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ime 43

1989

Number 1

ISSN 0024-0966

Journal

of the

Lepidopterists’ Society

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

3 March 1989

THE LEPIDOPTERISTS’ SOCIETY

Executive Council

Julian P. Donahue, President
Jerry A. Powell, Immediate Past
President

John N. Eliot, Vice President
Richard A. Arnold, Secretary

Members at large :

John W. Brown
Mogens C. Nielsen
Floyd W. Preston

George W. Gibbs, Vice
President

Ronald Leuschner, Vice
President

James P. Tuttle, Treasurer

Frederick W. Stehr
John E. Rawlins
Jo Brewer

M. Deane Bowers
Richard L. Brown
Paul A. Opler

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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 Lepidopterists’ Society

Volume 43

1989

Number 1

Journal of the Lepidopterists ’ Society
43(1), 1989, 1-10

SUNDRY ARGYNNINE CONCEPTS REVISITED
(NYMPH ALIDAE)

L. Paul 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 mormoma. 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-
miiller) 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 7iokomis 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 43, 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 leg. 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 (Guilder) is a junior synonym of S. z. 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 subspeeihc
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 (1983,
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 I
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. loashingtonia (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 43, 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, 1 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 laurina 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 43, 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).

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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. 42:1-13.

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 1

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-
bokovil) and A. clarkei (Figs. 3, 4, 13, 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 Yvretta, Polites, Ochlodes, Poanes,
and Paratrytone.

Many genitalic features divide Atalopedes from Hesperia (Burns

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Journal of the Lepidopterists’ Society

15 16 17 18

Figs. 1-18. Adults of Atalopedes (all xl) (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, Conceigao 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

15

Figs. 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

Figs. 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 — 37 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.

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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

3

28

11.0-13.4

12.28 ± 0.09

0.46

3.70

9

1

12.7

Cartagena, Colombia

3

8

11.0-13.0

11.88 ± 0.27

0.78

6.53

9

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. 3, 4 with 1, 2; and 13, 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 3 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 February] 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 Moreno 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. 5, 6, 15, 16, 21, 28, 29, 38, 39)

Thymelicyus [sic] bahiensis Schaus (1902:436).

[The brief, verbal original description of nothing but facies and wing-

Volume 43, 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 3, 4; and 15, 16 with 13, 14); chiefly in female, brown background encroaches upon
pale pattern elements (compare Figs. 15, 16 with 13, 14). Spots of the primary opaque
in female as well as in male, as in clarkei (but not campestris and meso gramma).

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

Figs. 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

21

Figs. 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

Figs. 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

Figs. 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

Journal of the Lepidopterists’ Society

24

Figs. 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

25

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
meso gramma 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

Figs. 36, 37. Female genitalia of Atalopedes clarhei 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 43, Number 1

27

Figs. 38, 39. Female genitalia of Atalopedes bahiensis from Conceigao 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. 3, 4, 13, 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

Figs. 40, 41. Female genitalia of Atalopedes mesogramma from Guantanamo Bay,
CUBA, 15 September 1943, “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

Figs. 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-

Fig. 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

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, pi. 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 43, 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

Burns, 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 pis.

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., pis. 54-88.
Gillham, N. W. 1954. The type of Hesperia horns Edwards (Lepidoptera: Hesperidae).
Psyche 61:162.

Lindsey, 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, pis. 1-6.

Lindsey, 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. 35:1-230.

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 pis.

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 pis.

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):341-353, pis. 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, pis. 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
HELICONIINE BUTTERFLIES (NYMPHALIDAE)

Woodruff W. Benson, Celio F. B. Haddad1

AND MARCIO ZlKAN

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
1983, 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 Raker (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 Heliconius .”
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 (Hub-
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 Itacaiunas. 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-l” (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-l 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

Volume 43, Number 1

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38

Journal of the Lepidopterists’ Society

'''Ks*v

?r ){

Fig. 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
V2 m below and behind. These tandem flights were often leisurely, and

Volume 43, Number 1

39

Table 2. 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

1

9

No chase or following

0

1

1

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 Vz 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 1 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 conspecihcs
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 43, Number 1

41

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 1 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-l were replaced in
3, 11, and <30 min, respectively. Three others taken from territories
at Areas 1 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 1 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 43, Number 1

43

HOUR

Fig. 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

Fig. 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.

deirao lacked larval food-plants, although at N-l 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
held 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-l 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 Dry as
iulia (Fabr.) at any locality.

Discussion

The heliconiines studied here are clearly territorial. Conspecihc males
rarely remained together in a promenade area longer than the few
seconds necessary for the resident to And 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 helieoniines 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 Coordena9ao de Aperfeicjoamento de Nivel Superior (CAPES) and logistic support
by the Companhia Vale do Rio Doce and Rio Doce Geologia e Mineragao 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.

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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.
Behav. 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. Midi. 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. 1983. Territorial defence and its seasonal decline in
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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 43, Number 1

51

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 0.5 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-

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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-

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53

ON ARRIVAL INCUBATED

Fig. 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

Fig. 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.

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 major similarities and some differences in the repro-
ductive tracts of monarch butterflies from the Mexican and Californian

Volume 43, Number 1

55

ON ARRIVAL INCUBATED

Fig. 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.

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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,

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57

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:
534-540.

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. S., 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.

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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 (0.3 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 ± 3°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 33.3°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 £ -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)
it is (P = 0.008).

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

Table 1. Larval and pupal duration of Lycaena phlaeas hypophlaeas indoors at 25°C.

Volume 43, Number 1

63

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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 cm2 (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.

Endo, K., Y. Maruyama & K. Sakai. 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). Kontyu 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 STEW ART I 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:

9/gfc/e Cc£. %o./<?/t/

Jinn . Cam. Jfl-aS.
Vo/. J<}z7

f/K/V/t. /,y,3 #

t^cu. (oii 'lAb. 5R%3>

/(e*t Cc£. Vio. /<?/*-/

Fig. 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 43, Number 1

67

Fig. 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, 108-113; 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. IB) 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. IB), 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. IB) appear to say “Loja” [Ecuador], compatible with data on two paratype
males (CMNH) labelled “Rio Bamba, Ecuador”.

Kurt 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,

SH1ROZUA 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, Shirozua 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):l-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 1

69

Fig. 1. Sequence of repeated copulations in S. janasi. M: male, F: female. Further
details in text.

pairs (No. 1; No. 2, 1st & 2nd copulations; No. 3, 1st & 2nd; No. 4, 1st; 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 1st 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.

Pair no.

Distance
from previous

Time after copulation began (min)

copulation (m)

0

l

3

5

10

30

150

1

C

UX

2

—

C

FF

FU

2*

1

R

F

UL

3

—

C

FU

3*

1

R

UL

4

—

C

F

U

4*

0

R

F

FX

5

—

C

FF

F

FF

FF

FFFU

5*

0

R

FF

U

5**

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 1st 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 1st 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

71

I thank B. A. Drummond and an anonymous referee for valuable comments on the
manuscript.

Hisashi Fujii, 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 EUP1THECIA (GEOMETRIDAE)

Additional key words: Chile, Eupithecia taracapa, E. tarapaca.

Prof. Raul Cortes, 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. Guenee, 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; pi. 46, fig. 14).

Prout (1913) chose Timandra over Calothysanis on the basis of Butler’s selection of
Acidalia imitaria Hiibner 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 Linnaeus 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 Hiibner 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
RRADYEPETES 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 Riology, 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. Rox 93, Faringdon, Oxfordshire SN7 7DR, England; Bioquip
Products, 17803 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 pis. 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 aur anticolor ata 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. elf a 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 green 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. elf a, 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 pis. 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 43, 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. Landry, 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 pis. 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. Cashatt, Illinois State Museum, Springfield, Illinois 62706.

Journal of the Lepidopterists ’ Society
43(1), 1989, 76

A Taxonomic Revision of the New World 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 2.8 x 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. 43, No. 1): 3 March 1989

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.

1961a. 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 con-
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
text. Vertical lines as well 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

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Page charges: For authors supported by institutions, page charges are $20 per Journal

page. For unsupported authors, page charges are $10 per Journal page with a $50
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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 1

Phylogeny and zoogeography of the bigger and better

genus Atalopedes (Hesperiidae). John M. Burns 11

Territorial behavior and dominance in some heliconiine
butterflies (Nymphalidae). Woodruff W. Benson, Celio
F. B. Haddad b Marcio Zikan 33

Reproductive tract development in monarch butterflies
OVERWINTERING IN CALIFORNIA AND MEXICO. William S.
Herman, Lincoln P. Brower b William H. Calvert 50

Instar number and larval development in Lycaena phlaeas
hypophlaeas (Boisduval) (Lycaenidae) . Gregory B.
Ballmer b Gordon F. Pratt 59

General Notes

Status of the papilionid types Papilio stewarti Avinoff and P. morrisi Ehr-
mann. Kurt Johnson, David Matusik 6- Rick Rozycki 66

Repeated copulation in an orange hairstreak, Shirozua janasi: A case of mate

guarding? Hisashi Fujii 68

An emended specific name in Eupithecia (Geometridae). Frederick H.

Rindge 71

The valid generic placement for “ Calothysanis” amaturaria (Walker) (Geo-
metridae, Sterrhinae). Charles V. Coveil Jr 72

Book Reviews

The moths of America North of Mexico (three fascicles). Charles V. Covell

Jr., J.-F. Landry <b Everett D. Cashatt , 73

A taxonomic revision of the New World moth genus Pero ( Lepidoptera :

Geometridae). Frederick H. Rindge 76

Immature insects, Vol. 1. William E. Miller 77

Feature Photograph

Douglas C. Ferguson 79

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

me 43

1989

Number 2

lU

'41

:rf.

ISSN 0024-0966

Journal

of the

Lepidopterists’ Society

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

t*>L

18 May 1989

THE LEPIDOPTERISTS’ SOCIETY

Executive Council

Julian P. Donahue, President
Jerry A. Powell, Immediate Past
President

John N. Eliot, Vice President
Richard A. Arnold, Secretary

George W. Gibbs, Vice
President

Ronald Leuschner, Vice
President

James P. Tuttle, Treasurer

Members at large :
John W. Brown
Mogens C. Nielsen
Floyd W. Preston

M. Deane Bowers
Richard L. Brown
Paul A. Opler

Editorial Board

Frederick W. Stehr
John E. Rawlins
Jo Brewer

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,
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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.

J OURNAL OF

The Lepidopterists’ 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 Geiger1

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 Hiibner 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

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 flow 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)
aldolase (ALD)
arginine kinase (APK)
fumarase (FUM)

glutamate-oxaloacetate transaminase
(GOT-1, GOT-2)

glutamate-pyruvate transaminase (GPT)
glyceraldehyde-phosphate dehydrogenase
(GAPDH)

cc-glycerophosphate dehydrogenase
(a-GPDH)

hexokinase (HK)
indophenol oxidase (IPO)
isocitrate dehydrogenase (IDH-1, IDH-2)
malate dehydrogenase (MDH-1, MDH-2)
malic enzyme (ME-1, ME-2)
phosphoglucomutase (PGM)
6-phospho-gluconate dehydrogenase
(6-PGD)

phosphoglucose isomerase (PGI)
pyruvate kinase (PK)

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 = 3);
Abra Infiernillo, 3300 m, i. 20.86 (n = 1); Tafi del Valle, 2100 m, i. 23-27. 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.23.86 (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: Bollene, vi.4.79 (n = 1); Dept. Bouches du Rhone: 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 = 3).

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-
lever’ difference. It is now well-established that some groups are much
more conservative electrophoretically than others, and that the relation

APK AK-1 AK-2 IDH1 IDH2 PGI PGM MDH1 MDH2 GOT1 GOT2 ALD ME-1 ME-2 GPT 6PGD GPDH FUM GAPD PK-1 HK IPO

Volume 43, Number 2

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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 “napf’-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 E. (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

1.0 , 1.0

.9
.8
.7
.6
.5
.4

Fig. 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.

r 1 — r

.84

I-:?!]

^69

,53

.63

.44

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. car ye 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 43, 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 (Tucuman, 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).

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Burns, J. M. 1975. Isozymes in evolutionary systematics, pp. 49-62. In Markert, C. L.

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Burns, J. M. & F. M. Johnson. 1967. Esterase polymorphism in natural populations
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Field, W. D. 1971. Butterflies of the genus Vanessa and of the resurrected genera
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Geiger, H. J. 1981. Enzyme electrophoretic studies on the genetic relationships of
pierid butterflies. I. European taxa. J. Res. Lepid. 19:181-195.

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Herrera, J. 1987. Biologia de Cynthia carye Hiibner 1812, especie criptica de C.
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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
1973, 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.

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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 43, 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.

Condition

June

1985

6

13

20

27

Fresh

1.00

1.00

1.50

1.63

Slightly worn

1.20

1.13

1.55

1.85

Intermediate

1.56

1.61

1.84

2.33

Very worn

2.00

2.10

2.12

2.62

Mean

1.48

1.53

1.89

2.30

Sample size

25

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

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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 mm3) result from males’ first matings and the smaller ones (about 4
mm3) 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 mm3) carried more than
one spermatophore compared with 63% of 70 females that had a small
first spermatophore (<4 mm3). This difference is significant (x2, P =
0.002, Table 2) although both samples had similar age distributions as
indicated by wing-wear classes (x2, 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 (%)

SPERMATOPHORE VOLUME (mm3)

Fig. 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.

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Journal of the Lepidopterists’ Society

Table 2. 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

1

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 (x2 = 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 43, Number 2

99

PERCENT MULTIPLE MATINGS

100

1-2 3 4

CONDITION CLASS

Fig. 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 1st spermatophore (stippled bars). Associated
contingency tests are in Table 2. Early-season females were captured 6-13 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 Dry as 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.

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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-l-1598 and #87-CRCR-l-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.

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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

Risa H. Rosenberg1

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: honey dew 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 m2 area
ca. 100 m away (and 90 m from the nearest territorial site) where

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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 43, 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 fly ways 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

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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
1963, 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.

Eickwort, G. C. & H. Ginsberg. 1980. Foraging and mating behavior in Apoidea.
Ann. Rev. Entomol. 25:421-446.

Fitzpatrick, S. & W. Wellington. 1983. Insect territoriality. Can. J. Zool. 61:471-
486.

Flemwell, G. 1914. Beautiful Switzerland: Lausanne and its environs. Blackie &
Son Ltd., London. 64 pp.

Fukuda, 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.

Murphy, 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.

Murphy, 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. Midi. 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). Kontyu, Tokyo 44:193-204.

Watt, 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 leaf tiers and leaf miners,
or in immature ovaries of developing fruit (Hodges 1978).

Clarke (1941a) 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 M3
and Cu: connate or stalked, rarely separate; forewing with Cux and Cu2
slightly downcurved from cell then parallel with M3, 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 43, Number 2

109

Fig. 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 2-3 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 = 3). 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

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Journal of the Lepidopterists’ Society

Fig. 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

3

h

4

Fig. 3. Eighth tergum of male Anoncia crossi (dorsal view). Scale line = 0.5 mm.
Fig. 4. Eighth sternum of male Anoncia crossi (dorsal view). Scale line = 0.5 mm.

Volume 43, Number 2

111

Fig. 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; IX-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.

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Journal of the Lepidopterists’ Society

Fig. 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:

23. Valvae short, broadly joined basally (text fig. 17b) 23'

Valvae longer, narrowly joined basally (text fig. 18d) 24

23'. Left valva with basal lobe setose apically diveni

- Left valva with basal lobe without setae crossi

39. Eighth abdominal tergum concavely excavated, genital plate at least V2 width

of 8th abdominal tergum (text fig. 28a) 39'

- Eighth abdominal tergum concavely excavated, genital plate less than V6 width

of 8th abdominal tergum (text fig. 26f) 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. 1941a. 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.

Hodges, 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. Fasc. 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,
Krai 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

115

Fig. 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 & Krai, 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 = 35). 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 R5 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 Ml5 M2, M3, and Cux. Ocelli in
M2 and M3 largest and most developed. Ocellus of cell Mx only faintly present. All 4 ocelli
have silvered pupils which are a series of flat, clear scales with silver sheen. Each 3 fully
developed ocelli have a thin ring of yellow buff scales with interior ground of black with
silver pupils. Ocellus in cell M3 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
R4 and meets 2nd submarginal band to form continuous band. Extracellular band then
flows diagonally to vein Mx ca. 2 mm from junction of M4 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

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Journal of the Lepidopterists’ Society

Table 1. Comparison of Neonympha mitchellii subspecies.

m. francisci (North Carolina)

m. mitchellii (Michigan & New Jersey)

Character

Male Female

Male Female

Left forewing length

Mean ± SD

16.7 ± 0.8

18.8 ± 0.8

16.4 ± 0.75

18.3 ± 0.5

Range

15.0-18.0

17.5-20.1

16.0-19.0

18.0-24.0

N

35

12

44

31

No. forewing ocelli

Mean ± SD

3.3 ± 0.7

4.0 ± 0.65

3.7 ± 0.5

5.9 ± 0.7

Range

2-5

3-5

2-4

5-6

N

35

12

44

31

No. hindwing ocelli

Mean ± SD

5.5 ± 0.6

5.75 ± 0.4

5.5 ± 0.55

5.9 ± 0.45

Range

4-6

5-6

5-6

5-6

N

35

12

44

31

3rd & 4th hindwing ocelli bifid

%

2

N

47

200

3rd & 4th ocelli set at obliq

[ue angle*

%

74

100

18

20

N

35

12

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 outer
margin. Cells R5, M1; M2, M3, Cul5 Cu2 have ocelli. Ocelli of cells R5, Mi greatly reduced
but retain silvered pupils. Ocelli in cells M2, M3 largest, best developed. All ocelli ovoid
and pointed distally. Third and 5th ocelli of cells M2, Cux nearly bifid with double silvered
pupils. Fourth ocelli in cell M3 double-pupiled but not bifid. Each ocellus with a thin
yellow buff circulus as in forewing. Ocelli of cells M2, M3 set at oblique angle, pointing
distally away from each other at 60°; ocellus of M2 pointing in anterior direction, ocellus
of cell M3 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 43, Number 2

117

Fig. 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 hindwing with oblique ocelli.

Museum of Natural History, New York; private collections of Thomas W. Krai, 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-

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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 hind wing
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 (1936) 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, Cary a 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. Iftner 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 43, 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 1943, 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 (1773) by original description.

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Journal of the Lepidopterists’ Society

Fig. 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.

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123

Fig. 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 entire wing 2

VHW postdiscal band distinctive only costad vein M3 or caudad vein M3 3

2 VFW postmedian line white or silver 5

VFW postmedian line black (without white) or faint to absent 4

3 VHW postdiscal band distinctive only costad vein M3 chileana, new species

VHW postdiscal band distinctive only caudad vein M3 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 absent telea

VHW limbal patch extending costad to M3; postmedian line not forming a “W”;
VFW postmedian line black maesites

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Journal of the Lepidopterists’ Society

Fig. 3. 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

simaethis

VFW, VHW band extremely thin, white; DHW with bright rufous limbal patch
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-

tized pads (Fig. 6A-F) simaethis

Ductus bursae inclined <60° or not at all; cervix bursae without sclerotized pads
(Fig. 6G-0) 2

2 Ductus bursae with cephalic tubular ductus and caudally flared antrum (Fig. 6G-

I) 3

Ductus bursae with caudally flared antrum only, antrum connected directly to
corpus bursae (Fig. 6N, O) 4

3 Ductus bursae inclined 30-60° 5

Ductus bursae not inclined or inclined <20°; length of antrum less than length of

ductus bursae (Fig. 6H, I) maesites

4 Terminus of antrum only slightly flared; lamella postvaginalis parabolic (Fig. 60)

chileana, new species

Terminus of antrum greatly flared; lamella postvaginalis distally lobate (Fig. 6N)
patagonia, new species

5 Length of antrum (Fig. 6 A) equalling or exceeding length of ductus bursae (Fig.

6G) telea

Length of antrum less than length of ductus bursae (Fig. 6M)

kuscheli, new combination

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-

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Journal of the Lepidopterists’ Society

Fig. 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).

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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
“ Theda phrutus” groups (Johnson 1986, Johnson & Matusik 1988), and
Parrhasius Hiibner 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, CMNH), 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-

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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) (name given in index published in
1773).

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, pi. 1), Godman & Salvin (1879-
1901:720, pi. 81), Moeschler (1889:301), Stahl (1882:93), Gundlach (1887:622), Draudt
(1919:798, pi. 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, 73; 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 Hiib-
ner 1818:75, no. 753), Dyar (1903:36), Draudt (1931:798), Barnes & McDunnough
(1917:13) (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); VFW, 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).

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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 VHW 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, pi. 1, fig. 7; holotype male, Fig. IB, 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 (1943) 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

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Journal of the Lepidopterists’ Society

Fig. 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.

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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 (9), Havana (<3), Guantanomo Bay (<3, 9). DOMINICAN
REPUBLIC: El Numero, Barahona Province, 3 July 1986 (3 <3, 2 9). HAITI: Petionville,
9 May 1930 (9), 24-29 January 1922 (<3). LESSER ANTILLES: St. Vincent (<3, 9); Dominica,
Canefields, 1-8 December 1933 (4 <3, 16 9), October 1919 (9); St. Kitts, Basseterre (<3, 9).
UNITED STATES: Puerto Rico, Cuamo Springs, 26 December 1914 (<3); Virgin Islands,
St. Croix, 14 March 1951 (5 <3, 6 9). C. s. jago : JAMAICA: primary types, paratypes with
same data except 23 December 1919 (<3), 28 January 1919 (9), 4, 22, 28 December 1919
(3 9), 4 November 1919 (9), Mt. Diablo, 5 March 1951 (<3), Constant Springs, 4 January
1924 (3 9).

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, BMNH).

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, 7 A) 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 Calif ornia-Baja California,
and Guatemala S through Panama. Males from S California- Baja California exhibit an
elongate vinculum, and females an unusually wide ductus (width V2 that of lamella;
normally about Vo,). 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).

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Journal of the Lepidopterists’ Society

Fig. 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.l., 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

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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 3, 1 2); Salta Province, Yariquarenda (1 <3, 2 2), Agua Blanca (2 3, 1 2), Mosconi
(2 <3, 2 2), Tartagal (1 3, 1 2), La Merced (1 <3, 1 2); Jujuy Province, San Pedro (2 <3, 2 2),
Rio Lazares (1 3, 1 2). BOLIVIA: Rio Surutu, 350 m, E Bolivia (1 <3, 1 2). BRAZIL:
highlands nr. Massaranduba Blumenau (<3); Annaburg, St. Catarina (6). COLOMBIA:
Cauca Valley, 3200 ft (975 m), 25 January 1935 (3 2). COSTA RICA: Turrialba, 29 May
1946 (3, 2). ECUADOR: Banos, February 1939 (6 3, 2 2); Duran, 400 ft (122 m), 24 June
1914 (1 3, 1 2); Aguarico, November 1979 (8 3, 6 2), Mishahualli (3); Latas, Oriente (2).
GUATEMALA: Guatemala City (3, 2). GUYANA: “British Guiana” (3); Bartica District,
Bartica (3). MEXICO: Baja California, Arroyo del Refugio, 5 May 1935 (2 3, 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 3, 2 2); Vera Cruz State,
Presidio (3, 2), Jalapa (4 3, 2 2); Colima State, Colima, April 1918 (1 3, 3 2); Tamaulipas
State, San Francisco, August 1964 (3, 2). PANAMA: La Boca, Canal Zone, 25 January
1908 (3, 2). PARAGUAY: Santissima Trinidad, Cordillera Province, June-July (2 3, 2 2).
PERU: Balsapuerto, Paranapura River, Loreto, June 1933 (3); Callao (BMNH) (3, 2);
Chanchamayo, Huanuco (BMNH) (3); Chosica, 850 m, January 1900 (BMNH) (2). TRIN-
IDAD-TOBAGO: Port-of-Spain, 1-9 April 1929 (2 3). UNITED STATES: Texas, Browns-
ville, 30 October 1965 (4 3, 5 2), Pharr (4 2), San Antonio (Comal Co.; TL), 29 October
1933 (3, 2); Arizona, Portal, 10 June 1958 (3); California, San Diego Co., 193- [sic] (2 3,
4 2). VENEZUELA: Caripito (3 3, 3 2); San Felipe Venezuela, 6 May 1938 (3). C. s.
rosario: I 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 (AMNH).

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. 1:143,
vol. 2:pl. 57, figs. 350, 351), Dewitz (1877:233, pi. 1), Godman & Salvin (1879-1901:
720), Moeschler (1889:301), Stahl (1882:93), Gundlach (1887:622), Draudt (1919:798,
pi. 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).

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Journal of the Lepidopterists’ Society

Thecla maesites telea: Comstock & Huntington (1943:73; 1958-64 [1964]: 123), Zikan &
Zikan (1968:57).

Eupsyche telea: Dyar (1902:36), Wolcott (1936:402), Grossbeck (1917:23).

Chalybs telea: Kaye (1921:103), 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; VFW post-
median line black (with only faint white borders, if any); VHW limbal
suffusion extending costad to Mx; 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 artif actual 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

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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 G 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 ( 3 , 2).
TRINIDAD-TOBAGO: Port-of-Spain (2). BRAZIL: Parana State, Caviuna (2); Santa Ca-
tarina State, highlands above Massaranduba, Blumenau (<3); Amazonas State, Obidos,
January 1936 (2). COLOMBIA: Caldaz, 14 May 1914 (3, 2). MEXICO: Vera Cruz State,
Presidio (3, 2); Colima State, Colima (3, 2); Tamaulipas State, San Francisco (6, 2). UNITED
STATES: Texas, Loredo (2). COSTA RICA: Turrialba (2 3, 2). GUATEMALA: Guatemala
City (2) (BMNH). PANAMA: La Boca, Canal Zone (3, 2).

Chlorostrymon maesites (Herrich-Schaeffer)

(Figs. 2C, D, 5F, G, 6H, I)

Thecla maesites Herrich-Schaeffer 1864:165. Dewitz (1877:233, pi. 1), Moeschler (1889:
301), Stahl (1882:93), Gundlach (1887:623), Wolcott (1936:402), Comstock & Hun-
tington (1943:72; 1958-64 [1961]: 158), Comstock (1944:487), Zikan & Zikan (1968:
57).

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 [1961J158 incorrectly attribute error to Draudt).

Thecla moesites Draudt (1919:798). Comstock & Huntington (1958-64 [1961J158, 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 Ms, postmedian line not making a
“W”; VFW postmedian line black (Fig. 2C, D). Genitalia differing

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Journal of the Lepidopterists’ Society

FREQUENCY (%)

Fig. 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, VFW with marginal and
submarginal areas, cells CuAj and M3, 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.l.)
to length of ductus (d) (d.s.l./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).

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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 mote 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 3x maximum width of vincular arc (in C. simaethis, 3.5-
4.0 x), and (b) with terminal lA 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 CuAl5 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

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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 (3, 9), North Caicos, 17-18 May 1983 (2 3, 1 9). CUBA: Guantanamo Bay. JAMAICA:
Port Antonio, 10 March 1954 (<3), Baron Hill, Trelawny, 16 February 1931 (3), Montego
Bay, 3 January 1965 (3), Reading, St. James, 27 March 1939 (9), Sandy Gully, St. Andrews,
20 June 1951, 3 December 1951 (2 3), 8 July 1951 (9). UNITED STATES: Miami, Florida,
various dates (7 3, 7 9), Brickell Hammock, Florida, 6 August 1914 (3, 9), Coamo Springs,
Puerto Rico, 26-29 December 1914 (2 3, 9). C. m. clenchi: types. DOMINICA: Roseau,
11 April 1929 (2 3, 3 9).

Chlorostrymon kuscheli (Ureta), new combination

(Figs. 3E, F, 5N, 6M)

Thecla kuscheli Ureta (1949:98, pi. 1, fig. 4), Comstock & Huntington (1958-64 [1961]:
58), Rojas (1964:103).

Diagnosis. FW 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. VFW, VHW
ground dull chartreuse; each wing with complete postmedial band, but
constricted as thin white lines. VFW 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. 3E), 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.

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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 (2).

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; VFW, VHW without bands, patterned
as short silver cellular streaks across VHW discal cell and cells CuA2 to
caudal M3 (limbal suffusion, dull rusty-red to grayish, generally re-
stricted to latter cells). Larger than C. telea and C. maesites : fore wing
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.
5H, L).

Description. Male. DFW, DHW bright iridescent red-violet, basal
area of costa widely folded and colored bright rufous. VFW 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-

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Journal of the Lepidopterists’ Society

red to slightly gray suffusion from cells CuA2 to caudal M3. Stubby tail,
terminus of vein CuA2. 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 ($), 3 December 1908 (<$) (C. S.
Larsen Collection in MNHN). Paratypes: MNHN — same data as allo-
type (<3), Mendoza, Argentina, 8 April 1907 (<$), 14 March 1907 (<$), 13
December 1906 (<3), all C. S. Larsen Collection; AMNH — same data as
allotype (5); MPM — Patagonia, August 1939, P. Gagarin Collection (2).

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 south westernmost, 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 urvey 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.

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141

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. VFW, 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 CuAx and CuA2. 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, VFW without pattern,
VHW with obsolescent postdiscal line, discal ceil costad to costal margin;
limbal area, cells CuAx and CuA2 slightly suffused reddish to grey distad,
silver basad; stubby tail at terminus of HW vein CuA2. 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

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Journal of the Lepidopterists’ Society

(4 <$, 1 9); BMNH — “Chili”, Walker, J. J. Joicey Collection, “ Thecla sp.
not in collection, S.G” (<$); AMNH — same data as primary types ($).

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. Bernard! 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.

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Holland, W. J. 1931. The butterfly book. Doubleday, New York. 382 pp.

Hubner, J. 1806-38 [1807]. Sammlung exotische [sic] Schmetterlinge 1. Privately print-
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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(l):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
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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.
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363 pp.

Kirby, W. F. 1871. A synonymic catalogue of diurnal Lepidoptera. John van Voorst,
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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
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1983. Papilionoidea, pp. 49-65. In Hodges, R. W. (ed.), Check list of the

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Schwartz, A. & C. J. Jimenez. 1982. The butterflies of Montserrat, West Indies. Bull.
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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. 633 pp.

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Puerto Rico. J. Dept. Agric. Puerto Rico 20(l):l-600.

Young, F. N. 1937. 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.
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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 Rutaceae (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 (Krai, 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).

I 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.

Locality

no.

Sample size*

Description

Mw

Mr

Fw

Fr

1

Montana, Missoula, elev. 1525 m

12

2

11

2

2

Alberta, Kananaskis Rd., Regal Ck., elev. 1525 m

2

6

2

8

3

British Columbia, Manning Park, Gibson Pass,
elev. 1370 m

7

13

5

14

4

British Columbia, Big Bar Creek, Poison Mt., elev.
2135-2195 m

9

5

9

4

5

British Columbia, Penticton, Mt. Apex, elev. 2190-
2247 m

24

8

4

6

* 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 M3 and Cu2 met the forewing margin) in which >25% (males) or >50% (females)
of scales were black. “Distal transparency” — proportion of distal fore wing area not cov-
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

PATCH WIDTH (%)

1 2 3 4 5

WING

LENGTH (mm)

1 2 3 4 5

DISTAL

TRANSPARENCY(%)

DISTAL

TRANSPARENCY(%)

1 2 3 4 5

Local i t y

Fig. 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

151

This study provides evidence of a genetic basis for interpopulation variation in three
of six phenotypic characters examined for F. 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 DFl’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 DFl’s) were taken from the common stock and divided into
four equal groups for 1, 2, 3, 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 (lODFl’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 1st- (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- (3-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.

Day after
oviposition

Age of eggs
(days)

Incubation period w

hen refrigerated for:

1 day

2 days

3 days

4 days

1st

0

12

12

13

13

(fresh eggs)

2nd

1

12

12

13

13

3rd

2

9

9

11

11

4th

3

9

9

10

10

5th

4

9

9

11

11

6th

5

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

50.12*

Error

18

1.71

0.09

—

Total

27

77.71

—

—

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. Dash, 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 pis., 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, Mexico, 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

Jean-Frangois 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
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
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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,

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Text: Manuscripts should be submitted in triplicate, and must be typewritten, en-

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Measurements should be given in metric units; times in terms of the 24-hour clock (0930 h,
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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.

1961a. 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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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 i? Hansjurg Geiger 81

Evaluation of spermatophore counts in studying mating
systems OF Lepidoptera. Robert C. Lederhouse, Matthew
P. Ayres ir J. Mark Scriber 93

Behavior of the territorial species Limenitis weidemeyerii
(Nymphalidae) within temporary feeding areas. Risa
H. Rosenberg ............... 102

A NEW NEOTROPICAL ANONCIA SPECIES (COSMOPTERIGIDAE).

David Adamski 108

A new subspecies of Neonympha mitchellii (French) (Sa-
tyridae) from North Carolina. David K. Parshall ir
Thomas W. Krai 1 114

Revision of Chlorostrymon Clench and description of

TWO NEW AUSTRAL NEOTROPICAL SPECIES (LYCAENIDAE).

Kurt Johnson || 120

General Notes

Papilio troilus L. on a new and rare larval food plant. M. Wayne Morris 147

Spencer collection given to Smithsonian. J. F. Gates Clarke 147

Evidence for genetic determination of variation in adult size and wing mel-
anism of Parnassius phoebus F. C. S. Guppy 148

Effect of refrigeration on egg incubation period of the tasar silk insect An-

theraea mylitta Drury (Saturniidae). B. K. Nayak <6- A. K. Dash 152

Book Review

The Saturniidae of America. Ceratocampinae. Frederick H. Rindge 154

Feature Photograph

Steven Passoa 155

Manuscript Reviewers, 1988 | 156

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

Volume 43

1989

Number 3

ISSN 0024-0966

Journal

of the

Lepidopterists’ Society

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

Q.L

5^1

L53

7 September 1989

THE LEPIDOPTERISTS’ SOCIETY

Executive Council

Jacqueline Y. Miller, President Niels P. Kristensen, Vice

Julian P. Donahue, Immediate Past
President

Richard Holland, Vice President
William D. Winter, Secretary

President

Donald J. LaFontaine, Vice
President

James P. Tuttle, Treasurer

Members at large :
Jo Brewer
Dale W. Jenkins
John E. Rawlins

John W. Brown
Mogens C. Nielsen
Floyd W. Preston

Editorial Board

Richard A. Arnold
Susan S. Borkin
David L. Wagner

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

The Lepidopterists5 Society

Volume 43 1989 Number 3

Journal of the Lepidopterists’ Society
43(3), 1989, 157-166

PRESIDENTIAL ADDRESS, 1988:
LEPIDOPTERISTS— COLLECTORS AND BIOLOGISTS?1

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

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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:

‘T 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 long 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.

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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. 35:76) 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

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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 (1983, 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;

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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).

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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 eclosion for ovarial study. Male-female pairs for
imbibing experiments were placed in 1-pint (0.48 1) 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 cm2 were
cut from the honeydew-laden leaves, misted with water just short of
runoff to form honey dew- water solution, and placed in moth containers

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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 cm2 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 25 x. 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
45 x. 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

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Journal of the Lepidopterists’ Society

Table 1. 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-
water group

(%)

Attribute

Plain water
[19 pairs]

Honeydew-water
[9 pairs]

Oviposition span, days1

8.8 ± 1.8 (6-12)

10.3 ± 4.1 (5—17)

17a

No. eggs and oocytes

Total

326 ± 46 (243-395)

339 ± 75 (204-446)

4

Matured

200 ± 46 (134-291)

245 ± 71 (109-350)

22b

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

264c

Unlaid immature

127 ± 28 (88-176)

94 ± 58 (0-184)

— 26a

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.
a P < 0.01, 1-tailed Wilcoxon 2-sample test.
b P < 0.05, 1 -tailed Student t-test.

c 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 cm2 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-
puted as c = (x — z)/[(y — x) + (x — z)].

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 Ftests. 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

Fig. 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 honey dew- 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

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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 1963).

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 bud worm pairs imbibing plain water and honeydew-
water in the main experiment. SD preceded by ±, range in parentheses.

Mean or

other value

Departure
from plain-
water group

(%)

Attribute

Plain water
[18 pairs]

Honeydew-water
[16 pairs]

Preoviposition period,

days

2.2 ± 0.7 (1-3)

2.3 ± 0.8 (1-4)

Lifespan, days

Female

10.7 ±

2.0 (7-14)

11.4 ± 3.4 (6-17)

6a

Male

9.4 ±

2.8 (5-14)

8.7 ± 3.8 (4-17)

-7

Oviposition period,

days

7.7 ±

2.2 (4-11)

9.5 ± 3.6 (4-15)

23b

No. eggs and oocytes

Total

238 ±

60 (104-332)

233 ± 80 (126-404)

-2

Matured

177 ±

43 (104-288)

214 ± 78 (111-372)

21

Laid

175 ±

43 (102-288)

210 ± 79 (104-370)

20

Change in no. laid (L)
as function of fore-

wing length (W)

25.0

74.4

198c

Viable

156 ± 36 (98-245)

181 ± 78 (59-325)

16

Unlaid immature

60 ±

41 (0-134)

19 ± 29 (0-104)

-68d

a P < 0.01, l-tailed Wilcoxon 2-sample test.
b P < 0.05, l-tailed Student <-test.

c P < 0.01, F-test. Values are slope coefficients from the regressions L = 25.0W - 106 (r = 0.36) and L = 74.4W -
646 (r = 0.81).

d P < 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

Day no. ol adulthood

Fig. 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, 2x2 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 imbibing1

3-4

5

60

5-6

15

80

7-8

30

97

9-10

16

69

11-12

13

54

1 Underlying frequencies significantly dependent on female age (P < 0.01, 2x5 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 = 37).
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 9.3-13.0) 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 > 0.13, one-tailed Student f -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

status

No.

observed

Mean age (days)

Mean forewing length (mm)

Amount imbibed (mg)

Plain-water imbibers

Fertile

9

8.1 ±

2.4 (5-12)

11.1 ± 0.3 (10.5-11.7)

3.9 ± 2.3 (0.9-8. 1)

Infertile

5

4.6 ±

1.8 (3-7)

10.7 ± 0.6 (9.8-11.2)

4.3 ± 1.5 (2.7-6.5)

Honeydew-water imbibers

Fertile

7

8.0 ±

1.0(7-10)

11.7 ± 0.8 (10.8-13.0)

5.2 ± 2.5 (1.7-10.0)

Infertile

7

5.1 ±

1.3 (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 (3-12)

11.1 ± 0.7 (9.8-13.0)

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

177

(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 = 0.035).

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 Laboratories 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. Houseweart. 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.

White, 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 43, 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 km2, the total land
area of the earth excluding Antarctica and inland waters, yields an
average density of 0.000134 species per km2. 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-

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Journal of the Lepidopterists’ Society

Table 1. Numbers of butterfly species by family and subfamily.

Family and subfamily

No. species

References

Hesperiidae

3592

Pyrginae

1193

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

1

Bridges 1988b

Lycaenidae1

4089

Lipteninae

527

Bridges 1988c

Poritiinae

52

Bridges 1988c

Liphyrinae

20

Bridges 1988c

Miletinae

111

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

Nymphalinae2

2500

Bridges 1988d, see text

Argynninae3

155

Warren 1944, 1955; dos Passos & Grey 1945;
Grey in lift.-, Common & Waterhouse 1972;
Brown 1981

Acraeinae

240

Pierre 1987

Calinaginae

16

Oberthur 1919, 1922; Wu 1938

Danainae4

462

Ackery & Vane-Wright 1984; Drummond &
Brown 1987

Apaturinae5

431

Stichel 1938, 1939; Le Moult 1950; Comstock
1961; van Someren 1975

Satyrinae6

2400

Gaede 1931; L. D. Miller, pers. comm.

Morphinae7

155

D’Abrera 1984; Parsons 1984

Brassolinae8

81

Stichel 1932

World total

17,280

present study

1 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 1930’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.

3 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 phytogeny
( fide Drummond). The Ithomiinae admirably show the gradual reduction in number 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
many widespread polytypic species in this group. There are 157 species of Danainae (s. strf (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 Gaeae (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 Antirrhea, and 3 Caerois. Smart (1977) estimates about 80 species of Morpho, probably too many (cf. D’Abrera
1984).

8 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.

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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-
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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 pis.

D’Arrera, 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. 10:11-32.

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., Ill, & 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.
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Fox, R. M. & J. W. Fox. 1964. Introduction to comparative entomology. Reinhold,
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Gaede, M. 1931. Satyridae. Lepidopterorum Catalogus 43, 46, 48.

Haensch, R. 1909. Familie: Danaidae, pp. 113-171, pis. 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
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Landing, B. H. 1984. Factors in the distribution of butterfly color and behavior pat-
terns— Selected aspects. Privately published, Los Angeles. 200 pp.

Le Moult, E. 1950. Revision de la classification des Apaturinae de Pancien monde.

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May, R. M. 1988. How many species are there on Earth? Science 241:1441-1449.
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1982. The growth of biological thought. The Belknap Press of Harvard Univ.

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1988. Toward a new philosophy of biology. The Belknap Press of Harvard Univ.

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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. Brown1

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 Cortes (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 Mexico. Except for butterflies,

1 Research Associate, Entomology Department, San Diego Natural History Museum.

Volume 43, Number 3

185

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 Mexico, 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-

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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 Mexico 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

base tip

distance from base

(degrees of latitude)

Fig. 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..

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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 (Cl); 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, ll-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, IF, 9-X-83, F. Andrews and D. Faulkner (SDNHM); Ricardo’s RV Park,
2 mi N Cabo San Lucas, 1M, 6-1-84, R. Wells (RW); 2 mi E El Triunfo, IF, 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', IF, 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 fore wing. 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

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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, IF, 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, ll-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 Jose del
Cabo, 1M (holotype), 26-XI-61, 1M (paratype), 27-XI-61, Cary-Carnegie Expedition
(CMNH).

6. Sphinx chersis (Hiibner) (Figs. 7, 28)

Lethia chersis Hiibner, 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, IF, 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 Jose del Cabo, 1M,
l-IX-59, Radford and Werner (CAS); 6 mi E El Aguaje, summit Canon Santo Tomas Rd,
3500', 1M, l-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.

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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, 7/8-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
General des Lepidopteres Heteroceres 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, IF, 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, ll-IX-61, D.
Patterson (CAS); Agua Caliente (San Carlos), 18.5 km E Maneadero, 1M, IF, 6-VII-73,
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, IF, 22-IV-63, H. Leech and P. Arnaud
(CAS); 3 mi S San Jose del Castillo, IF, 16- VI-63, E. Sleeper (CSULB). BAJA CALIFOR-
NIA SUR: 4 mi W summit, El Aguaje-Miraflores, Sierra de la Laguna, IF, 23-1-87, R.
Wells (RW); Arroyo San Bartolo, 4F, l-XI-61, 2F, 15-XI-61, Cary-Carnegie Expedition
(CMNH); Arroyo San Bernardo, 3F, 17-XI-61, Cary-Carnegie Expedition (CMNH); Puer-
to Chileno, IF, 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.).

11a. 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.):343.

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 Jose), 1M,
30- VI-68, 1M, l-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 Jose del Castillo, 4M, 3-IX-61, 1M, IF, 15- VI-61, E. Sleeper (CSULB);
3 mi S San Jose del Castillo, 1M, IF, 15- VI-63, E. Sleeper (CSULB); 10 mi S San Matias
Peak, Sierra San Pedro Martir, IF, 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).

lib. 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 Jose 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 Jose 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.

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Journal of the Lepidopterists’ Society

15. Pachylia syces syces (Hiibner) (Figs. 13, 36)

Enyo syces Hiibner, 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, IF, XI-67, V. Stuart
(RW); Hotel Hacienda, Cabo San Lucas, IF, 16-11-80, J. McBurney (LACM); Los Cabos
airport, 30 mi NE Cabo San Lucas, IF, 5-IX-83, E. Hawks (LACM); 6 mi W Los Barriles,
El Coro Rd, IF, 25-1-87, R. Wells (RW); Bahia de las Palmas, IF, 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, IF, 12-VIII-54,
alfalfa, Rohlf (SDHNM). BAJA CALIFORNIA SUR: 2 mi S La Paz, 2M, 6-VIII-66, J.
Chemsak (UCB), IF, ll-VIII-66, J. Powell (UCB); San Jose del Cabo, 1M, 11A6-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, IF, 30-IX-80, F. Andrews and D. Faulkner
(SDNHM); 27 km NE Todos Santos, 900', on flowers of Antigonon leptopus, IF, 8/9-X-
75, R. Snelling (LACM); Canon 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 Lepidopteres, l: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

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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, IF, 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, IF, 21-VII-80, J. Brown and D. Faulkner (SDNHM); 2-
5 km S El Condor, Sierra Juarez, IF, 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, IF, l-X-81,
F. Andrews and D. Faulkner (SDNHM); Punta Lobos, 1M, 20-VII-71, H. Real and R.
Main (CAS); Miraflores, 1M, l-VIII-71, H. Real and R. Main (CAS); Los Cabos airport,
30 mi NE Cabo San Lucas, IF, 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, 1U2-IX-85, J. and K. Donahue (#97,169, LACM).

20. Eumorpha achemon (Drury) (Figs. 16, 41)

Sphinx achemon Drury, 1773, 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, 21/29- 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 Jose del Cabo, IF, 25-X-61, Cary-
Carnegie Expedition (CMNH).

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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, IF, 20-III-36, C. Harbison (SDNHM).

Additional records: BAJA CALIFORNIA NORTE: Hiway 3, 4.7 mi N Valle de las
Palmas, 2M, 3F, 2Q23-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, ll-IX-83, J. and K. Donahue (LACM).

26. Hyles lineata (Fabricius) (Figs. 20, 47)

Sphinx lineata Fabricius, 1775, Systema Entomologiae, 541.

Celerio lineata; Cary, 1963:153.

Hyles lineata ; Hodges, 1971:153.

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 Cortes (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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Distribution of Agrius cingulatus. 4. Distribution of Manduca sexta.

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Journal of the Lepidopterists’ Society

Distribution of Manduca quinquemaculata. 6. Distribution of Manduca rustica.

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197

7. Distribution of Sphinx xantus (closed circles) and S. 8. Distribution of Sphinx libocedrus (open circles) and

chersis (open circles). S. perelegans (closed circles).

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Journal of the Lepidopterists’ Society

9. Distribution of Sphinx sequoiae (open circles) and 10. Distribution of Pachysphinx occidentalis .

Smerinthus cerisyi (closed circles).

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11. Distribution of Erinnyis ello. 12. Distribution of Erinnyis crameri (open circles) and

E. obscura (closed circles).

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Journal of the Lepidopterists’ Society

13. Distribution of Pachylia syces. 14. Distribution of Callionima falcifera.

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201

15. Distribution of Aellopos clavipes (closed circles) and 16. Distribution of Eumorpha satellitia (closed circles)

Hemaris diffinis (open circles). and E. achemon (open circles).

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Journal of the Lepidopterists’ Society

17. Distribution of Eumorpha vitis (closed circles) and 18. Distribution of Euproserpinus phaeton (open cir-

E. fasciata (open circles). cles) and Xylophones tersa (closed circles).

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19. Distribution of Xylophanes pluto. 20. Distribution of Hyles lineata.

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Journal of the Lepidopterists’ Society

Figs. 21-26. 21, Agrius cingulatus. 22, Manduca sexta. 23, Manduca quinque-

maculata. 24, Manduca rustica. 25, Sphinx xantus. 26, Sphinx perelegans.

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J 32 f 33 34

v

r

Figs. 27-34. 27, Sphinx libocedrus. 28, Sphinx chersis. 29, Sphinx sequoiae. 30,

Smerinthus cerisyi. 31, Pachysphinx occidentals. 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

Journal of the Lepidopterists’ Society

206

Figs. 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.

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207

Figs. 42-47. 42, Eumorpha vitis. 43, Eumorpha fasciata. 44, Euproserpinus phae-

ton. 45, Xylophones tersa. 46, Xylophones pluto. 47, Hyles lineata.

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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), P achylia 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 (Cl), 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 Institute 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.

Essig, E. O. 1926. Insects of western North America. Macmillan Co., New York.
1035 pp.

Hodges, 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. Ill (Insecta, Lepidoptera, familia Sphingidae).
Enumeracion de los Esfingidos Mexicanos (Insecta, Lepidoptera), con notas sobre su
morfologia y su distribution en la Republica. An. Escuela Nac. Ciencias Biol. 1(3 +
4):407-495 (incl. pis. 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. 1903. A revision of the lepidopterous family Sphingidae.
LCXXXV + 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.
13:57-73.

Taylor, R. J. & P. J. Regal. 1978. The peninsular effect on species diversity and the
biogeography of Baja California. Am. Nat. 112:583-593.

Wiggins, 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

211

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 4-1 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

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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 (IV2 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

Figs. 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 (A1 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

215

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 2Vz 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 hind wings. 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. Tong1 and Arthur M. Shapiro2

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
multi voltine (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 hawk weed ( 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

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 jul of glass-distilled water with a Teflon-coated tissue grinder.
Homogenate was absorbed onto 2x9 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

220

Journal of the Lepidopterists’ Society

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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

G3PD

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 /?-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.

Table 3. Electromorph frequencies in 10 Papilio zelicaon populations in California.

222

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Volume 43, Number 3

223

Table 4. G-test of genotype frequencies in populations >14.

G-statistics for populations

Locus (df)

Hemet

Orland

Butts

Canyon

Suisun

Rancho

Cordova

Castle

Peak

Average

PGI (1)

0

4.194*

6.605*

0

1.010

2.966

12.776*

PGM (6)

24.349*

20.526*

4.502

13.382*

21.838*

6.846

48.90*

MPI (10)

3.224

3.918

1.582

1.976

10.606

10.334

30.926*

* P < 0.05. G = 22 OBS In ; df = V2(n2 - n) (Ferguson 1980).

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

Washington

Hemet

Orland

Butts Canyon

Suisun

Blue Ridge

Rancho Cordova

Castle Peak

Auburn

0.97 0.96 0.96

— 0.98 0.97

— 0.99

0.95

0.96

0.96

0.98

0.97

0.98

0.99

0.99

0.99

0.99

0.99

0.99

—

0.99

0.99

—

0.98

0.98

0.97

0.98

0.98

0.98

0.99

0.99

0.98

0.97

0.99

0.98

0.97

0.99

0.99

0.97

0.99

0.98

0.96

0.99

0.99

0.97

—

0.99

0.98

—

0.98

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Journal of the Lepidopterists’ Society

Table 6. I-value matrix using only the three polymorphic (I = 0.867 ± 0.100) loci.

Wash-

ington

Hemet

Orland

Butts

Canyon

Suisun

Blue

Ridge

Rancho

Cordova

Castle

Peak

Auburn

Gazelle

.765

.713

.707

.656

.723

.676

.807

.730

.853

Washington

—

.855

.830

.842

.795

.812

.849

.850

.950

Hemet

—

.953

.962

.960

.982

.953

.985

.784

Orland

—

.930

.974

.953

.957

.973

.783

Butts Canyon

—

.900

.984

.901

.979

.808

Suisun

—

.952

.966

.964

.719

Blue Ridge

—

.942

.990

.754

Rancho Cordova

—

.757

.822

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 nonspecific,
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

Table 7. Observed genotype frequencies for the three polymorphic loci in populations
where n > 14.

Genotype

(Electromorph-

electromorph)

Populations

Hemet

Orland

Butts

Canyon

Suisun

Rancho

Cordova

Castle

Peak

PGI

13-13

0

0

0.14

0

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

0.17

0.08

0.19

0.11

32-20

0

0

0.17

0.08

0

0.09

26-26

0.11

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

0

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

0

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

0.11

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

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Journal of the Lepidopterists’ Society

Blue Ridge

Castle Peak
Hemet

Butt s Canyon
Orl and

Suisun

Rancho Cordova
Washi ngton

Auburn

Gazelle

0.800 0.900 1.000

I - V A L U E

Fig. 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

227

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 (1943a, 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.

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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-
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Brussard, P. F., P. R. Ehrlich, D. D. Murphy, B. A. Wilcox & J. Wright. 1985.
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phalinae). J. Kansas Entomol. Soc. 58:403-412.

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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
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1970. Is Papilio gothica a good species. J. Lepid. Soc. 24:230-233.

Coolidge, K. R. 1910. A California orange dog. Pomona College J. Entomol. 2:533-
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Creed, E. R., W. H. Dowdeswell, E. B. Ford & K. G. McWhirter. 1959. Evolu-
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363-391.

Dethier, V. G. 1941. Chemical factors determining the choice of food plants by Papilio
larvae. Am. Nat. 75:61-73.

Ehrlich, P. R. & P. H. Raven. 1969. Differentiation of populations. Science 165:1228-
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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
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Masuda, 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. Masuda. 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.

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(Lepidoptera: Papilionidae). Ph.D. Dissertation, University of California, Davis. 87
pp. Diss. Abstr. Order No. DA7815518.

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Genetics 28:139-156.

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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 (Muller). 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 Cu2; ground color fades
to whitish, then greenish gray in costal and apical regions, becoming strongly suffused
with melanized scales; eyespots absent. Ventral hind wing ground color brownish ochre
in discal region, fading to greenish gray beyond the median markings. Median band
whitish, well marked; absent only between Cuj and Cu2. 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 Cux or Cu2. 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 Cuj and Cu2, rarely
absent below M3 (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
Cujl and Cu2. 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 eunomia 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 Areata 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-

Figs. 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.

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Journal of the Lepidopterists’ Society

Fig. 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

233

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

0.97

I

0.98

_l

0.99

I

1.00

California

eryngii

yontocket

eunomia

0.03

0.02

0.01

I

0.00

Nei's (1978) unbiased genetic distance

Fig. 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

235

Table 1. Allelic frequencies of Coenonympha tullia-group taxa. Population localities
and locus abbreviations given in the text.

Taxon Taxon

Locus and Locus and

allele

California 1

eryngii 2

yontocket 3

allele

California1

eryngii2

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

no

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

no

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

0.111

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

111

0.028

0.069

GOT-2

114

0.194

0.056

0.027

100

0.944

0.944

0.973

117

0.014

112

0.056

0.056

0.027

121

0.042

a-GPDH

PGM

75

0.028

90

0.014

90

0.042

94

0.028

0.042

96

0.014

97

0.028

0.027

100

0.972

0.931

1.000

100

0.361

0.486

0.662

no

0.014

106

0.528

0.389

0.311

IDH-1

no

0.083

0.028

90

0.028

0.014

112

0.014

92

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

111

0.028

0.042

1 n = 18.

2 n = 36, except at AK-1, where n = 20.

3 n = 37, 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 = xalleles. Percent of loci polymorphic = P. Observed
heterozygosity = Hobs. Heterozygosity calculated from Hardy- Weinberg proportions =
Hexp. Standard errors in parentheses.

Population

Xalleles

p

Hobs

H„p

California

3.2 (0.5)

84.6

0.303 (0.090)

0.283 (0.080)

eryngii

3.5 (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 eunomia 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 43, Number 3

237

Table 3. 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.

McCorkle, 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 Benedictis1 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 (1935: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 monoethyl 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

241

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 methyl 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 methyl 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 43, 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
103:19-43.

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 + 435 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 EDIT HI A (NYMPH ALIDAE)

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 43, Number 3

245

Table 1. Linear regression of change in condition on number of handling events, by
days in residence (males).

Days in
residence

Sample size

Slope (m)

95% confidence limits
for the slope

Correlation
coefficient (R)

Power*
(1 - 0)

3

24

0.08

-0.30

0.47

0.10

0.12

4

30

0.00

-0.29

0.30

0.01

0.13

5

32

-0.18

-0.39

0.04

0.29

0.52

6

28

-0.21

-0.41

0.00

0.37

0.71

7

25

0.12

-0.13

0.37

0.20

0.25

8

27

0.07

-0.11

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 19:327-336; 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
residence

Sample size

Slope (m)

95% confidence limits
for the slope

Correlation
coefficient (R)

Power
(1 - 0)

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 — (3) 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

247

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

Figs. 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 jam); 2, Hades noctula, cluster of PCO’s around spiracle on A4, left side
(scale bar = 200 jwn); 3, H. noctula, individual PCO (scale bar = 10 jtm); 4, E. aurantiaca,
PCO in field of microtrichia, near spiracle on A4, left side (scale bar = 10 jam ); 5, E.
aurantiaca, sieve plate of PCO showing pores (scale bar = 4 jam).

number : number on right and left side; “?” denoting larva damaged and PCO’s present
but uncountable):

E. mystica: Al: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: Al:?,49; A2:55,52; A3:?, 45; A4:?,?; A5:?,35; A6:39,?; A7:53,?; A8:70,?.

H. noctula: Al: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 jam for E. mystica (on prothoracic shield), 27 jam
for E. aurantiaca (on A4), and 28 jam 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 II 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 II each genus will have an indication of its place in the higher classification that
will facilitate finding it in Appendix II, 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 43, Number 3

251

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 1 1 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

252

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 13, 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, pis. 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.

Marc 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 scientific
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 43, 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. 35:163f.]. 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. 301, 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

257

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 Jersey dale 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: V30 sec f4.0).

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.

Royce A. Drummond, Editor

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.

1961a. 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 well 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 157

Reproductive enhancement by adult feeding: Effects of

HONEYDEW IN IMBIBED WATER ON SPRUCE

bud worm. William E. Miller 167

World numbers of butterflies. Oakley Shields 178

The Sphingidae (Lepidoptera) of Baja California,

Mexico. John W. Brown {? Julian P. Donahue 184

Biology and immature stages of Schinia masoni (Noctui-

dae). Bruce A. Byers 210

Genetic differentiation among California populations of

THE ANISE SWALLOWTAIL BUTTERFLY, PAPILIO ZELICAON

Lucas. Mark L. Tong Arthur M. Shapiro 217

A NEW SUBSPECIES OF COENONYMPHA TULLIA (MULLER)

(Nymphalidae: Satyrinae) confined to the coastal
dunes of northern California. Adam H. Porter & Ster-
ling O. Mattoon 229

A PROCEDURE FOR EXAMINING THE GENITALIC MUSCULATURE OF

Lepidoptera. John A. De Benedictis ir Jerry A. Powell .. 239

General Notes

On the location of some H. A. Freeman skipper holotypes. (Hesperi-

idae). Hugh A. Freeman 244

Effects of handling on Euphydryas editha (Nymphalidae). Maria E. Orive

6“ John F. Baughman 244

Perforated cupola organs on larvae of Euselasiinae (Riodinidae) Donald J.

Harvey 247

Book Review

Catalogue of Lycaenidae 6- Riodinidae (Lepidoptera: Rhopalo-

cera). Donald J. Harvey 250

Papillons et chenilles du Quebec et de Vest du Canada [Lepidoptera and

larvae of Quebec and of eastern Canada]. Thomas C. Emmel 251

Slug and nettle caterpillars: The biology, taxonomy and control of the Li-
macodidae of economic importance on palms in southeast Asia. Marc
E. Epstein .... .. 252

The Guild handbook of scientific illustration. Charles L. Hogue 253

Butterflies of Nepal (Central Himalaya). Thomas C. Emmel Oakley

Shields 255

Feature Photograph

Split-level dining. Boyce A. Drummond III 258

Announcement

Color illustrations in the Journal 259

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

Volume 43

1989

Number 4

)

L-

m

f.

ISSN 0024-0966

Journal

of the

Lepidopterists’ Society

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

<£. cdcfHef. —

20 December 1989

j

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
Dale W. Jenkins
John E. Rawlins

John W. Brown
Mogens C. Nielsen
Floyd W. Preston

Editorial Board

Richard A. Arnold
Susan S. Borkin
David L. Wagner

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: Male Promethea moth (life size), Callosamia promethea (Dru.), a

subtly shaded member of the Saturniinae (Saturniidae). Submitted by Gerald P. Wykes,
2569 Reinhardt Road, Monroe, Michigan 48161.

J OURNAL OF

The Lepidopterists’ Society

Volume 43 1989 Number 4

Journal of the Lepidopterists’ Society
43(4), 1989, 261-273

APOSEMATISM OF URESIPHITA REVERSALIS
LARVAE (PYRALIDAE)

E. A. Bernays1 and C. B. Montllor

Division of Biological Control and Department of Entomology,

University of California, Berkeley, 1050 San Pablo Avenue,

Albany, California 94706

ABSTRACT. A description is presented of the basic biology of Uresiphita reversalis
and of the characteristics that indicate that the larvae are truly aposematic. Coloration,
position and movements on the plants, feeding activity, and the importance of host plant
alkaloids are described and discussed. Field observations and experiments with predators
indicate that the larvae are distasteful.

Additional key words: Cytisus monspessulanus, quinolizidine alkaloids, predation,

host plants, feeding stimulants.

Brightly colored insect larvae conspicuous to humans are often as-
sumed to be aposematic, even in the absence of information about the
supposed unpleasant attributes. Such is the case with caterpillars of the
pyralid moth Uresiphita reversalis (Guenee), which feeds on several
plant species in the tribe Genisteae (Papilionaceae), including Lupinus
spp. and Cytisus (—Genista) spp. (Munroe 1976). While the adult moth
is inconspicuous, with brown forewings and orange-brown hindwings,
the larvae (after the first instar) are brightly colored and highly exposed
on their host plants. The final (fifth) larval instar has obvious black and
yellow markings. It is a multivoltine species, commonly found in coastal
California but recorded widely in North America. In the San Francisco
Bay area, larvae may be found at any time of year, although growth
rate varies greatly, and by the end of winter they are rare. There are
four to five generations per year, and populations build up in summer,
reaching a peak in the early fall. Generation time varies from about
five weeks in summer to as long as 25 weeks in winter. A general
description may be found in Munroe (1976).

1 Present address: Department of Entomology, University of Arizona, Tucson, Arizona 85721.

262

Journal of the Lepidopterists’ Society

This study is a characterization of the particular features of U. rever-
salis that may allow it to be labelled aposematic: description of color-
ation; description of field behavior and degree of exposure on the plant;
feeding patterns on the plants in the field, and laboratory experiments
on food selection behavior; chemical protection with host plant chem-
icals; and levels of predation as determined from field and laboratory
experiments.

Coloration

Eggs are laid in batches of up to 70, and the groups of young larvae,
together with the damage they impart to leaves, make them conspicuous
to humans, even though first instar individuals are green and somewhat
cryptic. The later instars have a dark gold background color, and black
head and legs. On the sides of every segment are several black tubercles,
each with white patches and long white hairs. In addition there are
bright yellow patches laterally on each segment (Fig. 1). The interseg-
mental membranes of individuals are somewhat transparent so that as
ecdysis approaches, dull greenish bands alternate with the golden back-
ground. Thus, these larvae appear to have coloration patterns that are
conspicuous and consistent with aposematism (Rothschild 1972).

Position on the Host Plants

The positions of larvae on their host plants in the field is first deter-
mined by oviposition site. Egg masses found in the field are usually on
the abaxial (lower) surface of leaves 2 to 8 cm from the tips of branches
or branchlets. When larvae hatch, they usually feed first on the adaxial
(upper) surface of the leaf on which the egg mass was laid. They produce
copious amounts of silk, and as the leaf shrivels and curls, its edges are
loosely bound together by the silk to make a shelter. Larvae remain
together in the shelter, coming out only to feed. In the second instar,
larvae begin to disperse, often forming two or more subgroups on
adjacent branchlets. Contiguous leaflets or leaves are held together with
silk to make shelters, from which foraging occurs. Caterpillars rest and
molt in these shelters. By the third instar such shelters are no longer
used and groups of larvae feed and rest in groups inside a very loose
web of silk threads across branchlets.

On Cytisus monspessulanus, larvae of U. reversalis are almost in-
variably found on the upper third of the plants, usually on the distal
ends of branches. For example, on 4 September 1985, 57 larval groups
of different ages were observed on this plant species. All but five groups
were within the distal twenty percent of the length of the branch. The
remaining five were on young branchlets more proximally. Thus larvae

Volume 43, Number 4

263

Fig. 1. Drawing of fifth instar larva of Uresiphita reversalis to indicate markings.
Open areas, white; light dotted area, gold; densely dotted area, bright yellow; black as
shown. Hairs are white. Length 25 mm.

rest and feed on the upper periphery of the plant, utilizing the plant’s
youngest foliage. Populations observed on Lupinus spp. appear to have
similar relative positions.

There is a tendency for young larvae to move outwards, away from
the main stems. Egg masses or groups of newly hatched larvae from a
laboratory culture were placed about 15 cm from the tips of branches
of C. monspessulanus in the field during the summer of 1988 and
observed at intervals over the next three weeks. In all ten cases, the
first damage was to leaves distal to the placing, and over the first five
days all feeding was confined to this region. Only after such foliage
was severely damaged was there more proximal movement. Thereafter
larvae moved out to the tips of branchlets and across (usually up) to
other branches. During observations throughout the day, most move-
ment was associated with feeding, but dispersal bouts to new feeding
sites occurred within groups at intervals — usually between 0600 and
1000 h and between 1500 and 2000 h (Fig. 2).

Previous work in our laboratory has shown that larvae are attracted
to light and that, in choice tests, young leaves are strongly preferred
to old ones (Montllor & Bernays unpubl.). These factors may have a role
in determining where larvae are found on plants. An additional factor
concerns temperature regulation: larvae will be more exposed to the
sun when located on the top and the periphery of the plant, and may
be assumed under these circumstances to have body temperatures higher
than ambient. This provides an important potential advantage for ther-
moregulation. In the the San Francisco Bay area, this may be especially
important because the daily temperature maxima rarely exceed 25°C
(in the years 1984 through 1987, the mean number of days each year
on which temperatures rose to this level was only 25). Maximum rates
of growth and development of the larvae occur at over 30°C in the

264

Journal of the Lepidopterists’ Society

NUMBER

OF

LARVAE

WALKING

0500 2000

TIME OF DAY: HOURS

Fig. 2. Patterns of dispersal in five groups (a-e) of larvae (groups composed of 83%
third instar and 17% second instar) on 22 June 1988, Berkeley, California.

laboratory (Montllor and Bernays unpubl.) as expected by comparison
with other lepidopterans (e.g., Stamp & Bowers 1988).

Larvae are almost invariably found in groups. Initially, this reflects
the fact that eggs are laid in masses on the plant. Of the egg masses
found in the field during 1985-87, numbers of eggs varied from 23 to
70 (median 37, n = 13). Typically, groups of around 15-20 second and
third instar larvae are found, while groups of fourth instars and fifth
instars are smaller. It is not clear whether the larvae are actively gre-

Volume 43, Number 4

265

garious, or whether the stimuli for movement and direction of move-
ment cause similar displacements. In any case, the exposed positions in
which larvae are found, their clumped distribution, and their low rates
of movement are consistent with their being aposematic.

Feeding
Host Plants

Survival and growth of larvae on a range of potential host plants
(host records and related plants) in the San Francisco Bay area were
tested by giving twenty individual larvae single species diets in small
cages in the laboratory and examining survivorship and development
rates. These experiments were run simultaneously at laboratory tem-
peratures of 24-26°C and a 12:12 (L:D) cycle. Food was changed daily.
There was no feeding and rapid mortality on Medicago, Trifolium,
Vicia, and Pickeringia. There was limited feeding but no pupation on
Laburnum and Ulex. However, larvae fed extensively on Cytisus sco-
parius, C. striatus, Lupinus arbor eus, and C. monspessulanus. These
data together with data on rearing from museum collections indicate
that the main hosts of U. reversalis in California are species of Lupinus,
with some specimens from Cytisus. These plants all contain quinoli-
zidine alkaloids (Kinghorn & Balandrin 1984).

When given a choice in the laboratory of leaves and/or leaflets from
Cytisus monspessulanus or Lupinus arboreus, presented in an alter-
nating array in small cages over two hours, fifth instar larvae collected
from either plant in the field significantly preferred the lupine (n =
20, Wilcoxon’s Signed Rank Test, P < 0.01). The basis for host plant
preference is not yet known, but is assumed to involve quinolizidine
alkaloids, as a wide variety of these occur differentially in the hosts
(Johnson et al. 1987, Johnson & Bentley 1988, Kinghorn & Balandrin
1984, Wink 1987); for example, in a single sample of C. monspessu-
lanus, 27 different alkaloid structures were identified (Wink pers.
comm.).

Individual fourth or fifth instar larvae presented with an alternating
array of young trifoliates and more mature single leaflets (with similar
areas) of C. monspessulanus in Petri dishes in two-hour laboratory tests
preferred young leaves over old leaves (n = 20, Sign test, P < 0.001).
The basis for this preference is also not known, but by analogy with
other phytophagous insect species it can be expected that the tender
young leaves, rich in nitrogen, provide a better diet. Young leaves are
also particularly rich in quinolizidine alkaloids (Montllor unpubl.), con-
sistent with the assumption of the importance of these compounds to
larval preference.

266

Journal of the Lepidopterists’ Society

LARVAE FEEDING: %

TIME OF DAY! HOURS

Fig. 3. Patterns of larval feeding throughout the daylight hours on five separate days
during summer 1988, Berkeley, a, b = first instar; c, d = second instar and third instar;
e = third instar and fourth instar.

Feeding Patterns in the Field

Field observations indicate that larvae of U. r ever salts feed primarily
in the day. In July 1988, five groups were observed from dawn until
dusk on three separate days; in June five more groups were similarly
observed on two days. The numbers of individuals feeding were mon-
itored at intervals. At first light, young larvae began feeding within 30
minutes on four of the five occasions. On the fifth occasion, early

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267

Table 1. Effects of sparteine and cytisine on the palatability of sucrose-impregnated
glass fiber filters1: + = trend but not significant, ++ = significantly preferred at P <
0.05, + + + = significantly preferred at P < 0.01. At each concentration n = 14-19.

Concentration of alkaloid
(% dry weight)

0.1

0.5

1.0 2.0

All cones pooled

Sparteine

+

+

+ +

+ +

Cytisine

+ + +

+ +

+++ + + +

+ + +

1 5% dry weight of sucrose, larvae presented with a choice of alkaloid-treated (alkaloid plus sucrose) or control discs
(sucrose only). Preferences tested using Wilcoxon’s Signed Rank Test.

morning temperatures were similar (11-14°C and no direct sun), but
the conditions were very foggy and the plants wet; feeding began only
after the plant surface dried off at about 0900 h. Although there were
morning peaks of feeding, there were no other obvious patterns relating
to time, temperature, or sunshine (Fig. 3). Overall, it is clear that, as
typical aposematic caterpillars, U. reversalis larvae feed actively
throughout the day. This could be chemically significant; similar al-
kaloids are known to vary in concentration throughout the day in
Lupinus albus (Wink & Witte 1984).

Host Plant Alkaloids

Two host plant alkaloids common to some species of Cytisus and
Lupinus, sparteine and cytisine, were tested as potential phagostimu-
lants for larvae of U. reversalis. Individual larvae from a laboratory
culture were given a binary choice of glass fiber filters (GF/A) im-
pregnated with sucrose (5% dry weight) and similar glass fiber filters
with the sucrose plus sparteine or cytisine at one of several different
concentrations. Experiments were at 25°C with the filters in Petri dishes.
Amounts eaten were visually estimated after 24 h. The alkaloids were
shown to be phagostimulatory (Table 1), but they were not equally
effective as feeding stimulants, and it is conceivable that the actual
profile of quinolizidine alkaloids influences acceptability.

The host plant alkaloids are bitter to humans and are also known to
be poisonous to mammals, including livestock, causing vomiting, con-
vulsions and death (Kinghorn & Balandrin 1984). These compounds
are likely to have similar effects on birds. Actively feeding larvae, having
a preference for foliage with the highest levels of alkaloids, will contain
high doses of the toxins from the gut contents alone. In addition, larvae
transfer ingested alkaloids to the cuticle, although pupae contain no
detectable alkaloids (Montllor et al. in press). It seems likely that, in
common with a number of other species of Lepidoptera feeding on
alkaloidal plants, larvae gain protection from the ingested alkaloids.
Other cases of protection from predation due to ingestion of host plant

268

Journal of the Lepidopterists’ Society

NUMBER OF
LARVAE

DAYS

Fig. 4. Survivorship curves of two cohorts of 150 larvae from the time of hatching.
Total number of larvae from 5 original egg masses per cohort were counted on each date.
Observations were made during the period May-July 1988.

alkaloids involve mainly pyrrolizidine alkaloids (Rothschild 1972,
Schneider 1987). This seems to be the first example of the use of quin-
olizidine alkaloids for protection by caterpillars.

Predation

Field Observation in Relation to Invertebrate Predators

In two separate field sites in Berkeley, five groups of newly hatched
larvae were placed on appropriate branches of single C. monspessu-

Volume 43, Number 4

269

NUMBER IN GROUP

DAYS

Fig. 5. Survivorship curves of five fifth instar larval groups. Declines in numbers
marked with an arrow could be attributed with certainty to defoliation of host plant,
virus disease, and pupation as indicated. The otherwise constant numbers imply a lack
of predation. Observations were made during the period October-December 1986.

lanus plants in summer 1988. At each site, continuous observations were
made from dawn to dusk on activities of the larvae, and any predation
attempts on them noted. Similar observations were made on two sub-
sequent occasions during the second and third instars at one site, and
on four occasions (during second, fourth, and fifth instars) at the other
site. Total counts were made at both sites on four dates during devel-
opment. Each group had about 30 larvae at the start, and mortality of
early instars was high (Fig. 4). This was in marked contrast to the data
for fifth instars that was noted in an earlier year (Fig. 5). During the
periods of observation, Hemiptera (Anthocoridae) fed on first and sec-
ond instar caterpillars, and Hymenoptera (Vespidae) removed middle
instars. In one of these experiments, 80 larvae (53%) were lost in 17
days, giving an average daily loss of 4.4 larvae. About 40% of the loss

270

Journal of the Lepidopterists’ Society

Table 2. Numbers of invertebrate predators relevant for U. reversalis collected by
beating bushes of the host plant, Cytisus monspessulanus.

Sample date

Numbers per 60 bushes (6 sites)

Salticidae

Thomisidae

Heteroptera

Chrysopidae

18 July 1988

29

13

5

4

27 July 1988

22

6

3

0

18 August 1988

17

4

4

0

could be accounted for by observed predation levels. In the second
experiment, 15% of loss could be similarly accounted for.

During the period of held experimentation, censuses of potential
predators were made by beating bushes, collecting the arthropods, and
testing the predators among them with first and second instar cater-
pillars of JJ. reversalis as prey. Ants and coccinellid beetles did not
attack larvae that they were confined with in small containers in the
lab, while neuropterans ( Chrysopa ) and heteropterans (Anthoridae, Na-
bidae) did. Spiders in the families Salticidae and Thomisidae were
deemed relevant predators from daily held observations, and both of
these attacked larvae in the lab in no-choice tests. The census data
indicate that bushes of Cytisus averaged 0.4 to 0.8 relevant predators
per bush (Table 2). To test whether invertebrate predation is a signif-
icant cause of mortality, we conducted held experiments in which some
groups of larvae were protected from predation.

Survivorship of U. reversalis larvae placed on branches of C. mon-
spessulanus, with or without net bags over the branches, was compared.
In April-May 1988 approximately 30 neonate larvae were placed on
each of three plants at each of hve sites in the San Francisco Bay area.
Each plant had a protected and an unprotected group. A similar ex-
periment was done in June-July 1988 on a total of 21 plants at six sites.
In all cases, survivorship was considerably higher on bagged branches
(Table 3), and although there may be other unknown mortality factors,
it is likely that the exclusion of predators was the primary reason for
the difference. Fifth instar larvae in five natural groups at three sites
in Berkeley were monitored in fall and winter 1986. These represented
the last generation of the year and persisted as fifth instars over 5 to
12 weeks because of low temperatures. In all cases the population
declines could be attributed to causes other than predation (Fig. 5), in
spite of the exposed positions of the larvae and the abundance of birds.
Rejection behavior of birds was observed. On two occasions during four
all-day observations, a bird was observed to pick up a caterpillar, sub-
sequently drop it, and proceed to rub its beak on the branches of a
tree. Aposematism is most commonly associated with protection from

Volume 43, Number 4

271

Table 3. Survivorship of larvae in the field with or without protection from predators.
In all four comparisons, survival was significantly greater in the protected groups (Chi
Squared Test, P < 0.01).

Number of groups

Mean 9

l surviving (±SD)

(total number of larvae)

1-2 weeks

3 weeks

Spring 1988

Protected

15 (482)

76(15)

36 (15)

Unprotected

15 (499)

39 (4)

9(8)

Summer 1988

Protected

21 (530)

61 (18)

37 (23)

Unprotected

21 (503)

38 (17)

18(14)

predation by birds, and our limited field observations indicate that birds
do not prey on larvae of U. reversalis.

Predation Experiments

The possible defense against invertebrate predators was tested by
comparing the relative acceptability of U. reversalis to two invertebrate
predators in a situation where food deprivation was clearly not driving
indiscriminate predation. In the first case, the vespid wasp Mischocyt-
tarus flavitarsus (Saussure), a generalist predator, was employed (Ber-
nays 1988). Naturally hunting wasps from nests in a large greenhouse
were allowed to forage in open arenas (1m x lm x lm)in which
caterpillars of U. reversalis feeding on C. monspessulanus were avail-
able at the same time as Trichoplusia ni (Hiibner) (Noctuidae), Estig-
mene acrea (Drury) (Arctiidae) or Heliothis virescens (F.) (Noctuidae),
each feeding on its own host plants in close proximity. In each of these
three experiments, laboratory-reared larvae of U. reversalis and the
alternate species were presented at similar sizes and densities. In each
case the alternative prey were always attacked early in the experiments.

Table 4. Numbers of caterpillars taken by the wasp Mischocyttarus flavitarsus (Ves-
pidae) during choice trials. Experiments were terminated after approximately half of
either prey species was removed. In each of the three experiments the differences in
acceptability are significantly different (Chi Squared Test, P < 0.001).

Numbers of size-matched caterpillars
Nos. at start % taken

1.

Estigmene acreae (Arctiidae)

20

50

Uresiphita reversalis

20

0

2.

Heliothis zea (Noctuidae)

25

52

Uresiphita reversalis

25

0

3.

Trichoplusia ni (Noctuidae)

22

59

Uresiphita reversalis

30

0

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Journal of the Lepidopterists’ Society

Experiments were stopped when approximately half of one species was
taken. In no case were any U. reversalis larvae taken before half of the
alternate species was removed (Table 4).

Experiments on predation by Argentine ants, Iridomyrmex humilis
(Mayr) (Formicidae: Dolichoderinae), also showed that U. reversalis is
extremely unacceptable relative to many other species of caterpillars
(Bernays & Cornelius 1989).

Conclusions

Uresiphita reversalis larvae have all the characteristics of aposematic
caterpillars. They are brightly colored, relatively inactive, found in
groups, and they feed actively during the day. Their favored food is
rich in toxic quinolizidine alkaloids, which are phagostimulants. Al-
kaloids that are not excreted after ingestion are deposited in the cuticle.
Larvae are apparently distasteful to at least one avian species, wasps
and ants.

Predation data are limited, but on the basis of the observations pre-
sented here, it appears that in spite of being chemically defended, U.
reversalis larvae do suffer extensive predation. Losses to invertebrate
predators apparently account for a significant proportion of the mor-
tality in the early instars. Arthropods with piercing and sucking attacks
seem to be the dominant predators, perhaps circumventing any pro-
tection provided by the alkaloidal cuticle. The early larval instars are
much more vulnerable to predation than the last larval instar. Thus,
chemically defended aposematic species may nonetheless suffer high
mortality from predation.

Acknowledgments

R. V. Barbehenn, R. F. Chapman, M. L. Cornelius, J. Hamai, M. A. Graham, M. Guard
and N. E. Stamp helped in various ways. This research was funded by USDA CSRS grant
no. 88-38300-3628 to EAB.

Literature Cited

Bernays, E. A. 1988. Host specificity in phytophagous insects: Selection pressure from
generalist predators. Entomol. Exp. Appl. 49:131-140.

& M. L. CORNELIUS. 1989. Relative acceptability of caterpillars with different

host range to the generalist predator Iridomyrmex humilis. Oecologia 79:427-430.

Johnson, N. D. & B. L. Bentley. 1988. Effects of dietary protein and lupine alkaloids
on growth and survivorship of Spodoptera eridania. J. Chem. Ecol. 14:1391-1404.

, B. Liu & B. L. Bentley. 1987. The effects of nitrogen fixation, soil nitrate,

and defoliation on the growth, alkaloids, and nitrogen levels of Lupinus succulentus
(Fabaceae). Oecologia 74:425-431.

Kinghorn, A. D. & M. F. Balandrin. 1984. Quinolizidine alkaloids of the Legumi-
nosae: Structural types, analyses, chemotaxonomy and biological activities, pp. 118-

Volume 43, Number 4

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195. In Pelletier, S. W. (ed.), Alkaloids: Chemical and biological perspectives. Vol.
2. John Wiley and Sons, New York.

Montllor, C. B., E. A. Bernays & R. V. Barbehenn. The importance of alkaloids in
the biology of Uresiphita reversalis (Pyralidae: Lepidoptera). J. Chem. Ecol. In press.

Munroe, E. 1976. Pyraloidea (in part), p. 57. In Dominick, R. B., et al., The moths of
America north of Mexico. Fasc. 13.2A. E. W. Classey Ltd. & The Wedge Entomol.
Res. Found., London.

Rothschild, M. 1972. Secondary plant substances and warning coloration in insects,
pp. 59-83. In van Emdem, H. F. (ed.), Insect/plant relationships. Symp. Roy. Ento-
mol. Soc. 6. Blackwell Scientific Publications, Oxford.

Schneider, D. 1987. The strange fate of pyrrolizidine alkaloids, pp. 123-142. In Chap-
man, R. F., E. A. Bernays & J. G. Stoffolano Jr. (eds.), Perspectives in chemoreception
and behavior. Springer-Verlag, New York.

Stamp, N. E. & M. D. Bowers. 1988. Direct and indirect effects of predatory wasps
(Polistes sp.: Vespidae) on gregarious caterpillars ( Hemileuca lucina : Saturniidae).
Oecologia 75:619-624.

Wink, M. 1987. Chemical ecology of quinolizidine alkaloids, pp. 524-533. In Waller,
G. R. (ed.), Allelochemicals: Their role in agriculture and forestry. American Chemical
Society, Washington, D.C.

& L. Witte. 1984. Turnover and transport of quinolizidine alkaloids. Diurnal

fluctuations of lupanine in the phloem sap, leaves and fruits of Lupinus albus L.
Planta 161:519-524.

Received for publication 10 March 1989; accepted 28 August 1989.

Journal of the Lepidopterists’ Society
43(4), 1989, 274-288

BIOGEOGRAPHICAL AFFINITIES OF THE BUTTERFLIES
OF THE SOUTHWEST CARIBBEAN ISLANDS,

SAN ANDRES AND PROVIDENCIA

David Spencer Smith

Department of Zoology and Hope Entomological Collections,

The University Museum, Oxford 0X1 3PW, England

Eugene Munroe

Granite Hill Farm, RR2, Dunrobin, Ontario KOA 1T0, Canada

AND

Stuart J. Ramos

University of Puerto Rico, Mayagiiez, Puerto Rico 00708, USA

ABSTRACT. San Andres and Providencia are small islands east of Nicaragua. N. D.
Riley, in his Field Guide to the Butterflies of the West Indies (1975), included these
islands in his biogeographical definition of the West Indies. Subspecific analyses of butterfly
collections made in August 1986, however, show that, of 19 species collected, the 10 that
have one or more subspecies in the Antilles are all represented on San Andres and
Providencia by the subspecies occurring on the mainland. Thus, we conclude that this
area of the SW Caribbean shows no affinity with Antillean populations, but is clearly an
extension of the neighboring populations on the mainland of Central America. Two species
are newly recorded for any Caribbean island: Cymaenes odilia (Hesperiidae) on San
Andres and Chlorostrymon telea (Lycaenidae) on Providencia. Based on present, but
probably incomplete, sampling, it appears that San Andres and Providencia maintain
discrepant butterfly faunas. These islands are equidistant from a large donor source of
immigrants, but differ markedly in geological origin and present vegetation.

Additional key words: island biogeography, subspeciation, dispersal, Nicaragua.

The “West Indies” constitute an area of particular biogeographical
interest because they possess components of both the Nearctic (North
American) and Neotropical (Central and South American) faunas dis-
tributed across a chain of islands, varying greatly in size, topography,
and geological history. As a faunistic zone, this region is generally
considered to include the Bahamian islands to the north, the Greater
Antillean islands of Cuba, Hispaniola, Jamaica (with the small satellite
Cayman Islands), and Puerto Rico, and the Lesser Antillean chain from
the Virgin Islands east of Puerto Rico south to Grenada, but excluding
Trinidad and Tobago, which are insular faunistic extensions of the
adjacent continental mainland.

Riley (1975), in the only general account of the butterflies of this
area, extended the perimeter to the south and west to include three
islands: Swan (in line with the Islas de la Bahia chain off Honduras),
Old Providence (Providencia), and St. Andrew (San Andres). This report
is concerned with the biogeographic status of the butterflies of the last

Volume 43, Number 4

275

two islands and their supposed affinity with the West Indian (or An-
tillean) faunistic zone.

San Andres and Providencia are small isolated islands in the SW
Caribbean Sea, situated, respectively, ca. 197 km and 235 km due east
of the Central American mainland (Nicaragua) (Fig. 1). Providencia
lies ca. 87 km NNE of San Andres. The latter is situated ca. 365 km
due north of the coast of Panama, while Providencia is ca. 626 km from
Jamaica, the nearest neighbor among the Antillean Islands. Despite
their proximity, and their relative nearness to the continental landmass,
San Andres and Providencia are thought to have upheaved indepen-
dently in the early Tertiary and never to have had a land bridge either
between them, or between either island and the mainland (Parsons
1956).

The contours of Providencia and San Andres, with indication of
populated areas (and, for Providencia, the altitude of the central peaks)
are illustrated in Fig. 1. These maps are based on those in Parsons (1956)
and on a Colombian Government publication: “Mapa de la Intendencia
Especial de San Andres y Providencia” (1981). Parsons (1956) has pro-
vided a valuable and interesting account of the geology, history, and
ecological consequences of colonization, and of the social and cultural
geography of San Andres and Providencia. The following summary is
abstracted from that work, with some details obtained from Emmel’s
(1975) account.

Neither island has a known history of pre-Columbian settlement.
Following initial colonization in the early 17th century, much of the
islands5 original forests were removed for ship building and repair,
notably during the 18th century. The two islands differ strikingly in
terrain and in their present pattern of vegetation. Providencia is steeply
mountainous with igneous peaks (to 360 m) in the interior of the island.
Originally, Providencia was densely forested (Parsons 1956); today, the
island supports lowland agriculture, the higher areas being dominated
by grass and cattle grazing, extending much of the way to the peaks
where, in a few areas, the vegetation is described by Proctor (in Parsons
1956) as showing “distinctly Central American” affinities. San Andres
is a slender N-S oriented block of limestone (thought to overlie an
igneous base) with a single cliff-bound hill rising to ca. 100 m. Destruc-
tion of original forest occurred as on Providencia; a period of cotton
planting followed, and in the 1850’s the island was thickly planted with
coconut palms, a virtual monoculture that dominates much of the island
today, although the ancient palm stands are untended and of little
economic importance. Abandoned fields and paths through these plan-
tations provide the most ready access to the interior of San Andres.

Commercial development, primarily for the tourist industry, was

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Journal of the Lepidopterists’ Society

virtually non-existent on either island when Parsons wrote, but was
more extensive when Emmel visited the islands (in 1967-68) and clearly
continues. On San Andres (Fig. 1) development is centered primarily
around the town. On Providencia, however, other than expansion of
the main settlement and some development along the perimeter road,
little change has occurred since Parsons’ visit in 1953, and little en-
croachment into the interior has taken place.

Butterfly Faunas of the Islands

The only published records of the butterflies of these small islands
are those of Emmel (1975) who, with coworkers, collected on San Andres
during five visits, and on Providencia during two brief visits, in 1967
and 1968. Work on the former was in February-March and June-July,
while Providencia was sampled on one day in June, and two days in
July, 1968. Emmel documented 17 species from the two islands. Al-
though of considerable value, these records generally did not include
subspecific determinations, and thus afford little information on the
affinities of these island faunas to continental America and the Antilles,
where subspeciation has been a conspicuous factor (Riley 1975).

The present account is based on an intensive collecting visit by one
of us (DSS) to these two islands between 7-23 August 1986, apportioned
between San Andres (9 collecting days) and Providencia (5 collecting
days). Several species recorded by Emmel were not found in 1986 and,
conversely, a number of species were newly recorded (Table 1). The
extent of the discrepancy is noteworthy, and contributory factors may
include differing habitat sampling (even on small islands), seasonal
fluctuation in adult populations, presence or absence of occasional va-
grant species, and recent extinction and/or colonization.

All species collected were determined where appropriate to subspe-
cies, primarily to establish the southwestern limits of the Antillean
faunal area and to evaluate the appropriateness of Riley’s SW-directed
extension of this zone. Taxonomic determination of material collected
in 1986 was facilitated by comparison with often extensive series in the
Allyn Museum of Entomology, Sarasota, Florida, the Hope Entomo-
logical Collections, University of Oxford, and the British Museum (Nat-
ural History), London. The sequence of taxa follows that of Miller and
Brown (1981). All specimens collected in 1986 are deposited in the
Hope Entomological Collections, the University Museum, Oxford, Eng-
land.

The following list of species includes both Emmel’s published records
and our records from August 1986; those species recorded by Emmel
but not seen during this study are enclosed in parentheses. A compar-

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277

278

Journal of the Lepidopterists’ Society

Table 1. Butterflies collected in 1986 (this study) and in 1967-68 (Emmel 1975).

San Andres

Providencia

Hesperiidae

Hesperiidae

+ + Urbanus d. dorantes **

+ + Urbanus d. dorantes **

+ + Urbanus p. proteus

+ + Urbanus p. proteus **

+ + Cymaenes odilia trebius**

Hylephila phyleus

Panoquina sylvicola**

( Hylephila phyleus)

Papilionidae

+ + Battus p. polydamas **

Pieridae

Pieridae

+ + Ascia m. monuste**

Phoebis sennae

Phoebis sennae

+ + P. p. philea**

+ + P. p. philea**

Abaeis nicippe**

+ + Eurema venusta limbia**

( Phoebis statira)

( Eurema daira )
(E. lisa )

Lycaenidae

Lycaenidae

+ + Leptotes cassius striata

Chlorostrymon telea**

+ + Leptotes cassius striata
+ + Hemiargus hanno zachaeina**

Heliconiidae

Heliconiidae

+ + Agraulis v. vanillae

+ + Agraulis v. vanillae

( Dryas iulia)

(Dryas iulia)

Nymphalidae

Nymphalidae

— Vanessa cardui **

+ + Anartia jatrophae luteipicta**

Junonia genoveva **

+ + Anartia jatrophae luteipicta

( Historis odius)

Historis odius
( Junonia evarete)

( Siproeta stelenes)

Danaidae

Danaidae

(. Danaus plexippus )

(Danaus plexippus)

+ + Subspecies shared with Central America.

Species invariant in the area.

** Newly recorded in 1986.

( ) Recorded by Emmel (1975) but not seen in 1986.

ative summary of species recorded from both islands is given in
Table 1.

Hesperiidae

Urbanus p. proteus Linnaeus (n = 12). Emmel recorded this skipper as common on
San Andres, as it was during the 1986 survey, when a single specimen was also recorded
from Providencia. A recent revision of the proteus group (Steinhauser 1981) stresses the
external similarity of a number of related species occurring in Central and South America,
reliably separated only by genitalic characters. All specimens collected in 1986 were
dissected and confirmed as U. p. proteus : the generally large size of the forewing mac-
ulations shows the San Andres and Providencia populations to represent the Continental

Volume 43, Number 4

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nominate subspecies (Fig. 5A), rather than the Antillean U. proteus domingo Scudder
(Fig. 5B).

U. d. dorantes Stoll (n = 6). This species, absent from Emmel’s list, is newly recorded
from San Andres and Providencia, occurring frequently on both islands. All male spec-
imens collected were determined as U. dorantes, by comparison with Evans (1952). All
are clearly referable to the nominate subspecies occurring on the mainland of the Amer-
icas rather than to either of the Antillean subspecies ( JJ . d. cramptoni Comstock and
U. d. Santiago Lucas), which differ, respectively, in reduction of forewing maculation
(Comstock 1944) and underside markings (Riley 1975).

(Pyrgus oileus Linnaeus). This widely distributed skipper, recorded by Emmel as
sporadic around Big Pond (San Andres) in February 1968, was seen on neither island in
August 1986.

Cymaenes odilia trebius Mabille (n = 10). This skipper, in flight much resembling C.
tripunctus , was found commonly, but very locally on San Andres in August 1986 with
a pronounced mid-morning flight period, nectaring at Stachytarpheta flowers on an E-
W track crossing the island near the southern point. This subspecies (Fig. 6) is recognized
primarily by the extent of grey scaling on the forewing underside and by uniformity of
the ground color and the distribution and size of the pale spots on the hindwing beneath
(Evans 1955). It extends from southern Texas (Howe 1975) through Central America into
northern Colombia [Type locality Bogota]. This species has not previously been recorded
from any Caribbean island.

Hylephila phyleus Drury (n = 2). Perhaps the most widely distributed of New World
skippers, ranging from Canada to Argentina, H. phyleus was noted by Emmel as possibly
present on San Andres, but occurring commonly on Providencia. It was rare on San
Andres in August 1986, and not recorded from Providencia. This skipper has no described
insular subspecies in the West Indian region.

Panoquina sylvicola Herrich-Schaefifer (n = 1). Another widely ranging species (from
the southern U.S. south to Argentina and through the Antilles), P. sylvicola is newly
recorded from San Andres by a single specimen (August 1986).

Papilionidae

Battus p. polydamas Linnaeus (n = 7). This swallowtail is newly recorded from San
Andres, where it was seen, but never common, in August 1986 in scrubby beachside areas,
where Aristolochia was present. Riley (1975) lists 13 insular subspecific taxa sometimes
differentiated on island pairs separated by remarkably short distances. The San Andres
population represents the nominate subspecies (Fig. 2A), which Riley (1975) considered
confined to the mainland of Central and South America and not now present on any
Antillean island, “. . . though it seems to have been present earlier on the island of
Barbados.” A representative Antillean subspecies ( B . polydamas polycrates Hopffer from
Hispaniola and Puerto Rico) is illustrated in Fig. 2B. The island forms are distinguished,
one from another, by variations in a number of characters, including the shape, size and
tint of the pale upperside spots, and, on the underside, by the uniformity (or otherwise)
of the brown ground color of the fore- and hind wings, the size, shape and brightness of
the submarginal red markings on the hindwing and, where present, the size and shape
of the accompanying silver wedges and distal cream spots (Riley 1975). Each described
subspecies is distinctive and the extremes are dramatically different. The nominate sub-
species, ranging through Central and much of South America with little variation, cannot
be confused with any of the island races. The generally pale brown-grey color of the
hindwing beneath, the absence of submarginal pale spots and silver patches associated
with the submarginal red markings, and the reduction of the last two pairs of narrow
lunules, are diagnostic.

Pieridae

Ascia m. monuste Linnaeus (n = 7). This butterfly was first recorded from San Andres
in August 1986, when it was locally frequent in disturbed coastal waste land and abandoned

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281

fields. Males (all collected) are large, white beneath, with prominent marginal markings
(Fig. 3A), and are referable to the continental nominate subspecies which, according to
Riley (1975), elsewhere enters the Antillean area from South America via Trinidad into
the southern Lesser Antilles, possibly occurring further North as an occasional vagrant
(Comstock 1944). Despite its migratory proclivity, A. monuste has evolved two subspecies
in the West Indies: A. m. eubotea Latreille ( evonima Boisduval; see Comstock 1944) is
less heavily marked, pale yellow-ochreous on the hindwing beneath, and occurs from
Florida and the Bahamas through the Greater Antilles, and A. m. Virginia Godart, very
lightly marked on the upperside, occupying the area between the Virgin Islands and St.
Vincent. For comparison with A. m. monuste, subspecies eubotea is illustrated in
Fig. 3B.

Phoebis sennae Linnaeus (n = 35). This widely distributed sulfur was noted by Emmel
on each visit to San Andres and Providencia; in August 1986 it was common on the former
and occasional on the latter. Antillean and southern U.S. populations are usually regarded
as referable to the nominate subspecies, intergrading with the more northern P. s. eubule
Linnaeus. The mainland tropical subspecies, marcellina Cramer, is described (Howe 1975)
as showing a more orange and heavily patterned underside (male) and conspicuous
patterning and a ground color of pinkish orange (female) beneath. Klots (1960) regarded
this last subspecies as “of dubious worth.” Specimens collected in 1986 from San Andres
and Providencia are quite variable and fall within the range of variation of very extensive
series examined from the Antilles and the mainland of Central and South America. They
do not, therefore, yield reliable information about the affinities of the San Andres and
Providencia populations.

Phoebis p. philea Johansson. This large and (in the male) conspicuously colored species
is newly reported from both San Andres and Providencia (August 1986). Although no
specimens were collected, close sightings of males established the identity of the nominate
subspecies, which occurs from southern Florida through the mainland south to Brazil
(Riley 1975). The two Antillean subspecies, P. p. huebneri Fruhstorfer (Cuba) and P. p.
thalestris Illiger (Hispaniola) are readily distinguished from p. philea by a conspicuous
large black spot at the end of the forewing cell — certainly absent in specimens seen on
San Andres and Providencia.

( Aphrissa statira Cramer). This sulfur was reported by Emmel from two reliable
sightings on Providencia in July 1968. The nominate subspecies occurs on the mainland
and according to Klots (1960) migrates annually via Trinidad into the Lesser Antilles.
Two Antillean subspecies have been described, from Cuba and Hispaniola, but until
specimens from San Andres and/or Providencia are examined, the affinity of statira in
these islands cannot be assessed.

(Eurema daira Godart). Emmel recorded this species as “relatively rare” on San Andres
in July, but absent in February, noting that it is widely distributed in the Antilles and
the mainland of Central America (and indeed ranges from the SE U.S. to Brazil). It was
not found on San Andres or Providencia in August 1986.

Eurema venusta limbia Felder & Felder (n = 32). Very surprisingly, this “southern”
Eurema was not encountered by Emmel, yet was one of the most common butterflies on
San Andres in August 1986, notably in abandoned fields and neglected coconut plantations
in the southern part of the island. It was not found on Providencia. Eurema v. emanoma
Dillon, the Antillean subspecies (Fig. 9B), is found in the Lesser Antilles, from Grenada
north to Guadeloupe (Pinchon & Enrico 1969, Riley 1975). Specimens collected on San
Andres represent the more heavily marked continental subspecies E. v. limbia occurring
in Central America (Fig. 9A). The reported food plant, Mimosa (Riley 1975), is present
on San Andres, and the discrepancy between the 1968 and 1986 records may reflect a
recent successful colonization of the island from Central America.

(Eurema ( Pyrisitia ) lisa Boisduval & Leconte). As with E. daira, this small sulfur
was recorded by Emmel as relatively rare on San Andres in July and absent in February.
It was not found on either in August 1986.

Abaeis ( Eurema ) nicippe Cramer (n = 16). This species was not found on San Andres
in August 1986, yet was one of the most common butterflies on Providencia at a time
when few adult butterflies were seen. On Providencia, nicippe was generally distributed
along the road around the perimeter of the island. No subspecies have been described.

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Journal of the Lepidopterists’ Society

Fig. 7. Hemiargus hanno males, undersides. A: H. h. zachaeina, Providencia, August
1986. B: H. h. filenus, New Providence Is. (Bahamas), July 1980. (x2.0)

Fig. 8. Leptotes cassius males, undersides. A: L. c. striata, Providencia, August 1986.
B: L. c. theonus, Big Pine Key, Florida, September 1984. (x2.0)

Fig. 9. Eurema venusta males, uppersides. A: E. v. limbia, San Andres, August 1986.
B: E. v. emanoma, Bequia Is. (St. Vincent Grenadines), August 1987. (xl.4)

Lycaenidae

Hemiargus hanno zachaeina Butler & H. Druce (n = 10). This lycaenid was newly
recorded for Providencia in August 1986, where it occurred in the same localities and in
comparable numbers with Leptotes cassius, discussed below. Comparison between the
Providencia material and extensive series in the British Museum (Natural History) estab-
lishes the former’s grouping with the continental subspecies H. h. zachaeina [Type locality
Costa Rica]. Hemiargus h. zachaeina (Fig. 7A) is readily distinguished from the three
described island races (Fig. 7B) by the presence of two well demarcated eye-spots near
the anal angle of the hindwing beneath, rather than one.

Leptotes cassius striata W. H. Edwards (n = 12). Emmel noted L. cassius as present
on Providencia and distinctly colonial on San Andres. During the 1986 survey, it was
widespread and generally common on San Andres, while on Providencia, at a time when
adult butterflies were very scarce, it was the species seen most frequently. Three Caribbean
sub-species have been described: theonus Lucas from Florida, the Bahamas, and Greater
Antilles; and two subspecies from the Lesser Antilles (Riley). Reference to extensive
material in the British Museum (Natural History) clearly associates both San Andres and
Providencia specimens with the mainland subspecies L. c. striata [Type locality Bogota],
which ranges throughout Central America. Populations of L. cassius show considerable
variation, but L. c. striata (Fig. 8A) is distinguishable from other forms (Fig. 8B) by
reduction of the blue suffusion in the upper surface of the female and, in both sexes, in
the generally paler brown underside markings, and in the reduced size and more circular
shape of the hindwing eye-spots.

Chlorostrymon telea Hewitson (n = 1). A single worn female of this continental
hairstreak was collected on Providencia (17 August 1986), representing a new record for
a Caribbean island.

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283

Heliconiidae

Agraulis v. vanillae Linnaeus (n = 33). Emmel recorded this species as locally abundant
on San Andres and noted it from Providencia. In August 1986 it was locally common on
San Andres and generally distributed in lowland disturbed areas on Providencia. Emmel
described it as resembling “. . . typical Florida A. v. nigrior Michener, although the black
forewing spots often form a bar-like continuous row.” Material collected in August 1986
does not conform to this description: specimens are much more lightly marked than
nigrior or the widespread Antillean subspecies A. v. insularis Maynard, and correspond
to the nominate subspecies, which according to Riley (1975) enters the southern Lesser
Antilles from the mainland (Fig. 4).

( Dryas iulia Fabricius). Emmel found this widespread species (of which numerous
Antillean subspecies have been described) commonly on San Andres “throughout the
year” and also recorded it from Providencia. Surprisingly, D. iulia was not seen on either
island in August 1986.

Nymphalidae

(Junonia evarete Cramer). Emmel reported this species, as Precis evarete zonalis (C.
& R. Felder), to be relatively rare on San Andres, from one locality in February and
July, and did not report it from Providencia. The proposed reinstatement (Turner and
Parnell 1985) of Junonia evarete is adopted here for this species, which ranges widely
from S Texas and Florida through the West Indies and Central America into South
America. It was not seen on San Andres or Providencia in August 1986.

Junonia genoveva Stoll (n = 25). As for the preceding species, the terminology proposed
by Turner and Parnell (1985) is adopted here. It was not recorded by Emmel, but in
1986 it occurred in beach areas and abandoned fields on San Andres, but was not found
on Providencia. Specimens collected correspond well with Turner and Parnell’s figures
of Jamaican genoveva; this species shows considerable general and seasonal variation, but
the San Andres material falls within the range shown by series examined from Hispaniola,
Puerto Rico, Trinidad, Guatemala, Costa Rica, and Panama.

Anartia jatrophae luteipicta Fruhstorfer (n = 15). This is another nymphalid exhib-
iting incompletely documented general and seasonal variation. According to Riley (1975)
the nominate subspecies is primarily South American, extending into the Lesser Antilles
north to St. Kitts. A number of Antillean subspecies have been described (Munroe 1942,
Riley 1975), generally darker than populations on the mainland. Emmel found this species
to be fairly common on San Andres in July, though rare in February. In August 1986, it
was locally quite common on San Andres, especially around ornamental flowers in the
town and at the airport, and a single specimen was collected on Providencia. These
specimens correspond most closely (particularly in their generally light markings and in
displaying a conspicuously yellow distal margin of the hindwing) to the Central Amer-
ican subspecies A. j. luteipicta [Type locality Honduras], noted by Munroe (1942) as
ranging south from Mexico and merging with the nominate subspecies in Panama.

Vanessa cardui Linnaeus (n = 1). Howe (1975) suggests that this cosmopolitan and
invariant species is largely nonresident over much of its range in North and Central
America, dispersing widely from a few breeding areas. It has been recorded occasionally
from many of the Antillean islands (Riley 1975) and is newly recorded from San Andres
from a single worn specimen collected in August 1986.

(Siproeta stelenes Linnaeus). This conspicuous and widely ranging nymphalid was
regularly recorded by Emmel at one locality on San Andres during February 1968, and
more rarely in other months, but was not found on either island in August 1986.

Historius odius Fabricius. This species was noted by Emmel from both islands, but
rarely on San Andres (July). In August 1986, it was seen on several occasions, along E-
W tracks crossing the southern part of San Andres, but was not recorded from Providencia.
Emmel suggested that the few specimens encountered on San Andres may have been
vagrants from the sister island “or even the mainland,” but Cecropia, the larval food
plant, occurs on San Andres, and this insect may well be resident there. Because no

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Journal of the Lepidopterists’ Society

specimens were collected in 1986, it was not possible to determine whether the San Andres
population belongs to the nominate subspecies, which occupies the Greater Antilles, or
to H. odius orion Kaye, which ranges through tropical America to Argentina and (Riley
1975) enters the Lesser Antilles from the mainland.

Danaidae

( Danaus plexippus Linnaeus). Emmel recorded “scattered and relatively worn” spec-
imens on all visits to San Andres (February, March, June, July), and also on Providencia.
It was not seen on either island in August 1986, and its resident status is unknown. Emmel
found no Asclepiadacae on San Andres, but potential larval food plants could be present
in the shrubby untended coconut plantation understorey of that island, or in the grasslands
and disturbed areas of Providencia.

Discussion

An island’s observed fauna and flora may be regarded as the result
of a balance between addition by successful colonization from outside
and, potentially (in long established populations), by evolution of new
species and/or races, countered by extinctions (Munroe 1953, 1957,
1984, MacArthur & Wilson 1967, Williamson 1981). In the West Indian-
Caribbean region, evolution of butterfly species and subspecies, fre-
quently endemic to a single island, is a conspicuous feature (Riley 1975),
particularly on the large islands of the Greater Antilles (Cuba, Jamaica,
Hispaniola, and Puerto Rico). The present butterfly faunas of these
islands, together with the Bahamian chain, appear to have drawn pri-
marily from a Central American species pool, while the small islands
of the Lesser Antilles show more South American donor input (Fox
1963, Scott 1972, Riley 1975, Munroe 1984).

Scott (1972) has provided a survey of the affinities of Antillean but-
terflies with respect to individual islands and island groups, and the
continental land masses of North, Central, and South America. His
analysis makes use of an “index of faunal resemblance” (see also Munroe
1957) defined as “the percentage of species found on the island with
the smaller fauna which are the same as those on the island [or conti-
nental land mass] with the larger fauna.” Although providing a valuable
basis for analyzing faunal affinities across the entire West Indian region,
Scott’s index is of limited value when employed at the specific level to
assess short-range affinities, especially between an island pair of greatly
disparate area, or between a small island and a nearby continental land
mass. In the context of San Andres and Providencia Islands, Emmel
(1975) cites Scott’s recognition of the great dispersal ability of many
species found in the Antilles, and (of the 17 species he recorded) notes
that “All . . . are also found on the adjacent Nicaraguan mainland area,
and all represent species that are migratory or are known to be frequent
colonisers.”

At the level employed, this view is substantially accurate, with only

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285

Chlorostrymon telea (Providencia) and Cymaenes odilia (San Andres)
representing new Caribbean island records of continental species with-
out obvious dispersal potential. However, if the “higher resolution”
data provided by analysis of subspecies are taken into account, the
biogeographical status of Providencia and San Andres takes on a very
different aspect.

In the following discussion, only the 19 species recorded during the
1986 survey are considered, although Emmel’s records of species not
seen during the present field work raise the total species count for these
two islands substantially. Except where noted, the species records from
the two islands may be pooled for purposes of discussion of their affin-
ities. The two species noted above ( Chlorostrymon telea and Cymaenes
odilia ) are clearly derived from continental Central or South America;
Scott’s index would remain at 1.0 with respect to continental America,
and the addition of 2 species (of 19) not present in the Antillean islands
would reduce the index with respect to the Caribbean area from 1.0
(as implied by Emmel) to 0.89.

Several species recorded from San Andres and Providencia (Hylephi-
la phyleus, Panoquina sylvicola, Abaeis nicippe, Phoebis sennae, Va-
nessa cardui and Junonia genoveva) are considered to be invariant in
the area (Antilles and the adjacent tropical American mainland) and
thus provide no information on biogeographical origins of the islands’
populations. To this group must be added Historius odius, given the
absence of subspecifically determined material. The remaining 10 taxa,
determined to the subspecific level ( Urbanus d. dor antes, U. p. proteus,
Battus p. polydamas, Ascia m. monuste, Phoebis p. philea, Eurema
venusta limbia, Leptotes cassias striata, Hemiargus hanno zachaeina,
Agraulis v. vanillae, and Anartia jatrophae luteipicta) , in each instance
possessing one or more Antillean subspecies, are represented on San
Andres and/or Providencia by a subspecies in common with continental
tropical America. Thus, while the San Andres-Providencia butterfly
populations remain at 1.0 (Scott index) with respect to Central and
South America, this index drops to a maximum of 0.39 (7 of 19) with
respect to the Antillean islands; the balance being largely accounted
for by wide-ranging species that afford no affinity data.

It is now clear that the butterfly faunas of these islands, although
presumably much affected by destruction of native habitats, consid-
erable on Providencia and virtually complete on San Andres, represent
extensions of the adjacent Central American mainland with no docu-
mented Antillean component.

This not unexpected conclusion exemplifies the value of using sub-
specific information in approaching the question of affinity determi-
nation in island groups such as the Antilles, an approach for comparing

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Journal of the Lepidopterists’ Society

closely related faunas first proposed by Munroe (1957). Another recent
application of this approach documented the Puerto Rican, rather
than Hispaniolan, contribution to the present butterfly fauna of Mona
Island (Smith et al. 1988). In the case of San Andres and Providencia,
their proximity to the continental mainland evidently outweighs po-
tential wind-borne dispersal from the Antilles by predominantly easterly
winds (Fox 1963, Calvesbert 1973). In considering records for the two
islands, Emmel suggests that “ Every species found on San Andres must
occur on Providencia, and with the greatly increased diversity of vege-
tation [on the latter], one would presume that several additional species
of butterflies should occur,” a forecast that has received some confir-
mation here. However, we do not feel that one of a pair of small islands,
of approximately equal area and similarly positioned with respect to a
large and faunistically rich area, should necessarily support a butterfly
fauna fully represented on the other.

Even if islands in such a pair are ecologically similar, chance variation
in the distribution of incoming vagrant species, reaching one and not
the other, could well lead to discrepant faunas. Where the islands differ
substantially, as do San Andres and Providencia, the probability of
differences between the two faunas may well be enhanced by different
environmental spectra that engender differing probabilities of success-
ful colonization.

Present documentation of the butterfly faunas of San Andres and
Providencia is undoubtedly very incomplete, relying only on brief pe-
riods of sampling in 1967-68 and 1986. However, of the nine species
recorded in August 1986 on Providencia, three were not seen on San
Andres, of which Abaeis nicippe is widely ranging, while Chlorostry-
mon telea is not obviously dispersive. Hemiargus hanno, at the specific
level, is one of the widest-ranging lycaenids in the American tropics,
yet a continental subspecies is noted on only one island (Providencia).
Of the 16 species found during the same period on San Andres, eight
were not noted on Providencia (although sampling followed a prolonged
rainless period on that island). Of these, Vanessa cardui is a well known
wanderer, occasionally recorded as a vagrant on many Caribbean is-
lands: Panoquina sylvicola and Junonia genoveva are widely dispersed
residents in the area and beyond, as is Hylephila phyleus, found by
Emmel on both islands in 1967-68, but only on San Andres in 1986.
The remaining four butterflies ( Cymaenes odilia trebius, Eurema ve-
nusta limbia, Battus p. polydamas, Ascia m. monuste), at the subspe-
cific level noted, show either limited or no dispersive colonization of
the Antillean islands. Yet the last three, at the specific level, are amongst
the most widely distributed of the Antillean butterflies. While it is, of
course, possible that their absence from the Providencia list merely

Volume 43, Number 4

287

reflects our incomplete documentation, it seems as possible that this
reflects a real disparity between the butterfly faunas of Providencia and
San Andres.

The position of these ecologically dissimilar islands — about 87 km
apart, separated by long distances from a windward potential source
of immigrants (the Antilles), but approximately equidistant from a
much closer continental land mass — may be regarded as a “natural
experiment” from which further data pertinent to questions of general
biogeographical interest may be expected. Further field work on these
islands would be of great value, not only adding to our knowledge of
their butterflies, but in providing data which may refute, modify, or
substantiate the biogeographical hypotheses outlined above.

Acknowledgments

This work was supported by an Overseas Field Research grant from the Royal Society
(to DSS). We are grateful to Dr. Lee Miller and to Professor Sir Richard Southwood for
their critical reading of the manuscript. We are also grateful to Dr. Miller for providing
advice and help with the collections housed in the Allyn Museum of Entomology, Sarasota,
and similarly to Mr. Richard Vane-Wright and Mr. Campbell Smith for their guidance
with the collections in the British Museum (Natural History). Finally, we express our
thanks to Mrs. Deirdre Kincaid for her expert and patient help in preparing the manu-
script.

Literature Cited

Calvesbert, R. J. 1973. The climate of Mona Island. In Mona and Monito islands: An
assessment of their natural and historical resources. Estado Libre Asociado de Puerto
Rico, Office of the Governor: 2, Al-10.

Comstock, W. P. 1944. Insects of Porto Rico and the Virgin Islands: Lepidoptera
(suborder) Rhopalocera (superfamily) Papilionoidea (true butterflies) (superfamily)
Hesperioidea (skippers). Scientific Survey of Porto Rico and the Virgin Islands, Ann.
N.Y. Acad. Sci. 12:421-622.

Emmel, T. C. 1975. The butterfly faunas of San Andres and Providencia Islands in the
Western Caribbean. J. Res. Lepid. 14:49-56.

Evans, W. H. 1952. A catalogue of the American Hesperiidae indicating the classifi-
cation and nomenclature adopted in the British Museum (Natural History). Part II.
Pyrginae, Section 1. [C.13. Urbanus, pp. 85-101.] British Museum (Natural History),
London. 178 pp., plates 10-25.

1955. A catalogue of the American Hesperiidae indicating the classification and

nomenclature adopted in the British Museum (Natural History). Part IV. Hesperiinae
and Megathyminae. [J.27. Cymaenes, pp. 132-134.] British Museum (Natural His-
tory), London. 499 pp., plates 54-88.

Fox, R. M. 1963. Affinities and distribution of Antillean Ithomiidae. J. Res. Lepid. 2:
173-184.

Howe, W. H. 1975. The butterflies of North America. Doubleday & Co., New York.
633 pp.

Klots, A. B. 1960. A field guide to the butterflies of North America, east of the Great
Plains. Houghton Mifflin, Boston. 349 pp.

MacArthur, R. H. & E. O. Wilson. 1967. The theory of island biogeography. Princeton
University Press, Princeton. 203 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.

288

Journal of the Lepidopterists’ Society

Munroe, E. 1942. The Caribbean races of Anartia jatrophae Johansson (Lepidoptera:
Nymphalidae). Amer. Mus. Nov. 1179:1-4.

1953. The size of island faunas. VII Pacific Science Congr. 4:52-53 (Abstract).

1957. Comparison of closely related faunas. Science (Washington, D.C.) 126:

437-439.

1984. Biogeography and evolutionary history: Wide-scale and long-term patterns

of insects, pp. 280-304. In Huffaker, C. B. & R. L. Rabb (eds.), Ecological entomology.
John Wiley & Sons, New York.

Parsons, J. J. 1956. San Andres and Providencia, English speaking islands in the Western
Caribbean. Univ. Calif. Pubis, in Geography 12:1-84.

PiNCHON, R & P. Enrico. 1969. Faune des Antilles Franyais. Les Papillons. 1-258.
Fort-de-France.

Riley, N. D. 1975. A field guide to the butterflies of the West Indies. Collins, London.
224 pp.

Scott, J. A. 1972. Biogeography of Antillean butterflies. Biotropica 4:32-45.

Smith, D. S., S. J. Ramos, F. McKenzie, E. Munroe & L. D. Miller. 1988. Biogeo-
graphical affinities of the butterflies of a ‘forgotten’ island: Mona (Puerto Rico). Bull.
Allyn Mus. 121:1-35.

Steinhauser, S. 1981. A revision of the proteus group of the genus Urbanus Hiibner
Lepidoptera: Hesperiidae. Bull. Allyn Mus. 62:1-41.

Turner, T. W. & J. R. Parnell. 1985. The identification of two species of Junonia
Hiibner (Lepidoptera: Nymphalidae): J. evarete and J. genoveva in Jamaica. J. Res.
Lepid. 24:142-153.

Williamson, M. 1981. Island populations. Oxford University Press, Oxford. 286 pp.
Received for publication 27 June 1988; accepted 16 August 1989.

Journal of the Lepidopterists’ Society
43(4), 1989, 289-298

AN INEXPENSIVE PORTABLE TRAP FOR
MONITORING BUTTERFLY MIGRATION

Thomas J. Walker and Barbara Lenczewski

Department of Entomology and Nematology, University of Florida,
Gainesville, Florida 32611

ABSTRACT. Seven types of low-cost flight traps for monitoring migrating butterflies
were designed, built, and tested. The most efficient type caught ca. 7-55% of the migrants
crossing a 3- or 6-m line perpendicular to the direction of migration. Traps of polyester
mosquito netting, monofilament shrimp netting, or a combination of the two did not
differ significantly in capturing spring migrating Precis coenia. Trios of 3-m traps yielded
consistent estimates of variation in numbers of P. coenia migrating during spring. Portable
traps can be used in mark-and-release studies of migrating butterflies and for quantifying
distribution of migrants in time and space. Instructions for building the most efficient
type of portable trap are given in the Appendix.

Additional key words: Precis coenia, Phoebis sennae, phenology, Florida, trap effi-

ciency.

Migrating butterflies generally fly in a straight line within a few
meters of the ground. When they encounter an obstacle, their response
is to fly up and over rather than around (Williams 1930). This behavior
can be exploited in devising traps that intercept and capture migrants
(Walker 1978, 1985a, Gyotoku et al. 1987).

Early traps were inefficient and expensive to maintain; later ones
were elaborate, permanent structures costing ca. $500 each for materials
(Walker 1985a). Our long-term goal in the present study was to develop
an inexpensive, portable trap that would encourage and facilitate studies
of butterfly migration. For the short term, we needed traps to quantify
the phenology of butterfly migration at numerous stations on transects
along and across the Florida peninsula and to enable high school science
classes to catch migrants for marking and release.

Our study compared five variations of the most promising of two
prototype portable traps (Part I) and tested three fabric coverings of
the most promising variation (Part II).

I. Prototypes and Pilot Tests

During spring 1987, we built and tested two prototype traps that
contrasted in complexity and cost but that proved similar in efficiency.
That fall we tried five variations of the less expensive design.

Materials and Methods

All traps were of gray polyester mosquito netting supported by end
frames largely of thin-wall metal electrical conduit. Cross members and
guys were braided nylon rope. Cages that retained the trapped but-

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Journal of the Lepidopterists’ Society

Fig. 1. Standard traps. Trap in foreground is of polyester mosquito netting; the other
has the main wall and back of duct of monofilament shrimp net.

terflies were made of 6.4 mm (lA") hardware cloth held together by
pop rivets. Traps were generally 3 m long with openings 2 m tall.

During the 1987 fall migration, we tested these five variants of the
simpler prototype trap:

(1) Standard trap. This variant (Fig. 1) was most similar to the
prototype and is described in detail in the Appendix. It was designed
to catch butterflies coming from one direction.

(2) Two-way trap. This trap was two standard traps on opposite sides
of a shared main wall. The duct and cages were partitioned to segregate
butterflies entering the trap from opposite directions, thus permitting
calculation of net movement in the migratory direction (as in earlier
traps; e.g., Walker 1985a).

(3) Six-meter trap. This trap was identical to the standard trap except
it was 6 m instead of 3 m wide.

(4) Taller-opening trap. The roof of the standard trap sloped from
2.6 m to an opening 2.0 m tall. The roof of the taller-opening trap was
horizontal, producing a 2.6 m opening.

(5) Deeper trap. This trap had roof and side walls twice as deep as
in the standard design (3 m vs. 1.5 m for the roof). The opening was
still 2.0 m tall, reducing the slope of the roof to half that of standard.

In early October 1987, traps were erected randomly along an ENE-
WSW line in an 8-ha open field at the University of Florida’s Green

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291

Table 1. Relative trapping efficiencies of portable traps for Phoebis sennae : fall
migration, 1987.

Trap

Days3

Relative efficiencyb
n = 555c (501)

Standard

38

0.66d

Standard

30

0.83

Two-way

29

0.72

6-meter

31

0.95

Taller-opening

21

0.4

Deeper

11

(0.4)e

1st four combined

128

0.78

a Portable traps were in service for various periods during 1 Oct to 8 Nov.

b (Number caught flying southward per meter by portable trap)/(number caught flying southward per meter by
permanent trap).

c Total caught in north-facing portable traps. Value in ( ) is total for first four traps listed.
d Efficiencies with no ( ) are for >40 individuals caught in the portable trap.
e Efficiency based on 12 individuals caught in the portable trap.

Acres Farm (29°41'N, 32°20'W) near Gainesville. Two standard traps,
the two-way trap, and the 6-meter trap were run simultaneously for
29 days or more; the taller-opening and deeper traps were run se-
quentially (Table 1). All traps faced NNW, into the migratory stream
(Walker 1985b). Captures were recorded daily for all traps: after 1600
h or before 1000 h of the next day. For the standard traps, captives
were marked on the wings with a silver marking pen and returned to
the trap cages except on Mondays and Thursdays, when they were
permanently removed. Captives were removed daily from the remain-
ing traps. The marked captives were used to estimate the rate at which
butterflies disappeared from the cages by predation or escape. This was
an important consideration in evaluating the effects of reducing the
trap service interval to twice weekly.

Two permanent traps, Walker’s (1985a) model #3 and a minor mod-
ification of it, continuously monitored the migration across 12 m of
ENE-WSW line at a site 4.4 km ESE of the portable traps. For species
that cannot pass through 1.3 cm mesh hardware cloth, these traps catch
22 to 70% of migrants.

Results and Discussion

Numbers of fall migrants per meter caught by the various portable
traps were compared to the numbers caught per meter by the two
permanent traps (Table 1). This method of expressing results in terms
of relative trapping efficiency was used for two reasons. First, the
absolute trapping efficiencies of the model #3 trap had been estimated
through direct observations (Walker 1985a), making possible estima-
tions of the absolute efficiencies of the portable traps. Second, direct
comparisons among portable traps were complicated by small catches
and by different durations of service. The permanent traps ran contin-

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Journal of the Lepidopterists’ Society

uously and sampled more of the migratory front than any single type
of portable trap.

Phoebis sennae (L.) (Pieridae) was the only migrant caught by the
portable traps in substantial numbers. The taller-opening and deeper
traps ranked lowest in catching this species, and the 6-m trap ranked
highest (Table 1). When results from the four traps that were standard
in height of opening and depth of walls and roof were combined, the
estimate of relative efficiency became 0.78. If the rates of migration at
the portable and permanent traps are assumed to be the same, the
estimated absolute efficiency of standard portable traps in catching P.
sennae becomes 38 to 55%. (The permanent traps caught 49 to 70% of
P. sennae : Walker 1985a.)

The two-way trap caught 81 P. sennae flying south and 3 flying
north, for a southward consistency of 96%, not significantly different
from the consistency of 98% (447 south and 9 north) for the permanent
traps during the same 29-day period.

Marked butterflies disappeared from the cages of the standard traps
at an average rate of 12% per day (n = 147 butterflies and 284 butterfly-
cage-days). The rates of disappearance for the two traps were similar
(chi square: P > 0.50). Rates of disappearance between day 0 (=day
of marking) and day 1, between days 1 and 2, and between days 2 and
3 were similar (chi square: P > 0.95).

The standard trap equaled or exceeded the performance of its vari-
ants in the the fall 1987 tests — with the possible exception of the 6-m
trap, which caught 28% more P. sennae per meter than the mean of
the two standard traps.

II. Replicated Tests of Trap Fabrics

During the spring of 1988, we tested standard traps made of different
fabrics. This was prompted by two problems noted the previous fall:
polyester netting tore easily after a few months of exposure to sunlight,
and migrating butterflies seemed to detect and avoid portable traps
made of polyester more frequently than permanent traps made of 1.3
cm mesh hardware cloth.

Materials and Methods

The fabrics tested were polyester mosquito netting; shrimp net (9.5
mm square mesh made of knotted 0.28 mm diameter nylon monofila-
ment); and a combination of polyester netting and shrimp net. The
polyester traps were from the previous fall; the shrimp net traps were
made by stretching the monofilament netting on a frame and gluing
edges together with silicon caulking; the combination traps were made

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293

Table 2. Number of Precis coenia captured by standard portable traps of three fabric
types, Green Acres Farms: spring migration, 1988. a

Type

of fabric

Date

Block

Polyester

Combination

Shrimp net

Mean

6-22 April

(Blocks in
sequence: west
to east)

I

7

19

20

15

II

2

7

12

7

III

31

9

17

19

Mean

13

12

16

14

23 Apr-31 May

(Block I moved
to east end
of array)

I

18

19

13

17

II

4b

28

9

17

III

30b

25

11

19

Mean

17

24

11

17

6 Apr-31 May

Mean (n)

30.7

35.7

27.3

31.2

Mean (n/m)

9.7

11.3

8.7

9.9

a Analysis of variance for effect of fabric type showed no significance. (F value = 0.77, P = 0.46; model degrees of
freedom = 3 fabrics —1=2; error degrees of freedom = 504 trap-days — 1 = 503.)

b These numbers are for the traps, which were exchanged in position 11 May. The numbers for the trap positions are
14 and 20.

by cutting out the main wall and the back of the duct of polyester traps
and gluing shrimp net in their places.

Three traps of each material were erected along a 67-m ENE-WNW
line in the field at Green Acres Farm. The traps were assigned to three
randomized blocks with the constraint that the first trap in one block
could not be of the same material as the last trap in the previous block.
Traps were separated by 5 m, and the mouth of each trap was made
3.15 m wide.

Traps were serviced daily starting 6 April 1988. After 17 days, traps
in Block II had caught fewer than half the numbers of migrants caught
in the outside blocks, and the eastmost trap (Polyester III) had caught
>1X/2X as many migrants as any other trap. On 23 April, to test for
possible effects of relative or absolute trap position, we removed the
three traps of Block I from the west end of the array and erected them
east of Block III, thereby making Block II an end block and Block III
the middle block. By 10 May, Polyester II and Polyester III, which had
caught 2 and 31 butterflies, respectively, before Block I was moved,
had caught 3 and 19 more, indicating that relative position of blocks
had not caused the discrepancy. To distinguish between an individual
trap effect and an absolute position effect we exchanged the two traps.

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Journal of the Lepidopterists’ Society

% of Total Number
Northward

Fig. 2. Seasonal course of migration of Precis coenia, spring 1988, as recorded by
two permanent traps (black bars) and by three trios of portable traps (stippled bars,
representing traps of Blocks I, II, and III).

We concluded the test 31 May, when spring migration had ended. As
before, two permanent traps were operated continuously.

Results

As expected from previous studies (e.g., Walker 1985a), Precis coenia
(Hiibner) (Nymphalidae) was the only spring migrant numerous enough
to yield data useful in comparing trapping efficiency. Analysis of vari-
ance revealed no significant differences among fabric types (Table 2).

During 11-31 May, the Polyester II trap (moved to Block III) caught
1 migrant; Polyester III (moved to Block II) caught 11. These numbers
are similar in ratio to the total catches of 5 and 50 prior to swapping,
indicating that the trap rather than the position caused the variation.

The mean number of P. coenia caught per portable trap 6 April to
31 May was equivalent to 9.9 individuals flying south across each mon-
itored meter (mean no. caught per trap/meters per trap = 31.2/3.15).
During this time, the permanent traps caught 381 P. coenia or 31.8
per monitored meter. If the migration rate was the same at the two
sites, the portable traps were 31% as efficient as the permanent traps.
For the three most numerous migrant species, the permanent traps are

Volume 43, Number 4

295

22 to 70% efficient (Walker 1985a). Applying these figures to the present
data, the estimated absolute efficiency of the portable traps in catching
P. coenia is 7 to 22%.

The three trios of portable traps and the two permanent traps pro-
duced similar estimates of weekly changes in numbers of migrating P.
coenia. A chi square test of the weekly total catches for the five esti-
mators (Fig. 2) did not refute the hypothesis that all had come from
the same distribution (chi square = 19.2, df = 28, P > 0.75).

Discussion

Shrimp net did not result in a significant increase in the standard
trap’s ability to catch P. coenia, but it lasted longer outdoors and had
less wind resistance.

An unexpected result of the tests was that supposedly identical traps
of polyester differed greatly in numbers of migrating P. coenia caught:
6, 25, 61. The totals for the other traps were much more uniform —
viz., 34, 35, 38 for combination traps and 21, 28, 33 for shrimp net
traps. Inspection of the poorly performing polyester trap revealed that
its throat was narrowed at one end and that its catching cages fitted
loosely into the holding trays. Loose fitting catching cages were evident
on other traps, and on three occasions we watched a P. coenia escape
through a crack rather than enter a catching cage. Polyester III (n =
61) had a uniform throat and snug fitting cages.

The efficiency of the traps for smaller migrants (such as P. coenia )
could probably be increased and the variance in catches reduced by
redesigning the trays and building the cages to closer tolerances. A
substantial improvement might also be obtained by experimenting with
the width and uniformity of the throat. Traps of greater width, such
as the 6-meter trap, offer a cost-effective means of increasing the catch.

Unless the variance in catch among traps is reduced, groups of about
5 traps are needed to achieve a 95% chance of detecting a twofold
difference in numbers of migrants at two sites (Snedecor & Cochran
1980: 102; formula adjusted for variance being an estimate).

Standard portable traps have been used to capture fall migrants for
marking and release at four high schools in southern Georgia (J. J.
Whitesell pers. comm.) and to quantify fall migration at five stations
along a north-south transect from Valdosta, Georgia, to Lake Placid,
Florida (B. Lenczewski pers. comm.).

Acknowledgments

We thank J. J. Whitesell for help with construction of the combination and monofilament
traps and R. C. Littell and G. S. Wheeler for help with statistical analysis. The suggestions
of S. Malcolm, J. J. Whitesell, and Oakley Shields substantially improved the manuscript.
Florida Agric. Exp. Sta. Journal Series No. 9481.

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Journal of the Lepidopterists’ Society

3A FRAME 3R NETTING

tray

3C COLLECTOR

Fig. 3. Construction details of standard trap. (Dimensions are in inches: 1" = 2.54 cm.)

Appendix: Building a Trap

The standard portable trap (Fig. 1) consists of a frame, netting, and collectors that are
built separately and then assembled (Fig. 3). Total cost of materials (1988) is ca. $68 per
trap. (Most measurements are in inches: 1" = 2.54 cm.)

Volume 43, Number 4

297

Frame (ca. $23 for materials). Each end of the trap is supported by a 120" main post
(1" thin-walled metal electrical conduit = EMT), a 2 m eaves post ( V2 " EMT), and a 60"
rafter ( % " EMT) (Fig. 3 A). Attach 18" crosspiece of 1 x 1 x Vs" angle iron to top of each
main post with 5/16 x 2" bolt. Strengthen crosspiece with diagonal braces (Fig. 3A) or with
a welded 8" length of 1 x Vs" flat iron (Fig. 1). Secure bottom of braces to main post
with y4 x W2" eyebolt. At each end of each cross piece attach Va x 1" eyebolt. Attach
rafters to main posts at 102" with %6 x 3" bolts and to top of eaves posts with Va x 3"
eyebolts. Make eight guys (4 @ 120" and 4 @ 144") of Vs" braided nylon rope. Make
eight 15" guy stakes of V2" EMT; to keep rope from slipping off, put a Va" bolt through
each stake at 2" from top. Optional: Prepare four “feet” for the posts by cutting short
pieces of %" EMT that can be driven into the ground or welded to %2 x 2 x V2" pieces
of flat iron. The posts fit over or into the %" EMT.

Netting (ca. $33). (We used 653" of 64" wide “silvergray polyester mosquito netting”.)
The main wall, duct (except ends), and roof are two 222.5" lengths of fabric joined
lengthwise by sewing with nylon thread. At the indicated intervals (Fig. 3B), sew the
fabric around five 130" lengths of Vs" braided nylon rope. Cut two pieces for side walls
from 185" of netting (81" short side, 104" tall side) (Fig. 3B). In each, sew 1.5"-wide front
hem (for eaves post), 2.5" rear hem (for main post), and 2" top hem (for rafter). Cut two
20 x 20 x 18" triangles. Sew top hems of side walls to roof, triangles to ends of duct,
and wide hems of side walls to main wall. (The trap can be made wider by using more
than two 222.5" lengths of netting and longer ropes.)

Collectors (ca. $12). Each collector consists of a 10 x 11 x 16" cage resting in a 2 x
12 x 17" tray (Fig. 3C). To make the tray, join two pieces of W'-mesh hardware cloth,
cut and bent as shown (Fig. 3C: a + b), to leave a 4 x 17" opening in the bottom with
flaps of hardware cloth projecting downward for attachment of the netting. Make tray
anchors (c) of V16 x V2 x 3" aluminum pieces and rivet to each end of tray at center. Drill
Va" holes in tray anchors for attachment to frame. To make the cage, fold a 16 x 43"
piece of W'-mesh hardware cloth as shown (d) and use pop rivets with backup washers
to attach ends (e) made of two 11 x 12" pieces. If desired, cut 4 x 5" access in one end
and fit piece of hardware cloth to serve as closure. (Cages can be emptied through the
bottoms.) Longitudinally folded 1" strips of sheet metal may be crimped over principal
edges of the cages and trays for strength and safety. All other rough edges should be
taped with 1" pressure sensitive tape. Bolt to each cage 1" PVC elbow (f) with downward
projecting 30" shaft of 1" PVC pipe (g) to permit removing and replacing the cage from
the ground (Fig. 1). Make tool for servicing cages from 36" of 1" PVC pipe with female
fitting on one end, from which projects 4" of 1 x 8" dowel (not figured).

Erecting the trap. (Two persons recommended.) Determine the orientation and position
of the trap; place feet or markers at the four corners. Set stakes for guys in approximate
positions. Insert main posts, rafters, and eaves posts into appropriate hems. Attach rafters
to posts. Tie guy ropes to appropriate eye bolts. Tie suspension ropes to 1" eye bolts. Erect
one end of trap and temporarily guy; then erect other end. Tie ends of front and back
throat ropes around each main post. Tie ends of eaves rope to 2" eyebolts. Adjust ropes,
guys, and stakes until trap is trim. Make three Va x % x 2" wooden spacers with V16" holes
at each end. Sew across the throat at even intervals.

Once trap is up, install collectors from a tall step ladder. Cut fabric in top of duct and
insert flaps that surround slot in tray bottom. Insert shank ends of 1" eye bolts (Fig. 3 A)
into tray anchors and secure tray to main post crosspiece with two Va" nuts. Sew fabric
to flaps. Position cage in tray, making sure it fits snugly but is loose enough to service
from ground.

Literature Cited

Gyotoku, M., K. Ito & F. Nakasuji. 1987. Monitoring of lepidopterous insect migration
using the migration trap. Jpn. J. Appl. Entomol. Zool. 31:350-358.

Snedecor, G. W. & W. G. Cochran. 1980. Statistical methods. 7th ed. Iowa State
Univ. Press, Ames. 507 pp.

298

Journal of the Lepidopterists’ Society

Walker, T. J. 1978. Migration and re-migration of butterflies through north peninsular
Florida: quantification with Malaise Traps. J. Lepid. Soc. 32:178-190.

1985a. Permanent traps for monitoring butterfly migration: tests in Florida,

1979-84. J. Lepid. Soc. 39:313-320.

1985b. Butterfly migration in the boundary layer, pp. 704-723. In Migrations:

Mechanisms and adaptive significance. Contrib. Mar. Sci. 27 (Suppl.). University of
Texas at Austin, Port Aransas, Texas.

Williams, C. B. 1930. The migration of butterflies. Oliver and Boyd, Edinburgh.
473 pp.

Received for publication 22 November 1988 ; accepted 30 May 1989.

Journal of the Lepidopterists ’ Society
43(4), 1989, 299-304

BIOLOGY AND IMMATURE STAGES OF
POLIA LATEX (GUENEE) (NOCTUIDAE)

Peter S. Wood

Department of Extension and Continuing Education,

Gypsy Moth Office, Fairfax, Virginia 22030

AND

Linda Butler

Division of Plant and Soil Sciences, P.O. Box 6108,

West Virginia University, Morgantown, West Virginia 26506-6108

ABSTRACT. Polia latex was studied from 1984-86 at Cooper’s Rock State Forest in
northern West Virginia as well as under laboratory conditions. Adult flight period was
from 24 May to 29 August and larvae were collected from 21 June to 2 October. Thirteen
host plants and 14 species of parasites were recorded for P. latex larvae. The egg and
seven larval instars are described. Larval duration averaged 45.3 days at 24°C.

Additional key words: hardwood defoliator, West Virginia, parasites, Hadeninae.

Polia latex (Guenee) is a common hadenine (Noctuidae, Hadeninae),
the adults of which are well known and easily recognized. The adults
fly from May to August with melanics being common (Covell 1984).
The immature stages are less well known.

Larvae are found from early June to mid-August with a peak in early
August (Prentice 1962). Brief descriptions of larvae were given by
Crumb (1956) and Godfrey (1972). Godfrey figured the head, hypo-
pharyngeal complex, and mandible and illustrated habitus of the mature
larva.

According to Crumb (1956), the distribution of P. latex is restricted
to the northeast United States and adjoining Canada. Smith (1893)
recorded it from the northern, eastern, and middle states.

During 1984-86 P. latex was studied at Cooper’s Rock State Forest
in northern West Virginia, just west of the leading edge of the infestation
of the gypsy moth, Lymantria dispar (L.) (Lymantriidae). The objective
of the study was to obtain baseline data for native lepidopterous de-
foliators and their parasites before the build-up of gypsy moth to enable
evaluation of changes in native defoliator populations later as the gypsy
moth population increases. During this study, Polia latex was one of
the more abundant defoliators, but was producing no significant de-
foliation. Since little information is available on the biology of P. latex,
we recorded duration of larval development, host plants, and parasites
of this species. Voucher specimens are in the West Virginia University
Arthropod Collection.

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Journal of the Lepidopterists’ Society

Materials and Methods

The West Virginia University Forest at Cooper’s Rock State Forest
is located in Preston and Monongalia counties about 32 km east of
Morgantown, West Virginia. The area consists of 50- to 60-year-old
even-aged mixed mesophytic forest and has a mean elevation of 561
m (Carvell 1983). The most abundant tree species in the study area are
red maple ( Acer rubrum L., Aceraceae), white and red oak ( Quercus
alba L., Q. rubra L., Fagaceae), black cherry ( Prunus serotina Ehrh.,
Rosaceae) and black birch ( Betula lenta L., Betulaceae).

The flight period of Folia latex was determined by blacklight trap-
ping throughout the 1984 and 1985 seasons. A single gravid P. latex
female was live-trapped at a blacklight trap and caged with foliage of
maple, cherry, birch and oak on which to oviposit; the adult was pro-
vided with water and black cherry blossoms as a nectar source.

Larvae were reared in 150 x 25 ml plastic petri dishes. Fresh red
maple foliage was placed in clean petri dishes every other day. Larvae
were observed daily, and at each instar some larvae were killed in
KAAD and preserved in 80% ethanol for head measurements. Pupation
was in a layer of moist vermiculite 5 cm deep in quart canning jars.
All rearing was conducted at 24°C?and 12L:12D photoperiod.

Larval descriptions were based on laboratory reared specimens only.
The terminology used is that of Godfrey (1972). Eggs and larval head
capsule measurements were made with an ocular micrometer.

Field sampling of P. latex larvae was conducted in 1984 and 1985
by pruning foliage samples once each week during the growing season.
Foliage sampled was primarily Acer rubrum, A. saccharum Marsh., A.
saccharinum L. and A. nigrum Michx. f., Prunus serotina, Betula lenta,
Quercus alba, Q. rubra, Q. prinus L. and Q. velutina Lam. Some
additional host trees were observed during field sampling. During 1986,
larvae were sampled by burlap banding black cherry, black birch, red
maple, and mixed oaks (45 trees total). All field collected larvae were
reared in the laboratory on the appropriate host plants to isolate par-
asites.

Results

Phenology, Host Plants, and Parasites

Seasonal flight period of P. latex during 1984 was from 25 May-29
August; 128 specimens were trapped. In 1985, 79 specimens were col-
lected between 24 May-13 August. Peak trap numbers were on 6 July
1984 and 12 June 1985. Despite the long adult flight period, field
collections of larvae indicate that P. latex is univoltine.

In 1984, 91 larvae were collected on foliage in the field between 21

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301

June and 13 September; in 1985, 52 larvae were collected between 3
July and 12 September. In 1986, 255 P. latex larvae were taken between
11 July and 2 October from burlap bands placed on 45 trees. Thirty-
nine percent of the larvae were on birch (7 trees), 24% on black cherry
(12 trees), 19% on red maple (13 trees), and 18% on mixed oaks
(13 trees). Polia latex made up 86% of all lepidopterous larvae found
under burlap bands during that time period.

The larvae of P. latex are general feeders on deciduous broad-leafed
trees; the 13 host species found during this study were: Acer nigrum,
A. rubrum, A. saccharinum, A. saccharum (Aceraceae); Betula lenta
(Betulaceae); Nyssa sylvatica Marsh. (Nyssaceae); Magnolia acuminata
L. (Magnoliaceae); Prunus serotina (Rosaceae); Quercus alba, Q. pri-
nus, Q. rubra, Q. velutina (Fagaceae); and Sassafras albidum Nees
(Lauraceae). Additional host plants given by other authors include:
Betula nigra L. (Crumb 1956), B. lutea Michx. f. and B. papyrifera
Marsh. (Betulaceae) (Prentice 1962); Fagus grandifolia Ehrh. (Faga-
ceae) (Covell 1984); and Ulmus americana L. (Ulmaceae) (Coveil 1984).

Larval instar durations as determined from laboratory rearing are
summarized in Table 1; the larval period averaged 45.3 days at 24°C.

Parasites reared from field-collected P. latex larvae were: Hyphan-
trophaga virilis (Aldrich & Webber) and Lespesia stonei Sabrosky
(Diptera: Tachinidae); Alloplasta suberba (Provancher), Eutanyacra
improvisa (Cresson), Hyposoter fugitivus (Say), and Mesochorus pictilis
Holmgren (hyperparasite) (Hymenoptera: Ichneumonidae); Perilam-
pus sp. (hyperparasite) (Hymenoptera: Perilampidae); Eulophus neb-
ulosus (Provancher), Euplectrus sp. and Enplectrus maculiventris
(Westwood) (Hymenoptera: Eulophidae) ; Diolcogaster facetosa (Weed),
Microplitis hyphantriae (Ashmead), Microplitis sp. and Meteorus ba-
keri C. & D. (Hymenoptera: Braconidae).

Description of Immature Stages

Eggs: In the laboratory, the caged female oviposited, producing a one-layered egg mass
(129 eggs) and a three-layered egg mass (217 eggs). The eggs were spherical in shape,
0.66 mm (n = 50) in diameter. One egg observed by scanning electron microscope at
130 x was sculptured with 37 ridges and paralleled by transverse cross-striae forming
hexagons toward the anterior pole and more quadrate shapes toward the posterior; at
650 x, 16 micropyles were observed at the anterior pole.

Larvae: Instar 1 (n = 13): Head capsule 0.36 mm (range 0.33-0.36), light tan. Body
ground color transparent to yellowish with gut contents giving a basic green color; pinacula
black and prominent. Prolegs on abdominal segments 3 and 4 reduced with the greatest
reduction on abdominal segment 3. Prothoracic shield light tan and difficult to distinguish
from general body surface; lateral shields of anal prolegs tan and prominent.

Instar 2 (n = 14): Head capsule 0.58 mm (range 0.50-0.60), tan. Body ground color
yellowish-green. Pinacula prominent.

Instar 3 (n = 14): Head capsule 0.97 mm (range 0.90-1.00), tan with black reticulations.
Body color variable (olive green, light brown, grey, yellow); dorsal (D), subdorsal (SD),

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Journal of the Lepidopterists’ Society

Figs.. 1-5. Polia latex, mature larva: 1, head capsule, frontal view; 2, head capsule,
lateral view, 3, prothorax and mesothorax, lateral view; 4, abdominal segment 2, lateral
view; 5, abdominal segments 6-10, lateral view. Scale line = 1 mm for Figs. 1-2, 2 mm
for Figs. 3-5.

and lateral (L) lines white with the D line being most prominent; the body region between
the D and SD lines much darker than the body ground color.

Instar 4 (n = 16): Head capsule 1.49 mm (range 1.29-1.72), tan. A black pinaculum
around the D1 seta on each segment; abdominal segments 8 and 9 olive green to black,
the white D line ending on the anterior half of the 8th abdominal segment.

Instar 5 (n = 15): Head capsule 2.24 mm (range 2.00-2.57), with extensive reticulations

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303

Table 1. Larval duration of P. latex reared

on leaves of red maple at 24°C (n = 14).

Mean time ± SD

Instar

(days)

1

5.6 ± 0.5

2

4.2 ± 0.6

3

4.5 ± 1.3

4

5.9 ± 1.8

5

5.9 ± 1.5

6

7.0 ± 3.6

7

10.5 ± 3.8

Prepupal

3.8 ± 1.0

Total

47.4 ± 4.4

and submedial arcs. Body mostly dark green; lateral band creamy yellowish-white and
surrounding the lower half of black abdominal spiracles; venter pale green. White D and
SD lines still present; area between D and SD lines mottled green and white; area between
white SD and L lines dark olive, slightly lighter olive green just below SD line; dorsal
dark wedge-shaped markings on abdominal segments and black dorsal bars evident on
the 8th abdominal segment.

Instar 6 (n = 14): Head capsule 3.01 mm (range 2.86-3.29); reticulation and submedial
arcs similar to those of instar 5. Body dark with D, SD, and L lines mottled white and
green; the L line most distinct. Black dorsal bars more evident than in previous instar.

Instar 7 (n = 13): Head capsule 3.65 mm (range 3.29-4.15), dark brown with reticula-
tions covering the entire head. Lateral line consisting of a brown supraspiracular line and
a pinkish-peach subspiracular line; prothoracic and abdominal 8 spiracle set completely
within supraspiracular line; abdominal spiracles 1-4 and 7 located between the supra-
spiracular and subspiracular lines and spiracles 5 and 6 completely within subspiracular
line.

GENERAL: Body of mature larva 33 mm long and 4 mm wide; prolegs present on
abdominal segments 3-6, size increasing caudad; crochets uniordinal, 25-28 per third
abdominal proleg, 28 per fourth, 29-31 per fifth, 29-32 per sixth. All setae simple.
Chaetotaxy illustrated in Figs. 1-5.

DIAGNOSIS: Polia latex larvae may be distinguished from larvae of other Polia spp.
common in the study area on the basis of color pattern (Crumb 1956, Godfrey 1972).
Polia purpurissata (Grote) is larger and also differs in being yellow grey or violet grey
with obscure markings. The cervical shield, which is shiny brown in P. purpurissata,
lacks contrast in P. latex. Polia imbrifera (Guenee) is similar in size but is greyish with
distinct black oblique lines from D2 to SD1 on abdominal segments 1-8; these lines are
lacking in P. latex. Polia nimbosa (Guenee) also possesses distinct black oblique lines on
abdominal segments as well as a prominent segmental series of black, diamond-shaped
middorsal spots. Polia detracta (Walker) is smaller with head reticulation darker than
that of P. latex. The body of P. detracta is more reddish and has darker cervical and
anal plates than P. latex.

Acknowledgments

We thank the U.S. Forest Service for partial support of this research; also Vicki Kondo,
Lisa Bean, Ralph Iannone, Janice Jenkins, Robert Progar, Delray Shaffer, Judy Shaffer
and Laura Tate for field and laboratory assistance; James W. Amrine, Jr., John E. Hall
and Joseph E. Weaver for review of the manuscript and R. W. Carlson, E. E. Grissell,
P. M. Marsh, M. E. Schauff and N. E. Woodley of the U.S. Dept. Agr. Systematic
Entomology Laboratory, for parasite identifications.

Published with the approval of the Director of the West Virginia Agricultural and
Forestry Experiment Station as Scientific Article No. 2126. This research was supported
with funds appropriated under the Hatch Act and by a grant from the U.S. Forest Service.

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Journal of the Lepidopterists’ Society

Literature Cited

Carvell, K. L. 1983. A summary of 1973-1982 weather data from the West Virginia
University Forest. WVU Forestry Notes 10:13-16.

Covell, C. V. Jr. 1984. A field guide to the moths of eastern North America. The
Peterson Field Guide Series. Houghton Mifflin, Boston. 496 pp.

Crumr, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agr. Tech. Bull. 1135.
356 pp.

Godfrey, 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.

Prentice, R. M. 1962. Forest Lepidoptera of Canada. Vol. 2. Dept, of Forestry Bull.
128. pp. 77-281.

Smith, J. B. 1893. Catalogue of Noctuidae. U.S. Natl. Mus. Bull. 44. 424 pp.

Received for publication 10 August 1988; accepted 26 June 1989.

Journal of the Lepidopterists’ Society
43(4), 1989, 305-312

FOOD PLANT SPECIFICITY AND BIOLOGY OF
ITAME VARADARIA (WALKER) (GEOMETRIDAE),

A NORTH AMERICAN MOTH INTRODUCED INTO
AUSTRALIA TO CONTROL THE WEED
RACCHARIS HALIMIFOLIA L.

W. A. Palmer

North American Field Station, Queensland Department of Lands,

2801 Arrowhead Circle, Temple, Texas 76502

ABSTRACT. Itame varadaria (Walker) is an infrequently encountered ectophagous,
foliage feeding geometrid that occurs from South Carolina to Texas on Baccharis halimi-
folia L. Laboratory studies indicated that the moth completes its development in about
28 days, that there are five larval instars, and that females have an average potential
fecundity of 165 eggs. In Texas there are at least three generations per year and there
was no evidence of diapause. The moth was reared through a number of generations in
the laboratory using cut foliage as a food source. Food plant specificity studies were
conducted both in Texas and Australia. Moths were not specific in their oviposition
preference under cage conditions but larvae developed only on four species of Baccharis.
Itame varadaria is therefore sufficiently stenophagous for use as a biocontrol agent and
approval was granted for its release in Australia.

Additional key words: Texas, Queensland, Asteraceae, fecundity.

The woody shrub Baccharis halimifolia L. (Asteraceae: Astereae:
Baccharinae), introduced from North America, is a serious weed in
Queensland, Australia (Stanley & Ross 1986, Palmer 1987). The Queens-
land Department of Lands, through the Alan Fletcher Research Station
(AFRS), therefore initiated a long-range research program in 1962 to
find biological control agents in the New World for release in Australia.
This ultimately led to the establishment of the North American Field
Station (NAFS) at Temple, Texas, in 1982.

The insect fauna on B. halimifolia has been described by Palmer
(1987) and Palmer & Bennett (1988). A rather rare geometrid, Itame
varadaria (Walker) was listed in both these studies and was identified
as a possibly monophagous insect that should be investigated further
as a biological control agent.

This paper describes the experimental and field observations under-
taken to investigate the biology, phenology, and food plant specificity
of this moth prior to its release in Australia.

Life History

Eggs were green with a slightly flattened ovoid shape and sculptured
exochorion. Mean length and width of the eggs were 0.74 mm (SE =
±0.02; n = 6) and 0.40 mm (±0.02; n = 6) respectively. They were
attached to the leaf margin along the longitudinal axis of the egg and

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Journal of the Lepidopterists’ Society

Table 1. Life history of I. varadaria when reared at 28°C.

Stage

Head capsule width
mm (±SE; No. obs.)

Larval length range in
mm

Development
time days

Feeding damage

Egg

—

—

4

—

1st Instar

0.25 (0.01; 11)

2.2-3.3

3

pinholes

2nd Instar

0.44 (0.01; 12)

3. 0-6.0

3

pinholes

3rd Instar

0.71 (0.03; 5)

7. 0-8.5

1

from margin

4th Instar

1.1 (0.01; 24)

6.0-12.0

3

from margin

5th Instar

1.76 (0.04; 22)

10.0-21.0

6

whole leaf

Pupa

—

—

8

—

were usually oviposited singly or in groups of two or three, rarely in
larger clusters.

Larvae developed through five instars before pupation. A cohort of
larvae was kept at 28°C in the laboratory and observed daily. First and
second instars chewed tiny holes 0. 5-1.0 mm in diameter through the
epidermis to feed on underlying parenchyma tissue. Later instars fed
from the leaf margin inwards. Fifth instars consumed about a leaf a
day. Measurements of head capsule width and larval length were made
daily on a selection of 4-6 larvae from the cohort and these are given
in Table 1 together with the time spent in each development stage.
The insect completed its life cycle in approximately 28 days.

The mean head capsule widths for the five instars were consistent
with the “Brooks-Dyar” rule (cf. Daly 1985). Relatively constant growth
ratios of 0.57, 0.62, 0.65 and 0.63 can be calculated to describe the
width of head capsules as a proportion of the width of the following
instar.

Pupae were very difficult to sex and it was found more reliable to
wait for the emergence of the adult to determine sex. Mean weights
were 40 mg (±2; n = 6) and 55 mg (±2; n = 10), for male and female
pupae respectively.

Four single pair matings were made by confining pairs of moths in
10 oz. paper cups with moistened raisins. After the death of the female,
the eggs in the container were counted, the female dissected and eggs
remaining in the abdomen counted. The two counts were added to give
an estimate of the total potential fecundity. Mean total potential fe-
cundity was 165 eggs (±20; n = 4). Some 31% of the total egg count
remained in the insects’ abdomens.

Phenology and Range

7. varadaria occurs from South Carolina (Ferguson 1973) to Florida
(Palmer & Bennett 1988) and along the Gulf Coast to Houston, Texas
(Palmer 1987). Its only known larval food plant is B. halimifolia. Fer-

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307

guson (1973) noted at least two broods a year with the moths of the
spring generation being slightly larger, darker, and less brightly marked
than the summer ones.

An area to the west of Beaumont, Texas, was periodically surveyed
throughout 1986 and 1987 by sweeping the bushes with a heavy duty
net to recover larvae. Larvae were recovered as early as March and
were also seen as late as early December. It therefore appeared that
there could be up to 6 or 8 generations a year and certainly at least 3
generations per year.

Larvae were never very abundant and numbers showed little vari-
ation with season; an hour’s sweeping rarely resulted in the capture of
more than half a dozen larvae. At such low densities the insect was not
adversely affecting the plant.

Because only a few larvae were captured and these by sweeping, no
natural enemies were observed nor parasites reared from the immatures.

Rearing

A colony was maintained in a laboratory kept at 25-28°C and 40-
60% RH for more than four generations with the following procedure.
Batches of 10-15 unsexed pupae were placed in 10 oz. paper cups.
Emerging moths were offered soaked raisins (or a sugar-water wick)
and a piece of fine polyester netting (approx. 20-30 cm2). Moths suc-
cessfully mated in the cups and then laid most eggs on the netting but
some on the cup itself.

Neonate larvae were transferred to bouquets of foliage held in 1 oz.
souffle cups. The foliage of both B. halimifolia and B. neglecta Britton
(a species growing near the NAFS) were equally acceptable. The larvae
were transferred to fresh bouquets of foliage twice a week. On com-
pleting their growth, fifth instars usually wandered from the foliage
before becoming prepupae. Naked pupae were collected from the cage
floor and sometimes from the foliage.

An alternative method of rearing was also successful. Moths were
released into large cages containing potted plants held in an aircon-
ditioned greenhouse. Providing care was taken to replace plants if they
were defoliated, this system required little work and pupae or adults
could be recovered from the cage regularly.

Food Plant Specificity Testing

Oviposition preference. In an experiment with two replicates, mesh
cages (62 x 45 x 45 cm) were set up, each containing ten asteraceous
species in gallon pots. Three sugar-water wicks were placed in each
cage before the release of twelve unsexed moths from the laboratory

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Journal of the Lepidopterists’ Society

Table 2. Mean number of eggs oviposited by moths on various species of plants in
a multiple choice experiment replicated twice.

Plant species

Eggs (±SE)

Baccharis neglecta Britton

20.0 (±8)

Ageratum houstonianum Mill.

0.0

Aster novae-angliae L.

3.0 (±1)

Beilis perennis L.

1.0 (±1)

Callistephus chinensis (L.)

0.0

Chrysanthemum morifolium Rem.

0.0

Conyza canadensis L.

0.0

Dahlia pinnata Cav.

0.0

Parthenium hysterophorus L.

11.0 (±10)

Solidago altissima L.

0.5 (±0.5)

Cage walls

many (>10)

colony. Four and seven days later, all plants and the cages were very
carefully examined using a lighted magnifier and any eggs counted.

Although most eggs were laid on B. neglecta, some were also found
on other species and on the wire netting of the cages (Table 2). Under
these laboratory conditions the insect was not specific in its oviposition
behavior. This is not an uncommon finding when Lepidoptera are held
in close confinement.

Survival of neonate larvae on cut foliage. These tests, which con-
stituted the most comprehensive component of the food plant testing,
were conducted at both the NAFS in Texas and at the AFRS in Australia
because many of the plant species were available on only one continent.
However, the same procedure was used at both locations. The insect
was tested against 6 species of Baccharis, 13 genera in the Astereae
tribe, 26 genera in 10 other tribes of the Asteraceae and a further 24
genera in 18 other plant families. In all, a total of 72 plant species
(Table 3) was tested. Not all plants could be tested simultaneously so
they were tested in batches of about 20 plants. B. halimifolia was always
included as a control with each batch. The plants were randomly se-
lected for each batch but eventually each species was tested at least
three times by the following method. Except for a few cases, foliage
was taken from different potted plants for each replication.

Five unfed, neonate, laboratory reared larvae were placed in a glass
vial with a leaf of one plant species. The vials were observed daily and
wilted leaves replaced. Survival was assessed after 72 h and again after
144 h.

Larvae survived for 144 h only on 4 species of Baccharis. Mean
survival on B. halimifolia, B. neglecta, B. pilularis D.C., and B. saro-
throides Gray were 70%, 80%, 100% and 50%, respectively, and sig-
nificant feeding was observed in each case. Almost all larvae on plants

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309

Table 3. The plant list against which neonate larvae were tested in a cut foliage, no-
choice experiment. Species selected for additional experiments are also indicated (*f).

Anacardiaceae: Mangifera indica L.

Apiaceae: Apium graveolens L.

Asteraceae: Tribe Astereae: Aster novae-angliae L.*f, A. novi-belgii L., Baccharis bige-
lovii Gray, B. glutinosa Pers., B. halimifolia L.*t, B. neglecta Brittonf, B. pilularis

D. C., B. sarothroides Gray, Beilis perennis L.f, Brachyscome multi fida D.C.*f, Cal-
listephus chinensis (L.) Nees*f, Calotis cuneata (F. Muell. ex Benth.) G. L. Davis*t,
Chrysothamnus nauseosus (Pall.) Britt., Conyza canadensis L.f, C. sumatrensis (Retz.)

E. H. Walker*, Gutierrezia microcephala (D.C.) Gray, Isocoma wrightii (Gray) Rydb.,
Olearia subspicata (Hook.) Benth.*f, Solidago altissima L.f, Vittadinia sulcata N. T.
Burbidge*f. Tribe Heliantheae: Cosmos bipinnatus Cav.*f, Dahlia pinnata Cav., D.
variabilis Desf.f, Eclipta prostrata (L.) L.*f, Gaillardia aristata Purshf, Glossogyne
tenuifolium (Labill.) Cass.f, Helianthus annuus L.f, Iva frutescens L., Wedelia spi-
lanthoides F. Muell. f, Zinnia angustifolia H.B.K.*f. Tribe Inuleae: Cassinia laevis R.
Br.*t, Gnaphalium sphaericum Willd.f, Helichrysum bracteatum (Vent.) Andr*f. Tribe
Senecioneae: Emilia sonchifolia (L.) D.C.t, Flaveria australasica Hookf, Senecio lau-
tus G. Foster ex Willdf. Tribe Anthemideae: Artemsia frigida Willd., Chrysanthemum
carinatum Schousb.f, C. morifolium Remat, Cotula australis (Spreng.) Hook. F.*f.
Tribe Eupatorieae: Adenostemma lavenia (L.) Kuntzef. Tribe Vernonieae: Vernonia
cinerea (L.) Less*f. Tribe Lactuceae: Cichorium intybus L., Lactuca sativa L. Tribe
Cynareae: Carthamus tinctorius L.f, Cynara scolymus L.f. Tribe Calenduleae: Cal-
endula officinalis L.f- Tribe Tageteae: Tagetes lucida Cav.

Brassicaceae: Brassica oleracea L.

Caricaceae: Carica papaya L.

Chenopodiaceae: Beta vulgaris L.

Cucurbitaceae: Cucurbita maxima Duch.*

Fabaceae: Arachis hypogaea L.*, Medicago sativa L., Phaseolus vulgaris L.

Liliaceae: Allium cepa L.*

Malvaceae: Gossypium hirsutum L.*

Myrtaceae: Eucalyptus sp.

Passifloraceae: Passiflora edulis Sims.

Poaceae: Triticum aestivum L., Saccharum officinarum L., Sorghum vulgare L., Zea
mays L.

Proteaceae: Macadamia integrifolia Maid & Betche.

Rosaceae: Fragaria vesca L.

Rubiaceae: Coffea arabica L.

Rutaceae: Citrus sinensis (L.)

Solanaceae: Solanum tuberosum L., Lycopersicon lycopersicum L.

Vitaceae: Vitis vinifera L.

* Potted plants tested against nenonate larvae,
t Cut foliage tested against third instar larvae.

other than these four species were dead after 72 h and little evidence
of feeding was observed. The exceptions were two larvae on Cassinia
laevis R. Br. where a little feeding was seen, and one larva each on
Calotis cuneata (F. Muell. ex Benth.) and Olearia subspicata (Hook)
Benth. where no evidence of feeding was present.

Survival of neonate larvae on potted plants. To ascertain whether
larvae responded similarly on growing plants as they did on cut foliage,
neonate larvae were exposed to twenty plant species (Table 3). Two
potted plants of each selected species were available and these were

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Journal of the Lepidopterists’ Society

placed in an air-conditioned glasshouse. Ten neonate larvae were placed
on the foliage of each plant and a fine mesh bag placed over them to
confine them to a limited portion of the plant so that they could be
more easily observed and protected from marauding ants. The plants
were examined after 4 days and surviving larvae noted.

After 4 days 100% survival and substantial feeding were noted on
both B. halimifolia plants but no live larvae were found on any other
plant. Feeding marks were noted only on one Cassinia laevis plant.

Late instar survival on cut foliage. This experiment was conducted
to determine if late instar larvae have a wider food plant range than
neonate larvae. Neonate larvae were reared on B. halimifolia foliage
for 10 days at 22°C at which time they were third instars. Five larvae
were transferred to foliage of each of 25 species of Asteraceae (Table
3), cuttings of which had been placed in small bottles of water. These
cuttings were replaced after three and six days. The experiment was
evaluated on the 7th day and live larvae noted.

Survival of larvae on B. halimifolia and B. neglecta averaged 80%
and 100% respectively and extensive feeding occurred. No feeding
occurred on any other plant. Soon after their placement on the cuttings,
most larvae on plants other than Baccharis left the foliage and were
subsequently found on the bottom of the containers; most larvae had
died by the fifth day and all were dead by the seventh.

Discussion

Itame varadaria was tested very comprehensively against a wide
range of plant species with particular emphasis being given to closely
related species in the Asteraceae. It was evident that the larval food
plant range of this insect is confined to certain species of Baccharis.
The four species on which I. varadaria could be reared (viz., B. hal-
imifolia, B. neglecta, B. pilularis, and B. sarothroides ) must be phy-
tochemically very similar as other highly stenophagous species tested
at the NAFS have had the same host range [e.g., Prochoerodes truxaliata
(Guenee) reported by Palmer and Tilden (1987)]. Itame varadaria is
considered to pose no threat to existing flora in Australia.

Of the four plant species acceptable to I. varadaria larvae in the
laboratory, only B. halimifolia grows within the geographic range of
this moth. Therefore, environmental factors unsuitable to I. varadaria
probably explain why B. neglecta, B. sarothroides, and B. pilularis are
not natural larval food plants. However, two other species, B. glomeruli-
fora Pers. and B. angustifolia Michx., which are closely related to B.
halimifolia and B. neglecta, respectively, do occur within the insect’s
range and it would not be surprising if they are someday reported as
food plants.

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311

There was good agreement between the three feeding tests. Cut
foliage experiments are the simplest in design and have the advantage
that the insects are kept under maximum supervision. However the
question is sometimes raised that the insects may respond differently
on growing plants than on cut foliage. Similarly, while the insect can
not become established on a plant species if neonate larvae cannot feed
on it, there is sometimes concern, particularly with mobile species, that
late instar larvae might move onto non-target plants. In this case there
was no evidence that late instars had a broader food plant range. The
technique of using neonate larvae exposed to cut foliage accurately
determined the food plant range for this insect.

Although I. varadaria is not abundant in its native habitat, it has a
number of useful characteristics that might make it an effective agent.
It has a relatively short life cycle, is quite fecund, is easily reared in
the laboratory, occurs in areas with similar climates to southeastern
Queensland, and apparently has no diapause mechanism. Its success
will probably depend on the predation rate in its new environment.

Release in Australia

Permission to release I. varadaria in Queensland was granted in
December 1988 by the Australian Quarantine and Inspection Service
and the Australian National Parks and Wildlife Service, which admin-
ister the Quarantine Act of 1908 and the Wildlife Protection Act of
1982, respectively. The insect was then moved out of quarantine fa-
cilities at the AFRS and a mass rearing program commenced.

It is anticipated that the first field releases will be made in spring of
1989. Moths will be released at a number of sites in southeast Queensland
over the following twelve months. In some situations the moths will be
released into cages covering B. halimifolia bushes to give some measure
of protection against predators to the establishing colony.

Acknowledgments

I thank Dr. D. C. Ferguson, Systematic Entomology Laboratory, Agr. Res. Sta., U.S.
Dep. Agr., for expert identification and very helpful advice.

Mrs. J. Palmer and Mr. G. Paris conducted the laboratory trials at the NAFS and AFRS
respectively. This study was funded by the Rural Lands Protection Board, Queensland
Department of Lands.

Literature Cited

Daly, H. V. 1985. Insect morphometries. Ann. Rev. Entomol. 30:415-438.

Ferguson, D. C. 1973. The identity of Macaria inapta Walker and Itame varadaria
(Walker) (Geometridae). J. Lepid. Soc. 27:288-290.

Palmer, W. A. 1987. The phytophagous insect fauna associated with Baccharis hal-
imifolia L. and B. neglecta Britton in Texas, Louisiana and northern Mexico. Proc.
Entomol. Soc. Wash. 89:185-199.

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Journal of the Lepidopterists’ Society

Palmer, W. A. & F. D. Bennett. 1988. The phytophagous insect fauna associated
with Baccharis halimifolia L. in the eastern United States. Proc. Entomol. Soc. Wash.
90:216-228.

& J. W. Tilden. 1987. Host specificity and biology of Prochoerodes truxaliata

(Guenee) (Geometridae), a potential biocontrol agent for the rangeland weed Bac-
charis halimifolia L. in Australia. J. Lepid. Soc. 41:199-208.

Stanley, T. D. & E. M. Ross. 1986. Flora of southeastern Queensland. Vol. 2. Queens-
land Department of Primary Industries, Brisbane. Misc. Publ. QM84007.
623 pp.

Received for publication 2 March 1989; accepted 14 August 1989.

Journal of the Lepidopterists’ Society
43(4), 1989, 313-322

GENERIC REASSIGNMENTS FOR NEOTROPICAL
TORTRICID MOTHS (TORTRICIDAE)

John W. Brown

421 N. Kenneth Road, Burbank, California 91501

ABSTRACT. Based on current studies of neotropical tortricine moths, I propose
generic reassignments for 20 species, comprising 18 Euliini (Tortricinae), 1 Chlidanotini
(Chlidanotinae), and 1 Eucosmini (Olethreutinae), and identify characters that support
the new combinations.

Additional key words: Tortricinae, Euliini, Chlidanotini, Eucosmini, hairpencil.

Although nearly all species of Holarctic Tortricinae can be placed
confidently within our current generic and tribal framework (Powell
1983, Razowski 1987a), a comparable system for the neotropical fauna
has not yet been realized. The most significant contribution toward this
goal is Powell’s (1986) synopsis of the classification of the Neotropical
Tortricinae, in which all described genera are assigned to tribal cate-
gories compatable with those of the Nearctic and Palearctic faunas.
However, nearly 30% of neotropical species lack meaningful generic
assignments, and a large portion of the fauna is still undescribed. Through
the efforts of Razowski (1982, 1986a, 1987b, 1987c, 1988) and Powell
(1986), a generic framework is emerging, providing categories to which
previously “unplaced” species can be assigned. The purpose of this
paper is to transfer 20 neotropical species described prior to 1930 in
broadly defined, polyphyletic genera, such as Eulia Hiibner and Tortrix
Linnaeus, into well-defined, monophyletic genera. This will provide
appropriate generic combinations, and rationalizations for such, for a
forthcoming checklist of the Neotropical Tortricinae (Powell & Razow-
ski 1989).

Depositories abbreviated in the text are as follows: BMNH, British
Museum (Natural History), London, England; CMNH, Carnegie Mu-
seum of Natural History, Pittsburgh, Pennsylvania; NHMV, Naturhis-
torisches Museum Vienna, Austria; UCB, Essig Museum of Entomology,
University of California, Berkeley; USNM, United States National Mu-
seum of Natural History, Washington, DC.

Tortricinae: Euliini
Seticosta Razowski

Seticosta was described by Razowski (1986a) to accommodate Eulia
archnogramma Meyrick, Eulia tholeraula Meyrick, and Eulia homo-
sacta Meyrick. Males are characterized by elongate antennal cilia (1.5-
2.0 x segment diameter), a foreleg hairpencil (Brown 1989), narrow

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or subbasally constricted valvae, and a dense patch of long, spine-like
setae on the subbasal region of the costa of the valva. The latter character
appears to provide the most convincing synapomorphy for the genus.
Females have long, narrow apophyses, slender papillae anales, an elon-
gate, slender ductus bursae, and an accessory bursa arising from the
ductus bursae, unassociated with the ductus seminalis that usually arises
from the corpus bursae.

Seticosta aeolozona (Meyrick), new combination

Eulia aeolozona Meyrick, 1926, Exotic Microlepidoptera 3:252; Clarke, 1958, Cat. Type
Spec. Microlep. Brit. Mus. Descr. Meyrick 3:116.

The male holotype (BMNH) lacks the abdomen, prohibiting genitalic
comparison. However, aeolozona is extremely similar to several un-
described species of Seticosta (BMNH, USNM) in forewing shape, size,
and pattern. The holotype possesses a foreleg hairpencil consistent with
other species in the genus.

Seticosta hypsithrona (Meyrick), new combination

Eulia hypsithrona Meyrick, 1926, Exotic Microlepidoptera 3:251; Clarke, 1958, Cat. Type
Spec. Microlep. Brit. Mus. Descr. Meyrick 3:128:

The holotype male (BMNH) can be assigned to Seticosta on the basis
of the long, spine-like setae on the subbasal portion of the costa of the
valva, the narrow valva, the elongate labial palpi, and the long (1.5 x
segment diameter) antennal cilia.

Seticosta mirana (Felder and Rogenhofer), new combination

Tortrix mirana Felder and Rogenhofer, 1875, Reisse Novara, pi. 139:24.

The holotype (BMNH), reported to be a male from Venezuela, is a
female. In the genitalic preparation (BMNH slide no. 7811), the corpus
bursae has a pair of small triangular signa, indicating that this species
should be assigned to the Laspeyresiini (Olethreutinae). However, I
dissected an identical female (Venezuela, Aragua, Colonia Tovar, 6000-
7000', Holt C. M. Exped.; CMNH) and found the genitalia to be con-
sistent with other species of Seticosta, i.e., long, slender apophyses and
narrow papillae anales, and the presence of an accessory bursa unas-
sociated with the ductus seminalis. The female genitalia on BMNH
slide no. 7811 evidently are associated incorrectly with the holotype of
mirana. Seticosta mirana is superficially similar to other large species
in the genus (e.g., arachnograma and several undescribed species
[USNM]). A male from Rio de Janeiro (BMNH) has elongate labial
palpi, long antennal cilia, and a foreleg hairpencil. It lacks the abdomen,
prohibiting genitalic comparison.

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315

Seticosta multifidana (Zeller), new combination

Teras multifidana Zeller, 1877, Exotische Microlepid., Hot. Soc. Entomol. Ross. 13:47,
fig. 29.

This species is represented only by the holotype female (BMNH)
collected in Bogota. Although the genitalia (BMNH slide no. 8588) are
fairly divergent from other species in the genus, particularly the sparsely
setose, crescent-shaped papillae anales and the sclerotized patch of the
corpus bursae, multifidana is similar to other species of Seticosta in
the general configuration of the female genitalia and in superficial
facies. Three males from Carabaya and Cuzco, Peru, are associated
with the holotype (BMNH); although similar, they probably are not
conspecific with multifidana. The males possess several features typical
of Seticosta, namely elongate labial palpi, long antennal cilia, foreleg
hair pencil, and the unusual patch of spine-like setae on the costa of the
valva (BMNH slide no. 8767).

Seticosta sagmatica (Meyrick), new combination

Eulia sagmatica Meyrick, 1912, Trans. Entomol. Soc. London 1911:680; Clarke, 1958,
Cat. Type Spec. Microlep. Brit. Mus. Descr. Meyrick 3:136.

The holotype male (BMNH) from Colombia can be unambiguously
assigned to Seticosta on the basis of the spine-like setae on the subbasal
region of the costa of the valva (JFGC slide no. 6217). Other features
are consistent with this association, including elongate labial palpi, long
antennal cilia, and male foreleg hairpencil.

Seticosta versabilis (Meyrick), new combination

Eulia versabilis Meyrick, 1926, Exotic Microlepidoptera 3:251; Clarke, 1958, Cat. Type
Spec. Microlep. Brit. Mus. Descr. Meyrick 3:143; Razowski, 1986, Bull. Polish Acad.
Sci., Biol. Sci. 35:70.

Although superficially similar to some Chlidanotini (Razowski 1986b),
the female genitalia of versabilis are consistent with those of other
species of Seticosta. The apophyses are long and slender, the papillae
anales are narrow, the ductus bursae is long and slender and bears an
accessory bursa from the ductus bursae unassociated with the ductus
seminalis. In addition to the female holotype from Bolivia (BMNH),
there are two females and a male from Carabaya, Peru (BMNH), that
are almost certainly conspecific with versabilis.

Inape Razowski

Inape was described by Razowski (1988) to accommodate Eulia bi-
remis Meyrick and the type species, Inape penai Razowski. Synapo-
morphies include a pair of blunt, digitate processes submedially from

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the transtilla in the male, and complex signa in the female with a
longitudinally arranged, strongly spined signum and a large, marginally
dentate, more caudal signum. In some species one of the two signa is
reduced. Males of all species have a well developed foreleg hairpencil
(Brown 1989).

Inape auxoplaca (Meyrick), new combination

Eulia auxoplaca Meyrick, 1926, Exotic Microlepidoptera 3:255; Clarke, 1958, Cat. Type
Spec. Microlep. Brit. Mus. Descr. Meyrick 3:119.

I. auxoplaca is represented by the holotype male only (BMNH). The
unique modification of the transtilla (JFGC slide no. 6348) provides
evidence for its inclusion in Inape. In addition, the genitalia share many
symplesiomorphies with other species in the genus, e.g., large upturned
valvae with broadly rounded apex, short, stout aedeagus with two stout
cornuti, and unmodified uncus, gnathos and socius. I. auxoplaca also is
similar to other species in the genus in superficial facies. Given the
sexual dimorphism exhibited by several species of Inape, it is possible
that auxoplaca represents the male of I. biremis ; both were collected
at Mt. Tolima, Colombia, in October 1920.

Inape iantha (Meyrick), new combination

Cnephasia iantha Meyrick, 1912, Trans. Entomol. Soc. London 1911:684.

Eulia iantha; Clarke, 1958, Cat. Type Spec. Microlep. Brit. Mus. Descr. Meyrick 3:131.

I. iantha is represented by the holotype female (BMNH) and a second
female (USNM; same data as holotype). The characteristic longitudinal
signum with elongate spines confirms its membership in Inape. The
second signum is reduced to a patch of slender, parallel spines.

Inape xerophanes (Meyrick), new combination

Tortrix xerophanes Meyrick, 1909, Trans. Entomol. Soc. London 1909:15.

Eulia xerophanes; Clarke, 1958, Cat. Type Spec. Microlep. Brit. Mus. Descr. Meyrick 3:
143.

I. xerophanes was described from a single male taken in Aqualani,
Peru (BMNH). It is assigned to Inape on the basis of the digitate
submedial processes from the transtilla, the most convincing male syn-
apomorphy for the genus. It also shares many symplesiomorphies with
its congeners, including the upturned valva with broadly rounded apex,
stout aedeagus with two slender cornuti, and unmodified uncus, gnathos
and socius. A female specimen (BMNH) from Santo Domingo, Peru,
appears to be conspecific with the holotype. The genitalia of the female
(BMNH slide no. 8857) are typical of the genus.

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317

Inape zelotypa (Meyrick), new combination

Eulia zelotypa Meyrick, 1912, Trans. Entomol. Soc. London 1911:679; Clarke, 1958, Cat.
Type Spec. Microlep. Brit. Mus. Descr. Meyrick 3:143.

I. zelotypa was described from 15 specimens that appear to represent
at least three species (BMNH), all of which can be referred to Inape.
The two new species will be described in a revision of the genus (Brown,
in prep.). In I. zelotypa the digitate submedial processes of the transtilla
are extremely elongate, but other features of the genitalia (viz., valvae,
uncus, gnathos, socii, aedeagus) are consistent with other species of
Inape. In addition to the foreleg hairpencil, males of I. zelotypa have
an additional hairpencil arranged longitudinally along the costa of the
hindwing, densely covered by black scales. Three males (BMNH) of a
closely related undescribed species, included in the type series, have
both the foreleg and hindwing hairpencil, but lack the patch of black
scales on the hind wing.

Clarkenia Razowski

Clarkenia was described by Razowski (1988) to accommodate C.
superba Razowski and C. miramunda Razowski. Adults are large,
brightly colored, somewhat checkered moths. Males have large valvae,
short, oval socii, a long, narrow uncus, and two patches of distinctly
different types of cornuti in the vesica of the aedeagus; they lack the
foreleg hairpencil. Females have narrow, nearly parallel-sided papillae
anales, a short, slender ductus bursae, and an elongate corpus bursae.

Clarkenia lacertana (Zeller), new combination

Sciaphila lacertana Zeller, 1866, Stett. Entomol. Z. 27:151.

Eulia lacertana; Meyrick, 1926, Exotic Microlepidoptera 3:249; Clarke, 1958, Cat. Type
Spec. Microlep. Brit. Mus. Descr. Meyrick 3:140.

Although the holotype is apparently lost, two males, one from Co-
lombia and one from Venezuela, match the description of lacertana ,
and are identified as such in the collection of the BMNH. The specimens
are similar to other species of Clarkenia in superficial facies and char-
acters of the genitalia.

Clarkenia nivescens (Meyrick), new combination

Eulia nivescens Meyrick, 1926, Exotic Microlepidoptera 3:250; Clarke, 1958, Cat. Type
Spec. Microlep. Brit. Mus. Descr. Meyrick 3:132.

C. nivescens is represented by the holotype male (BMNH) collected
on Mt. Tolima, Colombia. The species is transferred to Clarkenia on
the basis of the similarity of the valvae, socii, uncus, and cornuti, to
those of C. superba. The two species also are superficially similar. C.

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nivescens differs from superba in its uniquely modified gnathos with
elongate lateral processes.

Clarkenia trilobopa (Meyrick), new combination

Eulia trilobopa Meyrick, 1926, Exotic Microlepidoptera 3:249.

Eulia triloba (misspelling) Clarke, 1958, Cat. Type Spec. Microlep. Brit. Mus. Descr.
Meyrick 3:140.

C. trilobopa was described from a single male (BMNH) from Rio
Grande do Sul, Brazil. The holotype is lacking the abdomen. Although
slightly smaller in forewing length, C. trilobopa is extremely similar in
superficial facies to other species in the genus. The holotype lacks the
foreleg hairpencil. In his description of Eulia trilobopa, Meyrick (1926)
indicated that it was “allied” to lacertana; both species herein are
transferred to Clarkenia. Razowski (1988) recognized the similarity of
trilobopa to Clarkenia superba, but incorrectly interpreted the figure
of trilobopa in Clarke (1958, pi. 70) as that of Eulia trapezoides Meyrick.
Consequently, he did not include trilobopa in Clarkenia because the
genitalia of trapezoides (which he believed belong to trilobopa) were
dissimilar to other species of Clarkenia.

Ernocornutia Razowski

Ernocornutia was described by Razowski (1988) to accommodate E.
catopta Razowski and E. capronata Razowski. Males are characterized
by the strong, slender sacculus reaching about three-fourths the distance
from the base to the apex of the valva, capitate uncus, and a small
funnel-shaped cornutus; all known species possess the male foreleg
hairpencil (Brown 1989).

Ernocornutia carycodes (Meyrick), new combination

Eulia carycodes Meyrick, 1926, Exotic Microlepidoptera 3:256; Clarke, 1958, Cat. Type
Spec. Microlep. Brit. Mus. Descr. Meyrick 3:120.

E. carycodes is represented by the holotype male (BMNH) from Mt.
Tolima, Colombia. It is similar to E. catopta, also from Colombia, in
several features, namely the narrow portion of the valva extending
beyond the sacculus, widely separated, distal processes from the gnathos,
and the general configuration of the aedeagus. E. carycodes can be
distinguished from E. catopta by the broad dorsal portion of the tegu-
men and by the presence of slender teeth-like cornuti in the distal
portion of the vesica.

Silenis Razowski

Razowski (1987) described Silenis for the single Bolivian species S.
senilis Razowski, known only from the male holotype (USNM). Several

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319

undescribed congeners are represented in the collections of the BMNH,
USNM, and UCB.

Silenis eurydice (Butler), new combination

Sericoris eurydice Butler, 1883, Trans. Entomol. Soc. London 1883:72.

The holotype male (BMNH) from Chile lacks the abdomen pre-
venting genitalic comparison. However, in superficial facies, S. eurydice
is nearly identical to several undescribed species (BMNH, USNM) that
are clearly congeneric with Silenis senilis.

Proeulia Clarke

Prior to the description of Proeulia boliviae (Razowski 1988), the
genus was considered restricted to Chile and its offshore islands. Clarke
(1962) described the genus for two species from Juan Fernandez Islands,
Obraztsov (1964) added nine species from central Chile, and Clarke
(1980) described two additional species from San Ambrosio Island.
Males have broad valvae with an upturned costa and a narrow sacculus,
short antennal cilia, a short, stout aedeagus with several large cornuti,
and a foreleg hairpencil. Females are characterized primarily by sym-
plesiomorphies, except for the short, broad ductus bursae, and the
unusual signum consisting of a nearly circular sclerotized patch bearing
a short, blunt thorn.

Proeulia hypochloris (Meyrick), new combination

Eulia hypochloris Meyrick, 1932, Exotic Microlepidoptera 4:256; Razowski, 1964, Polska
Akad. Nauk, Ann. Zool. 22:459.

Eulia hypochloris (holotype NHMV), which occurs from Costa Rica
to Brazil (USNM), is almost certainly congeneric with Proeulia boliviae,
to which the male genitalia are extremely similar. The female of hy-
pochloris lacks the signum and the male lacks the foreleg hairpencil.
The inclusion of hypochloris and boliviae in Proeulia significantly
broadens previous concepts of the genus both morphologically and
biogeographically; hence, it is possible that hypochloris and boliviae
represent an undescribed genus closely related to Proeulia. Conse-
quently, the assignment of these species to Proeulia may represent only
an interim solution.

Proeulia dives (Butler), new combination

Oenectra dives Butler, 1883, Trans. Entomol. Soc. London 1883:68.

The name dives is listed as a synonym of Cnephasia fulvaria Blan-
chard in the collection of both the BMNH and USNM. I examined the

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holotype of dives (BMNH), but I have been unable to locate the holotype
of C. fulvaria to confirm the synomymy. Powell and Razowski (1989)
assign fulvaria to Proeulia. The holotype of dives from Chile was
reported to be male; it is a female. The genitalia of the holotype and
two additional females (BMNH slides no. 7806, 8766, 8702) are similar
to other species of Proeulia. Although the single male from Valparaiso,
Chile (BMNH), matches the holotype in superficial facies, the genitalia
differ from other species of Proeulia in the spined transtilla, weakly
dentate gnathos, and absence of cornuti; the specimen lacks the foreleg
hairpencil. These characters are fairly divergent from other Proeulia
and either represent autapomorphies for the species, or evidence that
a new genus may be required for dives. Consequently, the transfer of
dives (and possibly fulvaria) to Proeulia is provisional.

Bonagota Razowski

Bona got a was described by Razowski (1986a) for Sciaphila bogotana
Zeller (Colombia), Ptherochroa cranaodes Meyrick (Argentina), Eulia
melanecta Meyrick (Ecuador), Cryptolechia penthinella Zeller (Co-
lombia), and Eulia salubricola Meyrick (Argentina). The genus is re-
markably homogenous in superficial facies, and extremely similar to
Apotomops Powell. Bonagota and Apotomops appear to be sister taxa
on the basis of the unique, slightly attenuate, accessory pouch from the
ductus bursae, and the similarity in wing venation (Powell 1986). The
male genitalia of both genera are characterized primarily by symple-
siomorphies. The valvae are large, weakly deflexed medially in the
distal third (inconspicuous in slide mounted preparations), and usually
with a distinctly sclerotized, strongly-arched costa. In Bono got a the
male antennal cilia and the uncus are unmodified, and the socii are
moderately digitate. In Apotomops the antennal cilia are extremely
short, the uncus is broadly capitate with a large basal expansion, and
the socii are greatly reduced and nearly entirely fused to the gnathos.

Bonagota wilkinsonii (Butler), new combination

Sericoris wilkinsonii Butler, 1883, Trans. Entomol. Soc. London 1883:41.

The holotype of B. wilkinsonii (BMNH) lacks the abdomen, but is
superficially similar to other species of Bonagota. A specimen matching
the holotype (BMNH) has genitalia that share many symplesiomorphies
with other species of Bonagota. They are also similar to “Acleris”
crocoptycha (Meyrick) from Argentina, and “A.” acmanthes (Meyrick)
from Chile, which appear to be congeneric with each other, and un-
related to the genus Acleris (Tortricini).

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321

Chlidanotinae: Chlidanotini
Monortha Razowski and Becker

Monortha was described in the Polyorthini for the species M. funesta
Razowski and Becker and M. corusca (Mey rick), known only from males
(Razowski and Becker 1981). Razowski (1986b) transferred the genus
to the Chlidanotini on the basis of the female genitalia, which share
several uniquely derived characters with other Chlidanotini, viz., strong
setae on the sterigma, ductus seminalis from near the junction of the
corpus and ductus bursae, a unique accessory bursa from near the
middle of the corpus, and the unusual “asteroid” signum of long spines.
Monortha is distinguished from other chlidanotines by the following
male genitalic characters: socii and hami fused, with stout, spine-like
setae, and similar spine-like setae from the venter of the uncus.

Monortha illaqueata (Meyrick), new combination

Capua illaqueata Meyrick, 1917, Trans. Entomol. Soc. London 1917:6.

Capua illaquaeta (misspelling) Clarke, 1958, Cat. Type Spec. Microlep. Brit. Mus. Descr.

Meyrick 3:71.

This species is represented only by the holotype female from French
Guiana (BMNH). The heart-shaped configuration of the papillae anales,
strong setae on the posterior edge of the sterigma, position of the ductus
seminalis, and putative possession of an accessory bursa, indicate that
illaqueata should be assigned to Monortha. The signum is greatly re-
duced to a patch of concentric, dimple-like sclerites, and the accessory
bursa is lacking in the genitalic slide (JFGC 6289), hence its position
in the Chlidanotini has not been recognized previously. Vestiges of the
ductus of the accessory bursa are present, and the dimple-like modi-
fication of the signum is seen to a lesser degree in other Chlidanotini.
In superficial facies illaqueata is most similar to M. corusca (holotype
BMNH).

Olethreutinae: Eucosmini
Eucosma torrens Meyrick, revised status

Eucosma torrens Meyrick, 1927, Exotic Microlepidoptera 3:334.

Eulia torrens; Clarke, 1958, Cat. Type Spec. Microlep. Brit. Mus. Descr. Meyrick 3:140.

The holotype of torrens (BMNH) lacks the abdomen. Clarke (1958)
transferred the species to Eulia on the basis of its superficial similarity
to other Neotropical species assigned to this large, polyphyletic genus.
However, the genitalia of an undescribed species (USNM) superficially
nearly identical to torrens indicate that the two are obviously olethreu-
tines, probably referrable to the Eucosmini.

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Acknowledgments

For allowing me to examine specimens in their care I thank: J. F. G. Clarke (USNM),

M. Lodi (NHMV), J. Powell (UCB), J. Rawlins (CMNH), and K. Tuck (BMNH). Jerry

A. Powell kindly reviewed the manuscript. An anonymous reviewer provided many

helpful suggestions for improvement of the manuscript. This work was completed under

the support of a Smithsonian Postdoctoral Fellowship.

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foreleg hairpencil in the Tortricinae (Lepidoptera: Tortricidae). Entomol. News 101.
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Clarke, J. F. G. 1958. Catalogue of the type specimens of Microlepidoptera in the
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1962. A new tortricid genus from South America. Proc. Biol. Soc. Washington

75:293-294.

1980. Two new species of Proeulia from the Desventuradas Islands (Tortricidae).

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Obraztsov, N. S. 1964. Neotropical Lepidoptera, V. Synopsis of the species of the
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Powell, J. A. 1983. Tortricoidea (Families Tortricidae and Cochylidae), pp. xvi-xvii,
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1986. Synopsis of the classification of the Neotropical Tortricinae, with descrip-
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372-398.

& J. Razowski. 1989. Tortricidae: Tortricinae. In Heppner, J. B. (ed.), Checklist

of Neotropical Lepidoptera. Vol. 2, Pyraloidea to Tortricoidea. W. Junk Publ., The
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Razowski, J. 1982. Description of the new Neotropical genus and species of Tortricidae
(Lepidoptera). Sci. Nat 34:3-4.

1986a. Description of new Neotropical genera of Archipini (Lepidoptera: Tor-
tricidae) and rectification of the Deltinea problem. Sci. Nat 52:21-25.

1986b. Neotropical Chlidanotini (Lepidoptera: Tortricidae). Bull. Polish Acad.

Sci., Biol. Sci. 35:61-71.

1987a. The genera of Tortricidae (Lepidoptera). Part I: Palaearctic Chlidano-

tinae and Tortricinae. Acta Zool. Cracov. 30:141-355.

1987b. New taxa of Archipini (Lepidoptera: Tortricidae) from South America.

Tinea 12 (suppl.):123-138.

1987c. Transtillaspis (Lepidoptera: Tortricidae) — New Neotropical Tortricinae

genus and its species. Bull. Polish Acad. Sci., Biol. Sci. 35:73-82.

1988. New genera and species of the Neotropical Archipini (Lepidoptera:

Tortricidae). Acta Zool. Cracov. 31:387-422.

& V. Becker. 1981. Brazilian Polyorthini (Lepidoptera, Tortricidae). Acta Zool.

Cracov. 25:389-404.

Received for publication 27 May 1989; accepted 30 August 1989.

Journal of the Lepidopterists’ Society
43(4), 1989, 323-326

NEOEROMENE LUTEA
(LEPIDOPTERA: PYRALIDAE; CRAMBINAE),

A NEW SPECIES FROM BRAZIL

David E. Gaskin

Department of Zoology, University of Guelph,

Guelph, Ontario NIG 2W1, Canada

ABSTRACT. Neoeromene lutea, a new species of diptychophorine Crambinae, is
described from three specimens from southern Brazil, raising the number of known species
in this genus to seven. Neoeromene lutea shares juxtal apomorphies with N. straminella
(Zeller), also of Brazil, but has less development in the costal region of the valva and in
the apical sclerotization of the aedeagus.

Additional key words: Neoeromene straminella, Santa Catarina.

The neotropical Crambinae assigned to Pareromene Osthelder by
Bieszynski (1967) were recently transferred to new genera, one of them
Neoeromene (Gaskin 1986). To prove the well-known unwritten rule
that a tidy revision of a taxonomic group is by far the best way to
stimulate discovery of all that other material the author missed, Julian
Donahue contacted me immediately and drew my attention to three
specimens from Brazil, taken by E. D. Jones, in the collection of the
Natural History Museum of Los Angeles County. This paper describes
these, which are conspecific and represent the only known specimens
of a hitherto unrecognized species of Neoeromene. The name lutea is
applied, with reference to the brilliant yellow-orange ground color of
the forewings. In the description, decimals indicate the position of a
feature, as a proportion of the total length of a structure or organ. In
the forewing, measurements along the costa are from the base, on the
termen from the apex, and along fascia from the costa. In the male
genitalia they are from the base of each structure.

Neoeromene lutea , new species

(Fig. 1)

Description. Forewing length 7.5 mm (N = 3). Head and thorax bright yellow, latter
with a black ring of scales anteriorly, patagia with a few scattered, large dark spots. Labial
and maxillary palpi pale yellow with blackish apices. Abdomen pale brown with some
scattered yellow scales, legs pale yellow banded with brown. Antennae with small bi-
pectinations and no macroscopic sexual differences. Male frenulum single, female double.
Ground color of forewings bright yellow, including apical and subterminal regions. Basal
region with three blackish brown spots in an oblique line towards base of antemedial
fascia which is at 0.4 from base, dark brown, narrow and tridentate in the form of a
crude Greek letter £. There is a small brown streak on the costa at 0.5 and some variable
dark streaks of scales in the reniform region of the disc. Postmedial fascia double, a pair
of brown waved lines with a few lustrous silvery scales between them from costa to about
0.3, fascia then diverging towards dorsum, proximal line dentate at 0.8 from costa. There
are three brown spots on the termen from 0.5-0. 8. Apical zone contains a narrow, elongate,

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Fig. 1. Neoeromene lutea n.sp. Male paratype from Santa Catarina, Brazil, and below,
female paratype, unlabelled but from the same series in the Natural History Museum of
Los Angeles County.

oblique silvery white streak from the costa, not reaching the termen. Subapical indentation
present at 0.3 from apex, and a secondary indentation at 0.5, with some silvery scaling
not quite reaching postmedial fascia. Ventral surfaces brownish, with apical and termen
markings repeated from dorsal surface. Hindwings lustrous off-white in both sexes, with

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Fig. 2. Neoeromene lutea. Male holotype. Genitalia in posterior aspect, with left
valva not shown (left) and aedeagus (right).

white cilia with brown bases, giving the appearance of a double brown marginal line.
The cilia of the forewings are lustrous gray-brown.

Male genitalia (Fig. 2) (N = 1). Uncus curved, with blunt apex; gnathos slightly shorter
than uncus, curved dorsad with apical region significantly swollen, bluntly pointed; teg-
umen squat with strong posterior margins; saccus broadly rounded, vinculum broadly
triangular. Juxta a large complex structure, with a spatula te apex equal in length to uncus,
and a broadly rounded basal region, the whole about 2 x uncus; below apical area a pair
of lateral swellings each have a sharp prong, about 0.5 x uncus, curved extrorse. Valva
about 2-2.5 x uncus [depending how measured, basally or along costa] with a sclerotized
costal region reaching the bluntly rounded apex. Aedeagus about 1.3 x length of valva,
sharply curved ventrad in proximal third, with a pair of elongate lateral striations and
some minute apical spines.

Types: Holotype <3; without label but probably same as for paratype male. Paratypes:
1 <3, BRAZIL, Santa Catarina, —4.1922, E. D. Jones; 1 9 without label. Both paratypes
lack their abdomen. Consequently, the only specimen with an abdomen has been des-
ignated as holotype, even though it has no locality label.

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Discussion. Neoeromene lutea resembles N. straminella (Zeller) in
most major forewing characters, but the yellow-orange ground colour
and the dark, strongly marked fascia immediately distinguish the for-
mer species. There is little variation between the three specimens ex-
amined. Like N. straminella, N. lutea has a proportionately huge,
apically bicornuate juxta. The form of the valva in both species is almost
identical except that N. straminella has a basal prong. Both differ from
all other species in the genus except N. felix (Meyrick) in having vir-
tually no sclerotization in the sacculus of the valva. Neoeromene lutea
has the least apical sclerotization of the aedeagus in the genus, repre-
sented only by a row of microscopic prongs. The apically swollen gna-
thos and the blunt, setulose uncus closely resemble those of N. parvi-
puncta Gaskin, also of Brazil. The early stages, food plants and duration
of the flight period (which included April), are unknown.

Acknowledgments

I thank Julian P. Donahue, Natural History Museum of Los Angeles County, for bringing
this material to my attention and arranging the loan, and two anonymous reviewers for
constructive criticism.

Literature Cited

Bkeszynski, S. 1967. Studies on the Crambinae (Lepidoptera). Part 44. New neotropical
genera and species. Preliminary checklist of neotropical Crambinae. Acta. Zool.
Cracov. 12:39-110.

Gaskin, D. E. 1986. New genera for the neotropical “Pareromene” species (Pyralidae:
Crambinae). J. Lepid. Soc. 40:271-288.

Received for publication 30 November 1988; accepted 15 May 1989.

GENERAL NOTES

Journal of the Lepidopterists’ Society
43(4), 1989, 327-328

DISCOVERY OF THE CARPENTER MOTH
MORPHEIS CLENCHI (COSSIDAE) IN MEXICO

Additional key words: Zeuzerinae, Sierra Madre Occidental.

When I described Morpheis clenchi (Cossidae: Zeuzerinae) from southern Arizona
(Donahue, J. P. 1980, J. Lepid. Soc. 34(2): 173-181), I predicted that this large, striking,
sphingid-like carpenter moth would undoubtedly be found in adjacent Mexico. This
prediction has been confirmed as a result of field work by members of the Sonoran
Lepidoptera Survey, in cooperation with Sonoran Arthropod Studies, Inc. (SASI) of Tuscon,
Arizona, and by Terry A. Sears et al., whose extensive moth collections from the Barrancas
del Cobre (“Copper Canyon”) region of the Sierra Tarahumara, Sierra Madre Occidental,
are deposited in the Bohart Museum of Entomology, University of California, Davis
(UCD). I thank Steve Prchal of SASI and R. O. Schuster and Adam Porter of UCD for
loaning the specimens reported on here.

I have now examined 11 adult male specimens of Morpheis clenchi from the Mexican
states of Sonora, Sinaloa, and Chihuahua, and which have been deposited in the following
institutions: SASI, UCD, Natural History Museum of Los Angeles County (LACM), and
Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico, D.F., Mexico
(UNAM). Seven localities are represented, all west of the continental divide in the Sierra
Madre Occidental; they are listed from north to south. Localities not readily found on
road maps are annotated. The female and the larval food plant of this species remain
unknown.

SONORA: lU mi (0.4 km) E of Rancho El Encino, elev. 3600 ft (1097 m), 29 July 1983,
Steve Prchal (1 8 SASI, 1 8 LACM). [This locality is in the Sierra La Madera between
Moctezuma and Huasabas, approx. 29°50'N, 109°20'W.]

SONORA: 17 mi (27 km) W of Moctezuma, to U.V. [light], 20 July 1984, Doug Mullins
(1 3 LACM).

SONORA: 13 mi (21 km) E of El Novillo, Study Canyon, elev. 3600 ft (1097 m), 3-4
Aug. 1983, Steve Prchal (1 8 SASI, 1 <5 UNAM). [This locality is in the Sierra La Campanera
between Mazatan and Bacanora, approx. 29°00'N, 109°30'W.]

SONORA: Trinidad Mine Canyon, at house, elev. 3425 ft (1044 m), 30 July 1984, Steve
Prchal (1 8 SASI); same locality and collector, 11-12 Aug. 1983 (1 8 SASI, 1 8 LACM).
[This locality is ca. 5 road mi (8 km) E of Santa Rosa, between Santa Rosa and Yecora,
approx. 28°27'N, 109WW.]

CHIHUAHUA: 3 mi (4.8 km) S Temoris, elev. 4700 ft (1433 m), 29 Aug. 1969, T. A.
Sears, R. C. Gardner, C. S. Glaser (1 8 UCD). [Temoris and nearby Temoris Station, where
the elevation is 3365 ft (1026 m), are at km 708 on the Chihuahua al Pacifico Railroad,
approx. 27°15'N, 108°15'W.]

CHIHUAHUA: Santo Nino, 9 Aug. 1969, T. A. Sears, R. C. Gardner, C. S. Glaser (1
8, UCD). [Approx, elev. 1800 ft (550 m), very near the Sinaloa state line at km 737 on
the Chihuahua al Pacifico Railroad, approx. 27°00'N, 108°22'W.]

SINALOA: 5.5 mi (8.9 km) NW Choix, 15 July 1968, T. A. Sears, R. C. Gardner, C.
S. Glaser (1 3, UCD).

All of these localities, and the known localities for this moth in Arizona, are in the
biotic community described as Madrean Evergreen Woodland, a mild winter /wet summer
community characterized by evergreen oaks ( Quercus ) or a mixture of oaks, juniper
(. Juniperus ) and Mexican pinyon ( Pinus cembroides ) (Brown, D. E., ed. 1982, Biotic
communities of the American Southwest — United States and Mexico, Desert Plants 4(1-
4):342 pp., and accompanying map by Brown, D. E. and C. H. Lowe 1980, Biotic

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communities of the Southwest, Gen. Tech. Report RM-78, Rocky Mountain For. & Range
Exp. Sta., Forest Service, U.S. Dept. Agric.).

Besides documenting the first, but not surprising, records of Morpheis clenchi from
Mexico, this note illustrates a distributional phenomenon that will be observed more
commonly as surveys of the Lepidoptera fauna of northwestern Mexico are intensified,
namely that species presently known only from the southwestern United States may
actually be widespread and common in adjacent northwestern Mexico. Similarly, with
more intensive faunal studies in the southwestern United States we may expect to find
species presently known only from northwestern Mexico, although perhaps only in isolated,
relict populations or as strays. I urge lepidopterists from the United States and Mexico to
pool their talents and efforts to conduct a joint survey of this intriguing border-straddling
region.

Julian P. Donahue, Natural History Museum of Los Angeles County, 900 Exposition
Boulevard, Los Angeles, California 90007-4057.

Received for publication 5 July 1988; accepted 21 July 1989.

Journal of the Lepidopterists’ Society
43(4), 1989, 328-331

A DESCRIPTION OF TOPOTYPICAL MALE
HEM1LEUCA DIANA (SATURNIIDAE)

Additional key words: Colorado, type locality, Hemileuca grotei, sister species.

Hemileuca diana Packard (Saturniidae) was first described in 1874 from a single female,
collected in 1873; the locality was cited only as “Plum Creek” in Colorado (Packard, A. S.
in Hayden, F. V. 1874, Annual Report of the United States Geological and Geographical
Survey of the Territories embracing Colorado, being a report of progress of the exploration
for the year 1873, Government Printing Office, Washington, D.C., 718 pp ). Prior to F.
M. Brown (1972, J. Lepid. Soc. 26:245-247) a more precise location of the type locality
of H. diana was largely in question because there are many streams in Colorado named
“Plum Creek.” According to the historical records of Hayden and A. C. Peale (in Hayden
op. cit.), the most probable Plum Creek would be the northward flowing tributary of the
segment of the South Platte River that flows through what is now Douglas County. The
most likely H. diana type locality on this tributary is along the headwaters of Plum Creek
in the area between Larkspur and Palmer Lake (Brown op cit.). This location is slightly
southwest of Castle Rock south of Denver, near present Interstate Highway 25. The study
area where this research was conducted is along Plum Creek about 12 km W of Castle
Rock at the junction of Douglas County roads 46 and 105 (Wolfensberger and Perry Park
roads, respectively) very near or in the Christy Ridge housing development. This is about
8 km N of the probable location cited by Brown and well within the presumed type
locality area.

Hemileuca diana is recorded from Colorado, Arizona, and Mexico (Ferguson, D. C.
1971, Bombycoidea, Saturniidae (in part), in Dominick, R. B., et al. (eds.), The moths of
America north of Mexico, fasc. 20.2A: 1-153, col. pis. 1-11, E. W. Classey, London; Tuskes,
P. M. 1986, J. Lepid. Soc. 40:27-35), but until quite recently no other specimens were
known to have been collected from the type locality. The biology of Arizona H. diana
has been described (Tuskes op. cit.), although comparison of southern Arizona males to
topotypical males was not made because such topotypical material was unknown at that
time. Topotypical H. diana males are now known. At the Christy Ridge housing devel-

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329

opment, between the dates of 28 September and 4 October 1987, 72 H. diana males were
collected by the authors and R. S. Peigler.

Hemileuca grotei Grote & Robinson is recorded from the Edwards Plateau in central
Texas and from New Mexico (Ferguson op. cit.; Kendall, R. O. & R. S. Peigler 1981, J.
Lepid. Soc. 35:41-50; and Tuskes op. cit.). Through comparison of topotypical H. diana
to Arizona and Mexico H. diana and to H. grotei populations found in Texas and New
Mexico, it is clear the two taxa are closely related, quite possibly sister species, and form
the closely knit H. diana complex (Tuskes, P. M. 1984, J. Lepid. Soc. 38:281-308, Table
1). Morphological differences of material sampled from various localities seem to indicate
that some populations of what are now called H. grotei and H. diana may be misidentified
or even undescribed taxa (see Tuskes 1986 op. cit.: p. 28, figs. 2c and 2d). An investigation
of the H. grotei complex is currently underway.

Topotypical H. diana males have been deposited in the Museum of Comparative
Zoology, Harvard University, in association with the holotype female; Natural History
Museum of Los Angeles County and San Diego Natural History Museum, California;
Denver Museum of Natural History, Colorado; National Museum of Natural History,
Smithsonian Institution, Washington, D.C.; American Museum of Natural History, New
York; and in the personal collections of R. S. Peigler, Colorado; K. L. Wolfe, M. J. Smith,
P. M. Tuskes, California; W. A. Nassig, West Germany; C. Lemaire, France; and the
authors.

The following description of a topotypical H. diana male is based on a specimen (Fig.
la) collected 4 October 1987 by the senior author and R. S. Peigler very near the presumed
type locality.

HEAD: Eyes dark brown near black. All hairs black. Labial palps absent. Antennae
black, bipectinate, 7 mm long, with 36 segments. THORAX: Collar off-white, thoracic
tufts light gray with black bases, pronotum clothed with short, light rust-red hairs
with mixed black to light gray bases, meso- and metathorax clothed with rust-red
tufts. LEGS: Femur covered with very long dark rust and white hairs, tibia with
shorter black and white hairs, tarsi black and sparsely covered with short black hairs,
claws brown. ABDOMEN: Abdominal segments I-VII covered dorsally with black
hairs, longer sparser hairs white. Caudal segments covered with long burnt orange
hairs. Ventrally segments I-IX covered with shorter black and white hairs. Interseg-
mental spaces black. FOREWING: Elongated, length 23 mm, ground color dark
brown with dark brov/n veins, antemedian band absent; yellow, bare, crescentic bar
in center of black discal spot, postmedian band white, originating at base of R5, 3
mm wide at R5, intersected proximally by discal spot; discal spot completely inter-
rupting postmedian band transforming it to two profound dashes, band terminating
at anal vein. Band 1-2 mm wider ventrally than dorsally. Postmedian area moderately
suffused with silver-gray scales. HINDWING: Length 18 mm, ground color light
black with light black veins, antemedian band absent, black discal spot partially
obscured by ground color and fading into postmedian band, postmedian band white,

3 mm wide at R, tapering sharply to tornus, and terminating at anal vein. Dorsal
and ventral postmedian band width the same. Postmedian band originating at Sc.
Proximal margin of postmedian band not sharply defined but grading from ground
color to white. Distal margin of postmedian band sharply defined. Anal margin heavily
clothed in long, light black hairs

Material examined (n = 50) demonstrates remarkably uniform morphological char-
acteristics; differences occur only in wing length, the amount of development of the fore-
and hindwing postmedian bands, and the suffusion of silver-gray scales in the forewing
postmedian area. The forewing is always elongated as in H. magnifica (Rotger) and H.
hera hera (Harris). The ground color of the fore- and hindwings remains constant while
forewing length varies between 21.0 mm and 24.5 mm (x = 23.3 mm), hindwing length
varies from 16.0 mm to 20.0 mm (x = 18.0 mm). Although the forewing postmedian
band is rarely reduced to two slight dashes, it is always conspicuously developed, varies

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Journal of the Lepidopterists’ Society

Fig. 1. a-d. Topotypical Hemileuca diana males, showing wing pattern variation.
All specimens from Plum Creek, Douglas County, Colorado; collected by S. E. Stone, D.
E. Bowman, and R. S. Peigler, 28 September-4 October 1987.

in width from 2 to 6 mm at R5, and is completely disrupted by the discal spot in 37
(74%) of the males examined (Fig. lb). In 13 (26%) of the males examined, the discal
spot penetrates the postmedian band but fails to disrupt it entirely; the band remains
connected distally by a thin (<1 mm) portion of the band (Fig. lc). The postmedian area
is suffused with silver-gray scales in 40 (80%) of the males examined; the amount of
suffusion varies from slight to very pronounced (Fig. Id). The hindwing postmedian band
is always present. It is well developed in 46 (92%), but reduced to a dash or dot in 4 (8%)
of the males examined.

We express our gratitude to R. S. Peigler for assistance in gathering data and reviewing

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331

the manuscript. We also thank C. Lemaire, M. J. Smith, and P. M. Tuskes for their reviews
of the manuscript and helpful suggestions. D. L. Vana-Miller provided assistance on the
laser printer and C. L. Fischer provided editorial comments on the manuscript.

Stephen E. Stone,1 18102 East Oxford Drive, Aurora, Colorado 80013.

Donald E. Bowman, 1602 Ulysses Street, Golden, Colorado 80401.

1 Natural Resource Specialist, U.S. Department of the Interior, National Park Service, Denver Service Center, 12795
W. Alameda Parkway, Lakewood, Colorado 80225.

Received for publication 5 December 1988; accepted 18 August 1989.

Journal of the Lepidopterists’ Society
43(4), 1989, 331

DIURNAL NECTARING BY A CATOCALA MOTH (NOCTUIDAE)

Additional key words: Manitoba, Canada, Catocala briseis, Cirsium arvense.

At about 1200 h on 4 August 1988, I was surprised to see a Catocala moth (Noctuidae)
nectaring on Canada thistle ( Cirsium arvense (L.) Scop.; Asteraceae) at the Whiteshell
Nuclear Research Establishment, 8 km south of Lac du Bonnet, Manitoba. The temper-
ature was about 21°C, with a slightly hazy sun and a light north wind. After feeding for
a few minutes, the moth flew to rest on the trunk of a cultivated poplar ( Populus sp.;
Salicaceae).

I identified the moth as C. briseis W. H. Edwards, based on illustrations in texts by C.
V. Covell, Jr. (1984, A field guide to the moths of eastern North America, Houghton
Mifflin, Boston, 496 pp.), W. J. Holland (1903, The moth book, 1968 reprint by Dover
Publications, New York, 479 pp ), and T. D. Sargent (1976, Legion of night: The underwing
moths, Univ. of Massachusetts Press, Amherst, 222 pp. ). The most distinctive features
were an irregular whitish band between the postmedian and subterminal lines on the
dorsal surface of the mainly blackish forewing, and a smaller whitish patch between this
band and the body. The hindwing was boldly banded with black and scarlet above, and
more subdued below. The only similar species, C. grotiana Bailey, is apparently unknown
in Manitoba (D. C. Hawks and R. R. Hooper pers. comm.).

Catocala moths are largely nocturnal, and are usually observed by day only if disturbed
(Sargent op. cit.). It is possible that this individual had been flushed by a predator, and
was subsequently attracted to the thistle flower before finding a new resting place. It may
well, however, have initiated feeding behavior without such stimulus. Diurnal feeding
has previously been observed in several Catocala species at “sugar” patches left over
from prior baiting trips (T. D. Sargent pers. comm.).

Although adults of many Catocala species visit bait readily (Sargent op. cit.), little is
known about their natural feeding habits. Sargent cites two species taken while nectaring
at night, both by M. C. Nielson in Michigan: C. unijuga Walker at milkweed ( Asclepias
sp.; Asclepiadaceae) and joe-pye-weed ( Eupatorium sp.; Asteraceae), and C. mira Grote
at wild bergamot ( Monarda sp.; Lamiaceae). In Manitoba and elsewhere, Canada thistle
is a favored nectar source for several butterflies (Opler, P. A. & G. O. Krizek 1984,
Butterflies east of the Great Plains, Johns Hopkins Univ. Press, Baltimore, 294 pp.).

I thank D. C. Hawks, R. R. Hooper and T. D. Sargent for information and helpful
comments.

Peter Taylor, P.O. Box 597, Pinawa, Manitoba ROE 1L0, Canada.

Received for publication 20 May 1989; accepted 23 July 1989.

332

Journal of the Lepidopterists’ Society

Journal of the Lepidopterists ’ Society
43(4), 1989, 332-333

MYRMECOPHILY AND LARVAL FOOD PLANTS OF
BREPHIDIUM ISOPHTHALMA PSEUDOFEA (LYCAENIDAE)
IN THE FLORIDA KEYS

Additional key words: Batis maritima, oviposition behavior, Salicornia virginica,

Tapinoma sessile, pupal stridulation.

G. W. Rawson (1961, J. N.Y. Entomol. Soc. 69:88-91) described and illustrated the life
history of Brephidium isophthalma pseudofea (Morrison) (Lycaenidae: Polyommatinae)
from a population near New Smyrna Beach, Florida. During a visit to the lower Florida
Keys, we made additional observations on the biology of this species.

On 23 June 1979, at the N end of Sugarloaf Key, Monroe Co., we found adults of B.
i. pseudofea in a semitidal flat where Salicornia bigelovii Torr., S. virginica L. (Cheno-
podiaceae), and Batis maritima L. (Batidaceae) grew. Fifteen larvae of B. i. pseudofea
were found on two adjacent plants of S. virginica and B. maritima. Between the bases
of the two plants was a nest of an ant, Tapinoma sessile (Say) (Formicidae: Dolicho-
derinae). These ants patrolled the stems of both plants, and avidly tended larvae of B. i.
pseudofea, stroking them with their antennae and feeding on secretions from the dorsal
nectary organ on abdominal segment (A)7. One pupa was found attached to a stem of

B. maritima. The ants also showed interest in this pupa, running over it and stroking it
with their antennae. We could not verify if they actually fed on any secretions from the
pupa, but later examination revealed a conspicuous scar on A7 corresponding to the dorsal
nectary organ on the larva. We placed a living pupa in a small glass vial, and, upon
tapping, the pupa stridulated, as has been reported for many lycaenids (Downey, J. C.
1966, J. Lepid. Soc. 20:129-155).

Several adults were seen near the plants with larvae. At ca. 1100 h, one female was
observed alternately flying around and walking over the plants for several minutes, in
the presence of the ants. We observed no aggressive behavior by the ants toward the
female. She eventually deposited a single egg on the tip of a leaf of B. maritima and
then flew away.

Rawson (op. cit .) raised B. i. pseudofea on Salicornia bigelovii, and suggested that
Batis may also be a larval food plant. We raised larvae from both S. virginica and B.
maritima to adults, thus confirming these species as larval food plants. Rawson made no
observations of ants associated with larvae or pupae.

A dorsal nectary organ on A7 and paired tentacle organs on A8 (terminology of Cottrell,

C. B. 1984, Zool. J. Linn. Soc. 79:1-57) are present in this species and many other members
of Polyommatinae. Myrmecophily has been reported for Brephidium exilis Boisduval
(Coolidge, K. 1924, Entomol. News 33:305-309), and for the African B. metophis (Wal-
lengren) (Clark, G. C. & C. B. C. Dickson 1971, Life histories of South African butterflies,
Purnell & Sons, Capetown, South Africa, 272 pp.). Ant association in B. i. pseudofea,
although previously unreported, is therefore not surprising.

The clumped distribution of immatures of B. i. pseudofea raises the possibility of ant
dependent oviposition. Although larvae and eggs were abundant on the two plants pa-
trolled by the ants, a search of nearby plants that appeared identical in general aspect,
but which lacked patrolling ants, yielded no larvae. This suggests that females may
preferentially oviposit on plants with ants (review in Cottrell op. cit.). The distribution
of ants may be a significant factor in the biology of B. i. pseudofea, which is extremely
sporadic in its appearance despite ubiquitous larval food plants in coastal habitats.

Preserved immatures are deposited in the Allyn Museum of Entomology, Florida State
Museum, Sarasota, Florida. We thank two anonymous reviewers for comments on this
manuscript.

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333

D. J. Harvey, Department of Entomology NHB Stop 127, Smithsonian Institution,
Washington, D.C. 20560.

J. Longino, Allyn Museum of Entomology, 3621 Bay Shore Road, Sarasota, Florida
33580.

Received for publication 3 October 1988; accepted 12 August 1989.

Journal of the Lepidopterists’ Society
43(4), 1989, 333-336

APPIAS PUNCTIFERA D’ALMEIDA (PIERIDAE) IN
THE DOMINICAN REPUBLIC AND PUERTO RICO

Additional key words: island biogeography, seasonal dimorphism, Greater Antilles,

Virgin Islands, Appias drusilla.

Appias punctifera is perhaps the least known and most elusive of endemic Antillean
pierids. It much resembles the familiar “Florida White” (A. drusilla Cramer) and most
obviously differs in possessing a black spot at the distal end of the forewing (FW) cell in
both sexes, more prominently in the female. It is rare in collections, and N. D. Riley
(1975, A field guide to the butterflies of the West Indies, Collins, London, 224 pp.)
mentioned that he had not seen a specimen.

The taxon punctifera was first proposed by R. F. D’Almeida (1939, Bol. Biologico, Sao
Paulo, 4(NS):50-66) as a subspecies of Appias drusilla, reducing to synonymy Tachyris
molpadia Dewitz, Tachyris margarita Dewitz, and Appias drusilla f. molpadia R fiber.
Appias punctifera was raised to specific status by W. P. Comstock (1943, Amer. Mus.
Novitates No. 1238, 1-6). D’Almeida based his description on four “all white” females
collected in Puerto Rico, the type locality. Both sexes were available to Comstock who
illustrated substantial differences between male genitalia of A. drusilla and A. punctifera,
notably in the shape of the valvae and aedeagus. He recorded from Puerto Rico one
female (Barros [sic], June), three females and one male (Coamo Springs, April) and one
female (Lajas, June). The known range of A. punctifera was extended eastwards (Com-
stock 1944, Ann. N.Y. Acad. Sci. 12 Pt. 4: Lepidoptera, 527-528) to include the American
Virgin Islands, by five males and one female (St. John, March) and one female (St. Thomas,
June). More recently, one of us (SJR) found A. punctifera to be rare but occurring
consistently in the Guanica xeric forest region of SW Puerto Rico, where it flies with the
common A. drusilla, from which is is indistinguishable on the wing. Additional records
(SJR) comprise one male (Toro Negro, October 1974), another male (Coamo, November
1974), and three males and two females (Ponce, Rt. 139, km 9.0, January 1987).

A. punctifera was first recorded from Hispaniola in a list of the butterflies of the
Dominican Republic published by O. Cucurullo (1959, Lista de Mariposas (Rhopalocera)
de Santo Domingo, published by the author, Santo Domingo, 1-13) without locality or
collecting data. Subsequently, it was mentioned by L. Marion (1975, Helios 3:42-49) but,
again, with no details. A. Schwartz (1983, Haitian butterflies, Editora R. Taller, Santo
Domingo, Dominican Republic, 69 pp.) does not list this species from western Hispaniola.
Here we report field observations on a population of A. punctifera in eastern Dominican
Republic, with comparative comments on this species in western Puerto Rico.

Puerto Rican specimens are housed in the Department of Biology, University of Puerto
Rico, Mayagiiez; all specimens collected by us from the Dominican Republic are deposited
in the Hope Entomological Collections, the University Museum, Oxford, England; the

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Journal of the Lepidopterists’ Society

Fig. 1. Appias punctifera male from Boca de Yuma, Dominican Republic: a, up-
perside; b, underside. Scale line = 1 cm.

Fig. 2. Appias punctifera female from Boca de Yuma, Dominican Republic: a, up-
perside; b, underside.

Fig. 3. Appias punctifera female from Ponce, Puerto Rico: a, upperside; b, underside.

Fig. 4. Appias drusilla boydi male from Boca de Yuma, Dominican Republic: up-
perside.

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335

male collected by L. D. and J. Y. Miller is in the Allyn Museum of Entomology, Sarasota,
Florida.

Between 4-10 January 1987, two of us (DSS, EWC) visited the Dominican Republic
and worked primarily near Boca de Yuma Dominican Republic (Altagracia Province).
The limestone surface of the plateau in this region is substantially under cultivation or
pasture, but with residual areas of xeric forest much resembling that of the Guanica
region of SW Puerto Rico. We found A. drusilla boydi Comstock in, and apparently
confined to, the forest patches, while in the cleared areas, A. punctifera was found. After
a chance capture of a worn male of A. punctifera, we particularly sought this species
and its relative A. drusilla and collected four further males and two females of the former,
all in fairly fresh condition. An additional male was collected by L. D. Miller and J. Y.
Miller, who revisited the site shortly after. All Appias collected in the fields and cleared
areas proved to be punctifera: most seen were flying out of reach (above 5 m) around
Ficus, lignum vitae ( Guiacum ), and other isolated trees, occasionally making excursions
to nearby ground scrub (where most specimens were collected), nectaring on Eupatorium,
Lantana, and other plants. We saw perhaps five specimens for each one collected, and
frequency of sightings did not appear to vary from early morning to late afternoon
through each day of observation. We cannot be certain that A. punctifera does not fly
with A. drusilla in the forest scrub, but all Appias collected there were of the latter
species.

Riley (op. cit.) did not illustrate A. punctifera, and male and female specimens from
Boca de Yuma are illustrated here (Figs. 1-2) with a male of A. drusilla boydi from the
same region (Fig. 4). Our specimens conform closely to Comstock’s (1943 & 1944 op.
cit.) descriptions, notably in the cell spot size and the conspicuous bright lemon-yellow
suffusion that occupies the basal third of the FW cell on the underside, and half or more
of the cell area in the female. The “glistening” or silky appearance of the underside
ground color in both sexes is striking. However, our material diverges slightly from
Comstock’s accounts in that we note the female upperside ground color as pale cream
on the forewing and deeper cream-yellow on the hindwing (rather than “white” and
“distinctly cream”) and, in fresh males, the upperside ground color is not “white” but
very pale eggshell green-blue, a tint that soon fades in dried specimens.

The original four males described by D’Almeida were “white” (with only a limited
black suffusion at the FW apex, as illustrated by H. Dewitz (1877, Entomol. Z. Stettin.
38:233-245) while Comstock (1944 op. cit.) described this sex as “white, with or without
a fuscous border,” but did not mention any seasonal distribution of the two forms in his
very small sample. The two female specimens in our Boca de Yuma sample both show
broad fuscous-black borders (Fig. 2), but sampling of the population over an extended
period is required to establish whether this dimorphism reflects seasonality, as has been
proposed (e.g., Riley op. cit.) for comparable variation in female markings in A. drusilla
poeyi Butler (Bahamas, Cuba and Cayman Is.) and A. drusilla boydi (Hispaniola, Puerto
Rico, the Virgin Is., and the northern Lesser Antilles), in which the more heavily marked
females have been associated with the wet season. Both females collected near Ponce
(Puerto Rico) at the time of our visit to the Dominican Republic represented the “white”
morph (Dewitz op. cit.) and one of these is illustrated in Fig. 3. In these specimens, the
fuscous FW border above is very reduced in width, to only ca. 1 mm at the apex. In
addition, the FW end-cell spot above is slightly narrower than in the more heavily marked
morph, although in each instance this spot occupies about two thirds of the distal end of
the cell. A male collected near Ponce is indistinguishable from specimens taken at the
same time in the Dominican Republic.

Unless future work in Hispaniola points to the contrary, it seems that the center of the
range of A. punctifera is Puerto Rico. Although present in W Puerto Rico and E His-
paniola, it has not been noted in the remarkably rich butterfly fauna of Mona Island,
almost equidistant from the two documented localities on either side of the Mona Passage.
On this small island (23 mi2/62 km2), largely covered with xeric forest, its congener A.
drusilla boydi is present in a very restricted area (Smith, D. S. et al. 1988, Bull. Allyn
Museum, No. 121, 1-35).

The larval food plant(s) and immature stages of A. punctifera are unknown. It is hoped

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Journal of the Lepidopterists’ Society

that this note will stimulate further documentation of the range, significance of female
dimorphism, and developmental biology of this species. Awareness of its possible presence
is important: except for the serendipitous recognition of a minute cell-end spot on a worn
male Appias we might well have overlooked a substantial population of punctifera.

David Spencer Smith, Hope Entomological Collections and Department of Zoology,
Oxford University, Oxford 0X1 3PW, England.

Eric W. Classey, Oxleaze Farm, Nr. Uffington, Oxon., SN7 7DR, England.

Stuart J. Ramos, Department of Biology, University of Puerto Rico, Mayagiiez, Puerto
Rico 00708, U.S.A.

Received for publication 27 June 1988; accepted 15 August 1989.

Journal of the Lepidopterists’ Society
43(4), 1989, 337-338

BOOK REVIEWS

Florida Butterflies, by Eugene J. Gerberg and Ross H. Arnett, Jr. 1989. Natural
Science Publications, Inc., Baltimore, Maryland, v + 90 pp., 105 color figs. Soft cover,
14 x 21.5 cm, ISBN 0-89140-031-1, $9.95.

Florida’s Butterflies and Other Insects, by Peter D. Stiling. 1989. Pineapple Press,
Sarasota, Florida. 95 pp., 155 color figs. Hard cover, 22 x 28.5 cm, ISBN 0-910923-54-
X, $24.95.

Florida has always occupied a special place in the minds of most North American
lepidopterists. Its subtropical southern counties and habitats, especially the Florida Keys,
serve as a breeding ground and landing place for exotic butterflies and moths not generally
encountered elsewhere in the eastern United States. The resulting attractiveness to trav-
elling collectors every year is almost paradoxical when one realizes that Florida has only
164 butterfly and skipper species, less than many central and most western states, and
approximately half the fauna of Texas, which has extensive subtropical areas adjacent to
the rich Mexican faunal regions.

The reasons for this exotic but relatively depauperate fauna are rooted in Florida’s
biogeographic history and in the current floral and climatic differences between the
northern and southern halves of the state. Most of Florida’s tropical species have, in fact,
arrived over water from the Caribbean islands and not (as in the case of Texas) via a rich
continental connection directly from Mexico. And the northern temperate species that
reach Florida are at the southern limits of their ranges when they hit subtropical conditions
and their northern plant hosts disappear.

But perhaps even more surprising is that this unique and attractive state of Florida has
received little attention in terms of books describing its lepidopteran fauna. The collector
has had to use Klots (1951, A field guide to the butterflies of North America, east of the
Great Plains, Houghton Mifflin Co., Boston, 349 pp.) or the poorly-illustrated monograph
by C. P. Kimball (1965, Lepidoptera of Florida. Arthropods of Florida and neighboring
land areas, Vol. 1, Division of Plant Industry, Florida Department of Agriculture, Gaines-
ville, 363 pp.) to identify his catch and learn a little about Florida butterflies and their
ecology. Now in quick succession, two new popular books have appeared in 1989 to
partially remedy the situation for the casual naturalist and lepidopterist.

Florida Butterflies by Eugene Gerberg and Ross Arnett provides a color illustration
and a half-page or so of text on each of 97 butterfly species found in Florida. The
illustrations are 6.5 cm square, two at the top of each page, and show normally only the
dorsal surface, occasionally both dorsal and ventral surfaces, of a pinned specimen on
black background. The brief species account includes forewing size (helpful, as the color
illustrations are not reproduced to relative scale) and a brief color description, and a little,
rather generalized information on known hosts, habitat, flight period, and distribution.
The skippers (species 98-164, representing an amazingly large proportion of the state’s
fauna) are given cursory telegraphic attention (e.g., “All of Florida. Larvae on various
grasses. ”) and only 8 skipper species are illustrated. A checklist of Florida butterflies (with
common name as well as scientific name) and a complete index to the butterfly names
are included. At the end of the text are sections on rearing butterflies and making a
collection. The introductory part of the book includes short sections on butterfly biology,
butterfly gardening, the butterfly families, and butterfly conservation. This attempt to
summarize the lives of butterflies is laudable, but, unfortunately, contains several mis-
leading statements, ranging from pointless (e.g., “their lives are free”) to erroneous (e.g.,
“the simple brain is without memory”). The brief descriptions of ten Florida life zones,
with five color illustrations, provide the first-time visitor with an idea of what to expect
and where most butterfly species may be found. In addition to the 164 breeding species,
25 stray species found in Florida are listed (but not described or illustrated). While some
glaring errors will be noted (e.g., butterfly eggs do not have an average size of “2-4 mm

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Journal of the Lepidopterists’ Society

in diameter,’’ most are less than 1 mm, and eggs of only a few neotropical satyrids and
Australasian Ornithoptera reach 2-5 mm in diameter), this pocket-sized field guide
provides a concise, color-illustrated guide to most of the butterflies that the visitor or
beginning student will encounter in Florida.

Florida’s Butterflies and Other Insects, by Peter D. Stiling, opts to present a cross-
section of Florida’s butterfly and skipper fauna with color photographs of “wild” specimens
(the great majority are posed dead or pinched specimens). Forty butterflies are illustrated;
each has an accompanying paragraph of text on that species. More information on Florida
food plants is usually included here than in Gerberg and Arnett’s book, and reference to
the original literature is often made following behavioral and ecological notations. In
addition to the butterflies, 22 photographs and species accounts of Florida moths (8
Saturniidae, 10 Sphingidae, 2 Arctiidae, and 2 Noctuidae) are included. Finally for the
lepidopterist, an interesting section of 17 photographs and species accounts covers butterfly
and moth caterpillars. The remainder of the book’s main text (pp. 50-84) covers other
insect groups. The author provides a valuable page on butterfly gardening, and three
pages listing nectar sources and larval food plants used in Florida. A brief section on
commercial butterfly gardens and state insect collections completes the text. An excellent
index to insects and plants, along with an extensive literature cited section, are included.
This book, published in 8V2 x 11" size, with a full-color hardbound cover depicting a live
Eumaeus atala, is not intended for field identification but is an attractive book for the
home library.

In summary, the intricacies of Florida butterfly biology, distribution, and ecology are
not covered by either of these two new books. For a thorough treatment of Florida’s
butterflies and skippers, we must await the publication of other, more-detailed books
currently being prepared by H. David Baggett, Marc C. Minno, John B. Heppner, and
others. However, the Gerberg and Arnett field guide will serve as a good introduction to
Florida’s fauna, and the Stiling book will interest the general beginner in other Florida
insects as well as in some of the state’s fascinating tropical Lepidoptera.

Thomas C. Emmel, Division of Lepidoptera Research, Department of Zoology, Uni-
versity of Florida, Gainesville, Florida 32611.

Journal of the Lepidopterists ’ Society
43(4), 1989, 338-339

Lepidopteran Anatomy, by John L. Eaton. 1988. Wiley-Interscience Series in Insect
Morphology, John Wiley & Sons, New York. 257 pp., 194 text figs. Hard cover, 17 x 24
cm, ISBN 0-471-05862-9, $49.95.

Here, finally, is a book dedicated solely to the somewhat disjointed subject of lepidop-
teran anatomy. The author has amassed a general collection of anatomical information
on the tobacco hornworm moth, Manduca sexta; information previously difficult to locate
amidst the scatterings of textbook chapters and original scientific articles, some of which
are difficult to obtain or are not available in English. Considering the growing number
of scientists using lepidopterans as experimental models, in addition to the multitude of
professional and amateur lepidopterists, there is potentially a large audience that would
appreciate a single source on lepidopteran anatomy.

The book is divided into eleven chapters. Each of the first four provides a general
description of one of the developmental stages. The chapters on the egg and pupa are
scant, consisting of fewer than three pages each, including drawings, while the two
chapters on the larva and adult are much more detailed, treating the complex subjects
of the exoskeleton and musculature. Each of chapters five through ten focuses on a
particular system: the nervous system and endocrine glands; the circulatory system; the

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339

respiratory system; the alimentary canal, salivary glands, and excretory system; the in-
ternal organs of reproduction; and the exocrine glands. Of these five, the chapter on the
nervous system and endocrine glands and the chapter on the adult exoskeleton and muscles
receive most attention, possibly because J. L. Eaton has published some worthy articles
on these topics himself. The last chapter of the book very briefly describes the organs of
sound production and hearing. As this is my own area of specialty, I must admit that I
find the author’s treatment of this subject to be somewhat cursory and outdated. Whether
this reflects the accuracy of other sections of the book, which I am unable to review with
such a critical eye, I cannot say.

The book concludes with two appendices and an index. Appendix 1 briefly describes
Eaton’s methods. I feel that this section would be more useful to the general reader if
the author had included formulae for the solutions used in dissections and an explanation
of the derivations of the nomenclature employed in the drawings. Appendix 2 defines,
in alphabetical order, the mnemonic abbreviations used in the book. The index contains
surprisingly few entries for a book on anatomy and includes only a fraction of the structures
treated in the text.

The book is exclusively devoted to descriptive anatomy. There are plenty of drawings
(although some were surprisingly simplistic) to guide the reader through each system. I
used the book as a dissection guide to the adult musculature and nervous system and felt
satisfied, as the book did not force me to consult other references to locate and identify
any particular structure. The book does not, however, go much beyond strict anatomical
description, and thus will disappoint those who are seeking a book that provides infor-
mation on the functional properties of a particular structure (as some morphology books
will do).

Personally, my main disappointment in this book stems from the expectations I had
after reading the title, Lepidopteran Anatomy. Since the Order Lepidoptera is extremely
rich in behavioral and morphological diversity, I expected a book that approached the
topic from a comparative standpoint. The author, in a sense, acknowledges this aspect of
his book in the preface, saying that he used Manduca sexta as a model for all Lepidoptera
and that “users of this book should be readily able to apply it to the anatomy of other
lepidopterans.” I agree with this only partially. For instance, there is considerable variation
in larval form, color, and ornamentation; in egg morphology; and in adult antennae and
feeding parts, which unfortunately receive no (or very little) mention. Perhaps the title
“ Anatomy of Manduca sexta ” would have been more appropriate. However, readers
interested in comparative morphology are not left entirely empty handed. At the end of
each chapter is a list of ‘selected references’ that covers at least some of the key references
to the anatomy of other lepidopteran species.

Despite these few shortcomings, the book is, in general, a good one, and, to my
knowledge, the only one of its kind. I recommend it as a useful addition to university
libraries and to the personal collections of lepidopterists or other scientists who have an
interest in morphology.

Jayne E. Yack, Department of Zoology, Erindale College, University of Toronto,
Mississauga, Ontario L5L 1C6, Canada.

Journal of the Lepidopterists’ Society
43(4), 1989, 340-343

OBITUARY

Gordon Burgess Small, Jr. (1934-1989)

Sometime during the evening of 22-23 January 1989, Gordon Small, a Life Member
of the Lepidopterists’ Society, passed away in his sleep in his apartment in San Francisco
where he had been residing for the past two years. His sudden death at a relatively young
age has left many of us in a state of shock. Indeed, the Society has lost one of its most
ardent collectors and accomplished lepidopterists.

Gordon Small in Panama, 1970

Gordon was born in Cambridge, Massachusetts, on 19 April 1934. He is survived by
two younger brothers, Chip and Peter. After graduation from Glenridge High School in
Cambridge in 1951, he entered Bowdoin College in Brunswick, Maine, graduating with
his B.A. in Mathematics in 1955. He obtained his Masters Degree in Mathematics at
Brown University in Providence, Rhode Island, in 1956 and began work toward a Ph.D.
He had completed most of the requirements for his doctorate except the dissertation
when he accepted a position with the Panama Canal Company as an instructor in their
Department of Education in the Panama Canal Zone in 1962.

Gordon’s interest in insects and natural history in general began at the age of 10. His
brother recalls an early photograph in which Gordon is chasing a bird in order to put
salt on its tail. His oldest spread butterfly specimen is a Compton Tortoiseshell taken in
Glen Ridge in 1950. Like many of us, he dreamed for years of collecting in the tropics.

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341

And suddenly, as if in answer to his prayers, the offer from the Panama Canal Company
provided an opportunity not only to collect in the tropics, but to live there and get paid
for it! It is not surprising then, that when the opportunity arose, Gordon took advantage
of it.

Gordon and I first corresponded in 1960. He wanted to know the exact whereabouts
of Problema bulenta (Hesperiidae), reported to be in or near Wilmington, North Carolina,
and knew through correspondence with others that I had collected it there. We eventually
met, collected a few times together prior to his departure for Panama, and began a
correspondence that continued for 28 years. We briefly discussed getting together for
some tropical collecting after he became settled in the Canal Zone. Both of us were very
interested in the Lycaenidae, particularly the genus Thecla, so it was exciting to think
of my getting some military leave to join him in Panama. Thus, after Gordon had spent
some 6 months in Panama, our day-dream became reality when we made our first
collecting trip together in February of 1963 — a memorable event indeed, and one that
was to be repeated time and time again for the next 20 years.

Collecting with Gordon was an excursion fortified with the highest expectation, punc-
tuated with surprises at every turn and with the never-to-be-forgotten exhilaration of a
new “find.” In those early years, when much of the territory was still unexplored, each
trip produced a number of species not taken previously. Gordon would spend weekends
searching out new places to collect, write long letters detailing his catch, and plan strategy
for our next joint venture. Our trips were taken in his beloved tomato-red VW Beetle.
In retrospect it proved to be one of the best vehicles for the purpose; few of the roads
we traveled in those early years were impassable, and the car’s light weight and maneu-
verability proved invaluable. In the tropics, passable roads do not necessarily mean good
roads; the Pan-American Highway between the Canal Zone and Chiriqui Province was
incredibly rough — passable, but just barely. But, bad road or not, we made the trip many,
many times.

The 1960’s were probably the most productive for Gordon because there was a seem-
ingly endless array of new localities to collect, each producing species that had not been
taken on previous trips or in other localities. It was during this period that he decided to
concentrate only on the butterfly fauna of Panama and to abandon the study of species
from other localities in the Neotropics. His original intent had been to concentrate on
the Lycaenidae and Riodinidae of the entire Neotropics, trading material with collectors
in other countries. But as he became more familiar and fascinated with all the Panamanian
fauna, it became clear that a choice between the two goals had to be made. His decision
resulted in the finest collection of butterflies from a single Neotropical country that has
ever been made. It is doubtful that such a feat will be duplicated.

Gordon’s initial assignment in the Canal Zone was as a mathematics teacher in the
Balboa High School. Eventually, he moved to the Balboa Junior College. With the ex-
ception of his teaching position, which, over time, he was able to arrange to better
accommodate a good field collecting schedule, there were no other responsibilities or
demands on his time. Gordon was not married. It is safe to say that, in reality, he devoted
most of his free time (and there was lots of it) to his collection. He was a wonderfully
meticulous technician, which is apparent to anyone who views his collection of superbly
spread and curated specimens. He often voiced regret over the lack of opportunity to
attend concerts or similar artistic endeavors. Panama is not a part of the world where the
arts flourish. To fill this void he purchased a beautiful baby grand piano that took up a
large section of his living room. He was an excellent pianist with a passion for the music
of J. S. Bach and he followed a rigorous practice schedule.

Gordon’s address in the Canal Zone became the focus of attention for collectors from
both North and South America. His unique location provided an ideal stopover for visitors
going north or south and he received a great many written requests for butterflies from
the Canal Zone. For more than 20 years he provided logistical support, transportation,
and a place to stay for a steady stream of visitors. Both professional and casual lepidopterists
sought information, help in collecting, or just a visit with this kind, knowledgeable young
man who lived where he could collect tropical butterflies virtually at his back door.

The decade of the 1970’s brought continuing challenges and new interest in families

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Journal of the Lepidopterists’ Society

other than his beloved Riodinids and Lycaenids. He became fascinated with the Ithomiinae
and other Nymphalidae. By the end of the decade, his collection of Panamanian butterflies
was unsurpassed. After it became increasingly difficult for him to find species new to his
collection, he purchased a four-wheel-drive vehicle to reach areas that were inaccessible
to his VW, which by this time was showing signs of distress and hard use. The new vehicle
provided Gordon with the opportunity to venture into remote areas recently penetrated
by new roads, as yet uncompleted and not yet open to normal traffic, the road into the
Darien being the most desirable. His summer vacations were now being spent in Costa
Rica and the areas adjacent to it in Chiriqui Province, collecting in the very high country
of the Volcan de Chiriqui.

By early 1980, Gordon was already beginning to see the end of life in Panama as he
had known it for nearly 20 years; he began to make plans to collect the final “unreachable”
collecting area in Panama — the Darien. In the first week of July 1981, Gordon hired a
small plane at considerable expense to take him, with some necessary supplies (rice, sugar,
salt, etc.) plus collecting equipment, to a small mining camp at the foot of a mountain
range far into the Darien Peninsula. Using a small building at the mine as a base of
operations, Gordon climbed some 1200 meters to the summit of a small mountain, where,
over a period of time, he cleared the top with a machete, creating his own “hilltop.”

His original intent was to stay only a few weeks, but collecting was so good he extended
his stay, spending 7 weeks in virtual isolation, climbing the 1200 meters to his mountain
top each day that weather conditions permitted. Due to the confusion following President
Torrijo’s death in August, the airplane that Gordon expected to take him back to civili-
zation never arrived. Accompanied by his young Indian guide, he had to carry out all
his gear and specimens on foot, walking over 40 kilometers through hilly jungle terrain
and swamp just to reach the bare edges of human habitation. In spite of the difficulties,
hardship, and personal expense, Gordon made four more collecting trips to his abandoned
mining camp during the next two years. The fourth trip was to be his last serious collecting
effort in Panama.

It is difficult to point out a single event that could be called more memorable than
many others, for so much of his collecting was into new and remote areas and each could
be called a memorable event. But the one incident that stays in my mind above all others
was Gordon’s encounter with a very large bushmaster snake in July 1969. The snake
struck him just above the knee; luckily he was near his car and, after identifying the
snake and making a brief effort to kill it, he drove as rapidly as possible to Gorgas Hospital,
which, fortunately, was only 30 minutes away. After intensive treatment, he was released
within a week. The experience never quelled his enthusiastic and determined efforts to
collect every remote area that he could reach. The Darien experience provides ample
testimony to his indomitable courage and dedication, for an encounter with a bushmaster
there would probably have been fatal.

It was during his work in the Darien that Gordon became interested in the study of
butterfly life histories. Encouraged by Annette Aiello of the Smithsonian Tropical Research
Institute, Gordon devoted most of his time and effort during his last three years of residence
in the Republic of Panama to the rearing of butterflies, especially the genus Anaea
(Nymphalidae). He reared about 150 species in all, mostly Nymphalidae, Pieridae, and
Papilionidae, but also Hesperiidae, Lycaenidae, and Riodinidae. His detailed rearing
notes, preserved immatures, and reared adults were added to his superb collection of over
50,000 specimens, all of which were sent to the Smithsonian Institution (Robbins, R. K.
& J. F. Gates Clarke 1986, J. Lepid. Soc. 40:106) just before he retired and moved his
residence from the Canal Zone to Panama City. Gordon moved from Panama to the
United States in 1986, spending most of that year on the east coast near his old home
while working on the Smithsonian butterfly collection. In the fall of 1986 he moved to
San Francisco, something he had dreamed about for many years. Here, surrounded by
the artistic and music community he had missed so much in Panama, and, enjoying the
mild climate, he lived the few remaining years of his life.

Gordon Small has left a rich legacy — a unique butterfly collection of over 50,000
specimens, extensive notes accompanying his reared material, volumes of personal cor-
respondence, and the meaningful way he touched the lives of all who knew him. These

Volume 43, Number 4

343

will remain forever a tribute to this intelligent, generous, wonderfully kind, and gentle
man.

Bibliography

Gordon was most reluctant to publish or become involved with it. He felt that he was
not well enough trained technically and that he could contribute more by concentrating
on his field work. A list of publications follows:

1. 1962. Notes on Eurystrymon Ontario Ontario and Satyrium caryaevorus (Lycaeni-
dae). J. Lepid. Soc. 16:195-196.

2. 1969. A new subspecies of Pyrrhopyge creon (Hesperiidae) from Panama. J. Lepid.
Soc. 23:127-130 (with S. S. Nicolay).

3. 1981. Illustrations and descriptions of some species of Pyrrhopyginae from Costa
Rica, Panama and Colombia (Hesperiidae). J. Res. Lepid. 19:230-239 (with S. S. Nicolay).

4. 1981. Wind dispersal of Panamanian hairstreak butterflies (Lepidoptera: Lycaeni-
dae) and its evolutionary significance. Biotropica 13:308-315 (with R. K. Robbins).

5. In press. Catalog of the Nymphalinae (Lepidoptera: Nymphalidae) of Panama. In
Quintero D., and A. Aiello (eds.), Insects of Panama and Mesoamerica: Selected studies.
Oxford Univ. Press, London (with Gerardo Lamas).

S. S. Nicolay, 1500 Wakefield Drive, Virginia Beach, Virginia 23455-4529.

Journal of the Lepidopterists’ Society
43(4), 1989, 343-345

I Remember Gordon

I remember a bus ride in 1976 bound from San Jose, Costa Rica, to Panama to meet
a man that several people had told me knew a lot about butterflies. Perhaps he could
help me identify the Costa Rican butterflies I had been working on was how my thoughts
ran at the time. Feeling as one can only feel after 28 hours on a bus, beyond weariness,
I eventually found myself in front of one of the Canal Zone houses, rapped on a door,
and found the face of Gordon B. Small smiling down the stairs at me.

I remember being filled with wonder when walking into his apartment for the first
time. Picture, if you will, a room furnished on one end with two chairs, a sofa with a
long, low table between, and two lamps. Only two other things were in that room, both
of which were of great importance to Gordon: long rows of insect cabinets and boxes
filled to capacity with what I have come to know as the finest collection of Central
American butterflies ever made, and a baby grand piano. The chair, lamps, and table
were obviously there to make looking at butterflies easier; the piano had its own seat and
lamp.

I remember that within a matter of hours Gordon and his collection profoundly changed
my life. Here was someone who knew, absolutely, the Central American butterfly fauna.
Over the course of a week he taught me not only the names of the species, but where
and how they flew, the problems with the revisions done by “experts,” and gave me
snippets of wisdom I still am puzzling over today. Gordon’s initial gift to me was that he
showed an unflagging interest in my meager observations on butterflies, and encouraged
me to keep working on this lunatic idea I had of doing a treatment of the Costa Rican
butterfly fauna. No matter that he was clearly the consummate master, and I the complete
novice.

I remember Gordon playing the piano. His sense of ordering complex patterns found
affinity within the music of J. S. Bach; for him, other composers apparently did not come
up to snuff. After playing some of “The Inventions,” Gordon made self-deprecating
remarks about his ability. When pressed, he reluctantly admitted his one-time desire to

344

Journal of the Lepidopterists’ Society

be a concert pianist, but felt he lacked the discipline to make it. I remember standing
next to that piano transfixed by what was clearly a magnificent performance — me a
hardened jazz fanatic. The man was a gifted player.

After a week of studying his collection in Panama, Gordon offered to drive me back
to Costa Rica where we could do some collecting enroute (and I could happily omit the
ordeal of another bus ride). I was struck by two things on that trip: the man’s knowledge
of butterflies in the field (he knew everything), and his patience with my repeated, naive
questions about how to distinguish the species of satyrines, charaxines, and ithomiines.
On the drive back to Costa Rica, I remember his wild-eyed excitement if I unwittingly
stumbled upon something interesting. After completely exhausting my recollection of
what the insect was doing, Gordon then worked relentlessly until he had found the species
and collected it. As comes natural to any great teacher, Gordon kindled a fanatical interest
in me by his example.

Some days later on a mud track through the forest in Costa Rica, I remember being
introduced to the quintessential Gordon Small. In the midst of conversation he suddenly
exclaimed, “Hey, there’s Napeogenes hemistictaV’ , jumped out of the jeep, opened the
back, attached several poles to his net, and caught a butterfly flying at a subcanopy level.
He was right, of course. It was hemisticta, and no matter that neither one of us had ever
been to this locality or seen the butterfly alive before, let alone pick it out of the other
mimetic species flying there at the time. This type of feat was commonplace with Gordon
and similar scenes were to be repeated virtually every time I was in the field with him.
Over the years Gordon would periodically visit me in Costa Rica to check on what I had
turned up and to talk butterflies, and we’d go off collecting together. Somehow he always
found new things, even at localities I had been pounding away at for years. Quintessential
Gordon Small.

When I left permanent residence in Costa Rica in 1980 to go to graduate school, Gordon
and I kept up a steady flow of letters concerned with all aspects of butterflies. My education
on butterflies continued through these letters. While I was at the British Museum writing
the text for my Costa Rican butterfly book Gordon was one of my most constant correspon-
dents. In letters he would query about the types of this or that species, inform me about
his work in the Darien and to proudly tell me that he had finally begun to rear caterpillars
and that he could see their importance to systematics. He provided a sounding board for
my many questions about certain species, and he often made sound suggestions about
the names of certain troublesome species, even though he had never seen the types. Even
from afar, sight unseen, Gordon was usually correct about the identity of the specimens
that sat in front of me. In fact, I remember Gordon being correct about most things
concerned with butterflies throughout the twelve years of our friendship. He had a gift
for this type of thing.

Gordon was a fanatically meticulous person who never relaxed his critical standards
for anyone or anything when it involved butterflies. I remember the last time I saw
Gordon. Standing around between sessions at the Lepidopterists’ Society meetings at
University of California, Berkeley, I saw him enter the building and make his way over
to me. Delighted to see him and keen to hear his response to my then just published book
(1987, The butterflies of Costa Rica and their natural history, Princeton University Press,
Princeton, New Jersey, 327 pp.) I immediately introduced him to someone near me as
the person who taught me everything I knew about butterflies. Gordon was cheerful and
shook off the compliment, but I knew him well enough to know something was up. When
I asked what was wrong he said, “Phil, you misrepresented me in your book.” Feeling
the bottom drop out of my stomach, I asked him falteringly, “I’m terribly sorry. How?”
Gordon stated in dead earnestness, “In your species account of Haematera pyramus. I
collected the species by clapping my net down over individuals visiting wet sand. Those
individuals never flew. The person who was collecting with me chased his down, and the
observation that they flew like Diaethria was his, not mine.”

I remember Gordon as a consummate butterfly biologist. Quite simply, Gordon B.
Small, Jr. knew more about butterflies than any person I have ever met, corresponded
with, or have heard about. Many of the places where he worked have vanished into the
oblivion of human “progress.” All that remains of those Panamanian habitats are the

Volume 43, Number 4

345

mute testimonies to tropical butterfly diversity now found in his collection, currently
housed at the Smithsonian Institution. We and future generations owe him a profound
debt of gratitude for singlehandedly learning the butterfly fauna of an entire country
and for making such a thorough and masterful collection. Death has silenced a truly great
naturalist, teacher, and extraordinary human being. After losing such a friend and source
of inspiration, some of us may feel left behind. But, as in so many things, I remember
that Gordon always was way ahead of the rest of us.

P. J. DeVries, Department of Zoology, University of Texas, Austin, Texas 78712.

Journal of the Lepidopterists’ Society
43(4), 1989, 346-350

INDEX TO VOLUME 43

(New names in boldface)

abundance, seasonal, 157
Adamski, D., 108
admiral butterfly, 102
aging, 244

alkaloids, quinolizidine, 261
amaretta, Godyris zavaleta, 258
amaturaria, Calothysanis, 72
amaturaria, Timandra , 72
American Museum of Natural History, 244
anise swallowtail, 217
annahella, Vanessa, 81
Anoncia
crossi , 108
diveni, 108

Antheraea mylitta, 152
aposematism, 261
appeasement behavior, 33
Appias

drusilla, 333
punctifera, 333
Argynninae, 1
aristata, Gaillardia, 210
Arizona, 79
arvense, Cirsium, 331
Asteraceae, 305, 331
Atalopedes, 11
clarkei , 11

A Taxonomic Revision of the New World
Moth Genus Pero . . . (book re-
view), 76

aurantiaca, Euselasia, 247
Australia, 305
Ayres, M. P., 93

Baccharis halimifolia, 305
Baja California, 184
Ballmer, G. R., 59
Batis maritima, 332
Baughman, J. F., 244
behavior, 157
appeasement, 33
feeding, 102, 167, 258, 331
mating, 68, 93
oviposition, 332
resource defense, 102
territorial, 33, 102
Benson, W. W„ 33
Bernays, E. A., 261
biogeography, 81, 120, 274, 333
biological control, 305
Boloria, 1

Bowman, D. E., 328
Brazil, 33, 323

Brephidium isophthalma pseudofea, 332

briseis, Catocala, 331
Brower, L. P. 50
Brown, J. W„ 184, 313
bud worm, spruce, 167
Burns, J. M., 11
Butler, L., 299

Butterflies of Nepal . . . (book review), 255
Byers, B. A., 210

California, 50, 59, 217, 229, 261
Calvert, W. H„ 50
Canada, 148, 251, 331
Caribbean, 274
carpenter moth, 327
carye, Vanessa, 81
Cashatt, E. D., 75
Catocala briseis, 331

Catalogue of Lycaenidae <6- Riodinidae . . .
(book review), 250

Chile, 71

chileana , Chlorostrymon, 120
Chlidanotini, 313
Chlorostrymon, revision, 120

chileana , 120
patagonia , 120

Choristoneura fumiferana, 167
Cirsium arvense, 331
Clarke, J. F. G„ 147
clarkei , Atalopedes, 11
Classey, E. W., 333
clenchi, Morpheis, 327
coenia, Precis, 289
Coenonympha tullia yontocket, 229
coffeae, Saissetia, 167
color illustrations in the Journal (announce-
ment), 259
Colorado, 102, 328
conservation, 178
copulation, 68, 79, 93
Cosmopterigidae, 108
Cosmopteriginae, 108
Cossidae, 327
Coveil, C. V., Jr., 72, 73
Crambinae, 323
crossi , Anoncia, 108
Cytisus monspessulanus, 261

Danaus plexippus, 50
Dash, A. K„ 152
De Benedicts, J. A., 239
DeVries, P. J., 343
diana, Hemileuca, 328
diapause, 50, 59, 157
digyna, Oxyria, 59

Volume 43, Number 4

347

dispersal, 274
dissection, 239

distributions, geographic, 184, 274, 327
diveni, Anoncia, 108
diversity, world, 178
dominance, territorial, 33
Dominican Republic, 333
Donahue, J. P., 184, 327
Drummond, B. A., Ill, 258
drusilla, Appias, 333

ecotypes, 217
Ecuador, 155, 258
editha, Euphydryas, 244
egg incubation period, 152
electrophoresis, 81, 217, 229
Emmel, T. C., 251, 255, 337
endangered, 114, 147
Epstein, M. E., 252
error aria, Glaucina, 79
Eucosmini, 313
Eueides tales, 33
Euliini, 313
Eumaeini, 120
Euphydryas editha, 244
Eupithecia
taracapa, 71
tarapaca, 71
Euselasia

aurantiaca, 247
mystica, 247
Euselasiinae, 247
evolution, 11

families, 178
faunal realms, 178
fecundity, 167, 305
feeding

effect on reproduction, 167
on sap, 102
stimulants, 261
Ferguson, D. C., 79
Florida, 289, 332

Florida Butterflies (book review), 337
Florida’s Butterflies and Other Insects
(book review), 337

food plants, 147, 157, 184, 210, 261, 299,
305, 332

francisci , Neonympha mitchellii, 114
Freeman, H. A., 244
Fujii, H., 68

fumiferana, Choristoneura, 167
functional anatomy, 239

Gaillardia aristata, 210
Gaskin, D. E., 323
Geiger, H., 81

Gelechioidea, 108
genera, 178, 313
genetic determination, 148
genetic differentiation, 81, 217
genitalia, 11, 239
genitalic musculature, 239
geographic variation, 148
Geometridae, 71, 72, 79, 305
Glaucina err or aria, 79
Godyris zavaleta amaretta, 258
Gonionota, 155

Great American Interchange, 81
Greater Antilles, 333
Grey, L. P„ 1
grotei, Hemileuca, 328
Guild Handbook of Scientific Illustration,
The (book review), 253
Guppy, C. S., 148

habitat, 114
Haddad, C. F. B„ 33
Hadeninae, 299
Hades noctula, 247
hairpencil, 313
hairstreak, 68, 120
halimifolia, Baccharis, 305
handling, effects of, 244
hardwood defoliator, 299
Harvey, D. J., 247, 250, 332
Heliconiini, 33
Heliconius
leucadia, 33
sara, 33

Heliothinae, 210
Hemileuca
diana, 328
grotei, 328
Herman, W. S., 50
Hesperia, 11
nabokovi, 11
Hesperiidae, 11, 244
Hogue, C. L., 253
holotypes, new location of, 244
honey dew, 167

host plants, 184, 210, 261, 299, 305
hypophlaeas, Lycaena phlaeas, 59

Immature Insects, Volume 1 (book re-
view), 77

immature stages, 210, 299

India, 152

instar number, 59

island biogeography, 274, 333

isophthalma pseudofea, Brephidium, 332

Itame varadaria, 305

Ithomiinae, 258

348

Journal of the Lepidopterists’ Society

janasi, Shirozua, 68
Japan, 68

Johnson, K., 66, 120

Krai, T. W„ 114

Landry, J. F., 74
larval development, 59
latex, Polia, 299
Lederhouse, R. C., 93
Lenczewski, B., 289

Lepidopteran Anatomy (book review), 338

lepidopterists, role of, 157

leucadia, Heliconius, 33

life histories, 210, 299, 305

Limacodidae, 252

Lindera melissifolia, 147

Limenitis weidemeyerii, 102

longevity, 157

Longino, J., 332

lutea , Neoeromene, 323

Lycaena phlaeas hypophlaeas, 59

Lycaenidae, 59, 68, 120, 250, 332

Lycaeninae, 59

Manitoba, 331

mark-release-recapture, 244

maritima, Batis, 332

masoni, Schinia, 210

mate guarding, 68

mate location, 33

mating, interfamily, 79

mating systems, 93

Mattoon, S. O., 229

Matusik, D., 66

melanism, 148

melanopis, Protorthodes, 79

melissifolia, Lindera, 147

Mexico, 50, 108, 184, 244, 327

migration, 289

Miller, W. E., 77, 167

mitchellii francisci, Neonympha, 114

Monroe, E., 274

monspessulanus, Cytisus, 261

Montllor, C. B., 261

Morpheis clenchi, 327

morphology, larval, 247

Morris, M. W„ 147

morrisi, Papilio, 66

Moths of America North of Mexico . . . ,
The (book reviews),

Fascicle 18.1 Geometroidea, Geometri-
dae (Part), 73

Fascicle 7.1 Gelechioidea, Gelechiidae
(Part), Dichomeridinae, 74

Fascicle 15.2 Pyralidae (Part), Phyciti-

nae (Part — Acrobasis and Al-
lies), 75

multiple mating probability, 93
musculature, genitalic, 239
mylitta, Antheraea, 152
myrmecophily, 332
mystica, Euselasia, 247

nabokovi, Hesperia, 11
Nayak, B. K., 152
Neoeromene
lutea , 323
straminella, 323

Neonympha mitchellii francisci, 114

Neotropics, 66, 108, 120, 313

Nepal, 255

Nicaragua, 274

Nicolay, S. S., 340

Noctuidae, 79, 210, 299, 331

noctula, Hades, 247

North America, 1, 305

North Carolina, 114

Nymphalidae, 1, 33, 50, 81, 102, 229, 244,
258

Oecophoridae, 155
Olethreutinae, 313
orange hairstreak, 68
organs, perforated cupola, 247
Orive, M. E., 244
overwintering, 50
oviposition behavior, 332
Oxyria digyna, 59

Palmer, W. A., 305
Papilio
glaucus, 93
morrisi, 66
stewarti, 66
troilus, 147
zelicaon, 217

Papilionidae, 66, 93, 147, 148, 217
Papillons et Chenilles du Quebec . . . (book
review), 251
parasites, 299
Parnassius phoebus, 148
Parshall, D. K„ 114
Passoa, S., 155

patagonia, Chlorostrymon, 120
peninsular effect, 184
perforated cupola organs, 247
phenology, 289
phenotype, 148

phlaeas hypophlaeas, Lycaena, 59
Phoebis sennae, 289
phoebus, Parnassius, 148
phylogeny, 11

Volume 43, Number 4

349

Pieridae, 289, 333
plexippus, Danaus, 50
Polia latex, 299

populations, genetic differentiation among,
217

Porter, A. H., 229
Powell, J. A., 157, 239
Pratt, G. F., 59
Precis coenia, 289
predation, 261

Presidential Address 1988, 157

Proclossiana, 1

Protorthodes melanopis, 79

Providencia Island, 274

pseudofea, Brephidium isophthalma, 332

Puerto Rico, 333

punctifera, Appias, 333

pupa, unusual, 155

pupal stridulation, 332

Pyralidae, 261, 323

Queensland, 305
quinolizidine alkaloids, 261

Ramos, S. J., 274, 333

refrigeration, effect on egg development,
152

reproductive enhancement, 167
reproductive tract development, 50
reversalis, Uresiphita, 261
Rindge, F. H„ 71, 76, 154
ringlets, 229
Riodinidae, 247, 250
Rosenberg, R. H., 102
Rozycki, R., 66

Saissetia coffeae, 167
Salicornia virginica, 332
San Andres Island, 274
Santa Catarina, Brazil, 323
sap feeding, 102
sara, Heliconius, 33
Saturniidae, 152, 154, 328
Saturniidae of America, Ceratocampinae,
The (book review), 154
Satyridae, 114
Satyrinae, 229
Schinia

masoni, 210
volupia, 210
Scriber, J. M., 93
seasonal abundance, 157
seasonal dimorphism, 333
Semnopsyche, 1
sennae, Phoebis, 289
sericulture, 152
sessile, Tapinoma, 332

Shapiro, A. M., 81, 217
Shields, 0., 178, 255
Shirozua janasi, 68
Sierra Madre Occidental, 327
silk moth, 152
sister species, 328
size variation, 148
skippers, 244

Slug and Nettle Caterpillars . . . (book re-
view), 252

Small, G. B., Jr. (obituary), 340
Small, G. B., Jr. (remembrance), 343
Smith, D. S„ 274, 333
Smithsonian Institution, 147, 244
South-east Asia, 252

Spencer, O. D., butterfly collection, 147
spermatophore
counts, 93
size, 93

Speyeria, 1, 147
Sphingidae, 184
spruce bud worm, 167
staining, 239
Sterrhinae, 72
stewarti, Papilio, 66
Stone, S. E., 328
straminella, Neoeromene, 323
stridulation, pupal, 332
subfamilies, 178
subspeciation, 274
systematics, 81, 114, 120

tales, Eueides, 33
Tapinoma sessile, 332
taracapa, Eupithecia, 71
tarapaca, Eupithecia, 71
tasar silk insect, 152

taxonomy, 11, 66, 71, 72, 114, 120, 229,
313, 323
Taylor, P., 331
territorial behavior, 33, 102
Texas, 305

threatened species, 229

Timandra amaturaria, 72

Tong, M. L., 217

topotype, 328

Tortricidae, 167, 313

Tortricinae, 167, 313

trap, butterfly, 289

trap efficiency, 289

troilus, Papilio, 120

tullia yontocket , Coenonympha, 229

type locality, 328

ultrastructure, 247

United States National Museum, 244
Uresiphita reversalis, 261

350

Journal of the Lepidopterists’ Society

Vanessa
annabella, 81
carye, 81

varadaria, Itame, 305
variation

body size, 148
geographic, 148

West Virginia, 299
wing

melanism, 148
wear, 244
Wood, P. S., 299

world numbers of butterflies, 178

wing melanism, 148
Virgin Islands, 333

Yack, J. E., 338

yontocket , Coenonympha tullia, 229

virginica, Salicornia, 332
volupia, Schinia, 210

Walker, T. J„ 289
weidemeyerii, Limenitis, 102
West Indies, 274

zavaleta amaretta, Godyris, 258
Zeuzerinae, 327
Zikan, ML, 33
zoogeography, 11

Date of Issue (Vol. 43, No. 4): 20 December 1989

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Natural Perspectives
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CONTENTS

Aposematism of Uresiphita reversalis larvae (Pyrali-

dae). E. A. Bernays ir C. B. Montllor 261

Biogeographical affinities of the butterflies of the
Southwest Caribbean Islands, San Andres and Provi-
dencia. David Spencer Smith, Eugene Munroe ir Stuart
J. Bamos 274

An inexpensive portable trap for monitoring butterfly

migration. Thomas J. Walker & Barbara Lenczewski 289

Biology and immature stages of Polia latex (Guenee)

(Noctuidae). Peter S. Wood {? Linda Butler 299

Food plant specificity and biology of Itame varadaria
(Walker) (Geometridae), a North American moth

INTRODUCED INTO AUSTRALIA TO CONTROL THE WEED BAC-

charis halimifolia L. W. A. Palmer 305

Generic reassignments for neotropical tortricid moths

(Tortricidae). John W. Brown 313

Neoeromene lutea (Lepidoptera: Pyralidae; Crambinae), a

new species from Brazil. David E. Gaskin 323

General Notes

Discovery of the carpenter moth Morpheis clenchi (Cossidae) in Mexi-
co. Julian P. Donahue 327

A description of topotypical male Hemileuca diana (Saturniidae). Stephen

E. Stone <b Donald E. Bowman 328

Diurnal nectaring by a Catocala moth (Noctuidae). Peter Taylor 331

Myrmecophily and larval food plants of Brephidium isophthalma pseudofea

(Lycaenidae) in the Florida Keys. D. J. Harvey 6- J. Longino 332

Appias punctifera D’Almeida (Pieridae) in the Dominican Republic and Puer-
to Rico. David Spencer Smith, Eric W. Classey i? Stuart J. Ramos 333

Book Review

Florida Butterflies and Florida’s Butterflies and other Insects. Thomas C.

Emmel 337

Lepidopteran Anatomy. Jayne E. Yack 338

Obituary

Gordon Burgess Small, Jr. (1934-1989). S. S. Nicolay 340

I remember Gordon. P. J. DeVries 343

Index to Volume 43 346

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