phyciodes and allies were omitted in the ex-
planation of Plate 2. These are as follows: 1) Chlosyne janais (Drury); 2) C. cali-
fornica (Wright); 3) Phyciodes (Eresia) claudina guatemalena Bates; 4) P. (Phyciodes)
tharos tharos (Drury) form “marcia” Edw.; 5) P. (P.) t. tharos form “morpheus”
F.; 6) P. (P.) batesii (Reakirt); 7) Chlosyne i. ismeria (Bdv. & LeC.); 8) P. (P.)
mylitta (Edwards); 9) P. (P.) campestris (Behr); 10) P. (Tritanassa) ptolyca
(Bates); 11) P. (Eresia)frisia (Poey); 12) P. (Tritanassa) myia (Hewitson); 13)
P. (Eresia) phillyra (Hewitson); 14) P. (Tritanassa) texana (Edwards).

The color representation of the boxed symbols, A-F, accompanying this plate is
as follows: A) Bright violet fluorescence; B) dull blue-violet; C) pale yellow; D)
pale blue; E) grayish green; F) pinkish (in the basal portion of nos. 5 and 11).

1968 Journal of the Lepidopterists’ Society 241

A TAXONOMIC LIST OF PHILATELIC LEPIDOPTERA

SIDNEY A. HESSEL
Peabody Museum of Natural History, Yale University, New Haven, Conn.

Many lepidopterists are also philatelists. This includes professional
entomologists, some of whom are those actually responsible as insti-
gators or consultants for the many butterfly and moth postage stamps
that have of late years appeared around the world.

The first philatelic lepidopteran was issued in 1890 as an ornament
in the hair of Hawaiian Queen Liliuokalani. Although one may speculate
that it is the beautiful Vanessa tameamea Esch., it was not until 1930
when Lebanon honored the silk industry that a definitely determinable
species was depicted. Stylized figures had appeared in the interval.
In these instances the insects were, of course, incidental. Sarawak in
1950 was the first with nomenclature, Troides brookiana Wallace, which
was figured unicolorous gray. It remained for the Swiss Pro Juventute
issue of 1950 to honor the insect exclusively and in full color. This was
largely the work of Dr. Loeliger, a member of our Society until his
death and an important force in the Pro Juventute youth movement.
The issue was accompanied by a brochure about the insects and was
a most noteworthy effort towards stimulation of interest in Lepidoptera
in that country.

From this beginning, at first slowly, but with accelerated frequency,
over 65 countries have “honored” species of Lepidoptera by 310 butterfly
and 115 moth stamps, a total of 425 major varieties by the end of 1966.
These embrace 248 species divided 181 and 67 respectively between
butterflies and moths. Papilio machaon L. with 12 instances leads the
list. In most cases the insect is well depicted and the species pertinent
to the issuing country. A few are monstrous viz. Lebanon and Togo
with Morphos and other distant species, San Marino also with inap-
propriate selections and Albania and Somalia with flying butterflies
both upside down and half “inside out.” This presents the showier
surfaces. Some fine endemics have been chosen (Madagascar, Jamaica
et al.). All families of butterflies are now represented together with
21 moth families.

Japan 1966 (879) is the only stamp with more than one species. It
is a large stamp artistically presenting a girl amidst a veritable swarm
of flying butterflies. Ten species can be recognized with reasonable
confidence and are included in the list although this stamp does not
truly serve as a satisfactory portrayal of the subject species.

It is apparent that the raison d’etre in may cases is the fiscal benefit
to the country of issue. Nevertheless, the popularization of the subject

242, Hesse: Philatelic list Vol. 22) eno

has worthy aspects. Flagrant abuses are discussed in the philatelic
press and need not be pursued here.

Only those species determinable, at least speculatively, are considered
in the body of the taxonomic list which follows. Only stamps recognized
by the International Postal Union are included.

As to be expected, nomenclature presents a troublesome problem,
the same species often appearing under different names. Frequently
a racial name has been elevated to specific status.

The issuance of the Cuban 1961 stamp portraying “Othreis toddi
Zayas (in litt.)” has a particular interest. Inasmuch as the species had
not theretofore been described it would appear to represent a most
novel medium for publication and be vested with priority standing.
Correspondence (January 1964) with Dr. E. L. Todd who is honored
in the naming of the striking new species advised that to the best of
his knowledge publication had not yet appeared in orthodox channels.
He goes on to state, however, that the illustration on the stamp does
not constitute one of the requirements for “availability” and quotes
the new Code of Zoological Nomenclature requiring that after 1930,
in addition to other requirements, there must be a statement of char-
acters differentiating the taxon, or at least reference to such (Article
13a). So the first sortee of philate-lepidopterology into the intricasies
of zoological nomenclature is adjudged invalid on a technicality.

There is, of course, difficulty in arranging a list embracing all world
faunal zones. Remington (1954) is followed to the family level, modified
by Ehrlich (1958) for butterflies to subfamily. For genera and species
the sequence follows Munroe (1960) for the Papilionidae, Rothschild
& Jordan (1903) for Sphingidae, Peters (1952) for African Rhopalocera,
Forster & Wohlfahrt (1955, 1960) for most European Lepidoptera, and
various sources including the many volumes of G. F. Hampson’s Cat-
alogue of the Lepidoptera Phalaenae in the British Museum and of
Macrolepidoptera of the World (edited by A. Seitz) for other groups
and regions. Nomenclature roughly follows these same sources but is
modified by work of more recent authors in limited categories and
faunas. The checklist is presumed complete through 1966.

Latin names in square brackets appear if nomenclature on the stamp
differs from that in the list. In such instances the inscriptions are not
necessarily deemed erroneous though such is usually the case. In any
event, somewhat arbitrary procedure cannot be avoided to conform to
the authorities chosen and to combine examples of the same species
under one name. Scott's Standard Postage Stamp Catalogue (1966)
numbers and monthly Journal for later assignments are indicated by
parentheses. Scott's numbers prefixed by “B” are semi-postal issues;

1968 Journal of the Lepidopterists’ Society 243

[Gee anmail KA’, postal tax stamps; J’, postage due: There are no
Scott's numbers for North Korea, Red China, Cuba or Mongolia for
recent years; numbers are published in foreign catalogues. The United
States Treasury Department through its Foreign Assets Control Section,
and Pres. Kennedy, by proclamation of Feb. 7, 1962 have forbidden
importation of stamps of these countries after that date. Numbers lacking
in other instances were not available at the time of preparation of
the list.

Grateful acknowledgment is made to Prof. Charles L. Remington of
the Department of Biology, Yale University, for his many helpful sug-
gestions, particularly concerning pertinent literature. Thanks are offered
also for his counsel in the matter of troublesome determinations. More
than once I observed him turning stamps over in his eagerness to detect
significant ventral characters.

MorTHs

CossIDAE

Cossus Pulchra Rothschild
PsYCHIDAE

Manatha microcera Bourgogne
GELECHIIDAE

Pectinophora gossypiella Saunders

ZYGAENIDAE
Zygaena carniolica Scop.

Arniocera ericata Btlr.

Erasmia pulchella Hope

Amesia sanguiflua Drury
CASTNIIDAE

Castnia eudesmia Gray
PYRALIDAE

Sylepta reginalis Cramer
GEOMETRIDAE

Dysphania militaris L

>)

Abraxas grossulariata L.
URANIIDAE

Chrysiridia madagascarensis Less.

Urania boisduvalii Guer.
DREPANIDAE

Epicampoptera strandi Bryk
BOMBYCIDAE

Bombyx mori L.

Spanish Sahara [C. pulcher] 1964 (143)
Mali 1964 (J14)
Central African Rep. 1965 [Platyedra] (55)

Switzerland 1956 (B258 )

Hungary 1966 (1730)

Mozambique 1953 (376)

China 1958 [E. p. chinensis] 1958 (1186)
Lebanon 1965 [Erasmia] (C434)

Chile 1948 (C124) (254) (255)
Cuba 1965

Laos 1965 (103)
Dubai 1963 (21) (C12)
Switzerland 1957 (B269 )

Malagasy Rep. [C. madagascariensis] 1960
(C64)
Cuba [Uranidia] 1961

Central African Rep. 1965 (53)

Lebanon 1930 (108-13)

Japan 1947 (383)

Trieste Zone B 1950 (30)
Zone A 1953 (187)—Overprint Italy
(640 )

Italy 1953 (640)

Afghanistan 1963 (640) (641) (C38)
(C40)

244 HESSEL:
BRAHMAEIDAE
Dactyloceras widenmanni Karsch
SATURNIIDAE
Saturniinae

Epiphora bauhiniae Guer.

Argema mittrei Guer.
mimosae Bdv.

Bunaea alcinoe Stoll

Attacus atlas L.

Nudaurelia hersilia Westw.

Athletes ethica Westw.
gigas Sonth.

Pseudaphelia apollinaris Bdv.

Saturnia pyri Schiff.

Gynanisa maja Klug
Gonimbrasia hecate Rougeot
Lobobunea phaedusa Drury
christyi Sharpe
LASIOCAMPIDAE
Lasiocampa quercus L.

SPHINGIDAE
Acherontia atropos L.

Polyptychus roseus Druce
Cephonodes hylas L.

Daphnis nerii L.

Celerio lineata Fabr.

NOCTUIDAE
Catocalinae
Mormonia dilecta Hbn.
Catocala fraxini L.

nupta L.
Egbolis vaillantina Stoll
Metopta rectifasciata Men.
[Othreis toddi Zayas]

AGARISTIDAE

Xanthospilopteryx mozambica Mab.

Aegocera frevida Wk.

Philatelic list

Romania 1963 (1582-4)
Lebanon 1965 (439-445 )
Libya 1964 (249-51 )
Afghanistan 1966 (731)

Central African Rep. 1960 (8)

Senegal 1963 (224)

Malagasy Rep. 1960 (65)
Mozambique ( Aigenia) 1953 (371)
Togo 1964 (466)

Rwanda 1966 (118A)

China 1958 (1185)

Ryukyu Islands 1959 (57)

Laos 1965 (C46)

Mozambique 1953 [N. h. dido M&W]

(370)
Mozambique 1953 (373)
Rwanda [South] 1965. (119)

Mozambique [P. pollinaris] 1953 (377)

Switzerland 1951 (B211)
France 1956 (790)
Romania 1960 (C89)
Jugoslavia 1964 (728)
Mali 1964 (J9)

Mali 1964 (J17)

Rwanda 1966 (116A)
Mali 1964 (J18)

Switzerland 1952 (B221)

Hungary 1959 (C207 )
Romania 1960 (C93 )
Poland 1961 (1036)
Albania 1963 (694 )
Mali 1964 (J7)

Central African Rep. [Cenophodes] 1965

(54)
Jugoslavia 1964 (726)
Mali 1964 (J8)
Israel 1966 (306)
Spanish Sahara 1964 (142) (144)

Bulgaria [Catocala] 1962 (1242)
Switzerland 1950 (B198)
Czechoslovakia 1961 (1088 )
Switzerland 1957 (B271 )
Mozambique 1953 (378)

South Korea 1954 (202A )

Cuba 1961

Mozambique 1953 (380)
Mozambique 1953 (383)

Vol. 22) now

1968 Journal of the Lepidopterists Society 24

ARCTIIDAE

Lithosiinae
Chionaema saalmeulleri Btlr.

Arctiinae
Carathis gortynoides Grt.
Holomelina heros Grt.

disparilis Grt.

Rhyparioides metelkana Led.
Pericallia matronula L.
Arctia caja L.

flavia Fuessl.
villica L.
Ammobiota festiva Hutn.

Panaxia dominula L.

quadripunctaria Poda
Amphicallia thelwalli Dre.
pactolicus Btlr.
NYCTEMERIDAE
Nyctemera leuconoe Hpffr.
PERICOPIDAE
Phaloe cubana H-S.
CTENUCHIDAE (SYNTOMIDAE )
Syntomis alicia Btlr.
Syntomidopsis variegata Wk.
Ctenuchidia virgo H-S.
Metarctia lateritia H-S.
LYMANTRIDAE
Lymantria monacha L.
dispar L.

HESPERIIDAE
Capila translucida Leech
PAPILIONIDAE
Parnassiinae
Parnassius phoebus L.
jacquemontii Bdv.
nomion Hbn.
apollo L.

mnemosyne L.
Allancastria cerisyi Gat.
Serecinus telamon Dyn.

Zerynthia hypermnestra Scop.

Ut

Malagasy Rep. [C. pauliani] 1960 (309)

Cuba 1965

Cuba [Eubaphe] 1965

Cuba [Eubaphe] 1965

Romania 1964 (1618)

Lebanon 1965 (C426B )

Switzerland 1954 (B238)

Yemen 1966

Switzerland 1955 (B250)
Czechoslovakia 1966 (1396 )
Hungary [Arctia hehe] 1959 (1269)
Bulgaria [Arctia hebe] 1962 (1243)
Hungary [Callimorpha] 1966 (1726)
Czechoslovakia 1966 (1395)
Albania [Callimorpha hera] 1963 (692)
Mozambique 1953 (365)

Rwanda 1965 (116)

Mozambique 1953 (381)

Cuba 1961

Ifni 1966 (137) (139) same design
Cuba 1965

Cuba 1965

Mozambique 1953 (379)

Switzerland 1953 (B228 )
Romania 1964 (1619)

BUTTERFLIES

Red China 1963

East Germany 1964 (684)

Red China 1963

North Korea 1963

Finland 1954 (B127)

Switzerland 1955 (B251 )

Poland 1961 (1038)

Czechoslovakia 1961 (1084)

Bulgaria 1962 (1238)

Germany 1962 (B380)

Czechoslovakia 1963 (1165), 1966 (1394)

Mongolia 1963

Jugoslavia 1964 (727)

Poland 1961 (1035)

Bulgaria [Thais] 1962 (1239)

North Korea 1962

South Korea 1966 (501)

Czechoslovakia [Z. hypsipyle Sch.] 1961
(1083)

246 HessEt: Philatelic list

Vol. 22, nom

North Korea 1962
Red China 1963:

Luehdorfia puziloi Ersch.
Bhutanitis thaidina Blanch.

Papilioninae
Lamproptera meges Zink.
Teinopalpus aureus Mell
Eurytides pausianus Hew.

Red China 1963
Red China 1963
Ecuador [Graphium] 1961 (680), 1964

molops R&J

protesilaus L.

Iphiclides podalirius L.

Graphium weiskei Ribbe

doson Felder
antheus Cramer

policenes Cramer
mandarinus Oberthur

Papilio memnon L.

elwesi Leech
euchenor Guer.
menestheus Drury
lormieri Distant

ophidicephalus Obert.

demodocus Esper

alexanor Esper
machaon L.

bianor Cramer
hoppo Mats.
blumei Bdv.
Plorquinianus Fldr.
ulysses L.

zalmoxis Hew.
antimachus Drury

(7A)

Ecuador [Graphium] 1961 (682), 1964
(CHI)

Venezuela [P. p. leucones R&J] 1966 (889)

Switzerland 1951 (B209)

Poland [Papilio] 1961 (1037)

Germany 1962 (B383)

Albania [Papilio] 1963 (691 )

Hungary 1966 (1729)

Czechoslovakia 1966 (1391 )

Indonesia 1963 (B158)

Papua & New Guinea 1966 (212)

Red China 1963

Mozambique [Papilio a. evombaroides
Him.) WOs3N Gres)

Guinea 1963 (294) (299) (C48)

Red China 1963

China 1958 (1188)

Togo 1965 (486) (489)

Japan 1966 (879)

China 1958 (1184)

Papua & New Guinea 1966 (216)

Guinea 1963 (297) (302) (C49)

Somalia 1961 (C77)

Central African Rep. 1963 (31)

Malawi [P. 0. mkuwadzi] 1966 (37)

Mozambique 1963 (364)

Guinea 1963 (292, 293) (298) (303)

Israel [P. a. maccabeus] 1966 (305)

Switzerland 1954 (B241)

Czechoslovakia 1955 (714)

Hungary 1959 (1268)

Romania 1960 (C92)

Czechoslovakia 1961 (1085 )

Mongolia 1963

East Germany 1964 (685)

Jugoslavia 1964 (729)

Lebanon 1965 (C431 )

Albania 1966 (927)

Yemen 1966

Japan 1966 (879)

Japan 1966 (879)

Red China 1963

Indonesia 1963 (B156)

San Marino 1963 (568)

Papua & New Guinea [P. u. autolycus]
1966 (209)

Central African Rep. 1963 (32)

Spanish Guinea 1953 (332) (B28)

1968 Journal of the Lepidopterists’ Society 24

dardanus Brown

phorcas Cramer

hesperus Westw.
jacksoni Sharpe

bromius Dbl.
magdae Gifford
nireus L.

androgeus Cramer

caiguanabus Poey
torquatus Cramer
zagreus Dbl.
homerus Fabr.

Parides gundlachianus Fldr.

alcinous Klug
Troides aeacus Fldr.

amphrysus Cramer

brookiana Wallace

Ornithoptera paradisea Stgr.

priamus L.
victoriae Gray

Battus lycidas Cramer
crassus Cramer

PIERIDAE
Dismorphiinae
Dismorphia cubana H-S

Pierinae
Aporia crataegi L.
Delias aruna Bdv.
Pieris brassicae L.

Ascia monuste L.

Anthocaris cardamines L.

a |

Central African Rep. [Drurya] 1960 (11)

Somalia 1961 (C78)

Mozambique [P. d. tibullus Kirb.] 1953
(369), (369) as stamp on stamp
(384) (385)

Central African Rep. 1963 (30)

Rwanda 1966 (117A)

Yemen 1966

Mozambique [P. p. ansorgei Rtsch.] 1953
(375)

Somalia [P. ansorgei] 1961 (C80)

Rwanda 1966 (114A)

Rwanda [P. j. ruandana Le Cerf] 1965
(GIST (a)

Rwanda 1965 (114)

Malawi 1966 (38)

Guinea 1963 (295) (300) (304)

Senegal 1963 (222)

Dominican Rep. [P. a. epidaurus] 1966
(C148) same overprinted 1966

Cuba 1958 (C185)

Ecuador [P. t. leptalea] 1961 (681)

Venezuela 1966 (891)

Jamaica 1964 (223), same overprinted
1966, (249)

Cuba [Papilio] 1961

Japan 1966 (879)

Red China 1963

Indonesia 1963 (B159)

Sarawak 1950 (180)

Dutch New Guinea [Papilio] 1960 (B23)

Papua & New Guinea 1966 (223)

Papua & New Guinea [O. p. poseidon]
1966 (215)

Br. Solomon Islands Prot. 1965 (140)
same overprinted 1966, (161)

Ecuador 1961 (683), 1964 (713)

Lebanon [Papilio c.] 1965 (C435)

Cuba 1965

Romania 1956 (1103)

Papua & New Guinea 1966 (218)

Switzerland 1956 (B261 )

Turkey 1958 (RA227 )

Czechoslovakia 1952 (512) (513)

Dominican Rep. [A. m. eubotea] 1966
(622) same overprinted 1966

Switzerland 1951 (B210)

Czechoslovakia [Anthocharis] 1961 (1082)

Albania [Euchloe] 1963 (695)

Lebanon [Aurore] 1965 (C432)

Hungary 1966 (1727)

Japan 1966 (879)

eupheno L.
Zegris eupheme Esp.
Colotis aurigineus Btlr.

zoe Grand.

danae Fabr.

antevippe Bdv.
euippe L.

eris Klug
Ixias pyrene L.
Coliadinae

Eurema lisa Bdv. & Lec.

proterpia Fabr.

Catopsilia florella Fabr.
Phoebis avellaneda H-S
sennae L.

Gonepteryx rhamni L.

mahaguru Gistel
Colias palaeno L.

berylla Fawcett
croceus Fourcroy
hyale L.
myrmidone Esp.
electo L.

Anteos chlorinde Godt.

Nathalis iole Bdv.

NYMPHALIDAE

Danainae
Danaus chrysippus L.

formosa Godm.

Euploea leucostictos Gmelin

callithoe Bdvy.

Philatelic list Vol. 22, no. 4

Ifni [Anthocharis] 1963 (111) (113)

Israel [Z. e. varda Hemm.] 1965 (307)

Rwanda 1965 (115)

Malagasy Rep. 1960 (306)

Senegal 1963 (223)

Mauritania 1966 (213)

Mali 1964 (J12)

Mozambique [Teracolus omphale Godt.]
1953 (368 )

Central African Rep. [C. evippe] 1963
(29)

Mali 1964 (J11)

Red China 1963

Cuba [Teria ebriola] 1958 (C187)

Cuba [Teria gundlachia Poey] 1958
(C186 )

Dominican Rep. [E. gundlachia Poey] 1966
(C146 )

Mali 1964 (J20)

Cuba 1961

Dominican Rep. /P. s. sennae] 1966 (624)
overprinted 1966

Turkey 1958 (RA225)

Czechoslovakia 1961 (1090)

Albania 1963. (693 )

Mongolia 1963

Great Britain 1963 (394)

Japan 1966 (879)

Switzerland 1950 (B200)

Czechoslovakia 1966: (1392)

Red China 1963

Switzerland 1957 (B268 )

East Germany 1964 (686)

Hungary 1966 (1734)

Albania 1966 (924)

Bulgaria [C. balcanica] 1962 (1244)

Albania 1966 (925)

Rwanda [C. e. pseudohecate Berger] 1965
(118)

Dominican Rep. [A. c. c.] 1966 (625)
overprinted 1966

Cuba [N. felicia] 1958 (C188)

Mozambique [Danais] 1953 (374)

Spanish Guinea 1958 3 vals., dif. designs
(B50) (B51) (B52)

Senegal 1963 (226)

Mauritania 1966 (215)

Ifni 1966 2 Vals., same design (138) (140)

Somalia [Danaida morgani] 1961 (C79)

Red China 1963

Papua & New Guinea [E. c. duerrsteini]
1966 (215A)

1968 Journal of the Lepidopterists’ Society 249

Amauris ellioti Btlr.
niavius L.

fenestrata Aur.

lobengula Sharpe
Lycorea ceres Cramer
Clothilda numida Hbn.

pantherata Mart.

cubana Salvy.

Ithomiinae

Hymenitis cubana H-S.
Satyrinae

Hipparchia semele L.

Ragadia crisilda Hew.

Melanargia galathea L.

Brintesia circe Fabr.
Morphinae

Caligo atreus Koll.

Morpho peleides Koll.

aega Hbn.
cypris Westw.
Taenaris catops Westw.
Stichophthalma neumoegeni Leech
Charaxinae
Charaxes varanes Cramer
antamboulou Lucas
azota Hew.
cynthia Btlr.
jasius L.
epijasius Reiche
ansorgei Roths.
ameliae Doumet

nobilis Druce

zingha Stoll
dehaani Dbl.
Anaea rufescens Btlr.
clytemnestra Cramer
Siderone nemesis Il.

marthesia Cramer

Prepona antimache Hbn.
Nymphalinae
Heliconius cyrbia Godt.
charitonius L.

Euxanthe wakefieldi Ward
Harma coccinata Hew.

Rwanda [Amaurina ellioti] 1966 (117B)

Guinea [Famille Papilionides] 1963 (291)
(296) (301) (C47)

Somalia 1961 (C75)

Malawi [A. crawshayi] 1966 (40 )

Cuba 1965

Cuba [Anetia numidia briarea (lLatr.) ]
1965

Dominican Rep. [C. p. p.] 1966 (C147)
same 1966, overprinted

Cuba [Anetia] 1965

Cuba 1965

Lebanon [Satyrus] 1965 (C430 )
Red China 1963

Switzerland 1952 (B219)
Romania [Kanetisa] 1964 (1620)

Venezuela 1966 (C917)

Br. Honduras 1953 (151), 1961, over-
printed (165)

Venezuela 1966 (840 )

Togo 1965 (511)

Lebanon 1965 (C433)

Dutch New Guinea 1960 (B26)

Red China 1963

Senegal 1963 (221)

Malagasy Rep. 1960 (C63)

Mozambique 1953 (382)

Somalia 1961 (C81)

Israel 1966 (304)

Mali 1964 (J13)

Rwanda [C. a. ruandana] 1966 (119A)

Central African Rep. 1960 (9)

Lebanon 1965 (436)

Central African Rep. [Charaxe mobilis]
1961 (5) (6)

Central Africa Rep. 1960 (10)

Indonesia 1963 (B157)

Venezuela [Hypna] 1966 (C916)

Cuba [A. c. iphigenia Luc.] 1965

Cuba 1965

Dominican Rep. 1966 (626), overprinted
1966

Venezuela [S. m. thebais Fldr.] 1966
(C915)

Cuba [P. a. crassina Fruhst.] 1965

Lebanon 1965 (C427)

Dominican Rep. [H. c. churchi] 1966
(623) same overprinted 1966

Mozambique 1953 (366 )

Central African Rep. [Symothoe sangaris]
1961 (4) (7)

HESSEL:

Neptis lucilla F.
Hamanumida chalsis F\dr.
Cyrestis camillus F .
Marpesia acilia Godt.

Salamis duprei Vinson
Precis hierta F\dr.

Hypolimnas dexithea Hew.
misippus L.

Doleschallia dascylus God. & Salv.

Kallima inachis Bdv.
?Nessaea obrinus L.
Parthenos sylvia Cramer
Apatura iris L.

ilia Schiff.

Apaturina erminea Cramer

Sasakia charonda Hew.
Limenitis populi L.

Vanessa cardui L.
atalanta L.

Aglais urticae L.

Kaniska canace L.
Inachis io L.

Nymphalis polychloros 1.

antiopa L.

Philatelic list

Vol. 22) mom

Cameroons [Cymothoe sangaris] 1962
(C42)

Albania 1966 (929)

Somalia | Euryphura] 1961 (C76)

Malawi [C. c. sublineatus] 1966 (53)

Papua & New Guinea [M. a. tervisia] 1966
(PALE)

Malagasy Rep. 1960 (308 )

Senegal [Junonia] 1963 (225)

Laos [P. cebrene] 1965 (102)

Malagasy Rep. 1960 (310)

Mali 1964 (J19)

Mauritania 1966 (214)

Japan 1966 (879)

Papua & New Guinea 1966 (222)

Ryukyu Islands 1959 (61), 1960-1 (79)

San Marino 1963 (565)

Red China 1963

Papua & New Guinea [P. s. pherekides|]
1966 (217)

Hungary [A. ilia] 1959 (1271)

Romania 1960 (C94)

Switzerland 1956 (B259)

Czechoslovakia 1966 (1393)

Papua & New Guinea [A. e. papuana
Ribbe] 1966 (220)

Japan 1956 (622), 1966 (886)

Romania 1960 (C90)

Mongolia 1963

Albania [Pyrameis] 1966 (922)

Switzerland 1950 (B197)

Hungary 1959 (C208)

Czechoslovakia 1961 (1089)

Hungary (C208) as stamp on stamp 1962
(B228 )

Romania 1962 (1511)

East Germany 1964 (683)

Lebanon 1965 (C428 )

Yemen 1966

Hungary 1961 (1394) 2 vars., silver, gold

Mongolia 1963

Germany 1962 (382)

Japan 1966 (879)

Switzerland 1955 (B248)

Turkey 1958 (RA228 )

East Germany 1959 (436)

Czechoslovakia [Nymphalis] 1961 (1086)

North Korea 1962

Albania [Vanessa] 1963

San Marino 1963 (564) (567)

Mongolia 1963

Jugoslavia [Vanessa] 1964 (724)

Japan 1966 (879)

San Marino 1963 (566 )

East Germany 1964 (687 )

Switzerland 1953 (B229)

1968

Eurzicy, P. R.

Forster, W., & G. WoOHLFAHRT.

Munprog, E. G.

Journal of the Lepidopterists Society 251

Terinos alurgis Godm.
Cethosia cydippe L.
biblis Drury
Pandoriana maja Cramer
Chlosyne perezi H-S

Acraeinae

Acraea hova Bdv.

LIBYTHEIDAE

Libythea geoffroy Godt.
celtis Feussl.

LYCAENIDAE
Lycaeninae

Shirozua jonasi Janson

Chrysozephyrus mushaellus Mats.

Hypokopelates otraeda Hew.
Myrina silenus F.

Epamera handmani Gifford
Axiocerses harpax F.
Lipaphnaeus leonina Sharpe
Heodes virgaureae L.

Lysandra argester Bergstr.
coridon Poda

Agrodiaetus damon Schiff.

Meleageria daphnis Schiff.

?Maculinea arion L.
Lycaena solskyi Ersch.
phoebus Fldr.
Thysonotis danis Cramer
Loxura atymnus Cramer

Riodininae

Dodona adonira Hew.

Czechoslovakia 1961 (1087 )

Bulgaria [Vanessa] 1962 (1241)

Germany 1962 (B381)

Jugoslavia [Vanessa] 1964 (725)

Hungary 1966 (1732)

Papua & New Guinea 1966 (213)

Dutch New Guinea 1960 (B25)

Laos 1965 (101)

Bulgaria [Argynnis pandora] 1962 (1245)
Cuba 1965

Malagasy Rep. 1960 (307)

Red China 1963
Hungary 1966 (1733)

Japan 1966 (879)

Red China 1963

Mali 1964 (J15)

Mauritania 1966 (212)

Malawi 1966 (39)

Mozambique 1953 (367 )

Mali 1964 (J16)

Hungary [Lycaena] 1959 (C206)
Romania [Chrysophanus] 1960 (C91)
Hungary [L. hylas] 1959 (1270)
Switzerland 1952 (B220)

Mongolia 1963

Bulgaria [Lycaena meleager] 1962 (1240)
Hungary 1966 (1728)

Turkey 1958 (RA226)

Red China 1963

Ifni 1963 (112)

Dutch New Guinea 1960 (B24)

Red China 1963

Red China 1963

LITERATURE CITED

1958. The comparative morphology, phylogeny and higher classi-

fication of the butterflies. Univ. Kansas Sci. Bull. 39: 305-70.

Fisk, F. M., B. E. MonrcoMery, K. P. Preuss, G. T. RmcEL, & R. W. Rincs. 1962.

A checklist of “entomological” stamps. Proc. No.-central Branch nt. Soc. Amer.,

17: 160-169.

Weios 126,239 pp:

standard postage stamp catalogue.

ro. 151, 1252 pp:

17: 51 pp.

1955, 1960. Die Schmetterlinge Mitteleuropas,

Harmer, G. R., E. N. Costates & J. B. HaAtcuer (editors). 1966. Scott’s
1966—122nd Ed. Scott, New York. Vols.

1960. The classification of the Papilionidae. Canad. Ent., Suppl.,

Peters, W. 1952. A _ provisional check-list of the butterflies of the Ethiopian

region. 201 pp. E. W. Classey, Feltham, England.

REMINGTON, C. L. 1954. Lepidoptera, in C. T. Brurs, A. L. MELANDER, & F. M.

ee) Masters: More Wisconsin records Vol. 22, nore

CarPENTER, A Classification of insects, Revised ed. 917 pp. Mus. Comp. Zool.,
Harvard Univ., Cambridge, Mass.
RoruscuiLp, W., & K. Jorpan. 1903. A revision of the lepidopterous family
Sphingidae, 1, 2. 972 pp. Zoological Museum, Tring, England.
SmiTH, M. E. 1954. Philatelic Lepidoptera. Lepid. News, 6: 13-16.
~ 1955. More philatelic Lepidoptera. Lepid. News, 9: 12.
1957. Philatelic Lepidoptera; 1954-1957. Lepid. News, 11: 221-224.

FIRST RECORDS OF TWO BUTTERFIES IN WISCONSIN
(NYMPHALIDAE, PIERIDAE )

I collected in northern Wisconsin in quick hit and run fashion on June 2nd, 3rd,
and 4th of 1967. My collecting objective was to document the widespread occurrence
of Erebia discoidalis (Kirby) in northern Wisconsin, but incidental to my objectives
I uncovered two butterflies previously unreported from the state.

3}

BouoriA FREIJA (Thunberg)

A single male of this species was taken in an open bog containing sphagnum,
labrador tea, cranberry, cottongrass and a sparse growth of black spruce on June 2.
After entering the bog, located on highway 17 about five miles north of Rhinelander,
Oneida county, I quickly collected three specimens of Erebia discoidalis and was
about to leave when I noticed and captured the freija. A second freija was sighted
but not captured. I had found freija very common in Minnesota bogs the previous
week and after capturing this specimen I expected to obtain others as I searched
other Wisconsin bogs, but I failed to do so. The abundance of freija in Minnesota
had been quite surprising since it was first reported from that state only two years
previously. The only other butterfly found in the bog was Incisalia augustinus (West-
wood ) which was common.

PIERIS VIRGINIENSIS Edwards

Later on June 2 I was collecting just south of Presque Isle, Vilas County, Wiscon-
sin, on a small side road leading into a rich maple forest and captured three specimens
(26 6, 19) of Pieris virginiensis. Other species taken in the same vicinity include
Euchloe olympia (Edwards), Papilio glaucus canadensis Rothschild & Jordan, Nym-
phalis j7-album (Boisduval & LeConte) and Polygonia satyrus (Edwards). My three
specimens of virginiensis are the first documented captures from Wisconsin; however
Mr. James R. Neidhoefer of Milwaukee reports (in litt) that he collected a specimen
(23-V-1961) near Hazelhurst, Oneida County—about 25 miles directly south of my
locality. Still later on June 2, 1967, I found P. virginiensis very abundant in a maple
forest about five miles south of White Pine, Ontonagon County, Michigan and cap-
tured about 40 specimens in an hour. These are the westernmost records for virginien-
sis in Michigan, which had previously been reported from Mackinac, Emmet and
Benzie counties by Voss and Wagner (1956).’

I am indebted to Dr. Alexander B. Klots of the American Museum for confirming
my determinations. The three specimens of Pieris virginiensis and the one of Boloria
freija have been donated to the American Museum Collection, New York.—Joun H.
Masters, Box 7511, St. Paul, Minnesota.

1 Voss, E. G. and W. H. Wagner, Jr. 1956. Notes on Pieris virginiensis and Erora laeta—two
butterflies hitherto unreported from Michigan. Lepid. News, 10: 18—24.

1968 Journal of the Lepidopterists’ Society ps5

TWO VARIANT FEMALES OF COLIAS (ZERENE) CESONIA
(PIERIDAE )

The southern dogshead, Colias (Zerene) cesonia (Stoll), is a fairly common visitor
in the Lubbock, Texas area. Fellow collectors and myself who have been collecting
in this area for years know the species well. I was quite surprised during the months
of October and November, 1966, to find two aberrant C. cesonia. Both were collected
in my back yard at flowers. The striking feature of both specimens was the great
reduction of black bordering on the dorsal side of the forewing. Females of this
species commonly have less black bordering than males, but the reduction is slight
and primarily in the border of the hind wings.

Colias (Zerene) cesonia (Stoll). Upper lefthand—typical male, upper and under-
side; upper right—aberrant female, upper and underside; lower right—aberrant fe-
male, upper and underside; lower left—location of capture.

The first of the two females (lower right in photograph) was captured on October
24, 1966; the weather was still very mild and that particular day was sunny and
warm. During the previous month we had experienced three mild frosts. October 23
I had taken a perfect male specimen (upper left) on the same flowers. The second
aberrant specimen (upper right) was taken 14 days later, November 7, 1966, follow-
ing another frost. The black border on this female’s forewings was even more greatly
reduced than in the first specimen. The variations could have been caused by genetic
mutations but more likely were related to environmental factors such as a rapid change
in temperature during pupation—Dwicnut BENNETT, 2808-1 Street, Lubbock, Texas
79415.

254 Scott: Sounds of Neptis Vol. 22, mom:

SOUNDS PRODUCED BY NEPTIS HYLAS (NYMPHALIDAE)

While in South Viet-Nam between 1955 and 1959, I witnessed two occasions on
which specimens of Neptis hylas (L.) produced noises which were not incidental to
- normal movement, as are the clicking sounds made during flight by some of the
robust-bodied Nymphalids such as Charaxes and Euthalia.

On the first of these occasions, in December of 1956, a specimen of Neptis hylas
alighted in my Saigon garden on a hibiscus leaf four feet from the ground and about
six feet away from me. It flattened its wings against the horizontal upper surface
of the leaf (a position commonly assumed by this species and others of the genus),
extended its reduced foremost pair of legs, and began to snap them together rapidly
so that the tarsi met to produce a sharp click which would have been audible even
at a distance several times greater than the two yards between myself and the butter-
fly. In quality, the sound was identical to that made by tapping a fingernail edge
against a sheet of paper resting on a resilient surface. The legs were partially ex-
tended on a horizontal plane, the angle between femur and tibia being about ninety
degrees, and the gesture which brought the tarsi together was similar to that of a
child clapping its hands.

The movement was very rapid, the insect giving three successive clicks within less
than a second; it paused for about two seconds and then repeated the series of three
clicks. After this it flew to a leaf on a level with my head and repeated the per-
formance for a third time before flying away.

The second occasion was three weeks later when another specimen (with fresher
coloring ) came to almost the same spot on the hibiscus hedge. It settled in the same
posture on a leaf five feet up and about eight feet away, turned itself through 180
degrees until it faced me, and produced three rapid clicks. In this instance, the
insect’s position and its greater distance from me made it difficult to be sure which
pair of legs was employed; however, the movement of the legs coincided with the
clicks. In an effort to approach more closely I frightened it away.

On neither day was I able to capture the butterfly, or to determine its sex. Both
of these incidents occurred between 10:00 and 11:00 A.M., on hot sunny mornings
with no wind. I never noticed the presence of a second specimen to which either
of the two could have been signaling, although this is inconclusive. Possibly, the
behavior was an attempt to dislodge clinging parasites. I subsequently examined
every fresh capture of this species for signs of ectoparasitism but found nothing.—-
FREDERICK W. Scott, P. O. Box 19, Chester, Nova Scotia, Canada.

1968

Journal of the Lepidopterists’ Society 255

INDEX TO VOLUME 22

(New names in boldface )

Ablepsis, 155
Abraxas, 243
Acherontia, 244
Achlyodes, 155
Acraea, 204, 251
Acronicta valliseola, 133
Actinote, 151
Adams, M. S., Variation in Catocala, 231
Adlerodea mineira, 16
Adopaea, 130
Aegocera, 245
Aeria, 151
Agaristidae, 245
Agathymus polingi, 177
Aglais, 250
Agraulis, 34, 77, 98, 152, 196, 229
Agriades glandon, 172
Agrodiaetus, 251
Aguna asander, 155
Allancasiria, 245
Amatidae, 37, 187, 245
Amauris, 249
Amblyscirtes, 170, 231
nysa, 231
simius, 170
Amesia, 243
Ammobiota, 245
Amphicallia, 245
Anaea, 153, 230, 249
Anagasta kiihniella, 174
Anartia jatrophae, 128, 152
Anatole, 153
Ancyloxypha, 169, 231
Anteos, 34, 154, 248
Anthocaris, 34, 97, 166, 247
lanceolata, 97
midea, 34
Sara O/. 266
Anthoptus, 155
Antigonus, 155
Apatura, 250
Apaturina, 250
Apodemia, 35, 97, 153, 163
mormo, 97, 163
Mais oD
Aporia, 125, 247
Appias. co, o4, 126, 154
Archonias, 154
Arctia, 245
Arctiidae, 197, 245
Argema, 244
Argynnis, 128

Argyreuptychia, 151
Arniocera, 243
Amold, A. & R. A., Effect of irradiation
on Papilio larvae, 173
Ascia monuste, 154, 166, 247
Asterocampa, 35, 120, 128, 230
Astraptes fulgerator, 155
Atalopedes campestris, 97
Athletes, 244
Atlides, 35, 97, 153, 163
Atrytone delaware, 170
Attacus, 244
Autochton, 155
Axiocerses, 251
Battus, 34, 77, 78, 82, 97, 154, 199, 247
Behavior, 125, 177
Behr, H. H., letters, 57
Bennett, D., Variant females of Colias
cesonia, 253
Bertoni, M. S., Variation in Catocala, 231
Bhutanitis, 246
Biblis, 153, 230
Blanchard, A., New moths from Texas,
133
Boloria, 61, 78, 86, 120, 160
Bombycidae, 243
Bombyx mori, 204, 243
Book Reviews, notice, 110, 187, 188
Brahmaeidae, 244
Brassolis, 151
Brazilian Lepidoptera, 1, 147
Brephidium, 35, 97, 230
exilis, 97, 230
isophthalma, 35
Brintesia, 249
Brown, F. M.,
Strecker, 57
American butterflies described by Lin-
naeus, 77
Fox Obituary, 192
Brown, K. S., Jr., Lepidoptera of central
Brazil, 147
Buckett, J. S., Description of Lithophane,
42
Bunaea, 244
Caicella, 155
Calephelis, 32, 153, 230
nilus, 153
virginiensis, 32
Caligo, 151, 249
Callicista, 153
Callicore, 152

Letters from Behr to

256

Callimormus, 155
Callophrys, 97, 1129, 164, 172, 225, 226
Callosamia, 185, 186
Capila, 245
Carathis, 245
’ Castnia, 243
Castniidae, 243
Catastica, 126
Catasticta, 154
Catocala, 58, 231, 232, 244
Catonephele, 152
Catopsilia, 121, 248
Celastrina, 97, 120, 129
Celerio, 244
Cephonodes, 244
Cepora, 121
Ceramidia viridis, 187
Cereyonis, 98, 1120; 127, 172, Né0
behri, 172
pegala, 98, 120, 127, 180
silvestris, 98
Cethosia, 251
Chalodeta, 153
Charaxes, 249, 252
Chiomara, 155
Chionaema, 245
Chlosyne, 152, 461, 162, 229, 237, 240,
Weyl
acastus, 162
californica, 240
damoetas, 162
fulvia, 161, 237
janais, 240
lacinia, 229
Chromatography, 27
Chrysiridia, 243
Chrysophanus titus, 129
Chrysozephyrus, 251
Clench, H. K., Mating behavior in butter-
flies, 125
Butterflies from Coahuila, Mexico, 227
Clothilda, 249
Cobalopsis, 156
Cobalus, 156
Codatractus, 155
Coenonympha, 31, 98
Cogia, 129
Colaenis, 152
Colias, ol, 32) O47 Oe pele
166, 176, 196, 248, 253
alexandra, 196
cesonia, 166, 253
eurydice, 97
eurytheme, 31, 32, 34, 97, 120, 127,
158

interior, 34

Index to Volume 22

Vol, 22> nent

philodice, 120
scudderi, 166
Colobura, 153
Coloradia, 58
Colotis, 248
Copaeodes aurantiaca, 231
Cossidae, 243
Cossus, 243
Covell, C. V., Jr., Book review, 110
Cressida, 121
Ctenuchidia, 245
Cupha, 121
Cybdelis, 152
Cymaenes, 11, 13, 155
laza, 11
chapa, 13
“ibn, 2
Cynea conta, 17
Cyrestis, 250
Dactyloceras, 244
Danaidae, 32, 34, 64, 78, 84, 97, 108, 120,
121, 125, 127, Uses oeZ02e 229:
248
Danaus, 32, 34, 63, 78, 84, 98, 120, 125,
127, 151, 159, 188, 204, 229/248
Daphnis, 244
Dardarina para, 6
Delias, 121, 247
Diaethria, 152
Diaeus, 155
Dimock, T., Aberration of Vanessa cardui,
146
Dione, 152
Dircenna, 151
Dismorphia, 154, 247
Dodona, 251
Doleschallia, 250
Doxocopa, 153
Doyle, J. F., III, Foodplant of Everes
comyntas, 122
Drepanidae, 243
Dryas julia, 75
Dynamine, 152
Dryadula, 152
Dysphania, 243
Eff, D., New Colorado butterfly records,
159
Egbolis, 244
Elbella, 154
Ellis, S. L., New Colorado butterfly rec-
ords, 159
Emesis, 153
Emmel, J. F. & T. C., Life history of
Papilio indra martini, 46
Epamera, 251
Epargyreus clarus, 120

1968

Epicampoptera, 243
Epiphile, 152
Epiphora, 244
Episcada, 151
Epistor lugubris, 32
Erasmia, 243
Erebia magdalena, 172
Erynnis, 31, 97, 120, 130, 168, 179
Eryphanis, 151
Euchloe, ausonides, 40, 97
hyantis, 166
Eucosma graziella, 143
Eumaeus atala, 35
Eunica tatila, 35
Euphydryas, 61, 98, 120, 160, 200
anicia, 61, 160
chalcedona, 98
editha, 98, 161, 200
phaeton, 120
quino, 61
Euphyes conspicua, 130
Euploea, 121, 248
Euptoieta, 128, 152, 229
Fuptyehia, 32, 34, 120, 127, 159, 172
areolata, 34
cymela, 34, 120, 127
dorothea, 159, 172
hermes, 32, 127
pyracmon, 159
Burema o4, 121 197, 154, 299.948
daira, 127
dina, 34
lisa, 127, 229
mexicana, 34, 229
nicippe, 229
nise, 229
Euristrymon favonius, 35
Eurybia, 153
Eurytides, 246
Euselasia, 153
Eustixia pupula, 157
Euthalia, 252
Euxanthe, 249
iveres) oo, 97, 120, 122° 129° 165
Evolution, 197
Evonyme, 153
Feniseca tarquinius, 120
Fox, Richard Middleton, obituary, 192
Funk, R. S., Overwintering of monarch,
63
Galleria mellonella, 198
Gelechiidae, 198, 243
Geometridae, 37, 110, 243
Gindanes, 155
Glaucopsyche, 60, 97, 120, 165
Godartiana, 151

Journal of the Lepidopterists’ Society

bo
Ol
“I

Gonepteryx, 248

Gonimbrasia, 244

Gorelick, G., Biology of Vanessa tame-
amea, 111

Grais, 155

Graphium, 34, 83, 154, 246

Grotella margueritaria, 142

Gynanisa, 244

Habrodais grunus, 97

Haetera piera, 109

Hamadryas, 153

Hamanumida, 250

Hamearis, 153

Hannemann, H. J., Hering obituary, 124

Hardwick, D. F., An efficient light trap
for noctuids, 65

Harma, 250

Haywardina, 151

Heliconius, 32, 34, 152, 159

Heliopetes, 97, 155, 168

Heliothis zea, 200

Hemiargus, 33, 35, 97, 153, 230

Heodes, 251

Hering, Erich Martin, obituary, 123

Hermeuptychia, 151

Hesperia, 97, 130, 169, 170

Hespenidac. 1 20, 213i, 32:35, 78,
Sees 0S TAI VOOR oper 68.9 let
MIG 30 245

Hesperocharis, 154, 189

Hessel, S. A., Philatelic Lepidoptera, 241

Hipparchia, 249

Holomelina, 245

Hoyme. ie At;
cardui, 118

Hydroecia auripurpura, 136

Hylephila, 35, 97, 130, 156, 231

Hymenitis, 249

Hypanartia, 152

Hypaurotis crysalus, 129, 172

Hypokopelates, 251

Hypoleria, 151

Hypolimnas, 121, 250

Hypothyris, 109, 151

Icaricia icarioides, 59, 62

Inachis, 250

Incisalia, 26, 225

Iphiclides, 246

Irwin, R. R., Thymelicus lineola in Illi-
nois, 21

Phillips obituary, 209

Ithomia, 151

Ithomiidae, 108, 151

Ixias, 248

Junonia, 32, 35, 98, 120, 152, 162

Kallima, 250

Migration of Vanessa

258

Kaniska, 250
Kolyer, J. M., Eclosion of Pieris rapae,
211
Kricogonia, 229
Krivda, W. V., Survival of Pieris rapae
. in Manitoba, 191
Lambremont, E. N., Unidirectional flight
of Phoebis, 182
Lamproptera, 246
Langston, R. L., Contra Costa butterflies,
89
Lasiocampa, 244
Lasiocampidae, 244
Leaf-mining habit, 123
Lemmon, Helen Lee, obituary, 196
Lephelisca, 35
Leptotes, 97, 153, 230
Lerema veadeira, 15
Lerodea eufala, 97, 156
Lethe, 34, 120
Leucidia, 154
Leucochimona, 153
Libythea, 251
Libytheana, 35, 153, 230
Light trap, 65
Limenitis, 98, 120, 128, 152, 250
Linnaeus butterflies, 77
Lipaphnaeus, 251
Lithophane gausapata, 42
Lobobunea, 244
Loxura, 251
Luehdorfia, 246
Lycaeides melissa, 129, 172
Lycaena, 58, 97, 120, 129, 164, 172, 251
Lycaenidae, 26, 32, 35, 58, 60, 97, 120,
121 22129) aS eG Sel 25222.
226, 230, 251
Lycorea, 108, 151, 249
Lymantria, 245
Lymnas, 153
Lyropteryx, 153
Lysandra, 251
Maculinea, 129, 251
Manatha, 243
Maniola, 127
Marpesia, 152, 162, 250
Marsden, D., Collecting
Queensland, 121
Masters, J. H., Incisalia henrici records,
26
Collecting Ithomiidae with Heliotrope,
108
Aberrant Colias from Minnesota, 158
Hesperocharis longstaffi in Venezuela,
189
New butterflies for Wisconsin, 252

in northern

Index to Volume 22

Vol. 22, mem

Matine, 125, 177, 197

Mechanitis, 109, 151

Megathymus, 171, 177

Melanargia, 31, 128, 240, 249

Meleageria, 251

Melete, 154

Melitaea, 98, 120, 172

Mestra, 153, 230

Metamorpha, 78, 152

Metarctia, 245

Metopta, 244

Mielke, O. H. H., New Hesperiidae from
Brazil, 1

Lepidoptera of central Brazil, 147

Migration, 118, 182

Miller, L. D., Mating behavior in butter-
flies, 125

Miltomiges, 156

Mimoniades, 155

Mithras, 153

Moeris, 156

Moiz, S. A., On Polydorus aristolochiae,

WS, 13}

Mormonia, 244

Morpho, 150, 241, 249

Morvina, 155

Morys sobra, 15

Muller, J., Variation of Cercyonis pegala,
180

Munshi, G. H., On Polydorus §aristo-
lochiae, 115, 183

Mycalesis, 121

Mylon, 155

Myrina, 251

Myrinia catua, |

Myscelus, 155

Mysoria, 155

Narope, 151

Nathalis iole, 229, 248

Neominois ridingsii, 60, 172

Neophasia terlooti, 60, 166

Neperigea mephisto, 138

Neptis, 121, 250, 254

Nessaea, 250

Nisoniades, 155

Noctuidae, 32, 42, 58, 65, 133, 200, 231,
244

Nomenclatural notice, 195

Nudaurelia, 244

Nyctemera, 245

Nymphalidae, 32, 33, 34, 59, 75, 77, 86,
98, 111, 118, 120) 12 eaezsaaAG:
150, 151, 1153; 159) i72eersoeeZ 00;
904, 229, 240; 241 FAG eee

Nymphalis, 78, 86, 98, 120, 128, 230,
250

1968

Nymula, 153

Oarisma edwardsii, 169

Ochlodes, 97, 170

Oeonus brasus, 17

Oeneis, 31, 34, 60, 159

Oleria, 151

Onaphas, 156

Oncocnemis teddi, 135

Opler, P. A., Contra Costa butterflies, 89

Book review, 188

Opsiphanes, 151

Ornithoptera, 121, 247

Othreis, 242, 244

Ouleus fridericus candangus, 5

Oxycnemis franelemonti, 140

Pachlioptera, 121

Palmer, H. B., Eclosion of Pieris rapae,
211

Panaxia, 245

Pandoriana, 251

Panoquina confusa, 20

panoquinoides, 32

Rapier. 34. 46, 53, 77, 81, 84, 97,
IO E26, 154 166, 173, 187, 199,
229, 241, 246

Papilionidae, 32, 34, 46, 53, 77, 81-84,
Pee nO tks, 120, 121. 196; 154,
166, 173, 183, 187, 199, 204, 299,
PAY, 245

Paratrytone melane, 97

Pareuptychia, 151

Parides, 247

Parnassius, 34, 204, 245

Parthenos, 250

Paryphthimoides, 151

Pease, R. W., Jr., Multiple pairing, 197

Peba striata, 9

Pectinophora gossypiella, 198

Pereute, 154

Pericallia, 245

Perichares, 156

Pericopidae, 245

Peridroma saucia, 200

Perkins, E. M. & S. F., Life history of
Papilio oregonius, 53

Perkins, O. A., Addenda to butterflies of
Michigan, 119

Phaloe, 245

Phanes, 155

Pharneuptychia, 151

Phillips, L. S., obituary, 209

Philotes, 59, 97, 165, 172

Phocides pigmalion, 35, 155

Phoepis, o2, 34, 78, 154, 166, 182, 299,
248

Pholisora, 97, 120, 168, 169, 180, 230

Journal of the Lepidopterists’ Society

259

Phyciodes: 35:57, 96, 120, 198-157 162:
230, 240
Physiology, 173, 183, 191, 211
Pierella, 128, 151
Pieridae, 31, 32, 34, 40, 60, 78, 97, 120,
PO tO LO Gen kots 166s 172, 176.
1625 ALGO IO 196 21 990, 2999:
DAT 25D O58
Pieris; 34,197, 120;=426- 166, 191; 211,
229, 247
Pigments, 27
Placidula, 151
Plebejus, 59, 97, 129, 165, 172
Poanes hobomok, 120
Polites, 97, 130, 156, 170
Polyctor, 155
Polydorus, 115, 183
Polygonia, 35, 98, 119, 120
Polyommatus, 129
Polyptychus, 244
Polythrix, 129
Powell, J. A., Callophrys iroides food-
plants, 225
Precis, 121, 250
Prepona, 153, 249
Pseudaletia unipuncta, 200
Pseudaphelia, 244
Pseudolycaena, 153
Pseudopieris, 154
Pseudoscada, 151
Psychidae, 243
Pteronymia, 151
Pyle, R. M., Butterfly swarm in Colorado,
N72;
Lemmon obituary, 196
Pyralidae, 157, 174, 198, 243
Byreus)) Sl, 32, 97, l5>..165, 168, 179,
230
Pyrrhopyge, 154
Pythonides, 155
Quadrus, 5, 6, 155
zolus, 5
u-lucida parabus, 6
Racheospila gerularia, 37
Radiatus bradus, 7
Ragadia, 249
Rawson, G. W., Fluorescent pigments, 27
Addendum, 240
Rekoa, 153
Rhyparioides, 245
Rickard, M. A., Life history of Dryas
julia, 75
Riodina, 153
Riodinidae, 35, 97, 153, 163, 230, 251
Riotte, J. C. E., Euchloe ausonides in
Ontario, 40

260

Saiseeliol

Salamis, 250

Saliana longirostris, 156

Sasakia, 250

Saturnia, 244

- Saturniidae, 58, 185, 204, 244

Satyridae, 31, 32, 34, 60, 97, 109, 120,
PU PA IWSIk., WSS), I, ISO

Satyrium, 59, 97, 129, 163

Scolitantides piasus, 59, 165

Scopula, 110

Scott, F. W., Sounds produced by Neptis
hylas, 254

Scott, J. A., New Colorado butterfly rec-
ords, 159

Life history of Chlosyne fulvia, 237

Serecinus, 246

Shapiro, A. M., Eustixia pupula on Cru-
ciferae, 157

Notes on three Pyrginae, 179

Shininger, F. S., Life history of Papilio
oregonius, 53

Shirozua, 251

Shull, E. M., Thymelicus lineola in In-
diana, 20

Siderone, 249

Smerinthus ophthalmicus, 59

Sophista, 155

Sostrata, 155

Spathilepia, 155

Speyeria, 59, 61, 62, 98, 120, 128, 159,
OG

Sphingidae, 32, 59, 244

Stalachtis, 153

Stichophthalma, 249

Stinga morrisoni, 169

Strecker, H., letters, 57

Strymon, 35, 97, 120, 230

Sylepta, 243

Synchlora denticularia, 37

Syntomeida epilais, 37

Syntomidopsis, 245

Syntomis, 245

Systasea evansi, 230

Taenaris, 249

Index to Volume 22

Vol. 22; no, 4

Taractrocera, 121

Taygetis, 151

Techniques, 27, 65, 108, 173

Teinopalpus, 246

Temenis, 152

Terimos, 251

“Theela,” 153

Thorybes, 35, 120, 179

Thymelicus lineola, 20, 21, 120

Thysonotis, 121, 251

Tilden, J. W., Exotic Lepidoptera in Cali-
fornia, 187

Timochares, 155:

Tithorea, 151

Tmolus azia, 164

Toliver, M., Partial courtship between
two Megathymus, 177

Tortricidae, 143

Weir, Jays

Tristyla alboplagiata, 135

Troides, 241, 247

Urania, 243

Uraniidae, 243

Urbanus, 35, 78, 155

Utetheisa ornatrix, 197

Vanessa, 35, 98, 111, 118, 120, 128) 146,
152, 2308 2420

Vehilius, 155

Vettius, 156

Vidius felus, 11

Viola, 155

Wallengrenia, 156

Wielgus, R. S., Biology of Vanessa tame-
amea, 111

Xanthopastis timais, 32

Xanthospilopteryx, 245

Xenophanes tryxus, 155

Young, A. M., Variant of Callosamia
promethea, 185

Yphthimoides, 151

Yvretta rhesus, 165, 169

Zegris, 248

Zoological Nomenclature, 195

Zopyrion, 155

Zygaena, 243

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1940a. Notes on the early stages of Xanthothrix ranunculi. Bull. So. Calif. Acad.
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Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964)
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA

by Cyrit F. pos Passos
Price: Society members—$4.50, others—$6.00; uncut, unbound signatures

available for interleaving and Ry ate. binding, ie prices; hard cover bound,
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1968 Journal of the Lepidopterists’ Society Vol. 22, no. 4

TABLE OF CONTENTS

The evolution and biological significance of multiple
pairing in Lepidoptera
by Roger ‘Wi Pease): Jr. S22 197-209

The effect of barometric pressure and other factors on
eclosion of the cabbage butterfly Pieris rapae (Pieridae)

by Ji..M. Kolyer ‘and’ H. B.\Pabner 911-225
Foodplants of Callophrys (Incisalia) iroides (Lycaenidae)

by Jerry Al ‘Powell) icc Go.) es UN 225-226
Butterflies from Coahuila, Mexico

by “Harry (K. Clerielis: 20 hea Nh 227-231:

Continuous variation in related species of the genus
Catocala (Noctuidae )

by M.S. Adams'and M.S: Bertoni (200 231-236
The life history and habits of Chlosyne fulvia (Nymphalidae)
by James A. Scott 2 20 ee 237-240

A taxonomic list of philatelic Lepidoptera
by’ Sidney: A. Hessel! i ee 241-252

FIELD NOTES

First records of two butterflies in Wisconsin
(Nymphalidae, Pieridae )
by John H. Masters: (sicko Cu 252

Two variant females of Colias (Zerene) cesonia (Pieridae)
By, Dwight Bennett: 2080 ee TE 253

Sounds produced by Neptis hylas (Nyphalidae)

by Frederick) W. Sce@bty (iu ou ln 254
LEONARD STEVENS PHILLIPS (1908-1968 )

by’ ‘Roderick: Ry Urwin a ea a) a 209-210

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