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ISSN 0024-0966

JOURNAL

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LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

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

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5 June 1990

THE LEPIDOPTERISTS’ SOCIEFY

EXECUTIVE COUNCIL

JACQUELINE Y. MILLER, President NIELS P. KRISTENSEN, Vice
JULIAN P. DONAHUE, Immediate Past President

President DONALD J. LAFONTAINE, Vice
RICHARD HOLLAND, Vice President President
WILLIAM D. WINTER, Secretary JAMES P. TUTTLE, Treasurer

Members at large:

Jo BREWER JOHN W. BROWN RICHARD A. ARNOLD
DALE W. JENKINS MOGENS C. NIELSEN SUSAN S. BORKIN
JOHN E. RAWLINS FLOYD W. PRESTON Davip L. WAGNER

EDITORIAL BOARD

PAUL A. OPLER (Chairman), FREDERICK W. STEHR (Member at large)
Boyce A. DRUMMOND (Journal), WILLIAM E. MILLER (Memoirs), JUNE PRESTON (News)

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

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

Active members—annual dues $25.00
Student members—annual dues $15.00
Sustaining members—annual dues $35.00
Life members—single sum $500.00
Institutional subscriptions—annual $40.00

Send remittances, payable to The Lepidopterists’ Society, to: James P. Tuttle, Treasurer,
3838 Fernleigh Ave., Troy, Michigan 48083-5715, U.S.A.; and address changes to: Julian
P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, California
90007-4057 U.S.A. For information about the Society, contact: William D. Winter, Sec-
retary, 257 Common St., Dedham, Massachusetts 02026-4020, U.S.A.

For back issues of the Journal and News, write the Publications Coordinator at the
address below about availability and prices. Prices of The Lepidopterists’ Society com-
memorative volume 1945-1973 are $12.00 ($8.00 to members and subscribers); of A
catalogue/checklist of the butterflies of America north of Mexico, clothbound, $19.00
($12.00 to members and subscribers), paperbound, $10.50 ($7.00 to members and sub-
scribers). Order from the Publications Coordinator, Ronald Leuschner, 1900 John St.,
Manhattan Beach, California 90266-2608, U.S.A. Make remittance payable to “The Lep-
idopterists’ Society.”

Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for
$40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’
Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los
Angeles, California 90007-4057. Second-class postage paid at Los Angeles, California and
additional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’
Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, California 90007-
4057.

Cover illustration: Female ceanothus moth, Hyalophora euryalis (Boisduval), resting
on a branch of immature apricots shortly after emerging from her cocoon. Submitted by
John Cody, 2704 Woodrow Court, Hays, Kansas 67601.

JouRNAL OF
Tue LepiporpreristTs’ SOCIETY

Volume 44 1990 Number 1

Journal of the Lepidopterists’ Society
44(1), 1990, 1-10

PRESIDENTIAL ADDRESS, 1989: WHY CAN’T
LEPIDOPTERISTS BE MORE LIKE BOTANISTS?!

JULIAN P. DONAHUE

Natural History Museum of Los Angeles County,
_ 900 Exposition Boulevard, Los Angeles, California 90007-4057

Additional key words: _ identification manuals, faunal inventory, biodiversity, National
Lepidoptera Agenda.

When members of The Lepidopterists’ Society elected me to be their
President in 1988 I was both honored and flattered. And honor is the
appropriate term: the Society’s Secretary, Treasurer, and Editors do
most of the work and provide continuity to our ongoing operations,
while the President has only three principal functions—all of which
occur within a four-day period at the end of a year of doing not much
of any significance. One function is to chair a meeting of the Executive
Council. That’s done. The second is to pass on the symbols of office to
the next President. That’s coming up. The third function is to present
a Presidential Address. That’s right now. And it’s the toughest part of
the job, because the subject of the address should be one of general
interest to the members of the Society, rather than my research, cur-
atorial, and public service activities in which I am intimately involved
on a daily basis—and about any of which I could speak for hours with
few or no notes. The task is made easier, however, by my being able
to say what I think without having to prove it, and by not having to
subject my thoughts to the peer-review filter.

The importance of habitat preservation, for the sake of our natural
environment, and the importance of collecting now, for the sake of
advancing our knowledge, are two subjects very important to me, but

* Based on a draft composed in the snug comfort of a van camped at 8400 feet in New Mexico’s Sacramento Mountains
two days prior to the Society’s 40th Annual Meeting in Albuquerque, July 1989. A family emergency necessitated the
author's premature departure from the meeting, and the address was never delivered. The author suggests that any
errors in logic or lucidity be attributed to a temporarily oxygen-starved brain rather than to any permanent organic
disorder

2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

I chose not to address them further since several of my presidential
predecessors have already done so quite adequately.

I considered talking about the fact that human overpopulation is the
compelling and ultimate environmental issue, from which virtually all
others flow—but I realized that I would only be telling you something
you already know, particularly since our eminent fellow lepidopterist,
Paul Ehrlich, is one of the most outspoken and eloquent proponents of
this view.

On my way to this meeting I spent several days exploring the cultural
and natural riches of this magnificent state of New Mexico. Besides
learning that Deming is the home of “clean water and fast ducks,” in
the course of some very fruitful botanizing I realized that the state
flower, Yucca elata, is clearly one of the most spectactular and striking ~
plants in the American Southwest.

Ever since my college days in Michigan I’ve been fascinated by plant
identification, not only as a direct adjunct to the study of Lepidoptera
and the food they eat, but also as a tool for understanding and describing
habitat differences and, frankly, just for the fun of learning the name
of another organism I’ve met in my travels. Now, no matter where I
go, I am able to recognize a few old friends, while encountering a lot
of new plants I’ve never seen before. I still remember my surprise when
I discovered that my midwestern botanizing had left me totally un-
prepared for a woody composite, which I encountered in abundance
soon after moving to the Southwest.

My old friends, Gray's Manual of Botany and the Spring Flora of
Wisconsin, were useless in my new environment, as I found myself
having to build an entirely new botanical reference library to help me
cope with all the surprises that awaited me. And I was not disap-
pointed—plant books by the score abounded: handbooks on trees and
shrubs of southern California; trees, shrubs, and wildflowers of the Sierra
Nevada; cacti of California; wildflowers of the California desert; keys
to the flowers and shrubs of the desert; a manual of southern California
plants; a manual of plants for the entire state of California—my shelves
groaned, and sagged even further as my travels and botanical interests
took me farther afield into adjacent states and Mexico.

But the point is that I have been able to find a book to identify
virtually any plant nearly everywhere my travels have taken me—and
yet I live in a state with at least 3000 species of moths and nary a list
of what those species are, much less where and when they occur within
the state and how to identify them.

Had I stumbled upon the Great American Anomaly? Or some Griev-
ous Inequity? I finally had a subject worthy of Presidential exploration.

Was this abundance of plant manuals peculiar to California? A stroll

VOLUME 44, NUMBER 1 3

through the Museum’s botany library quickly convinced me that this
was indeed not the case. A sample of some North American titles I
found: A Flora of Tropical Florida (Long & Lahela 1976); Manual of
the Plants of Colorado (Harrington 1954); Arizona Flora (Kearney &
Peebles 1951—treating 3370 species!); Flora of Alaska (Anderson 1959);
Wild Flowers of the United States (Rickett, various dates; multi-volume
regional work heavily illustrated with color photos); Illustrated Flora
of the Pacific States (Abrams 1940-1960); Trees, Shrubs, and Woody
Vines of the Southwest (Small 1972); Gray's Manual of Botany (Fer-
nald 1950—treating 5523 species of the northeastern U.S.); Manual of
the Vascular Plants of Texas (Correll & Johnston 1970—treating nearly
5000 species!); A Utah Flora (Welsh et al. 1987— treating 2572 species);
Intermountain Flora (Cronquist et al., various dates); Aquatic and
Wetland Plants of Southwestern United States (Correll & Correll
1975); and Flora of Baja California (Wiggins 1980—treating 2705
species).

Furthermore, I found numerous more specialized manuals and mono-
graphs, including Moss Flora of North America (Grout 1972); Agaves
of Continental North America (Gentry 1982); Atlas of North American
Astragalus (Barneby 1964), and Manual of the Grasses of the United
States (Hitchcock 1951). Wow!

How does this abundance of botanical manuals compare with iden-
tification tools available for Lepidoptera? Although I shall be concen-
trating on comparisons within the North American flora and fauna,
with which I am most familiar, similar comparisons most likely can be
made everywhere else on earth with the possible exception of western
Europe.

Butterfly people have it easy. Many state and regional manuals have
been published, with many still in print (e.g., New York, Michigan,
Missouri, Georgia, Oregon, Indiana, Rocky Mountains), not to mention
several readily available continent-wide identification manuals by Scott,
Howe, Ehrlich and Ehrlich and, for the skippers, by Evans, Freeman,
and Lindsey, Bell, and Williams, PLUS a comprehensive bibliography
of all state and regional butterfly lists ever published for North America
(Field et al. 1974). Most regional manuals have been written as a labor
of love by non-professionals (that is to say, by people who aren’t paid
to be lepidopterists).

Identifying a moth in North America is another story altogether, and
I think that our scarcity of identification references is the fundamental
reason we don’t have more moth collectors. To test this view I examined
the moth handbooks in my office and found deficiencies in virtually all
of them. Here are some examples from the North American fauna for
continent-wide works:

4 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

1. Noctuidae, one of the largest moth families, with at least one widely collected genus
(Catocala):

a. Hampson (1903-1913), Catalogue of the Lepidoptera Phalaenae in the British
Museum, Vols. 4-18: treated all world species known at the time, including all
North American species; now out of date, expensive, and not fully illustrated.

b. Seitz (1923), Macrolepidoptera of the World, Vol. 7: never completed, expensive,
out of date.

c. Barnes & McDunnough (1918), Illustrations of the North American species of
the genus Catocala: out of print, hard to find, expensive, out of date.

d. Lafontaine (1987), Euxoa, Moths of America North of Mexico, Fascicle 27.2:
treats only a single (but large) genus, expensive.

2. Phycitine Pyralidae: Heinrich (1956), American Moths of the Subfamily Phycitinae:
excellent genitalic figures for virtually all species in the Western Hemisphere, but
not a single photograph of an adult moth!

3. Acrolophus: monographed by Hasbrouck (1964), male genitalia well figured, but
lacks figures of female genitalia and has no photos of adults.

4. Pterophoridae: monographed by Barnes & Lindsey (1921): surprisingly useful but

now outdated and hard to find.

Olethreutine Tortricidae: monographed by Heinrich (1923, 1926): genitalia well

illustrated but not a single figure of wing pattern, now hard to find and somewhat

outdated.

Ol

Although The Moths of America North of Mexico series is slowly
filling the void in identification manuals for the North American fauna,
and doing it in a superb fashion, it is beset with at least three lingering
problems: it is unquestionably slow (begun 18 years ago), it is expensive,
and it tends to be biased toward eastern collections.

With very few exceptions, the regional identification manuals for
North America are similarly limited:

1. Forbes (1954), Lepidoptera of New York and Neighboring States, Part III, is the
only comprehensive state faunal identification manual that contains keys, but it
suffers from a lack of photographs of adult moths.

2. Kimball (1965), Lepidoptera of Florida, is an illustrated and annotated checklist,
not an identification manual.

3. Covell (1984), Field Guide to the Moths of Eastern North America, is the best
regional moth manual we have, but it does not cover the entire fauna.

Every single one of the references cited above is diminished by one
or more limitations of being out of date, out of print, incomplete,
inadequately illustrated, or too expensive.

Before a comprehensive identification manual can be produced, you
first need to have a list of species present in the area to be covered.
Several such lists have been published, including:

1. Hodges (1983), Check List of the Lepidoptera of America North of Mexico [North
American faunal list].

2. Moore (1955), Annotated List of the Moths of Michigan [excluded the “microlepi-
doptera ’].

3. Kimball (1964), Lepidoptera of Florida.

4. Forbes (various years), Lepidoptera of New York and Neighboring States: a com-
prehensive work that embraced all species known to occur in the state, thus serving
as a state list.

VOLUME 44, NUMBER 1 5

In addition, there are a number of lists that inventory, in varying
detail, a portion of the moth fauna occurring in a given area, such as
those by Tietz (1936, The Noctuidae of Pennsylvania); Selman and
Barton (1971, The Sphingidae of Northeast Arkansas); Metzler (1980,
Saturniidae of Ohio); and Godfrey, Cashatt, and Glenn’s fascinating
1987 annotated checklist of the 30,000 “‘microlepidoptera”’ that Murray
O. Glenn, a farmer, collected as a hobby in a small section of Illinois.

The search through my library did produce, however, a very few
works that approach my concept of the ideal identification manual in
that they are current, complete, contain keys and/or diagnoses, and are
well illustrated. However, they all have one drawback: they are gov-
ernment or organizational publications unobtainable through normal
bookselling channels—you have to scrounge the publisher’s name and
address, and enter into correspondence to ascertain availability and
price. Three examples are McGuffin’s series, Guide to the Geometridae
of Canada (Entomological Society of Canada); Pogue and Lavigne’s
The Tortricinae of Wyoming (1981, University of Wyoming Agricul-
tural Experiment Station); and the unquestioned cream of the crop, to
the best of my knowledge the best regional moth identification guide
ever published in North America, William E. Miller's Guide to the
Olethreutine Moths of Midland North America (1987, U.S.D.A. Forest
Service, Agric. Handbook 660). This last publication could easily serve
as an idealized model for the scores of similar manuals we need—the
use of color photos is the only improvement it could have used.

I am amazed, or perhaps stunned is a more appropriate word, that
neither of the two states with biological or insect survey programs, each
having published major works on their insect faunas (Illinois Biological
Survey and California Insect Survey), has published a single work on
its moth fauna, not even at the family level. Texas, with one of the
richest Lepidoptera faunas in North America, has not even produced
a guide to its butterflies, not to mention its moths!

This appalling disparity—the abundance of plant identification man-
uals and the paucity of moth identification manuals—led me to inves-
tigate the cause of this discrepancy, with the hope of finding some clue
that we lepidopterists can use to our advantage.

At first, I thought that the botanists had it easier because they don’t
have to deal with the overwhelming numbers of species we moth people
face. Wrong! Published estimates for the number of known, named
species of Lepidoptera in the world range from 113,000 to 150,000 or
more (with some estimates for the actual total world fauna, both named
and unnamed, of ten times or more this number)—for the sake of this
discussion I shall use an estimate of 140,000 known species of Lepi-
doptera in the world. By comparison, there are some 225,000 species

6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

of flowering plants (dicots and monocots) in the world, or 61% more
species of plants than there are of Lepidoptera. In North America north
of Mexico, there are more than 11,000 named species of Lepidoptera,
compared with twice that number of species of plants (22,200).

Considering the fact that most Lepidoptera feed, as larvae, on only
one species of plant, or on a few closely related species of plant, and
further taking into account that most plants probably serve as host to
more than one species of lepidopteran, I think it is reasonable to expect
that, on average, there is at least one species of lepidopteran for every
species of flowering plant (disregarding, for the moment, the species
whose larvae feed on lower plants, detritus, etc.). This translates to a
potential North American Lepidoptera fauna of over 22,000, a reason-
able figure when one considers the large numbers of new species being
discovered in poorly studied families (e.g., Gelechiidae, Scythrididae).
Using these same assumptions for the world Lepidoptera fauna, we can
reasonably expect the potential world fauna to be about a quarter of
a million species, or about twice the number of species we have suc-
ceeded in describing in the last 200 years!

No, the botanists have not been so successful because their job has
been easier. The real reason that botanists know their business so well
is because of their subject matter: plants are necessary for converting
solar energy to a form that can be utilized by animals, and are thus
fundamentally essential for human existence and survival. Homo sapi-
ens has a vested and direct interest in plants, not just for food, but for
fiber, shelter, medicine, ornamentation, etc. Human survival, popula-
tion expansion, and colonization all depend on our having an intimate
knowledge of botany.

So of course we’re going to know far more about plants than about
any other component of our natural environment, including a lot of
apparently non-essential incidental botanical knowledge of no imme-
diate practical advantage acquired during the course of applied re-
search.

What, exactly, do I think is so special about botanists? The two
qualities of botany and botanists that I admire most, and which I
perceive to be deficient or lacking among lepidopterists, are (1) the
availability of an identification manual for virtually any group of plants
virtually anywhere on earth, and (2) the ability to agree on the im-
portance of floristic studies, and then to cooperate in providing the
funds and staff to accomplish the goals.

Why are botanists so much more successful in understanding their
discipline than we lepidopterists? And here I am addressing not the
practical, applied aspects of botany, but basic knowledge of natural
history, taxonomy, and distribution.

VOLUME 44, NUMBER 1 7

First, there are simply more of them than us! The Lepidopterists’
Society has just 1500 members in 50 nations, and is the only organization
in North America devoted entirely to Lepidoptera. In contrast, just look
at some of the botanical organizations in the United States: Botanical
Society of America, American Society of Plant Taxonomists, American
Fern Society [there are only 345 species of ferns in North America north
of Mexico!], Phycological Society of America (algae), Mycological So-
ciety of America (fungi, 1600 members), North American Mycological
Association, American Orchid Society, the Bromeliad Society, and
doubtless others, not to mention native plant societies, garden clubs,
and special interest groups devoted to limited subjects such as bonsai,
roses, insectivorous plants, etc.

Secondly, they are publishing fools! Every one of the organizations
I just listed has its own publication. Furthermore, numerous other pub-
lications emanate from free-standing institutions, such as the Missouri
Botanical Garden (Annals), New York Botanical Garden (a slew of
titles, including Brittonia, Botanical Review, Economic Botany, Mem-
oirs, Flora Neotropica, etc.), Field Museum of Natural History (Field-
iana, with ongoing series on Ferns & Fern Allies of Guatemala, Flora
of Peru, Flora Costaricensis, etc.), and Rancho Santa Ana Botanical
Garden (Aliso), to mention just a few.

The study of plants has certain obvious advantages over the study of
Lepidoptera, and I think this is another reason why botanists far out-
number us. Specifically, plants are a lot easier to study—although they
are often seasonal (as are leps), they stand still for prolonged scrutiny
and manipulation, and can be found in exactly the same spot the next
day; they are (usually) diurnal, conspicuous, attractive and esthetically
pleasing, and easily observable; many wild species can be adapted for
domestic cultivation. Although butterflies (and some moths) share some
of these characteristics, the fact that they don’t allow a close approach
and manipulative examination (without stalking and netting them) makes
them unworthy of study by the casual naturalist. And moths—well, you
can forget about the general public having an intellectual interest in
any moth smaller than a bat, and usually not even then. (Interestingly,
the sedentary nature of caterpillars, on the other hand, lends them to
be observed in a more leisurely, plant-like, fashion.)

The availability of plant identification manuals seems to have a cat-
alytic effect on a person’s interest in botany—being able to readily learn
the name of one plant leads to a desire to learn the names of additional
plants. In fact, the late Harry Clench, co-founder of our Society, ob-
served a similar positive reinforcement phenomenon when he attributed
two spurts in the growth of our domestic membership to the ‘‘Klots
factor” and the “Ehrlich factor,” otherwise unexplained surges in mem-

8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

bership a year after the publication of two popular butterfly field guides—
collectors had found a way to identify their captures, at the same time
discovering the existence of an organization of similarly inclined nat-
uralists.

There thus seems to be a number of fundamental differences between
the study of plants and the study of Lepidoptera, so that the two can
not be compared fairly. However, I have left until last one final dif-
ference that I think we can do something about.

After discussing the matter with entomologists and botanists, I have
concluded that entomologists in general, and lepidopterists in particular,
tend to be aloof, egotistical, self-centered, selfish, secretive, individu-
alistic, and unwilling or unable to agree on a cooperative national/
international agenda. The entomological community appears to have
a low general regard for faunistic studies and basic taxonomic research,
so that grant proposals in these disciplines receive low marks when
competing against sexier or more high-tech fields of entomological
inquiry. In the United States I am aware of only one major grant-
funded moth faunal survey (in Costa Rica, possibly funded because of
the superior reputation of the investigator rather than the significance
of the survey itself) and not a single similarly funded taxonomic study.
(It is encouraging that there are more and more locally funded, locally
focused studies, such as inventories of the butterfly fauna of natural
areas, but I am concerned here with the negative national attitude
toward such studies.)

Botanists, in contrast, seem to have no difficulty agreeing on the
importance of floral surveys and taxonomic research both at home and
abroad, and in providing the manpower and finances to accomplish the
task. Because botanists agree on the necessity of this type of research,
this view is reflected in positive anonymous peer reviews of grant
proposals, which are then funded.

Botanists can justify their existence because they study this planet’s
energy source. We entomologists, and particularly lepidopterists, can
justify our existence because we study the primary herbivores, those
animals at the bottom of the food chain that benefit man and the
ecosystem by providing food for birds, fish, herptiles, and even man,
and by recycling nutrients and enriching the soil, by pollinating plants,
and so forth, in addition to some of them just being lovely to look at;
comparatively few species actually compete with man for food or fiber.

The naked truth of the matter is that we are still very ignorant about
one of the best known groups of insects! Not only have we named only
about half the species in existence, but we don’t know what most of
them eat, much less their role in the grand scheme of things. Even
more frightening is the fact that they are surely becoming extinct as

VOLUME 44, NUMBER 1 9

rapidly as any other group of organisms as their habitats and hostplants
are being destroyed.

It is sad and unfortunate that entomologists as a group are perceived
by the general public as harmless at best, insane at worst; while butterfly
collectors are frivolous airheads and moth collectors have most likely
lost all touch with reality. Burdened as we are with these misperceptions
of our worth, it is difficult for us or the object of our inquiry to be taken
seriously. The fact that we like our work is complicated by the fact
that the animals we study are actually beautiful; it is even more com-
plicated if we work on small species, because the general public equates
size with significance.

I have now completed my examination of “Why can’t lepidopterists
be more like botanists?” There may still be some hope for us! The Moths
of America North of Mexico project and the Atlas of Neotropical
Lepidoptera project are positive steps in the direction I feel we should
be following, but both suffer from a shortage of funds and a severe
lack of specialists to participate. (The shortage of taxonomists is a general
one, not limited to lepidopterists; the ultimate reason is slashed funding
for jobs in systematics as a result of a perceived relative lack of signif-
icance of this discipline.) Pending legislation for a national biological
inventory would be a major step forward if it receives adequate funding.

The most important basis for our Society, and the one that makes us
so special, is that we are organized to serve as the meeting ground for
amateurs and professionals, for the mutual benefit of both and for the
advancement of lepidopterological knowledge. The “professionals,” un-
fortunately, are a vanishing race, torn between the conflicting goals of
producing the taxonomic and faunistic monographs and manuals we
need so badly on the one hand, and providing guidance and assistance
to the amateur community on the other hand. We have seen examples
of outstanding work by our amateur members, and I know that many
more members are capable of similarly fine contributions if only they
could receive a little encouragement and help. Without that help they
are left to their own devices, leading to results that may be less than
satisfactory.

Short of a miraculous but unlikely increase in funding for taxonomic
research, if we are to make any significant progress in understanding
the taxonomy and distribution of our Lepidoptera fauna it is absolutely
essential that we adopt a plan, a National Lepidoptera Agenda, that
will make the most efficient possible use of our biggest resource, our
members. The formalization of this Agenda will require a lot of thought
by all our members and particularly by the Executive Council, our
elected representatives. Some suggestions to consider in the implemen-
tation of the Agenda include:

10 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

identification and techniques workshops at national meetings

traveling workshops and seminars to regional meetings

use of the Publication Fund to publish identification and techniques manuals

a techniques video (an advanced techniques video could be filmed in a lab or at one
of the workshops)

a newsletter of Lepidoptera taxonomy, a periodical directory of who is working on
what group, and who has material available for study

a directory of taxonomic “holes”: a guide to taxa in need of specialists and/or more
specimens

a similar directory of regional/habitat “holes”: a guide to areas with poorly known
faunas

participate in national discussions on biological surveys and standardization of databases;
contract/cooperate with The Nature Conservancy's Natural Diversity Data Base
program

disseminate information to members on curatorial standards and database management
systems

publish bibliographies of taxonomic revisions (perhaps providing copies of original
works on microfilm, computer disc, video disc, or xerographic copies)

establish a “specialist network” to make critical identifications for faunistic publications

compile a directory of “visiting specialists” willing to provide on-site taxonomic assis-
tance to those paying expenses

publish a directory of research collections willing to accept and curate voucher spec-
imens from faunal surveys

publish a leaflet on how to ship specimens safely

publish a handbook or video on rearing techniques

promote preservation of larvae and parasitoids by publishing a leaflet on proper tech-
niques

reinstate/resurrect the larval voucher repository program

commission/solicit and publish a compendium of hostplant data for North American
“microlepidoptera” (to complement data in Tietz); consider funding or seek grant
for development of a database for this purpose

consider establishing a network of semi-autonomous regional branches or affiliates
(including existing regional groups), to foster communication, standardization, and
unification of purpose

Proposing suggestions is easy; implementing them is the tough part.
Although the Society has an abundance of very talented and capable
members, fewer than 10 of them—all volunteers—are responsible for
our day-to-day operation. With a little leadership and guidance from
the Executive Council, let us hope that more of our members will
become personally involved in helping to make the National Lepidop-
tera Agenda become a reality. Perhaps lepidopterists can be more like
botanists.

Received for publication 10 February 1990; accepted 10 February 1990.

Journal of the Lepidopterists’ Society
44(1), 1990, 11-27

AMBLYSCIRTES: PROBLEMS WITH SPECIES,
SPECIES GROUPS, THE LIMITS OF THE GENUS,
AND GENUS GROUPS BEYOND—A LOOK AT WHAT IS
WRONG WITH THE SKIPPER CLASSIFICATION OF
EVANS (HESPERIIDAE)

JOHN M. BuRNS

Department of Entomology, NHB 169, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560

ABSTRACT. In detecting and correcting errors at all these taxonomic levels, I lean
heavily on genitalia. Two similar, closely related, ostensibly allopatric differentiates treated
by some as species and by others as subspecies are indeed separate species that are barely
sympatric: Amblyscirtes celia Skinner and A. belli Freeman. Most closely related to this
pair is the mainly Mexican complex A. tolteca Scudder/prenda Evans, rather different
in facies and currently misplaced in a different species group of Amblyscirtes. Another
species that looks very like an Amblyscirtes—simius Edwards—assuredly is not! Although,
like simius, A. alternata (Grote & Robinson) has a short, blunt antennal apiculus that is
“wrong for Amblyscirtes, alternata clearly belongs. Placed by Evans (1955) in his N or
Lerodea group of American hesperiine genera and said to be allied to Atrytonopsis,
Lerodea, and Oligoria, Amblyscirtes actually has close ties with various neotropical genera
in Evans's J or Apaustus group: Remella, Mnasicles, and Callimormus! By extrapolation,
much of Evans's taxonomic system just below the level of the subfamily may be invalid.
Ironically, a Guatemalan skipper that Bell (1959) described in the J group genus Moeris
(with which Evans erroneously synonymized Remella) is really an Amblyscirtes: A.
patriciae, new combination. This species clusters with A. folia Godman, A. immaculatus
Freeman, and A. raphaeli Freeman, which come from southern Mexico.

Additional key words: genitalia (male and female), Callimormus, Mnasicles, Re-
mella, Moeris.

Arbitrary change in the rank of a species can sow confusion. Merely
by listing Wallengrenia egeremet (Scudder) as a variety of W. otho
(Smith), Edwards (1877) launched a systematic muddle lasting nearly
a century and culminating in a published load of bioillogicalities (details
in Burns 1985). Unless we ignore arbitrary action from the outset (and
lepidopterists seldom do), it will sooner or later require critical response.

With just 7 specimens of Amblyscirtes belli Freeman from northern
Texas and 11 of A. celia Skinner from southern Texas, Evans (1955)
made them subspecies. Freeman (1941) had described A. belli, in terms
of facies, from 109 specimens from Dallas County in northeastern Texas
which he compared with 37 specimens of A. celia, the most similar
species, noting four facies differences (not altogether valid). Skinner
(1895) had described A. celia, also in terms of facies, from an unstated
number of specimens from Blanco, Comal, and Nueces counties in
southcentral Texas.

Close similarity plus apparent or actual allopatry do not, of them-
selves, warrant reduction to subspecific rank. Careful analysis of various

12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

skippers has revealed monophyletic groups of closely related but largely
or entirely allopatric species—see, for example, the superspecies in
Erynnis and Atrytonopsis (Burns 1964, 1983).

Evans’s (1955) action has been divisive: supported most notably by
dos Passos (1964), Harris (1972), and Scott (1986), with echoes to the
present (Watson & Hyatt 1988); and opposed by Freeman (1973),
MacNeill (1975), and Miller and Brown (1981, 1983), as well as by others,
with some waffling—Opler and Krizek (1984:259) treated A. celia and
A. belli separately but allowed that “Celia’s Roadside Skipper is some-
times considered to be the same species as Bell’s Roadside Skipper.”

Facies differences between A. celia and A. belli (unevenly dealt with
in many of the works cited above) are subtle and variable but pervasive
enough to suggest different species in the context of the genus Am-
blyscirtes. Still, rank is arguable. New information is needed.

Freeman (1978, and again in Irwin & Downey 1973), in defending
the rank of species, cited an instance of sympatry between A. celia and
A. belli near the type-locality of the latter in Dallas County, Texas.
Because I found celia common 265 km to the southsouthwest at Austin,
Travis County, Texas, in 1966 and 1967, contact seemed plausible. But
because Freeman mistook a few of my 56 Austin specimens of celia
for belli at that time, I was dubious—until 1989, when he graciously
collected, mounted, and forwarded 8 6 3 2 of celia and 6 4 1 2 of belli
that were flying together at Garland, Dallas County, Texas, during
August and September, and when I discovered, among Amblyscirtes
in the National Museum of Natural History (USNM), a misplaced worn
female of belli taken at Austin in May 1980. Like many other pairs of
differentiates, these two skippers overlap narrowly in eastcentral Texas.

Without giving specifics, Freeman (1973:54) added, “There are slight
differences [between celia and belli] in the genitalia, however genitalic
determinations in the genus Amblyscirtes are practically impossible
with most species due to the fact that the basic pattern is very similar.”
The qualification is disturbing. Early in the same review of Amblys-
cirtes, Freeman (1973:41) put it this way: “... there is a remarkable
similarity in the male genitalia of all of the species. Often worn spec-
imens are very difficult to identify even after an examination of the
genitalia due to this great consistency in basic form.” Long before,
having reprinted the Skinner and Williams (1928) figures of the male
genitalia of Amblyscirtes from the United States and Canada, Lindsey
et al. (1931:126) observed, “The genitalia of all of these species are
remarkably uniform in structure.” I am skeptical, then, when Freeman
(1973:45, 48, 50, 51, 54) repeatedly asserts—for five pairs of species
whose rank has been (or, in one case, may be) questioned—‘“‘there are

VOLUME 44, NUMBER 1 13

differences in the genitalia,” especially since he never says what any
of those differences are.

Considering the genitalic conservatism, intrageneric taxa showing
distinct genitalic differences will almost certainly be species rather than
subspecies. However, I must stress that in Amblyscirtes, as elsewhere,
genitalia vary individually so that, even in a local population of a single
species, no two tails are exactly alike; and the detection of “distinct
genitalic differences” entails much dissection and comparison. In com-
paring genitalia of different species in this and similar taxonomic papers,
the reader should keep in mind that many of the differences between
figures are individual instead of interspecific and that the angle of
illustration (and hence of observation) is crucial.

A Tail of Two Species

At certain angles, the male genitalia of the taxa in question clearly
differ. From above, the middle of the distal end of the uncus looks
convex in A. celia (Fig. 1) but concave in A. belli (Fig. 4). In all species
of Amblyscirtes the simple, roughly rectangular valva ends in a prom-
inent, pointed, dorsally directed terminal process that is slightly set off
from the body of the valva and slightly medial to it (Figs. 3, 6, 13, 20,
plus figures in many other sources, especially Godman & Salvin 1879-
1901, Skinner & Williams 1923, Lindsey et al. 1931, and Evans 1955).
A posterior look at the distal end of the valva shows—at the level of
the base of the dorsally directed terminal process—a major, modestly
dentate, medial expansion in A. celia (Fig. 2) which, by contrast, is
relatively low and smooth in A. belli (Fig. 5).

The distal location of these genitalic characters often makes them
accessible in situ. Using both wet and dry dissection, I have examined
them critically in 43 males of A. celia from Dallas, Travis, Bastrop,
Comal, Bexar, Kendall, Kerr, Harris, Hidalgo, and Cameron counties,
Texas, plus Ciudad Victoria and Ciudad Mante, Tamaulipas, Mexico;
and in 40 males of A. belli from Dallas and Tarrant counties, Texas,
Garfield County, Oklahoma, Carroll, Faulkner, and Pulaski counties,
Arkansas, Labette County, Kansas, Barry and St. Clair counties, Mis-
souri, Fulton and McCracken counties, Kentucky, Madison County,
Tennessee, Fulton County, Georgia, and Pickens County, South Car-
olina. The genitalic differences hold over the range of each species,
even where the two are sympatric or geographically close (nearly half
the celia genitalia examined come from Austin, Texas, and nearly a
fifth of the celia, and more than half the belli, from around Dallas,
Texas).

Though highly variable, the female genitalia also differ in a distal

14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

I, Behe

Fics. 1-3. Male genitalia of Amblyscirtes celia from Austin, Travis Co., Texas, 15
May 1967, J. M. Burns (genitalic dissection no. X-2528) (USNM). 1, Tegumen, uncus,
and gnathos in dorsal view; 2, Distal ends of valvae and aedeagus in posterior view; 3,
Complete genitalia (minus juxta and right valva) in left lateral view.

character (mere removal of terminal ventral abdominal scales will re-
veal it). The sclerotized posterior margin of the lamella postvaginalis
(just ventral to the ovipositor lobes) is, in A. celia, widely but decidedly
V-shaped, with the point of the V anterior, midventral, and more or
less rounded into a small midventral notch (Fig. 7). In A. belli this
sclerotized posterior margin varies from shallowly U-shaped (Fig. 9) to
shallowly W-shaped to virtually straight. I have compared this feature
again and again in 33 females of A. celia from Dallas, Travis, Bexar,
Harris, San Patricio, Hidalgo, and Cameron counties,Texas; and in 25

7 Ze

Fics. 4-6. Male genitalia of Amblyscirtes belli trom the vicinity of Irving, Dallas Co.,
Texas, 28 July 1984 (X-2529) (USNM). 4, Tegumen, uncus, and gnathos in dorsal view;
5, Distal ends of valvae and aedeagus in posterior view; 6, Complete genitalia (minus
juxta and right valva) in left lateral view.

VOLUME 44, NUMBER 1 15

females of A. belli from Travis and Dallas counties, Texas, Pulaski
County, Arkansas, Labette County, Kansas, St. Clair and Cape Gi-
rardeau counties, Missouri, Fulton County, Georgia, and Pickens Coun-
ty, South Carolina.

In both species the sclerotized ductus bursae—which angles to the
left (Figs. 7, 9)—is not a plain tube: approximately the anterior third
bears a wide, deep groove in the left side which, posteriorly, becomes
shallow as it twists to the dorsal side and disappears (Figs. 7-10). Details
of expression vary greatly. Because most of this variation is individual,
no consistent interspecific differences emerge.

Disruption of Species Groups

Both A. celia and A. belli display a striking asymmetry of the ae-
deagus. (This feature, too, is distal and thus visible without dissection
whenever the superlengthy shaft projects from the end of the abdomen.)
Along the left side of the aedeagus, just before its end, a large, thin,
triangular plate with a long base curves outward and upward forming
a conspicuous pointed titillator (Figs. 2, 3, 5, 6).

No one has ever mentioned it. Skinner and Williams (1923:144, fig.
24) and Lindsey et al. (1931:pl. 31, fig. 24), in illustrating the right
valva and the distal end of the aedeagus of what they called A. celia,
either got the wrong species or missed the titillator, which, given its
size and the amount of detail in their figures, seems unlikely. On the
other hand, Evans (1955:pl. 82, fig. N.2.18), without comment, caught
the titillator in his caricature of distal portions of A. celia genitalia.

Nothing like it appears in any of his 22 other genitalic caricatures
of Amblyscirtes species—not even the one for A. tolteca Scudder (Evans
1955:pl. 82, fig. N.2.11). Ranging through most of Mexico, A. tolteca
apparently gives way in the west and northwest (especially in Sonora
plus southern Arizona) to what Evans (1955) briefly described (in terms
of size and facies) as subspecies prenda, which Freeman (1973) raised
to the rank of species without adequate justification. Though lacking
enough material to resolve the status of tolteca vis-a-vis prenda, I can
definitely say that both have a titillator like that of A. celia and A. belli
and, furthermore, that no other known species of Amblyscirtes does.

Using a few characters of facies, Evans (1955) divided the sizeable
and superficially variable genus Amblyscirtes into four species groups
which Freeman (1973), in his review of the genus, endorsed. Freeman
also followed Evans’s sequence of species—as have most workers, with
little or no deviation, from dos Passos (1964) on. This arrangement puts
A. celia plus A. belli far from A. tolteca/prenda in a different species
group.

The distinctive titillator unites them in a close-knit assemblage (above

16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 7, 8. Female genitalia of Amblyscirtes celia from Austin, Travis Co., Texas, 25
May 1967, J. M. Burns (X-2452) (USNM). 7, Sterigma, bursa copulatrix, and part of the
ductus seminalis in ventral view; 8, Sterigma and bursa copulatrix in right lateral view.

VOLUME 44, NUMBER 1 Me

Fics. 9,10. Female genitalia of Amblyscirtes belli from Vickery, Dallas Co., Texas,
30 August 1952, H. A. Freeman (X-2692) (USNM). 9, Sterigma, bursa copulatrix, and
part of the ductus seminalis in ventral view; 10, Sterigma and bursa copulatrix in right
lateral view.

18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

a superspecies since A. celia and A. tolteca overlap broadly in Mexico).
Though all have a fundamentally similar spot-pattern on the ventral
secondary, the four taxa run a gamut in average spot expression from
weak in belli and rather better in celia to strong in tolteca and very
strong in prenda; and tolteca/prenda stand out especially on account
of a bold double spot in the cell and a sharp spot in space 1b,, on both
surfaces of the primary, plus several spots on the dorsal secondary—
all of which A. celia and A. belli lack. (In these species of Amblyscirtes,
males express spots better than females.) In addition, primaries are
more pointed in tolteca/prenda than in celia and belli. (And primaries
are more pointed in males than in females.)

Banishing the Imposter

Originally described in Amblyscirtes, simius Edwards (1881) was
moved by Barnes and McDunnough (1916) to Chaerephon Godman—
which Hemming (1935) pegged as a junior homonym and renamed
Yvuretta—from which Freeman (1943) moved simius back to Amblys-
cirtes, where it remains (uncomfortably). (Draudt [1924] put it in Sto-
myles, which has long been considered a synonym of Amblyscirtes.)

Barnes and McDunnough justified removal by citing the third palpal
joint and the antennal club. The third joint of the palpus is relatively
short and directed forward to moderately upward in simius but long,
slender, and about vertical in most species of Amblyscirtes; the antennal
club looks nearly blunt with an exceedingly short and stubby apiculus
in simius but conspicuously pointed with an abruptly constricted and
attenuate apiculus in Amblyscirtes. Also mentioning the peculiar stig-
ma, Barnes and McDunnough (1916:125) went so far as to say, “it may
be necessary to erect a new genus for” simius; but conservatively, and
tentatively, they placed it in Chaerephon—which is wrong (compare
the figures of the male genitalia of simius with those of the other two
species under Chaerephon in Skinner & Williams 1923, reprinted in
Lindsey et al. 1931).

Proclaiming that “Edwards was correct when he described this species
in the genus Amblyscirtes,’ Freeman (1943:75) argued his opposite
case using the very same characters—third palpal joint and antennal
club—which he misrepresented. Freeman (1943:76) admitted that
“genitalically this species is not like other members of the genus Am-
blyscirtes.”

Evans (1955), MacNeill (1975), Stanford (1981), and Scott (1986)
reiterated that the genitalia of simius are aberrant for Amblyscirtes;
and the three Americans added, on this and other (especially behavioral)
grounds, that simius may belong in another genus. Having shown con-
clusively that “‘genitalic characters, generally so useful in differentiating

VOLUME 44, NUMBER 1 19

D7 oa

Fics. 11-13. Male genitalia of Amblyscirtes vialis from Lexington, Middlesex Co.,
Massachusetts, 2 June 1975, J. M. Burns (X-2516) (USNM). 11, Tegumen, uncus, and
gnathos in dorsal view; 12, Distal ends of valvae and aedeagus in posterior view; 13,
Complete genitalia (minus juxta and right valva) in left lateral view.

species, are also exceptionally valuable at the generic level in skippers”’
(Burns 1987:173), I wish, once and for all, to banish simius from Am-
blyscirtes.

In both sexes the genitalia of A. celia and A. belli (Figs. 1-10) are
obvious variations on the theme of the type-species, A. vialis (Edwards)
(Figs. 11-15). In males this singular theme boasts several salient ele-
ments: a wonderfully long, narrow aedeagus (Figs. 3, 6, 18) split distally
into two, more or less parallel, linearly toothed ends (Figs. 2, 3, 5, 6,
12, 13); a correspondingly long, narrow saccus (Figs. 3, 6, 18); and a
tegumen with a delicate, middorsal, caudally arching prong supporting
membrane over a variously oval to cordate to triangular dorsal opening
(Figs. 1, 3, 4, 6, 11, 13).

The male genitalia of simius (Figs. 16, 17) differ radically: the ae-
deagus is relatively short and stout, without distal modification, but
with a pair of simple cornuti (lacking in Amblyscirtes); the saccus,
though somewhat long, is not half what it is in Amblyscirtes; and the
tegumen, uncus, and gnathos are all utterly distinct, both in form and
in relative proportions, from their counterparts in Amblyscirtes. As in
many other hesperiine skippers, the paired distal tips of the gnathos
are completely free of the overlying uncus instead of being individually
joined to its underside, as they are in Amblyscirtes. Any fancied resem-
blance between the valvae in lateral view is doubtless convergent.

The female genitalia of simius stand apart at the grossest level: after
4 or 5 minutes of boiling in 10% KOH, they are virtually membranous
throughout—even the lamella postvaginalis and the ductus bursae. These
parts are always well sclerotized in Amblyscirtes (Figs. 7-10, 14, 15),
even after 7 or more minutes of boiling.

20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. 14, 15. Female genitalia of Amblyscirtes vialis from Jacksonville, Windham
Co., Vermont, 5 June 1963, J. M. Burns, E. D. Hanson, and D. W. Hottenstein (X-2519)
(USNM). 14, Sterigma, bursa copulatrix, and part of the ductus seminalis in ventral view;
15, Sterigma and bursa copulatrix in right lateral view.

Where does simius go? Not in any named nearctic genus, but con-
ceivably in a neotropical one. After all, a basically tropical skipper
genus can produce a species that breaks the distributional mold by
occurring primarily, and extensively, in North America north of Mex-

VOLUME 44, NUMBER 1 21

Fics. 16, 17. Male genitalia of ? simius from Horsetooth Mountain Park, 1800 m,
Larimer Co., Colorado, 14 June 1987, P. A. Opler (X-2509) (USNM). 16, Tegumen, uncus,
and gnathos in dorsal view; 17, Complete genitalia (minus right valva) in left lateral view,
with vesica everted to show cornuti well.

ico—witness Epargyreus clarus (Cramer). To describe a new genus for
simius still strikes me as premature (as it did Barnes & McDunnough
in 1916). For the present I advertise its tail (Figs. 16, 17) so that others,
too, may look for a possible match. Though I cannot fix its genus, better
that simius float than clutter the wrong one.

Sometimes It’s Okay To Be Different

The sole excuse for its erroneous placement is that simius looks like
an Amblyscirtes—but we know that facies can converge and appear-
ances, deceive. As discussed above, not only are the entire genitalia of
simius wrong for Amblyscirtes but so are such prominent cephalic
appendages as the palpus and, particularly, the antenna, whose apiculus
is a blunt fraction of what it ought to be.

In this connection, it is instructive to point out for the first time that
A. alternata (Grote & Robinson) has a similarly short, blunt apiculus!

22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

BA,

Fics. 18-20. Male genitalia of Amblyscirtes alternata from 5 km N Panacea, Wakulla
Co., Florida, 5 April 1980, J. M. Burns (X-2457) (USNM). 18, Tegumen, uncus, and
gnathos in dorsal view; 19, Distal ends of valvae and aedeagus in posterior view; 20,
Complete genitalia (minus juxta and right valva) in left lateral view.

Nevertheless, alternata is a true Amblyscirtes: the third joint of its
palpus is long, slender, and erect; and (what is really crucial) its genitalia
(Figs. 18-20) are a manifest variation on the Amblyscirtes theme (com-
pare Figs. 1-6, 11-13). Note that the aedeagus carries a subterminal
titillator in the form of a modest midventral keel (Figs. 19, 20). Both
this keel and the much reduced apiculus are unique in the genus Am-
blyscirtes.

Biologic variation may sharply change the singular genitalic theme
itself. In three species, A. nereus (Edwards), A. eos (Edwards), and A.
nysa Edwards, the delicate, middorsal tegumen prong—one of the
“salient elements’’—totally disappears. But the underlying dorsal open-
ing persists, as do all other salient elements. Nature simply makes it
harder to generalize.

Neotropical Ties and Disruption of Genus Groups

With simius out, the rest of the 30 species currently in Amblyscirtes
(Freeman 1973) comprise a related lot sharing more or less similar
genitalia. Still, the genus is mixed with respect to such features as facies,
the length of the apiculus, the third joint of the palpus, and, most
important, the stigma of the male. Not just the species groups of Am-
blyscirtes but the exact limits of the genus may need to be reworked.

But larger, more basic problems loom. When he arranged the genera
of American hesperiines in 8 groups (lettered H to O), Evans (1955:
7-8, 383) put Amblyscirtes in the N or Lerodea group (the smallest),
observing that, “The 4 genera placed in this group [Atrytonopsis, Am-
blyscirtes, Lerodea, and Oligoria] appear to be more or less allied and
would be out of place in any other group.”’ Actually, the nearest relatives
of Amblyscirtes are far removed in Evans’s J or Apaustus group!

VOLUME 44, NUMBER 1] 23

Support for this startling assertion comes from figures of male gen-
italia in two classic works treating skippers at opposite ends of the
neotropics (Mexico and Central America; Argentina). The critical fig-
ures (Godman & Salvin 1879-1901:vol. 3, pl. 99, fig. 3, pl. 103, figs.
26, 27, 31, 33 [all reprinted in this paper as Figs. 21-25]; Hayward 1950:
pl. 8, fig. 3, pl. 13, figs. 5, 11) show salient elements of the singular
Amblyscirtes theme—especially those relating to the aedeagus and the
saccus, but, in one case, even the delicate tegumen prong—coupled
with valvae loosely reminiscent of Amblyscirtes valvae. According to
Evans (1955), those figured genitalia belong to 5 species in 3 genera—
Moeris remus (Fabricius), Mnasicles geta Godman, M. hicetaon God-
man, Callimormus juventus Scudder, and C. alsimo (Méschler)—and
those genera (all polytypic) constitute, respectively, numbers 33, 6, and
2 of his Apaustus group (which contains 53 genera in all).

At this point I must sunder the Evans combination Moeris remus
and restore remus to its proper genus. Without any question, the figures
of the male genitalia of the type-species of Godman’s new genera
Perimeles (Godman & Salvin 1879-1901:vol. 3, pl. 99, fig. 3 [Fig. 21
in this paper]) and Moeris (vol. 3, pl. 100, fig. 2 [Fig. 26 in this paper])
reflect very distinct genera, which Evans (1955:146) wrongly lumped.
The genitalia of the type-species of Perimeles—remus Fabricius—
suggest Amblyscirtes, while the genitalia representing Moeris do not.
There has been a legitimate name change, though: Perimeles remus
became Remella remus when Hemming (1939) saw that Godman’s
Perimeles is a junior homonym and replaced it with Remella.

Having examined a number of genitalic dissections of Remella, Mna-
sicles, and Callimormus for myself, I reiterate that those small neo-
tropical skippers are phylogenetically close to Amblyscirtes. To see—
almost at a glance—that the male tails of Atrytonopsis, Oligoria, and
Lerodea depart much farther from those of Amblyscirtes, compare
relevant figures in Godman and Salvin (1879-1901), Lindsey et al.
(1931), Hayward (1950), and Burns (1982, 1983).

Blatant genitalic heterogeneity in both the J and the N groups of
Evans probably exceeds what I have indicated. Worse yet, it may occur
in other groups, invalidating much of Evans’s taxonomic system just
below the level of the subfamily.

I have a final irony in the fire. In the course of checking out possible
neotropical relatives of Amblyscirtes, I studied the male holotype (the
only known specimen) of what Bell (1959) designated Moeris patriciae
(taken by Patricia Vaurie at Salama, elevation 3000 ft [915 ml], in the
middle of Guatemala on 22 July 1947), including the slide Bell had
made of its genitalia. From this—as well as from Bell’s (1959:figs. 9,
15) illustrations of the whole animal and its genitalia—the skipper

24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

25

Fics. 21-25. Godman’s figures of male genitalia of various neotropical relatives of
Amblyscirtes; complete genitalia (minus juxta and left valva) in left lateral view. Note
that, because Godman removed the left valva, his figures show the inner surface of the
right valva, whereas mine of Amblyscirtes show the outer surface of the left valva. 21,
Remella remus; 22, Mnasicles geta; 23, Mnasicles hicetaon; 24, Callimormus juventus;
25, Callimormus alsimo.

VOLUME 44, NUMBER 1] 25

26

Fic. 26. Godman’s figure of the male genitalia of Moeris striga; complete genitalia
(minus juxta and left valva) in left lateral view. Even in a limited lateral comparison, this
tail differs grossly from that of Remella remus (Fig. 21): the aedeagus is short and simple
but equipped with cornuti, the saccus is very short, and the distal ends of the valva are
much produced.

obviously belongs in Amblyscirtes as then and now conceived. Am-
blyscirtes patriciae (Bell), new combination, clusters with A. folia God-
man, A. immaculatus Freeman, and A. raphaeli Freeman, which come
from southern Mexico.

For Amblyscirtes, these species are large to extremely large in size
(the length of the male primary ranges from about 15 to 20 mm). Males
have a well-developed, conspicuous, linear stigma, in three sections,
consisting mainly of fine, dense, short, brown, hairlike scales. The long-
est (and uppermost) section begins at the origin of veins 3 and 4, runs
along the lower side of the cubitus, diverges from it well before the
origin of vein 2, and extends down to that vein. The second and third
sections pick up below vein 2 as a pair of dashes or dots, much more
nearly vertical in orientation, that extend to, or toward, vein 1. Although
tripartite, this stigma looks more or less continuous to the naked eye.
The apiculus of the antenna (unfortunately missing from the specimen
of A. patriciae) is perceptibly longer and more delicate than it is in
other Amblyscirtes. The third segment of the palpus is relatively short,
rather than long and slender. A pale area (cream or tan or yellowish)
tends to develop on the ventral primary in the distal half of space 1b.

ACKNOWLEDGMENTS

My thanks to F. H. Rindge for his generous hospitality during repeated visits to the
American Museum of Natural History, New York, including access to Bell’s and Freeman’s
various genitalic preparations and their Amblyscirtes types; to Rindge and C. V. Covell
Jr. for lending specimens of Amblyscirtes and to H. A. Freeman, P. A. Opler, and C. N.
Watson Jr. for donating them; to Adrienne Venables for 123 precise KOH dissections of
male and female genitalia of Amblyscirtes, its relatives, and ? simius and to Young Sohn
for superb drawings of some of them; to F. D. Godman for figuring male genitalia of
skippers before it was fashionable and to Victor Krantz for reproducing a few of Godman’s
figures; and to Sarah Burns for catching Amblyscirtes, recording data, reading manuscript
and proof, and giving all sorts of general support.

26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

LITERATURE CITED

BARNES, W. & J. H. MCDUNNOUGH. 1916. Notes on North American diurnal Lepi-
doptera. Contributions to the natural history of the Lepidoptera of North America.
3:51-156, pls. 4-11. Review Press, Decatur, Illinois.

BELL, E. L. 1959. Descriptions of some new species of neotropical Hesperiidae (Lepi-
doptera, Rhopalocera). Am. Mus. Novit. No. 1962. 16 pp.

BuRNS, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif.
Publ. Entomol. 37:1-217.

1982. Lychnuchoides frappenda from central Mexico joins lunus and zweifeli

in a lunus group of Atrytonopsis (Lepidoptera: Hesperiidae: Hesperiinae). Proc.

Entomol. Soc. Wash. 84:547—567.

1983. Superspecies Atrytonopsis ovinia (A. ovinia plus A. edwardsi) and the

nonadaptive nature of interspecific genitalic differences (Lepidoptera: Hesperiidae).

Proc. Entomol. Soc. Wash. 85:335-358.

1985. Wallengrenia otho and W. egeremet in eastern North America (Lepi-

doptera: Hesperiidae: Hesperiinae). Smithsonian Contrib. Zool. No. 423. 39 pp.

1987. The big shift: nabokovi from Atalopedes to Hesperia (Hesperiidae). J.
Lepid. Soc. 41:173-186.

bos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. Soc.
Mem. No. l. v + 145 pp.

DrauptT, M. 1924. Vol. 5. The American Rhopalocera. In Seitz, A. (ed.), The Macro-
lepidoptera of the world. Alfred Kernen Verlag, Stuttgart.

Epwarps, W. H. 1877. Catalogue of the diurnal Lepidoptera of America north of
Mexico. Trans. Am. Entomol. Soc. 6:1-68.

1881. Descriptions of new species of diurnal Lepidoptera found within the
United States. Trans. Am. Entomol. Soc. 9:1-8.

EvaANs, W. H. 1955. A catalogue of the American Hesperiidae indicating the classifi-
cation and nomenclature adopted in the British Museum (Natural History). Part IV.
Hesperiinae and Megathyminae. British Museum, London. 499 pp., pls. 54-88.

FREEMAN, H. A. 1941. A new species of Amblyscirtes from Texas (Lepidoptera, Rho-
palocera, Hesperiidae). Entomol. News 52:50-51.

1943. New Hesperioidea, with notes on some others from the United States

(Lepidoptera, Rhopalocera). Entomol. News 54:72-77.

1973. A review of the Amblyscirtes with the description of a new species from
Mexico (Hesperiidae). J. Lepid. Soc. 27:40-57.

GoDMAN, F. D. & O. SALVIN. 1879-1901. Biologia Centrali-Americana; Insecta; Lep-
idoptera-Rhopalocera. Vol. 2, 782 pp.; Vol. 3, 113 pls.

Harris, L., JR. 1972. Butterflies of Georgia. Univ. Oklahoma Press, Norman, Oklahoma.
xxii + 326 pp.

HAYWARD, K. J. 1950. Genera et species animalium argentinorum. Vol. 2. Insecta,
Lepidoptera, Hesperiidae, Hesperiinae. G. Kraft Ltd., Buenos Aires. 388 pp., 26 pls.

HEMMING, F. 1935. Notes on seventeen genera of Rhopalocera. Stylops 4:1-3.

1939. On five genera in the Lepidoptera Rhopalocera at present without valid
names. Proc. Roy. Entomol. Soc. London, series B 8:39.

IRWIN, R. R. & J. C. DowNeEy. 1973. Annotated checklist of the butterflies of Illinois.
Illinois Nat. Hist. Survey, Urbana, Ill. Bio. Notes No. 81. 60 pp.

Linpsey, A. W., E. L. BELL & R. C. WILLIAMS, JR. 1931. The Hesperioidea of North
America. Denison Univ. Bull., J. Sci. Lab. 26:1-142.

MACNEILL, C. D. 1975. Family Hesperiidae, pp. 423-578. In Howe, W. H. (ed.), The
butterflies of North America. Doubleday & Co., Inc., Garden City, New York.
MILLER, L. D. & F. M. BRown. 1981. A catalogue/checklist of the butterflies of

America north of Mexico. Lepid. Soc. Mem. No. 2. vii + 280 pp.

1983. Hesperiidae, pp. 42-48. In Hodges, R. W. (ed.), Check list of the Lepi-

doptera of America north of Mexico. E. W. Classey Ltd. and The Wedge Entomo-

logical Research Foundation, London.

VOLUME 44, NUMBER 1 27

Oper, P. A. & G. O. KrizEkK. 1984. Butterflies east of the Great Plains. Johns Hopkins
Univ. Press, Baltimore, Maryland. 294 pp., 54 pls.

Scott, J. A. 1986. The butterflies of North America. Stanford Univ. Press, Stanford,
California. xiii + 583 pp., 64 pls. ;

SKINNER, H. 1895. Notes on Rhopalocera, with descriptions of new species. Entomol.
News 6:112-114.

SKINNER, H. & R. C. WILLIAMS JR. 1923. On the male genitalia of the Hesperiidae of
North America, Paper III. Trans. Am. Entomol. Soc. 49:129-153.

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.

WarTSsON, C. N., JR. & J. A. Hyatr. 1988. Butterflies of northeast Tennessee. J. Lepid.
Soc. 42:19-31.

Received for publication 30 July 1989; revised and accepted 6 January 1990.

Journal of the Lepidopterists’ Society
44(1), 1990, 28-31

A NEW SPECIES OF PIRUNA FROM
MEXICO (HESPERIIDAE)

HUGH AVERY FREEMAN
1605 Lewis Drive, Garland, Texas 75041

ABSTRACT. Piruna kemneri is described from Oaxaca and Puebla, Mexico, the type
series consisting of 18 males collected by John Kemner and one male collected by H. A.
Freeman. The new species is differentiated from its most similar ally P. haferniki Freeman
1970 by morphological and genitalic characters. Holotype and genitalia of a paratype
are illustrated.

Additional key words: Piruna kemmneri, P. haferniki, P. brunnea, P. pirus, P. cy-
closticta.

During the summers of 1987-88 John Kemner collected rather ex-
tensively over Mexico. Among the many interesting species of Hesper-
iidae that he collected in the state of Oaxaca was an undescribed species
of Piruna Evans that is described here. According to Evans’ (1955)
arrangement this species belongs in Group H of the Hesperiinae.

Piruna kemneri Freeman, new species

(Figs. 1, 2, 8)

Male upper side (Fig. 1). Primaries black, with seven variable white, hyaline spots; a
round or linear spot in space 2 and, midway between this spot and base of wing, another
spot in same space, which may be absent in some specimens. In space 3, a small round
or linear spot situated directly under apical spot in space 6. Three, small, apical spots, in
line, with the one in space 7 being minute and sometimes absent. A small, round, upper
cell spot. Fringe black becoming whitish at the tips. Costal margin slightly concave.
Secondaries black, unmarked. Fringe black, becoming lighter at tips, uncheckered.

Male under side (Fig. 2). Primaries brown, lightly overscaled with golden scales, with
space | being lighter in coloration than the rest of the wing. All spots are white, hyaline,
and better defined than on upper side. Fringe dark, brownish black, slightly checkered.
Secondaries brownish black, with a heavy overscaling of golden scales over the basal half
of the wing and extending from anal fold to space 3. There is a heavy concentration of
golden and white scales midway between the apex and base of the wing near the costa,
forming an irregularly shaped blotch. The remainder of the wing brownish black. No
hyaline spots present. Fringe dark brownish black, uncheckered.

Body. Thorax black, heavily overscaled with brown hairs above, lighter beneath due
to brown and golden hairs present. Abdomen dark brownish-black above, lighter beneath.
Head black, covered with brown hairs. Palpi black above, sordid white at base below,
gray at termen, dark grayish on lateral sides. Legs brown. Antennae, both shaft and club,
dark brownish black above, lighter beneath, club pale beneath, yellowish. Apiculus gold.

Wing measurements. Holotype. Primaries: base to apex, 10.5 mm; apex to outer angle,
7 mm; outer angle to base, 9 mm. Secondaries: base to end of vein 3, 8.5 mm; center of
costa to anal angle, 7 mm. Total expanse: 22 mm. Average total expanse of paratypes:
22 mm (n = 17).

Types. Holotype, male, Mexico: Oaxaca: 5 miles north of Oaxaca, Hwy. 175, ca. 1800
m elev., 21 July 1987 (leg. John Kemner) in the American Museum of Natural History,
New York. There are 17 male paratypes, 16 same location and collector as holotype,
collected during July and August 1987-88, at present in my collection, and one male
paratype from Mexico: Tehuacan, Puebla, 14 August 1964 (leg. H. A. Freeman) in the
American Museum of Natural History.

VOLUME 44, NUMBER | 29

Fics. 1,2. Upper side (Fig. 1) and under side (Fig. 2) of Piruna kemneri Freeman,
holotype, male, Mexico: Oaxaca, 5 miles north of the city of Oaxaca, ca. 1800 m elev.,
21 July 1987 (leg. John Kemner).

Fic. 3. Piruna kemneri, male genitalia of paratype (Genitalia Vial H-904), same
location and collector as holotype, 11 July 1988. a) tegumen, uncus, gnathos, and associated
structures in lateral view; b) right valva in lateral view of interior, upper part ampulla,
lower part harpe; c) same as (a), in ventral view; d) aedeagus in lateral view.

30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Etymology. I take great pleasure in naming this new species for my good friend John
Kemner who collected most of the type series.

This new species is a member of the brunnea complex (Freeman
1970) of Pirwna, which contains brunnea (Scudder), cyclosticta (Dyar),
pirus (Edwards), and haferniki Freeman, and is characterized by having
no spots on either the upper or under side of the secondaries. All other
Piruna have either spots or streaks on the upper or under side of the
secondaries. The male genitalia of members of this complex have the
terminal end of the harpe smooth with no indication of being serrate
or having spines, which are present in most of the other species of
Piruna. The members of this complex are briefly described as follows:
brunnea (Scudder) 1872—Average expanse of primary 13 mm. Black.
Two apical spots in spaces 6 and 8. Discal spots in spaces 2 and 8. Cell
spot may or may not be present. No basal spot in space 2. Under side
of secondaries chocolate brown. pirus (Edwards) 1878—Average ex-
panse of primary 11 mm. Brownish-gray. Three apical spots. Discal
spots in spaces 2 and 3 may or may not be present. Small cell spot
usually present. No basal spot in space 2. Under side of secondaries
ferruginous. cyclosticta (Dyar) 1902—Average expanse of primary 11
mm. Brownish-gray. Three apical spots. Discal spots in spaces 2 and 3
well defined. Cell spot always present. Basal spot in space 2 always
present and well defined. Under side of secondaries dull brown. haferni-
ki Freeman 1970—Average expanse of primary 1] mm. Blackish-brown.
Three large apical spots, in line. Discal spots in spaces 2 and 3 well
developed. Cell spot large. Basal spot in space 2 always present and
well developed. Under side of secondaries overscaled over basal and
submarginal areas purplish-gray leaving the center of the wing dark
brown.

The most closely related species to kemneri is haferniki, from which
it can be separated by the following features. In haferniki the apical,
discal, and cell spots on the primaries are much better developed than
in kemneri. Perhaps the easiest way to separate the two species is to
compare the under side of the secondaries. P. haferniki has the unusual
purplish-gray overscaling over the basal half of the wing and extending
from the anal fold along the outer margin to space 5, leaving the discal
area from space | to costa a broadly triangular area darker than the
remainder of the wing, which lacks overscaling. P. kemneri has a heavy
overscaling of golden scales over the basal half of wing and extending
from space 1 and anal fold to space 3, leaving the remainder darker,
except for the large irregularly shaped, lighter blotch midway between
the apex and base on the costa, produced by a heavy concentration of
golden and white overscaling. The costal margin of the primaries of P.
kemneri is slightly concave, in contrast to all other species of Piruna.

VOLUME 44, NUMBER 1 31

The male genitalia of kemneri (Fig. 3) differ from haferniki in the
following ways: the tegumen (lateral view) is broader; terminal end of
divided uncus is straight and not curving laterally like haferniki (ventral
view); ampulla is narrower at posterior end than in haferniki and the
anterior end is broader and more upturned than in haferniki (lateral
view); and the harpe is slightly narrower at the terminal end than in
haferniki (lateral view). These genitalic differences are based on the
examination of four kemneri paratypes.

ACKNOWLEDGMENTS

I thank Dr. Frederick H. Rindge of the American Museum of Natural History, New
York, for the loan of Piruna specimens and for making the photographs of the holotype
used in this article.

LITERATURE CITED

Evans, W. H. 1955. A catalogue of the American Hesperiidae indicating the classifi-
cation and nomenclature adopted in the British Museum (Natural History). Part IV.
Hesperiinae and Megathyminae. British Museum, London. 499 pp., pls. 54-88.

FREEMAN, H. A. 1970. A new species of Piruna from Texas (Hesperiidae). J. Lepid.
Soc. 24:247-249.

Received for publication 3 February 1989; revised and accepted 13 October 1989.

GENERAL NOTES

Journal of the Lepidopterists’ Society
44(1), 1990, 32

PREDATION OF FIVE SPECIES OF NOCTUIDAE AT ULTRAVIOLET
LIGHT BY THE WESTERN YELLOWJACKET
(HYMENOPTERA: VESPIDAE)

Additional key words: Nephelodes minians, Pseudaletia unipuncta, Heliothis zea,
Trichoplusia ni, Catocala faustina.

Between 23 September and 26 October 1989, I observed several western yellowjackets,
Vespula pensylvanica (Saussure), preying on five species of Noctuidae at an ultraviolet
light in central Colorado.

Vespula pensylvanica ranges from Hawaii, east across the Rockies onto the high plains
of central Colorado (Akre, R. D., A. Greene, J. F. MacDonald, P. J. Landolt & H. G.
Davis. 1980, Yellowjackets of America north of Mexico, U.S. Dept. Agric. Handbook No.
552, pp. 69-71). This predation was observed in Greenwood Village, a suburban area
southeast of Denver, in Arapahoe County. Wasps began arriving at the ultraviolet light
about 15 min before sunrise and returned continually until later afternoon, searching for
moths. The wasps patrolled even on days when the ultraviolet light was not used the
previous night. Although only one wasp was observed at a time, it is likely that more
than one individual was involved in the attacks.

Moths attacked by Vespula pensylvanica were resting either on the white sheet behind
the ultraviolet light, or on the house wall adjacent to the sheet. The wasp attacked by
stinging a moth once in the abdomen. Some moths jumped and flew away, but others
fell to the ground, flapping their wings. The wasp then followed the moth to the ground
and stung it several more times. After the moth stopped moving, the wasp bit the moth’s
wings off at the base, and ate the body. Usually (n = 5), the abdomen and most of the
thorax was eaten by the wasp, leaving only the tougher parts of the thorax and head. In
one case, the wasp flew off with the prey.

The five species observed to be victimized by V. pensylvanica were: Nephelodes
minians Guenee., Pseudaletia unipuncta (Haworth), Heliothis zea (Boddie), Trichoplusia
ni (Htibner)(identified in Covell, C. V., 1984, A field guide to the moths of eastern North
America, Houghton Mifflin, Boston, pls. 22, 29, 31), and Catocala faustina Strecker
(identified in Holland, W. J., 1968, The moth book, Dover, Toronto, pl. 33). The successful
attack on the Catocala species was surprising because C. faustina is much larger than
the wasp. Other moths present but not attacked included several larger Catocala species
and one unidentified, blackish Noctuidae.

Two other published observations of vespid wasps preying on adult Lepidoptera were
recorded by S. H. Scudder (1889, The butterflies of the eastern United States and Canada,
with special reference to New England, published by the author, Cambridge, MA), who
observed a Liminitis arthemis (Drury) in the clutches of a large vespid wasp that had
seized its prey as it sunned on a road (p. 1612). Scudder (p. 1217) also recounted an
earlier observation made in England by G. Newport (1863, Trans. Entomol. Soc. Lond.
1:228-230), who reported a successful attack by Vespa vulgaris on Pieris rapae. Although
this is not the first published observation of a vespid wasp preying on adult Lepidoptera,
it is the first observation of a vespid preying on moths that have been attracted to UV
lights (Frank, K. D., 1988, J. Lepid. Soc. 42:63-93). (For further observations of inver-
tebrate predation of Lepidoptera, see Nielsen, M. C., 1977, The Great Lakes Entomol.
10:113-118.)

I thank Richard S. Peigler, Denver Museum of Natural History, for his invaluable help
and advice in preparing this paper. Five voucher specimens of V. pensylvanica were
deposited in the Denver Museum of Natural History.

ANDREW D. WARREN, 9951 East Ida Place, Englewood, Colorado 80111.
Received for publication 9 November 1989; revised and accepted 14 February 1990.

VOLUME 44, NUMBER 1 33

Journal of the Lepidopterists’ Society
44(1), 1990, 33-34

LYCAENA HYLLUS (CRAMER) (LYCAENIDAE): NEW HOST AND
MATING BEHAVIOR IN A DROUGHT-INDUCED
POPULATION EXPLOSION

Additional key words: dispersal, parasitism, Aprostocetus, Eulophidae, Polygonum
natans.

On 24-26 August 1988, we observed a large population of the bronze copper, Lycaena
hyllus (Cramer), in a wet meadow approximately 4—5 ha at the north end of Carnelian
Lake, Stearns County, Minnesota. The lake has no natural outlet and fluctuations in the
lake level are due to differences between precipitation and evaporation. The area was
observed again in August 1989. Weather conditions during both years were warm and
sunny with daytime high temperatures of 23-26° C.

In 1988 the wet meadow inhabited by L. hyllus consisted of a dense herbaceous
community of Polygonum coccineum Muhl., P. natans Eat., P. persicaria L., Rumex
spp., grasses, and sedges. P. coccineum grew in dense stands up to 1.5 m tall. The shorter
P. natans covered areas as large as 9 x 15 m. First instar as well as older larvae and
pupae of the bronze copper were found on P. natans. This is a new larval foodplant
record for L. hyllus. Plants previously reported as larval hosts are Rumex crispus L., R.
obtusifolius L., R. longifolius, R. patientia L., and P. coccineum (Scott, J. A., 1986, The
butterflies of North America: A natural history and field guide, Stanford Univ. Press,
Stanford, California, 583 pp.). Both P. coccineum and P. natans exhibited extensive
feeding damage assumed to have been caused by larvae of the bronze copper. We did
not observe females ovipositing on either plant but one was seen crawling about the base
of P. natans plants. On 13 September 1988, 35 adult parasitoids of an undetermined
species of Aprostocetus (Hymenoptera: Eulophidae) emerged from two pupae collected
on P. natans. This is the first report of L. hyllus as a host of any species of Aprostocetus
(M. E. Schauff, Systematic Entomology Laboratory, PSI, pers. comm.).

Males and females of the bronze copper nectared on the abundant flowers of P.
coccineum. Both males and females rested and basked on P. natans with open wings;
the males apparently engaged in perching behavior to await females. Perching behavior
is typical of species in which the distribution of foodplants throughout the normal flight
area is spotty (Scott, J. A., 1975, J. Res. Lepid. 14:1—40). Males in the vicinity of P. natans
behaved markedly differently from those on the taller P. coccineum. Large numbers of
males, estimated between 15-25/10 m2, were observed patrolling 0.5-1.5 m above large
patches of P. natans. Many fewer females were observed in these areas. Males, which
did not interact outside of P. natans patches, sometimes chased each other in groups of
10 or more and often reached heights of 2-3 m. Males flew into and crawled beneath P.
natans plants and several were observed to land on pupae attached to leaves, suggesting
that females probably are mated soon after emergence.

Courtship behavior and mated pairs were observed from 0930-1200, but were more
common from 1300-1500. Males sometimes chased females, but behavior that culminated
in mating was not observed. Only 10 mated pairs were encountered in this gigantic
population, suggesting intense competition among males for receptive females. Males that
attempted to mate landed next to a resting female and both sexes began to flutter their
wings rapidly. The male moved rapidly around the female and, when behind her, placed
his head and antennae beneath her wings, i.e. male nudging (Scott, J. A., 1973, J. Res.
Lepid. 11:99-127; Scott, J. A., 1974, Pan Pac. Entomol. 50:9-22). The female’s abdomen
remained horizontal to the leaf surface throughout the entire courtship. Attempts by the
male to copulate by moving the abdomen laterally were apparently blocked by the open
position and fluttering of the female’s wings (Scott, J. A. & P. A. Opler, 1975, J. Lepid.
Soc. 29:63-66). Female fluttering is a rejection dance in Lycaena (Scott, J. A., 1974, J.
Lepid. Soc. 28:64-72), indicating either that females were not old enough to mate or that
they had previously mated. Occasionally the male closely followed the female down into

34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

the vegetation. After approximately 1 min of this rejection dance by the female, the male
flew. Five males in succession were rejected by one female. Males that landed near a
mated pair flew off after several seconds of wing fluttering by all three.

Estimates of the density of adult butterflies in the wet meadow in August 1988 ranged
from 1 to 25/10 m? (1000 to 2500 adults/ha) in areas with large P. natans patches. The
population explosion of L. hyllus may have resulted from the effects of the extreme
drought conditions throughout central Minnesota during the spring and summer of 1988,
which exposed shoreline that was colonized by P. natans and P. coccineum. These
Polygonum have both terrestrial and aquatic forms (Gleason, H. A. & A. Cronquist, 1963,
Manual of vascular plants of northeastern United States and adjacent Canada, D. Van
Nostrand Co., Princeton, NJ, 810 pp.) enabling them to invade rapidly and dominate.
We did not observe large numbers of bronze coppers in 23 previous years of normal and
above normal precipitation. The presence of several female bronze coppers visiting flowers
approximately 1 km from the meadow suggests that dispersal may be an important means
by which this butterfly exploits temporary wetland habitats.

After twelve additional months of below average precipitation, the lake level had
dropped several more feet by 20 August 1989, exposing additional shoreline. Areas that
had been newly exposed in 1988 were extremely dry. Plant diversity had increased at
the expense of the Polygonum species. The dense stands of P. coccineum observed in
1988 were greatly reduced except for newly exposed areas of the slough and shoreline.
Searching the area on 20, 21, 22, and 26 August 1989 revealed only 10 adult male L.
hyllus. Explanations of the population crash may include parasitism and a reduction of
suitable larval foodplants.

We thank Dr. M. E. Schauff, Research Entomologist, Systematic Entomology Labo-
ratory, Plant Sciences Institute, USDA, Beltsville Agricultural Research Center, Beltsville,
Maryland for identification of the Aprostocetus sp. parasitoid and Dr. James A. Scott,
Lakewood, Colorado, for reviewing the manuscript.

DavipD B. MACLEAN, Department of Biological Sciences, Youngstown State Univer-
sity, Youngstown, Ohio 44555 AND BONNIE K. MACLEAN, Department of Biology, Thiel
College, Greenville, Pennsylvania 16125.

Received for publication 21 November 1988; revised 18 September 1989; accepted 15
January 1990.

Journal of the Lepidopterists’ Society
44(1), 1990, 34-36

SEASONAL VARIATION OF OCCURRENCE OF DEFORMED COCOONS
OF THE TASAR SILK MOTHS ANTHERAEA MYLITTA (DRURY)
AND ANTHERAEA PAPHIA (L.) (SATURNIIDAE) IN INDIA

Additional key words: bipupate, bishellate, flimsy cocoons.

The tasar silk moth, Antheraea mylitta (Drury), is trivoltine in India and is reared in
the Rainy season (July-August), Autumn (September—October), and Winter (November—
December). By contrast, Antheraea paphia (L.) is reared only during the Autumn season;
although it multiples in nature during the Rainy and Winter seasons, its economic per-
formance at these times is poor. Tasar culture is an age-old practice in Orissa and good
cocoons are more highly valued in tasar commerce than deformed cocoons because of
the former’s better reelability (ability to be spun onto a reel with a continuous filament
for production of fine fabrics). Although the occurrence of deformed cocoons in the tasar
crop is a common phenomenon, the seasonal variation of their occurrence has not been

VOLUME 44, NUMBER 1 35
TABLE 1. Seasonal variation in percentage occurrence of deformed cocoons of A.
mylitta' and A. paphia.!

; : J Deformed cocoons (%)
Tasar species with commercial

Season names of the seasonal crops Bipupate Bishellate Flimsy Total
Rainy A. mylitta (Ampatia) 0.047 0.698 6.130 6.875
A. paphia (Godamodal) 0.008 0.154 0.125 0.287
Autumn A. mylitta (Daba) 0.060 1.073 25.912 27.045
A. paphia (Bogei) 0.021 1.868 10.412 12.301
Winter A. mylitta (Jadei) 0.131 1.479 47.559 49.169
A. paphia (Patrajhada) 0.005 0.132 0.122 0.259
2N = 10,000

documented, which creates confusion in commercial sectors during grading and trading.
This study documents the relative abundance of deformed cocoons produced in different
rearing seasons of the year.

Because of the availability of both A. mylitta and A. paphia, the important tasar zones
of Singda, Kendujuani, Thakurmunda, Sarat, and Kaptipada in the Mayurbhanj district
of Orissa were chosen for this study. Ten thousand cocoons of both A. mylitta and A.
paphia were collected at random in five equal groups from the above five localities during
each of the three rearing seasons in 1987. The Rainy season and Winter cocoons of A.
paphia were collected from natural populations in the forests of the study sites. All other
samples were taken from commercial cultures. Deformed cocoons were sorted from good
cocoons in each sample group.

Deformed cocoons were classified as either bipupate (a single cocoon having two pupae),
bishellate (two cocoons joined by their shells, which interrupts the continuity of the silk
filament), or flimsy (cocoons having thin and unsubstantial shells without strength or
solidarity) (Nayak, B. K., M. L. Gupta, B. C. Guru & B. N. Satpathy, 1987, Towards
classification of cocoons in tasar silk insect Antheraea mylitta Drury (Lepidoptera: Sa-
turniidae), Sericologia 27:505-512). The mean percentages of each class of deformed
cocoons during each season were calculated and the data were analyzed by season and
by species (Table 1).

In A. paphia, the percentage of deformed cocoons was highest (12.30%) during the
Autumn Season, followed by Rainy Season (0.29%) and Winter Season (0.26%) (Table 1).
In A. mylitta, the highest percentage (49.17%) of deformed cocoons appeared during the
Winter season, followed by Autumn season (27.05%) and Rainy season (6.88%). Trends
in the occurrence of deformed cocoons of both A. paphia and A. mylitta were found to
be similar with respect to each type of cocoon deformity.

The percentage of flimsy cocoons in A. mylitta increased dramatically from the Rainy
season (6.13%) to Winter Season (47.60%), whereas the number of flimsy cocoons was
negligible (0.12%) in natural populations of A. paphia during the Rainy and Winter
seasons. The maximum number of deformed cocoons of A. paphia occurred during Au-
tumn Season and was probably due to its rearing in semidomesticated conditions. Such
rearing is against its natural tendency, as it usually grows only during the Rainy and
Winter seasons. In the case of A. mylitta, the percentage of deformed cocoons was much
greater and might result from the frequent handling of the silkworm during the semi-
domesticated rearing.

The higher frequency of deformed cocoons in the Winter Season might result from
physioclimatic stress at the onset of Winter in the places of study. Nayak et al. (Nayak,
B. K., A. K. Dash, P. K. Das, A. K. Sasmal & B. N. Satpathy, 1986, Sex association in
doubled cocoons of tasar silk insect Antheraea mylitta Drury (Lepidoptera: Saturniidae),
Sericologia 26:285-290) assumed that physiological, physical, or environmental stress
causes formation of bipupate (double) cocoons. However, studies should be carried out

36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

to determine the effects of varying climatic conditions and of handling of silkworms
during rearing and spinning on the frequency of deformed cocoons.

A. K. DasH* AND B. K. Nayak, State Sericultural Research Station, Baripada-757001,
Orissa, India.
* Present address: Dept. of Zoology, Dr. J. N. College, Salt Road, Balasore-75601, India.

Received for publication 26 March 1989; revised 12 June 1989; accepted 2 September
1989.

BOOK REVIEWS

Journal of the Lepidopterists’ Society
44(1), 1990, 37-38

CIE GUIDES TO INSECTS OF IMPORTANCE TO MAN. 1. LEPIDOPTERA, by J. D. Holloway,
J. D. Bradley, and D. J. Carter (C. R. Betts, ed.). 1987. CAB International Institute of
Entomology, London. Distributed by CAB International, 56 Queen’s Gate, London SW7
5JR, United Kingdom; and 845 North Park Avenue, Tucson, Arizona 85719, USA. 262
pp., many unnumbered illustrations, 19.3 x 26.7 cm. Hard cover, ISBN-0-85198-594-6,
£25.00 ($52.25); spiral bound, ISBN-0-85198-594-7, £14.00 ($29.00).

Fortunately for lepidopterists, the first guide produced by the Commonwealth Institute
of Entomology (CIE) treats the Lepidoptera. It is a good, generalized introduction to the
order that addresses characterization and recognition for the world fauna of higher taxa
down to the family level, and for several large families, to the subfamily level. In this
effort the work is a boon to students and nearly unique for contemporary treatments. It
was developed as a manual for students of the CIE training course for recognition and
handling of Lepidoptera.

The introduction treats characterization of the order, discussion of injurious and ben-
eficial species, problems associated with identification, collection and preparation of spec-
imens, and a checklist of higher taxa. I found myself agreeing with the caveat concerning
the need to exercise care to ensure that specimens are properly handled, mounted, and
labeled so that as many characters as possible are retained. Directions for dissecting and
mounting genitalia on slides are perhaps not quite adequate for the uninitiated who does
not have access to a tutor, but they suffice. Euparol is indicated as the mounting medium
of choice; Canada balsam should be recognized as another long-term mounting medium.
A checklist of higher taxa presents useful information on numerical size of families, their
geographic distribution, and their current placement within superfamilies, infraorders,
and suborders.

Suborders, infraorders, and divisions are briefly characterized following N. P. Kristensen
(1984, Steenstrupia 10:141-191). Note that 99% of the Lepidoptera are in the division
Ditrysia of the infraorder Heteroneura and suborder Glossata. Categories to division are
keyed in four couplets; several taxa are grouped in the same couplet. Adult ditrysian
Lepidoptera are keyed to family using characters of the head, thorax, wings, and abdomen
when possible. Occasionally, genital characters are used. All characters are illustrated
throughout the key. Inevitably, some specimens will not pass through the key “correctly,”
and considerable practice with specimens and access to an array of identified specimens
are useful adjuncts to the process. Identification of Lepidoptera is not easy—it requires
considerable experience—but it is made easier by concentrating on a limited geographic
area.

Most of the text is devoted to characterization of families and subfamilies; John Bradley
is responsible for the “microlepidoptera” and Jeremy Holloway for the “macrolepidop-
tera.” Each does a fine job of summarizing characters, capturing nuances of significant
features that are useful but not invariably diagnostic, balancing various authors’ views of
classification, illustrating useful characters, citing pertinent literature [however, Der-
zhavets (1984) cited on page 147 was omitted from the references section], and noting
representative species. Many will be surprised that nearly all examples are drawn from
faunas other than the New World. David Carter developed the illustrated key to last
instar larvae and the family characterizations based on this stage. The key is usable but
could afford a few more illustrations of characters and character states. Because I identify
1500+ mostly non-North American microlepidopteran larvae each year, I have become
addicted to the illustrated key by F. W. Stehr and P. J. Martinat (1987, pp. 306-340, in
Stehr, F. W., ed., Immature Insects), which is particularly well illustrated, as a preliminary
source for family level identification.

A short glossary is helpful. The 21-page list of references (nearly all post 1950) is very
useful for anyone attempting to learn more about classification of the order.

38 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

I strongly recommend this book to those lepidopterists who want to stray beyond the
butterflies and skippers for a general introduction to this extremely diverse group.

RONALD W. HonpGEs, Systematic Entomology Laboratory, USDA, % U.S. National
Museum of Natural History, MRC-168, Washington, D.C. 20560.

Journal of the Lepidopterists’ Society
44(1), 1990, 38

NORDEUROPAS PRYDVINGER (THE OECOPHORIDAE OF NORTHERN EUROPE), by Eivind
Palm (in Danish with brief English summary for each species). [DANMARKS DYRELIV,
vol. 4]. 1989. Fauna Boger, Copenhagen. Distributed by Apollo Books, Lundbyvej 36,
DK-5700 Svendborg, Denmark. 247 pp., 214 text figs., 199 maps, 25 color plates. Hard
cover, 17.5 x 25 cm, ISBN 87-87738-08-6, Danish kroner 420 (plus postage) (about $56.00).

This work is a manual of the Oecophoridae of Scandinavia, areas immediately adjacent
to the Baltic Sea, and northern Netherlands with particular emphasis on Denmark. It has
an introduction to the family, its zoogeography, morphological characters of adults,
immatures, ecological habitats, phenology, and economic importance. It has a map of
the region to show the counties/districts of the countries and one at a larger scale of
Denmark. Subfamilies and genera have identification keys, but species do not. For genera
the treatment consists of a brief summary of distribution, number of species, diagnosis,
differentiation from closely allied taxa, discussion of immature stages and hosts; for species
it consists of differentiating characters, extensive statement of distribution, and bionomics.
Each species has a dot map to indicate verified distribution data by county /district for
the entire region, and for 80 species there is an additional map showing more specific
localities within Denmark. Illustrations generally are very good, particularly the eight
color plates of adults. Each species has at least one specimen shown in color, sometimes
more when the species is variable or when the sexes are dimorphic or dichromatic.
Differences between closely similar species are illustrated by male and/or female genitalia,
monochrome plates of wings with key characters indicated, or line drawings of specific
structures. Occasional habitat photographs to indicate where a species has been found
are interesting but not always top quality.

Scandinavia has an oecophorid fauna of 119 species as compared with approximately
226 in America north of Mexico. Within the family, representation by subfamily (number
of North American taxa in parentheses) differs greatly between the two areas: Depres-
sariinae 74 (108), Ethmiinae 6 (50), Peleopodinae 0 (2), Stenomatinae 0 (24), Oecophorinae
35 (41), Deuterogoniinae 1 (0), Chimabachinae 3 (1), reflecting both Scandinavia’s north-
ern location relative to the Palearctic Region and the development of Stenomatinae and
Ethmiinae in the northern Neotropical and southern Nearctic Regions. The distribution
maps show very clearly that most species occur within the southern %-% of the area;
only Agonopterix arctica (Strand) occurs in the north and not in the south, and only a
limited number of species occur throughout Scandinavia.

Perhaps because the volume is a manual, complete lists of synonyms and literature
citations to them are missing. I was puzzled by the “Literature” section not being al-
phabetically arranged throughout. It appears as though items were found after the initial
numbered list was developed and added at the end; thus, there are four alphabetically
arranged lists within this section.

The English summary for each species treats distribution, frequency of occurrence,
habitat(s), and when and where adults may be collected. Although the text is in Danish,
reference to the illustrations, similarity of many words to English cognates, and equivalents
in scientific terms, make the content generally available to an inquiring English reader.

I recommend this work to everyone interested in gelechioid moths in particular, and
Lepidoptera in general. The author and editor are to be commended for this fine volume
and for the execution of the series.

RONALD W. HODGES, Systematic Entomology Laboratory, USDA, % U.S. National
Museum of Natural History, MRC-168, Washington, D.C. 20560.

VOLUME 44, NUMBER 1 39

Journal of the Lepidopterists’ Society
44(1), 1990, 39-40

SATURNIIDAE: ECOLOGICAL AND BEHAVIORAL OBSERVATIONS OF SELECT ATTACINI, by
Robert D. Weast. 1989. Published privately by the author, whose address is: 5324 NW
78th Court, Johnston, Iowa 50131. 53 pp., 3 color plates, 1 text fig. Soft cover, 21.5 x 28
em, no ISBN number, $16.90 (postpaid).

With the increasing availability of word processing programs and desk-top publishing
on personal computers, it came as no surprise to see this vehicle used to publish an
entomological work. Such methods need to be used cautiously to publish scientific work,
and apparently the result does not constitute valid publication under the International
Code of Zoological Nomenclature (1985, Articles 8 & 9). Since Weast did not propose
any new names, the question of whether this book (some may prefer to call it a booklet)
is a valid publication is not so critical. However, should this outlet become commonly
utilized, the tempting and obvious opportunity to circumvent peer review and editorial
input of the manuscript can lead to reduced quality of scientific literature. Even where
manuscripts are refereed by other scientists, I would recommend that authors abstain
from using desk-top publishing for work that is intended to have widespread and per-
manent application.

The vinyl cover, textured bond paper, and plastic spine are of good quality. The three
color plates are glossy photographic prints measuring 18 x 25 cm and are attached loosely
onto the pages with rubber cement. They should have been dry mounted. I will protect
my copy by sheathing those three pages individually in clear plastic. The 13 photos in
these three plates have great aesthetic and scientific value. The book will be fine for
personal libraries, but less suitable for loaning out of public libraries.

The text provides discussions filled with details pertaining to field observations, rearing
procedures, geographical localities, and ecological aspects. I learned a lot. I am sure that
the great majority of statements in the text are accurate and factual. Weast generously
gives credit throughout the text to all his correspondents for their ideas and discoveries;
many authors are much less willing to credit others. Weast has made many trips to study
Saturniinae, and in this book shares a lot of his personal observations made in southern
Texas, Arizona, France, etc., effectively conveying his life-long enthusiasm for these big
moths. He has carried out various experiments to introduce/colonize certain species (Samia
cynthia, Callosamia promethea, Automeris io) into urban and rural locales, mostly in
Iowa, where they did not occur formerly. Some data are qualitative, but he provides
enough quantitative data in tabular form, combined with insightful discussion, that I
believe this work will be useful to persons studying saturniid population ecology. Weast
is to be commended for disproving the myth that everything is already known about the
large, common saturniids.

The organization is weak in some ways. Many disjunct points are scattered throughout
the text. Some paragraphs consist of a single sentence. Some are arranged under inap-
propriate subheadings. Certain examples detract from the scientific image of the work;
these include referring to vertebrates as vertebrae (in a discussion on predation) and
misusing the word speciation to convey a taxonomic rather than an evolutionary concept.

I was pleased to find satisfactory answers to several questions I have long puzzled over.
These include: Why has Samia cynthia died out in certain cities and not others? Does S.
cynthia really occur in Savannah, Georgia, and throughout most of Ohio and Indiana?
Is the widely published report of an urban population of C. promethea specializing on
lilac in Milwaukee, Wisconsin, really valid? What is the western range of C. promethea
along the region between Minnesota and central Texas?

Most errors in the text are minor, but I feel the need to rectify a few. Weast consistently
misspells Lemaire as LeMaire and Veracruz as Vera Cruz. The 1906 and 1907 citations
in his bibliography were published by Caroline Soule, not Fred Tepper. Israel is incorrectly
referred to as Palestine. Lastly, I suspect it unwise to publish the particulars when your
friends repeatedly break federal law by collecting in national parks.

I consider the book to be overpriced, but the price is still generally affordable and
substantially lower than the cost of most scientific books on the market. Despite the

40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

shortcomings I have outlined above, it offers a lot of value and interest. I recommend it
to the many lepidopterists who are enthusiastic in their rearing and field collecting of
larger Saturniidae.

RICHARD S. PEIGLER, Department of Zoology, Denver Museum of Natural History,
2001 Colorado Boulevard, Denver, Colorado 80205.

Journal of the Lepidopterists’ Society
44(1), 1990, 40

THE BUTTERFLIES OF MANITOBA, by Paul Klassen, A. Richard Westwood, William B.
Preston, & W. Brian McKillop. 1989. Manitoba Museum of Man and Nature, Winnipeg,
Manitoba, Canada. vi + 290 pp. with 27 color plates, distribution maps, and various text
illustrations. Soft cover (plastic laminate), 20.5 x 25 cm, ISBN-0-920704-16-6; $18.66
(U.S.) + $1.70 shipping.

This book is a comprehensive treatment of the butterfly fauna of Manitoba. The
- introductory pages include such topics as butterflies in perspective, nomenclature, the
differences between butterflies and moths, butterfly life histories, Manitoba as an envi-
ronment for butterflies, Manitoba biomes, butterfly distribution, conservation, and study,
with a brief historical perspective on early lepidopterists in Manitoba. The scientific
treatment begins with the Hesperioidea and concludes with the Danaidae. Then follows
a listing of potential and dubious species, a comprehensive list of references, and a glossary
of terms. The six appendices include butterfly collecting methods, a list of larval and
adult foodplants, a checklist of Manitoba butterflies, geographic locations (latitude and
longitude) of collection sites with a map of major areas, summary of flight periods, and
the Lepidopterists’ Society’s “Statement of the Committee on Collecting Policy.” An index
of species concludes the book.

Each species entry includes information and topics in the following order: common
name, scientific name (author and date) with basic synonyms, plate and figure numbers,
description, life cycle, habitat, range, Manitoba records, subspecies, general remarks. A
dot distribution map accompanies each entry, with the dots color-coded for each family.
The color plates are well done, and, for the most part, the specimen colors are natural
(for museum specimens), while the plate backgrounds vary from white to antique-white.
Specimens are shown life-size with three illustrations for each species: male dorsal, female
dorsal, and ventral (male or female). Scientific nomenclature follows Klassen’s 1984 Man-
itoba checklist published in J. Lepid. Soc. (38:32-39), and differs significantly in some
groups from the 1981 Miller and Brown catalogue and other recent publications. The
Miller and Brown names, however, are shown as synonyms. This is the only area in which
I offer adverse criticism, as yet another set of scientific names is introduced, which only
further muddles nomenclature.

Manitoba encompasses six vegetation zones ranging from tall grass prairie through
boreal coniferous forest to arctic tundra. Consequently there is wide diversity among the
144 butterfly species recorded from this province. Although several subspecies have been
described from Manitoba, there are no unique species. The dominant groups are Nym-
phalidae (40 species), Hesperioidea (33), Lycaenidae (27), and Satyridae (19).

The Butterflies of Manitoba, with an overall format of 8 x 10’, is nicely produced
and has a pleasing aspect regarding the general layout. The text provides a wealth of
information about Manitoba butterflies and is certainly “must” reading for any collector
wishing to visit this Canadian province. This book is the most recent addition to the
collection, from various publishers, of North American regional butterfly works that have
appeared during the past several years. It will be a valuable addition to the bookshelf of
any collector with an interest in the North American fauna.

CLIFFORD D. FERRIS, Bioengineering Program, University of Wyoming, P.O. Box
3295, University Station, Laramie, Wyoming 82071-3295.

VOLUME 44, NUMBER 1 4]

Journal of the Lepidopterists’ Society
44(1), 1990, 41

MARIPOSAS DE VENEZUELA, by Theophile Raymond. 1982. Ediciones Corpoven, Caracas,
Venezuela. 275 pp., 53 plates. Hard cover, 29.5 <x 23.7 cm, approx. $30.00.

Theophile Raymond was a classic naturalist about whose life we know very little, except
for his scientific accomplishments during the late 19th and early 20th centuries. His
research in natural history was diverse and notable and his role in the early scientific
development of Caracas is well recognized, but he is best known for his studies on
butterflies.

Young Raymond arrived in Venezuela from Martinica, probably in the early 1890's,
and it is presumed that he collected butterflies ever since. His premature death, in 1922,
left fatherless perhaps one of the best and largest butterfly collections in Venezuela.
Nothing is known about what happened to this collection, and the reason why it was not
preserved by any of his notable scientist coworkers remains an enigma. Today there is
no doubt that it was lost forever.

His manuscript Fauna Entomoldégica Venezolana. Lepidépteros Rhopaloceros, written
in 1902, which remained undiscovered for many decades, now is available to lepidopterists
in the present excellent facsimile edition entitled Mariposas de Venezuela. Included in
this edition is an interesting biographical sketch of Raymond, written by the now deceased
Dr. Francisco Fernandez-Yépez, a pioneer Venezuelan lepidopterist. Fernandez-Yépez
also made modifications in the original text, added new identifications, and corrected
errors. Fifty-three full-color original plates, drawn and painted by Raymond, are included.
The first 47 plates represent the most common butterflies (and some Geometridae and
Uraniidae) of Venezuela. The remaining six plates illustrate members of the family
Sphingidae. Only the Hesperiidae and some members of the Lycaenidae and Satyridae
are not classified, because Fernandez-Yépez believed that more time and study were
required to properly identify the specimens illustrated.

The original text of 1902 was hand written in beautiful classic script, providing a strong
indication of the care and dedication he put into his scientific work. The calligraphy of
the latter pages tends to be notoriously impaired, possibly due to his battle with tuberculosis
and his interest in concluding the work rapidly. A sanitary seal printed in the original
book is witness to his lost battle. Despite the fact that it represents an unfinished work
and that it far from covers the entire butterfly fauna of Venezuela, it remains a valuable
scientific and historic contribution to Venezuelan and neotropical entomology. It is a
unique and rare documentation of the butterfly fauna of Caracas and neighboring areas
of yesteryear, before strong demographic influences began to change the abundance and
diversity of this group of colorful insects.

Mariposas de Venezuela, published in 1972 by Michael Schmidt, is primarily directed
at amateur lepidopterists and provides but a poor description of the abundant and rich
fauna of this country. Raymond’s work is a much more professional representation, even
though it was written much earlier. It covers both practical and theoretical lepidopter-
ology, and contains an exceptional recount of the classic taxonomic methods of Linné,
Dégeér, Scopoli, Fabricius, Latreille, Godart, and many other naturalists. Finally, Ray-
mond describes the natural history of thirteen species of Papilionidae, demonstrating a
good capacity for such detailed biological studies.

Undoubtedly, this book is a bibliographic curiosity, but it is also an important work
that reflects both scientific and artistic facets of lepidopterology.

ANGEL L. Vitoria, Museo de Biologia, Facultad Experimental de Ciencias, Univer-
sidad del Zulia, Apartado 526, Maracaibo 4001, Zulia, Venezuela.

42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Journal of the Lepidopterists’ Society
44(1), 1990, 42-43

ATLAS OF THE JAPANESE BUTTERFLIES, edited by Toshio Inomata. Color photographs by
Katsuji Iwao. 1986. Take Shobo Co. Ltd., 7-3, lidabashi 2, Chiyoda, Tokyo, 102 Japan.
500 pp., numerous text figs., 86 color plates. Hard cover, 21.5 x 35 cm, no ISBN number,
£440 (about $704.00).

This authoritative and beautiful work surely must be the ultimate regional butterfly
book. The authors, editor, and publisher have combined the best features one could dream
up, and even a few that you would likely never have envisioned yourself, in creating this
masterpiece on the Japanese butterfly fauna. Under the editorship of Toshio Inomata,
five authors have joined with Katsuji Iwao’s photographic talents to produce this intel-
lectual and visual feast.

The book opens with 86 outstanding color plates of greatly reduced adult figures
illustrating all of the species, subspecies, and forms of the butterflies of Japan. What is
extraordinarily innovative in this work is the lavish use of these color plates to illustrate
the total known range of minute variations within each species. For example, variation
in the first papilionid alone, Leuhdorfia japonica, is illustrated across three and a half
plates, utilizing some 368 (!!) adult specimens of this single species. Male and female,
dorsal and ventral surfaces, are illustrated for all species and subspecies. Lest one feel
that these figures may be too small to depict all essential details, this reviewer may assure
you that he has never seen finer studio photography and photographic reproduction of
adult butterfly figures. All details can be seen accurately. Additionally, the publishers
have reprinted each plate in a huge oversized poster print of approximately 51 x 71 cm,
with the same immaculate care of reproduction shown in the original plates! These 86
poster-size plates come included with the case-bound volume of the book, and are placed
in a special protective clothbound wooden case with its own lock and appropriate fasteners!
To top all of this, an acrylic and aluminum frame is provided (stored inside this large
plate case) to mount your “print of the month” on the wall of your office or living room
to admire and enjoy! Perhaps the reader of this review is now beginning to realize why
this book costs over $700 U.S.

The text of the Atlas is presented in four principal parts. Part I consists of a synonymic
catalogue of the butterflies of Japan by Toshio Inomata and Yuji Itagaki. This excellent
section of 72 pages is in English and includes all names that have been applied to genera,
species, and subspecies. It also includes a line or two on the distribution of each species
and subspecies throughout Japan and the rest of the world. From all indications, the
coverage is complete and without any obvious errors.

Part II is a detailed discussion of the species of Japanese butterflies, authored by Toshio
Inomata, Masumi Ikeda, Yasunori Kishida, and Yoshitaka Kanada. This section is almost
entirely in Japanese; only the scientific names and authors’ names and dates, along with
plate and figure numbers, are given in English. The authors include synonyms and present
a detailed discussion of distribution, including excellent distribution maps for all the
islands of Japan.

Part III, by Toshio Inomata and Yuji Itagaki, consists of one of the most innovative
features in any general work on butterflies: namely, a treatment of the type specimens
of the Japanese butterflies and their institutional depositories, with monochromatic plates
of the types and of aberrant forms. Besides illustrating both dorsal and ventral surfaces
of 257 type specimens, with most specimens enlarged beyond life size, the authors provide
an exhaustive table, mostly in English, listing the number of types, kind of types (holotype,
syntype, lectotype, neotype), number of males and females, type locality, date of types,
and location of types. This invaluable section will be of use to all museums and systematists
working on Asian butterflies.

Part IV, entitled “Butterflies and International Code of Zoological Nomenclature, ” is
in Japanese and is written by Toshio Inomata. It not only discusses the I.C.Z.N., but shows
how our nomenclature has changed from the time of Linnaeus to today’s detailed no-
menclatural descriptions, construction of synonymies, and other useful tools of the modern
systematist.

VOLUME 44, NUMBER 1 43

At the end of the book is a list of new names and other nomenclatural changes in
the work. The new genus described here is Sibatinozephyrus Inomata, of which the type
species is Zephyrus fujisanus Matsumura, 1910 (Lycaenidae: Theclinae). Its distribution
is confined to Japan. Finally, an extensive section in Japanese presents locality data for
the specimens in the 86 color plates. A complete index to Latin scientific names and to
Japanese names is also included.

As treated in this book, the Japanese butterfly fauna includes 19 species of Papilionidae,
30 Pieridae, 77 Lycaenidae, 2 Libytheidae, 22 Danaidae, 63 Nymphalidae, 29 Satyridae,
and 37 Hesperiidae. The rich Japanese fauna of 279 species of butterflies is therefore
comparable to what occurs in large mountainous temperate-zone sections of western
North America, such as the states of California or Colorado. The remarkable richness of
the Lycaenidae and the Satyridae in Japan is especially notable. The Atlas authors and
illustrators have made it possible for workers in the Palearctic and the Nearctic to learn
much about their favorite groups by the study of this incredibly complete and magnifi-
cently wrought volume and its accompanying plates.

Even for the marvelous books on butterflies now being published around the world,
this volume from Japan sets a new standard to which publishers may aspire. While the
extremely high price puts the volume out of reach of most individual lepidopterists, the
work definitely should be purchased by institutional libraries and museums around the
world to make its uniquely complete coverage of the Japanese butterfly fauna generally
available to researchers and students of Lepidoptera.

THOMAS C. EMMEL, Division of Lepidoptera Research, Department of Zoology, Uni-
versity of Florida, Gainesville, Florida 32611.

Journal of the Lepidopterists’ Society
44(1), 1990, 43-44

BUTTERFLIES OF Laos, by Akira Motono and Noriko Negishi (supervised by Tadahiro
Takakura). 1989. Kirihara Shoten, 2-44-5, Koenj-minami, Suginami-ku, Tokyo 166, Japan.
215 pp., numerous text figs., 6 color plates, 118 half-tone plates. Hard cover, in slip case,
20 x 27 cm. No ISBN number. $50.00 (seamail), $60.00 (airmail) from publisher.

In recent years, a number of books and other publications have appeared on the
butterflies of various countries in Southeast Asia. Several good books are available on
butterfly faunas of Taiwan, India, Malaysia, Thailand, and most recently Nepal. Addi-
tionally, Karal Spitzer and his colleagues in Czechoslovakia have been publishing notable
papers on butterfly communities in southern Vietnam from 1983 to date. The accumu-
lating data on faunistic composition and habitat preferences in the tropical habitats of
these areas are providing important information on the evolution and ecology of tropical
Lepidoptera, in particular pointing out the importance of contiguous undisturbed forest
tracts for some species, and the role of fragments of forests (or natural clumps of trees
and shrubs) as the main and constant sources of species diversity in some of the Southeast
Asian biomes, including even open savanna (Spitzer, K., J. Leps & T. Soldan, 1987, Acta.
Entomol. Bohemoskov. 84:200-207). Thus, besides providing richly illustrated catalogs of
these highly diverse tropical butterfly faunas, these publications are beginning to play an
important role in conservation biology.

Now Japanese lepidopterists once again have proven that their intensive field work
throughout Southeast Asia in pursuit of Lepidoptera, combined with the outstanding
quality of printing offered by a number of Japanese publishers, can produce superb
references on Southeast Asian butterflies. This book by Motono and Negishi is the first
significant work on Laotian butterflies, supplementing the substantial work already pub-
lished on the butterfly faunas of adjacent Thailand and Vietnam. The Butterflies of Laos
illustrates and describes very briefly 512 species, including color plates of some of the
larger and more common butterflies of Laos. The text and monochrome plates include

44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

44 species of Papilionidae, 39 Pieridae, 20 Danaidae, 123 Nymphalidae, 38 Satyridae, 9
Amathusiidae, 4 Libytheidae (this must be a record for the number of snout butterflies
found in one fairly small geographic area!), 6 Riodinidae, 131 Lycaenidae, and 98 Hes-
periidae. Of these, 62 are pictured in color (dorsal surfaces only), and all species are
pictured in half-tone plates (usually the dorsal and ventral surface of the male of each
species, or occasionally the female, but rarely both sexes). Each section of the text is
printed in both English and Japanese.

The book begins with several pages of introduction to the history of collecting in Laos
and discusses the geography, seasonality, and topography of that country. A complete list
of the 512 species in the butterfly fauna covered in this book is then given, with an index
to the illustrations. The species included are only those that are in the authors’ collections
and in those of their friends, so the list is admittedly incomplete. In the explanatory notes
on each species that follow, the name, dates of capture, frequency of occurrence, and
collection localities in Laos are given. Although extremely telegraphic in format, this
procedure does allow a great deal of information to be conveyed in the space of relatively
few pages. The half-tone illustrations are excellent and readily allow identification of
virtually all the species of butterflies and many of the skippers figured. The color plates
offer a selection of some of the more beautiful and unusual species found in the country.

‘The book ends with an index to the genus, species, and subspecies names, a two-page list
of references, and a brief biography of each of the two authors and the editorial supervisor
of the project (all three are members of the Lepidopterological Society of Japan). The
book is nicely produced with clear typography, 10 half-tone illustrations of various habitats
in the country, and a map of Laos and adjoining areas.

The map shows that the Lao People’s Democratic Republic occupies a key biogeographic
position between Thailand and Cambodia (Kampuchea) on the south, Vietnam on the
southeast and east, and China and Burma to the northeast and northwest, respectively.
It is thus situated in the center of the Indo-Chinese Peninsula and extends over 1000 km
between latitudes 14° and 22.5°N. The highest point in the country is 2800 m and these
northern central highlands slope gently downward in central and southern Laos to the
east and the west. The country has a marked alternation of wet and dry seasons, with
the rainy season beginning in May and lasting until October. In March and April, before
the start of the rainy season, the weather can be quite warm (22° to 39°C). In December,
January, and February, the temperature drops to between 10° and 29° in southern Laos
and far cooler in upper Laos. This book will help a visiting lepidopterist to plan the best
time to visit the country for a maximum diversity of species, which seems to appear in
March and April as the rains begin, although a great many species are also found from
August through November.

In Macrolepidoptera of the World (1908-28), A. Seitz discussed only 30 species from
Laos. Brief papers since that time, primarily by Japanese workers, have added some
species to this list. This is the first comprehensive and systematic study of the butterfly
fauna of Laos ever published, and, although Laos still remains relatively unexplored, is
an important contribution to our understanding of the distribution and ecology of South-
east Asian butterflies. Obviously, many additional butterflies, probably including unde-
scribed subspecies or even new species, await discovery in Laos as the country becomes
more accessible to future research. This book will do much to stimulate further interest
in the butterflies of the Indo-Chinese Peninsula and deserves a place in the library of
every lepidopterist and institution interested in Old World tropical butterflies.

THOMAS C. EMMEL, Division of Lepidoptera Research, Department of Zoology, Uni-
versity of Florida, Gainesville, Florida 32611.

Journal of the Lepidopterists’ Society
44(1), 1990, 45-55

OBITUARY

JAMES WILSON TILDEN (1904-1988)
A REMEMBRANCE

J. W. “Bill” Tilden passed away on 27 December 1988. He will be missed in many
ways by all who knew him. Bill was a walking encyclopedia of entomological information.
A simple question about insects from student or colleague would invariably bring an
informed and detailed response replete with facts and figures. Arnold “Bud” Applegarth
reported that his stock answer for students asking difficult questions was always, “Go
down the hall and ask Dr. Tilden. He'll be able to tell you that.” Bill’s expertise was not
confined to insects, however. He was also knowledgeable about other invertebrates, birds,
plants, rocks and minerals, and vector control as well.

J. W. “Bill” Tilden, 1986 (photograph by Hazel Tilden).

Bill was possessed of an inquiring mind from an early age and his curiosity knew no
bounds. His personal living “computer” was loaded with information on many subjects
beyond biology. How broad the spectrum and how readily facts were recalled was
impressed upon my wife Eileen and me during the final months of work on A Field
Guide to Western Butterflies. During this period Bill drove down from San Jose to
Watsonville almost every day and we put in long hours on the manuscript, legends, and
plates. However, we did take time out to eat and butterflies were ignored for a time.
Lunchtime conversation ranged far and wide. Among the favored topics were the big
bands and their vocalists, early radio programs, classical music and opera, and movies of
the 20's, 30's, and 40's. Early writers and their publications came in for discussion often.
Because of the fruit we were eating for lunch, old, rare varieties of apples, pears, plums,
and peaches were remembered. Bill had been a “fruit tramp” for several years starting

46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

in the late 20’s and I was now growing many of those old varieties remembered from
childhood in Placer County.

Regardless of the subject that came up, Bill was able to reach back into the deep recesses
of his memory and resurrect the basic facts about the specific subject under discussion.
For a popular song of the 20’s, 30's, or 40’s he might recall such things as the band and
vocalist responsible for its popularity, the year it was introduced or revived, the names
of the members of the band and instrument each played, and in many cases the name
of the hotel or nightclub where the group played regularly. Bill learned to play the slide
trombone in the early 20's, joined the San Francisco Musician’s Union in the late 20’s,
and during the 30’s and 40’s played professionally from time to time with jazz bands in
the Monterey Bay area, the San Francisco Bay area, and at Lake Tahoe.

Rarely did a meal go by without Bill being reminded of a poem or a passage from a
writer such as Burns, Kipling, Tennyson, Whittier, Shakespeare, or perhaps Robert W.
Service. After a moment Bill would recite the passage word for word. As a result of this
indulgence in nostalgia, Eileen and I began to keep what we called our “Ask Bill” list of
questions on a pad near the phone. So we, along with his many friends, miss Bill in many
ways.

Details concerning the Tilden family, Bill's early life, education, and professional career
have appeared in the March/April 1989 issue of the News of the Lepidopterists’ Society
(Smith, A. C., 1989, No. 2, p. 38) and will not be repeated here.

Bill started collecting Lepidoptera seriously in Santa Cruz in 1927 through the influence
of E. A. Dodge and J. P. Strohbeen. His first exchange of specimens took place with Hugh
Gibbon of Miniota, Manitoba, Canada a few years later. When I started corresponding
and exchanging with Bill in 1933 he sent me his “Exchange list of duplicates,” which
then consisted of 158 species of butterflies, 10 moths (saturniids and sphingids), many
skippers, and many noctuids, geometrids and micros. At that time he was also corre-
sponding and exchanging specimens with W. D. Field and V. F. Calkins in Kansas, A.
E. Brower and L. P. Grey in Maine, O. E. Booth in Iowa, H. Glazbrook in Texas, and
Lowell Hulbirt and Dale Bulgrin in California, among others. I first met Bill in the fall
of 1933 when he was working in a cannery in San Leandro.

In 1933 Bill and his brother Tom went to work collecting insects of all orders in large
numbers for L. M. McQuesten of the National Insect Company, Davis, California. Mr.
McQuesten then made up a set of ten large Riker mounts to demonstrate the different
orders of insects, which he sold to schools. Bill continued to supply insect specimens to
McQuesten from time to time for several years.

Again in the summers of 1963 and 1964 Bill took on a professional collecting job for
the newly established California State College (now University), Hayward. He signed a
contract to provide a synoptic collection of mounted and identified San Francisco Bay
Region insects, and large numbers of mounted and preserved (in alcohol) specimens in
certain orders for classroom use, about 4000 specimens altogether. For the most part,
though, Bill collected butterflies, moths, and beetles for his own research and enjoyment
and to discover and publish new information about their distribution, life history, ecology,
habits, and habitats.

Bill had planned to go to the University of California in Berkeley when he graduated
from high school in 1922 but that did not work out. Nevertheless he continued to hope
that it would be possible to continue his education. Over the years several older friends,
including Charles Bowles, M. Soutar, and Dr. Holman suggested repeatedly that he should
get a college education. In the spring of 1938 Bill stopped by our apartment in San Jose.
During the visit he asked us what we (I am not using the editorial plural, but the twin
“we,” as all events prior to 1944 included my twin brother, Edgar A. Smith) thought of
San Jose State and especially its entomology courses. Needless to say, we praised everything
about it highly, especially the teacher, Dr. Carl D. Duncan. We went on to tell him that
the Pacific Coast Entomological Society was having its annual field trip meeting at Alum
Rock Park, hosted by San Jose State’s Comstock Entomology Club on the following
Saturday
introduced him, and he and “Dunc” spent a good part of the afternoon getting acquainted.
They took to each other at once. Bill was favorably impressed and soon made his decision

VOLUME 44, NUMBER 1 47

Spring collecting trip to Patagonia Mountains, Arizona, 1939. From left to right: Ed
Smith. J. W. “Bill” Tilden, Darrel Welch, and Art Smith, standing in front of the 1937
Chevrolet that was flooded while crossing the Sonoita River (photograph by Art and Ed
Smith).

to attend San Jose State. We invited Bill to share our apartment and he moved in with
us for his freshman year and our senior year.

In the late 20’s, the 30’s, and early 40’s, Bill managed to collect throughout most parts
of California. By the late 30’s he was also collecting in the adjacent states. As a matter
of fact, one collecting expedition to Arizona during spring vacation in 1939 came very
close to ending the careers of Bill Tilden as well as Art and Ed Smith and Darrel Welch,
a volunteer assistant. We had had excellent weather and very good collecting in Sabino
Canyon, Madera Canyon, and in the Nogales area. We then headed up a dirt road from
Sonoita that wound up into the Patagonia Mountains along the Sonoita River. It was warm
and sunny when we started and the stream we crossed and recrossed was only a few
inches deep. As we proceeded higher into the mountains the sun disappeared, we saw a
huge black cloud ahead to the south, and the stream crossings seemed to be a few inches
deeper. Thunder and lightning accompanied by light rain then began. We came to a
very broad crossing, but were cautious enough to stop, wade out and check the water
depth. It only seemed deep enough to reach the running board at that time so we proceeded
slowly in low gear. About midstream we could see the water rising rapidly on all sides.
Suddenly the engine drowned out and we sputtered to a halt. We quickly put Darrel
(the lightest person) behind the wheel with instructions to keep it in low gear and keep
pressing the starter (in those days you pressed it with your foot). The remaining three of
us had jumped out and were frantically pushing as hard as we could. By then the water
was above our knees and a few moments later a crest brought it above our hips. With
the combination of pushing and aid from the starter we finally made it to the other side
just before the highest crest of the flash flood arrived. A few minutes later and we certainly
would have been swept away and washed down the canyon. An hour later the sun was
out and drying our belongings, which were spread all over the landscape. A ring of mud
on the inside of the car right up to the lower edge of the windows showed how deep the
water had been and how fortunate we had been!

Bill soon began to collect throughout the country and, in fact, everywhere he traveled.
After signing the contract for the Peterson field guide to western butterflies, Bill began
a program of extensive trips to visit the areas covered by the book—from the middle of
Kansas to Hawaii, and from the Mexican border to Alaska and western Canada. During
the period from 1969 to 1986 Bill, alone or with Hazel and sometimes Bruce as well,

48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

traveled more than 210,000 miles in their van on collecting and field survey expeditions.
This included three trips to Alaska, and numerous trips to the states along the Mexican
border, the northwest, the Rocky Mountains, and the prairie states. He visited every state
and province in the area designated at least once and most of them many times.

During the 60’s and 70’s Bill collected many interesting butterflies in the Lower Rio
Grande Valley of Texas. Among them were a number that represented new United States
records for Mexican species. A representative collection of all those taken at the Santa
Ana National Wildlife Refuge, Hildago County, Texas was deposited with the Refuge
and has now been added to the National Museum of Natural History collection in Wash-
ington, D.C.

After Bill’s heart surgery in 1971, I visited him at Stanford Hospital. He looked up at
me from his hospital bed and said, “Well, Art, I guess I'll never collect another butterfly.”
And he didn't for some time. Then one day, parked by the roadside, Bill saw something
“choice” (as he would say) fly across the road. He grabbed a net and said to Hazel, “If
I can't go after that butterfly, life just isn’t worth living.” And off he went. He continued
to collect until his legs failed to cooperate in 1986.

The J. W. Tilden Lepidoptera Collection was left to the California Academy of Sciences
in San Francisco. Actually, Bill had been transferring portions of his collection to the
Academy since 1946. The total number of Lepidoptera donated to the Academy to date
is 43,547. The collection is very strong in North American butterflies, with Bill’s favorite
families, the Hesperiidae and the Lycaenidae, especially well represented. In the moths,
the Saturniidae, Sphingidae, Arctiidae, and Noctuidae (especially the underwings) are
most numerous. Many exotics from Southeast Asia, Europe, Japan, and Latin America
are also included. Unmounted butterflies and moths in papers, numbering between five
and six thousand specimens will also be turned over to the Academy. Insects of other
orders and miscellaneous other terrestrial arthropods add another 1648 specimens to the
collection.

The Tilden collection includes 95 paratypes in the families Papilionidae, Nymphalidae,
Satyridae, Hesperiidae, Pieridae, and Lycaenidae. One is Mexican, five Canadian, and
the remainder are from the United States.

The specimens (representing over 500 species) used to make the photographic illustra-
tions for the plates in the Houghton Mifflin book, A Field Guide to Western Butterflies
by J. W. Tilden and Arthur C. Smith, are being kept in a separate collection. Plate
specimens from the Smith Collection, including the rare (only 10 others known) hybrid
between the West Coast Lady and the Red Admiral, will go to the Academy at a later
date.

The J. W. Tilden Coleoptera Collection of some 8000 specimens is now on permanent
loan to the California Academy of Sciences and title will be transferred to the Academy
at a future time. Approximately the same number of beetles had been presented by Bill
to the Academy in earlier years, starting in 1946.

Bill was active in many organizations, environmental and historical, as well as biological
in nature. In addition to The Lepidopterists’ Society (Charter Member, 1947; President,
1978) he belonged to the Pacific Coast Entomological Society (Life Member, 19438; Pres-
ident, 1960), California Academy of Sciences (Elected Life Fellow, 1968), Entomological
Society of America, Coleopterists’ Society, Xerces Society, California Native Plant Society,
Santa Cruz Bird Club (Honorary Member), Santa Clara Valley Audubon Society, Point
Reyes Bird Observatory, Santa Cruz Museum Association, Sierra Club, Bohart Museum
Association, Coyote Creek Riparian Station, San Jose Historical Museum Association,
California Pioneers of Santa Clara County, Santa Cruz County Historical Trust, National
Trust for Historical Preservation, Sigma Xi, and the Explorer’s Club, as well as others.

NEw TAXA DESCRIBED BY JAMES WILSON TILDEN
Lepidoptera—Lycaenidae

1. Schellbach’s Copper. Tharsalea arota schellbachi Tilden. 1955. TL: North Rim,
Grand Canyon, Arizona.

2. Lembert's Hairstreak. Callophrys lemberti Tilden. 1963. TL: W. above Tioga Pass,
Yosemite National Park, California.

VOLUME 44, NUMBER 1 49

oo

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be |

. Mansfield’s Hairstreak. Mitoura siva mansfieldi Tilden. 1951. TL: Seven miles west

of Simmler, San Luis Obispo Co., California.

. Pallid Blue. Euphilotes rita pallescens (Tilden & Downey). 1955. TL: Little Granite

Mtns., Dugway Proving Grounds, Tooele Co., Utah.

. Boharts’ Blue. Philotiella speciosa bohartorum (Tilden). 1968. TL: Briceberg, Mariposa

Co., California.

. Behr’s Blue. Glaucopsyche lygdamus incognita Tilden. 1974. TL: Alum Rock Park,

Santa Clara Co., California.

TAXA NAMED IN HONOR OF JAMES WILSON TILDEN

Lepidoptera~Nymphalidae

. Basilarchia tildeni Field. 1934. TL: Vic. Lawrence, Kansas. [Now a synonym of ar-

themis astyanax (Fabricius)].

Lepidoptera—Hesperiidae

. Hesperia comma tildeni Freeman. 1956. TL: Cherry Flat Reservoir, Santa Clara

County, California.

Lepidoptera—Lycaenidae

. Euphilotes enoptes tildeni (Langston). 1964. TL: Del Puerto Canyon, 22 mi. W.

Patterson, Stanislaus County, California.

Coleoptera—Cerambycidae

. Tanyochraethes tildeni Chemsak & Linsley. 1965. TL: Welder Wildlife Refuge, San

Patricio County, Texas.

Lepidoptera—Gelechiidae
New Genus, Tildenia Povolny. 1967. Type Species—Tildenia glochinella (Zell. 1878).

Hymenoptera—Andrenidae

Andrena tildeni Ribble, 1974. TL: Soboda Hot Springs, Riverside County, California.

Lepidoptera—Gracillariidae

. Cameraria tildeni Opler & Davis. 1981. TL: Kings Mountain, San Mateo County,

California.

Additional new insect taxa associated with the name of J. W. Tilden will continue to

appear in the literature for a time as work in progress is completed. These names will
include patronyms and taxa carrying the name Tilden as co-author.

bes CY) OV oe

DES

PRESENTATIONS BY J. W. TILDEN AT MEETINGS OF THE
PACIFIC COAST ENTOMOLOGICAL SOCIETY: SUMMARIZED IN THE
PROCEEDINGS, AND PUBLISHED IN THE PAN-PACIFIC ENTOMOLOGIST

1942. Tilden, J. W. and E. G. Linsley. Range Extensions in Pleocoma. 18:39.

1949. The Insect Fauna of Baccharis pilularis. 25:42. (Formal address.)

1954. Philotes rita associated with Eriogonum wrightii. 30:84.

1959. Alpine Butterfly Associations at Tioga Pass, Yosemite National Park. 35:56.
(Formal address. )

1964. Stylopized Wasp. 40:61.

1967. Food Plant of Vanessa tameamea. 43:91 .

1967. The Kauai Green Sphinx Moth, Tinostoma smaragditis. 43:91.

50

10.

VA

12.

13.

14.

15.

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

SCIENTIFIC PAPERS, NOTES, AND BOOKS By JAMES WILSON TILDEN
1940

. Preliminary List of the Butterflies and Skippers of Santa Clara and Santa Cruz

Counties, California (mimeographed). Natural Science Department, San Jose State
College, San Jose, California.

194]

. Collectors’ Calendar for Santa Clara and Santa Cruz Counties, California, and

Preliminary List of the Butterflies and Skippers of Santa Clara and Santa Cruz
Counties. Entomologists’ Exchange News 6(4):1-6.

. Tilden, J. W. and G. S. Mansfield. Some Observations and Captures of California

Lepidoptera. Pan-Pacific Entomologist XVII (3):113-114.

1942

. Erynnis lacustra (Wright) from Near The Type Locality (Lepidoptera, Hesperiidae).

Pan-Pacific Entomologist X VIII(1):8.

. Tilden, J. W. and G. S. Mansfield. A Capture of Lerodea eufala Edwards at Light

(Lepidoptera, Hesperiidae). Pan-Pacific Entomologist XVIII (1):26.

. An Unusual Record for Ochlodes yuma (Edwards) (Lepidoptera, Hesperiidae). Pan-

Pacific Entomologist X VIII(1):29.

1944

. Tilden, J. W. and G. S. Mansfield. Notes on Three Species of Coenonycha Horn

(Coleoptera, Scarabaeidae). Pan-Pacific Entomologist XX(3):115-117.

. Breeding of Phoebis sennae marcellina Cram. in San Jose, California (Lepidoptera,

Pieridae). Pan-Pacific Entomologist XX(3):115-117.

1945

. Notes on Redwood Cerambycidae (Coleoptera). Pan-Pacific Entomologist XXI(1):

30-31.

. Notes on Some Moths of the Family Saturniidae (Lepidoptera). Pan-Pacific Ento-

mologist XXI(1):32-33.

1946

An Occurrence of Schizopus in Monterey County, California (Coleoptera, Bupres-
tidae). Pan-Pacific Entomologist X XII(2):60.

1947

Mansfield, G. S. and J. S. Tilden. An Additional Locality for Aulicus terrestris
(Coleoptera, Cleridae). Pan-Pacific Entomologist X XIII(1):34

An Occurrence of the Pupa of Glaucopsyche lygdamus behrii (Edwards) in an Ant
Nest (Lepidoptera, Lycaenidae). Pan-Pacific Entomologist X XIII(1):42-43.

1948

Aestivation in Larvae of Arachnis picta picta Packard (Lepidoptera, Arctiidae). Pan-
Pacific Entomologist XXIV(1):31.

The Insect Community on Baccharis pilularis De Candolle. Doctoral Dissertation,
Stanford University. 408 pages. Copies at Stanford University and in Tilden Library.

1949

Notes on Parasites of Certain Microlepidoptera (Gracilariidae). Pan-Pacific Ento-
mologist XX V(1):27-28.

VOLUME 44, NUMBER 1 51

38.

A Note on Caicella mysie (Dyar) (Lepidoptera, Hesperiidae) with a figure of the
Male Genitalia. Bulletin of the Southern California Academy of Sciences XLV111
(1):4-6.

7. Note on Tragidion armatum (Lee) (Coleoptera, Cerambycidae). Pan-Pacific Ento-

mologist XX V(1):37.

. Occurrences of Diurnal Lepidoptera at Light. Pan-Pacific Entomologist XXV(2):

94-96.

. Oviposition of Cryptocephalus confluens Say (Coleoptera, Chrysomelidae). Ento-

mological News LX:151-154.

. Oviposition and Hatching of Pselliopus spinicollis Champion (Hemiptera, Redu-

viidae). Pan-Pacific Entomologist XX V(4):190-192.

1950

. Microlepidoptera Associated with Baccharis pilularis. 1. Pterophoridae. The Was-

mann Journal of Biology 8(1):81-88.

. Biological Notes on Corythuca morilli O. & D. (Hemiptera, Tingidae). Entomolog-

ical News LXI:135-137.

. Notes on Oxygrillus ruginasus (Lec.) (Coleoptera, Scarabaeidae) and Xylotrechus

undulatus (Say) (Coleoptera, Cerambycidae). Pan-Pacific Entomologist XX VI:37, 46.

. Oviposition and Behavior of Liriomyza pusilla (Meigen) (Diptera, Agromyzidae).

Pan-Pacific Entomologist XX VI(3):119-121.

. Notes on the Habits of Phylobaenus scabra (Le Conte) (Cleridae). The Coleopterists’

Bulletin IV(2):17-19.

. The Feeding of Podabrus pruinosus Le Conte (Cantharidae). The Coleopterists’

Bulletin IV(6):92.

1951

. Notes on Chrysobothris bacchari Van Dyke (Buprestidae). The Coleopterists’ Bulletin

V(1):1-3.

. Observations on Rhopalomyia californica Felt (Diptera, Itonididae). Pan-Pacific

Entomologist XX VII(1):20-22.

29. The Insect Associates of Baccharis pilularis De Candolle. Microentomology 16(1):

149-188.

. Microlepidoptera Associated with Baccharis pilularis. Il. Tortricidae, Phaloniidae,

Gelechiidae. The Wasmann Journal of Biology 9(2):239-254.

. Biological Notes on Two Species of Chrysopidae (Neuroptera). Entomological News

LXII:224—226.

. On the Distribution of Heliopetes domicella (Erichson) (Lepidoptera Hesperiidae).

The Lepidopterists’ News V(6-7):70.

. A New Subspecies of Mitoura siva Edwards (Lepidoptera, Lycaenidae). Bulletin of

the Southern California Academy of Sciences 50(2):96-98.

. A Note on the Manner of Feeding of Agulla adnixa Hagen (Raphidiodea, Raphi-

diidae). Pan-Pacific Entomologist XX VII(4):192.

. Concerning the Identity of Mitoura nelsoni muiri (Lepidoptera, Lycaenidae). The

Lepidopterists’ News 6(6—8):95-96.

1952

. Range Extension of Pholisora libya Scudder (Lepidoptera, Hesperiidae). Pan-Pacific

Entomologist XX VIII(2):92.

7. Observations on the Habits of Certain Syrphids (Diptera). Entomological News

LXII1:39-43.

1953

Observations on the Biology of Psoa maculata Le Conte (Psoidae). The Coleopterists’
Bulletin VII(2):9-12.

52

39.

40.

41.

42.

43.

44.

45.

46.

47.

48.

49.

50.

ol.

o2.

53.

o4.

55.

36.

57.

58.

59.

60.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Biological Notes on Trirhabda flavolimbata (Chrysomelidae). The Coleopterists’
Bulletin VII(3):20-23.

1954

Tilden, J. W. and J. C. Downey. A New Species of Philotes from Utah (Lepidoptera,
Lycaenidae). Bulletin of the Southern California Academy of Sciences 54(1):25-29.
Microlepidoptera Associated with Baccharis pilularis. II. Aegeriidae, Coleophori-
dae. The Wasmann Journal of Biology 12(1):43-52.

New Santa Clara County Mosquito Records. California Vector Views 1(5):3.

1955

Interspecific Cannibalism in Brennus Mots. (Coleoptera, Carabidae). The Coleop-
terists’ Bulletin [X(1):10.

Collecting Acmaeodera in Santa Clara County, California (Coleoptera, Buprestidae).
The Coleopterists’ Bulletin [X(2):23-25.

A Revision of Tharsalea (S.Str.), with Description of a new Subspecies (Lepidoptera,
Lycaenidae). Bulletin of the Southern California Academy of Sciences 54(2):67-77.
Tilden, J. W. and J. St. Germaine. Flies of Public Health Interest in Santa Clara
County. 1. Chicken Manure. California Vector Views 2(3):12.

. Flies of Public Health Interest. 11. Walnut Hulls. California Vector Views
2(4): 18.

. Flies of Public Health Interest. 111. Cow Manure. California Vector Views

2(5):25.

Common Pest Ants. In Proceedings, California Structural Pest Control Operators
Annual Conference, 1955. Chapter 7, 11 pages. California Polytechnical College,
San Dimas, California.

1956
San Francisco's Vanishing Butterflies. The Lepidopterists’ News 10(3—4):113-115.

1957

A Record of Stanleya as a Food Plant of Pieris beckerii (Lepidoptera, Pieridae).
Pan-Pacific Entomologist XX XIII(4):169.

Taxonomic History and Distribution of Ochlodes yuma (Hesperiidae). The Lepi-
dopterists’ News 11(4—5):151-152.

Carterocephalus palaemon in California (Hesperiidae). The Lepidopterists’ News
11(1-8):48.

Flies from Major Sources. California Vector Views 4(4):24, 27.

1958

Notes on the Life History of Euphydryas editha bayensis. The Lepidopterists’ News
12(1-2):33-35.

Tilden, J. W. and Carl D. Duncan. The Monarch Butterfly and its Migrations. Turtox
News 36(9):203-204.

Distributional Oddities. The Lepidopterists’ News 12(5-6):203-204.

1959

Microlepidoptera Associated with Baccharis pilularis. 1V. Gracillariidae. Lyoneti-
idae, Oinophilidae. The Wasmann Journal of Biology 17(1):43-54.

The Butterfly Associations of Tioga Pass. The Wasmann Journal of Biology 17(2):
249-271.

Schoening, Ernest H. and J. W Tilden. Anoplodera laetifica Le Conte and Ergates
spiculatus Le Conte from Knob-cone Pine (Coleoptera, Cerambycidae). Pan-Pacific
Entomologist XXX V(3):167-168.

VOLUME 44, NUMBER 1 53

6l.

63.

64.
65.

66.

79.

A Record of Pyramidobela angelarum Keifer (Lepidoptera, Ethmiidae) from Santa
Cruz, California. Pan-Pacific Entomologist XXX V(4):211.

2. Book Review—Laboratory Manual for Introductory Entomology. Pan-Pacific Ento-

mologist XXX V(4):211.

1960
An Additional Note on the Life History of Mitoura spinetorum (Lepidoptera,
Lycaenidae). Pan-Pacific Entomologist XXX VI(1):40.

1961

Certain Comments on the Subspecies Problem. Systematic Zoology 16(1):17-23.
Studies in the Genus Ochlodes Scudder. II. The Type Material of the North American
Species (Lepidoptera, Hesperiidae). Entomological News LX XII(2):37—45.

1962

General Characteristics of the Movements of Vanessa cardui (L.) (Nymphalidae).
Journal of Research on the Lepidoptera 1(1):43-49.

. Additional Records of Satyriwm behrii (Lycaenidae) from Oregon. Journal of the

Lepidopterists Society 16(3):199-200.

1963

. The Argynnis Populations of the Sand Creek Area, Klamath Co., Oregon. Journal

of Research on the Lepidoptera 1(2):109-1138.

. An Analysis of the North American Species of the Genus Callophrys. Journal of

Research on the Lepidoptera 1(4):281-300.

. Garth, John S. and J. W. Tilden. Yosemite Butterflies. Journal of Research on the

Lepidoptera 2(1):1—96.

1964

Two Species of Hesperiidae Previously Unrecorded from the United States. Journal
of the Lepidopterists’ Society 18(4):214-216.

1965

. Urbanus procne and Urbanus simplicius Hesperiidae. Journal of the Lepidopterists’

Society 19(1):53-55.

73. A Note on Pyrgus communis and Pyrgus albescens (Hesperiidae). Journal of the

Lepidopterists’ Society 19(2):91-93.

74. The Genus Panoquina Occurring in Texas. Journal of Research on the Lepidoptera

4(1):37-40.

75. Butterflies of the San Francisco Bay Region. California Natural History Guides: 12.

88 pages, 8 plates. Berkeley: University of California Press.

76. An Apparently New Dragonfly Record for California (Odonata, Libellulidae). Pan-

Pacific Entomologist 41(3):179.

1967

. A Previously Unrecognized Subspecies of Philotes speciosa. Journal of Research on

the Lepidoptera 6(4):281-284.

. Obituary—Carl Dudley Duncan. Pan-Pacific Entomologist 43(2):97-99.

1968

Records of Two Species of Exotic Lepidoptera Captured in California. Journal of the
Lepidopterists’ Society 22(3):187.

54

80.

81.

82.

83.

84.

88.

89.

90.

Oil

92.

97.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Cultivated Snapdragon a Host of Tobacco Budworm. Pan-Pacific Entomologist 44(3):
253
Tilden, J. W. and Bruce A. Tilden. Predation by Adelocera and Alaus (Coleoptera,
Elateridae). Pan-Pacific Entomologist 44(3):258.

1969
Concerning the Names and Status of Certain American Members of the Genus
Phyciodes. Journal of Research on the Lepidoptera 8(3):94—98.

1970
Comments on the Nearctic Members of the Genus Precis Huebner. Journal of
Research on the Lepidoptera 9(2):101-108.

1971
Aguna claxon (Hesperiidae) New to the United States. Journal of the Lepidopterists’
Society 25(4):293.

1973

. Specific Entities of the Subgenus Icaricia Nabokov (Lycaenidae). Journal of Research

on the Lepidoptera 12(1):11-20.

. A Name for Glaucopsyche lygdamus behrii Auct., not Edwards 1862. Journal of

Research on the Lepidoptera 12(4):213-215.

. Junonia and Precis—A Correction. Journal of Research on the Lepidoptera 12(4):

216.

1974

Unusual and Interesting Butterfly Records from Texas. Journal of the Lepidopterists’
Society 28(1):22-25.

Melitaea pulchella Boisduval, 1852, a Replacement Name. Journal of the Lepidop-
terists’ Society 28(4):252-253.

1975

An Analysis of the W. G. Wright Butterfly and Skipper Plesiotypes in the Collection
of the California Academy of Sciences. Occasional Papers of the California Academy
of Sciences, No. 118, 44 pages.

Hesperiid Notes. Journal of the Lepidopterists’ Society 29(1):60-62.

1976

A Proposed Terminology for the Types of Diapause Occurring in the Order Lepi-
doptera. Journal of Research on the Lepidoptera 15(1):33-39.

3. Urbana simplicius (Stoll), a New Record for California. Journal of Research on the

Lepidoptera 15(1):40.

. Observations of Predation on Lepidoptera in Alaska. Journal of Research on the

Lepidoptera 15(2):100.

. Obituary—Peter J. Herlan (1909-1977). Journal of the Lepidopterists’ Society 31(3):

213-214.

. Tilden, J. W. and David H. Huntzinger. The Butterflies of Crater Lake National

Park, Oregon. Journal of Research on the Lepidoptera 16(3):176-192.

1978

Obituary—James H. Baker (1910-1978). Journal of the Lepidopterists’ Society 32(3):
240.

VOLUME 44, NUMBER 1 eye)

98.

99.

100.

101.

1979

Attempted Mating Between Male Monarchs. Journal of Research on the Lepidoptera
18(1):2. ;

1986

Garth, John S. and J. W. Tilden. California Butterflies. Berkeley: University of
California Press. 246 pages, 24 plates.

Tilden, J. W. and Arthur C. Smith. A Field Guide to Western Butterflies. Boston:
Houghton Mifflin Company. 370 pages, 48 plates.

1987

Palmer, W. A. and J. W. Tilden. Host Specificity and Biology of Prochoerodes
truxaliata (Guenee) (Geometridae), A Potential Biocontrol Agent for the Rangeland
Weed Baccharis halimifolia L. in Australia. Journal of the Lepidopterists’ Society
41(4):199-208.

Acknowledgments

I am especially grateful to Hazel Tilden and to Bill’s brother Tom for much detailed
information about Bill’s interesting life and career. I am greatly indebted to Paul H.
Arnaud, Jr., C. Don MacNeill, Paul A. Opler, and Jerry A. Powell for reviewing the
manuscript and making many valuable suggestions. I also wish to thank those listed below
for providing additional information and assistance: Arnold G. Applegarth, Charles A.
Bridges, John A. Chemsak, Thomas Dimock, John Lane, Sterling O. Matoon, Edgar A.
Smith, and Matthew F. Vessel.

ARTHUR C. SMITH, 1536 Green Valley Road, Watsonville, California 95076.

FEATURE PHOTOGRAPHS

Journal of the Lepidopterists’ Society
44(1), 1990, 56-61

BUTTERFLY PHOTOGRAPHY IN THE TROPICS

Butterfly photography in tropical habitats is usually a much greater challenge than
photographing butterflies in temperate North America. Here, I briefly explain some of
the techniques I have developed in the tropics, and recount some personal experiences
that emphasize the points I wish to make.

Since 1973, when I first became interested in photographing butterflies, I have visited
several tropical countries, including Malaysia, East Africa, Mexico, Costa Rica, Venezuela,
and Brazil (twice). The examples in this article are from my most recent trip to Brazil,
in November 1989, led by Thomas C. Emmel for Holbrook Travel of Gainesville, Florida.

Equipment and Techniques

In the tropics, I use the same basic equipment that I use to photograph butterflies in
North America (Opler, P. A. & G. O. Krizek, 1984, Butterflies east of the Great Plains:
An illustrated natural history, Johns Hopkins Univ. Press, 294 pp.): a 35 mm single lens
reflex camera with a 200-mm macro lens and a flash unit mounted directly to the hot
shoe connection on the camera body. I shoot Ektachrome film (Eastman Kodak Company,
Rochester, NY 14650) for color transparencies (ASA 200) with the camera’s shutter speed
set at %4,oth of a second and the lens set to a small aperture (=high f-stop). The combination
of flash and small aperture produces well-lit close-up photographs with acceptable depth
of field. Batteries in the flash unit should be changed about every third roll, and sweat
should be wiped off camera equipment each evening to insure proper operation.

I recommend that the photographer work at a given tropical locality for a number of
days in a row. In that way familiarity with the community of butterflies can be gained
and the photographer can work on photographing particular species. On a two-week trip
I can usually count on obtaining good to very good photographs of 50-70 butterfly species.

Ideally, I try to take many photographs of the same butterfly using several lens aperture
settings. On my camera these are f32, 22, and {16. Such a range of exposures gives me
the best chance of having at least one properly exposed and correctly focused photograph.
I try to collect the subjects photographed so that I can identify them after my return
home. Often, however, Iam able to take only one exposure and am unable to capture
the butterfly before it flies away. Even more discouraging is the fact that several times
on each trip I see the photo of a lifetime take off into the forest before I am able to take
a single exposure. On my last trip this happened when a freshly emerged Pierella (Sa-
tyrinae) with iridescent blue upper hindwings perched in front of me and then took off
one second before I could press the shutter release.

Of course, sometimes my luck is good. Once, as I followed a patrolling Haetera piera
(Satyrinae), it disappeared around a tree trunk, and when I came around the tree I
discovered a mated pair, with male and female leaning in different directions (Fig. 1).

Butterfly Behavior

In many tropical habitats, butterfly behavior is sufficiently different to cause problems
for the photographer. One of the standbys of the temperate zone photographer—butterflies
visiting low flowers—is usually not available. Instead, tropical butterflies must be pho-
tographed while perching, while mating, or while feeding on rotting fruit, animal dung,
or bird droppings. Many butterflies perch too high in trees to be photographed or perch
on the underside of leaves. Furthermore, tropical forests are filled with would-be pre-
daceous birds, large ants, anoles, and large insects. As a result, most tropical butterflies
are very nervous and are apt to land only for a second or two.

Attractive nymphalids usually perch with their wings closed and open them only
periodically, and then only for a brief moment. The photographer must be aware of this
and must wait patiently with the camera in hand ready to take an exposure. In this way

VOLUME 44, NUMBER | 57

Fic. 1. Haetera piera L. (Satyrinae), pair in copula. Brazil: Rondonia, 60 km S of
Ariquemes; 9 November 1989.

I have been able to photograph the iridescent blue upper wing surfaces of morphos
(Morpho helenor /achilles) and preponas (Prepona demophon) (Nymphalidae) (Plate 1).
The use of baits is a good way to attract these butterflies to a convenient area for
photography. The best baits are feces, rotting meat, or rotting fruit. African collectors
have told me that excrement from big cats such as leopards is the best bait for Charaxes
spp. (Charaxinae). I often use human excrement or rotting fruits such as mangos or

Fic. 2. Eurybia lycisca (Ww) (Riodinidae). Costa Rica: Cartago Province, Turrialba;
15 May 1985.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 3. Morpho helenor/achilles (Morphinae), rare behavior by an otherwise shy
species, exploring the author’s left index finger with its proboscis. Brazil: Rondonia, 60
km S$ of Ariquemes; 10 November 1989.

Fic. 4. Batesia hypochlora Felder (Nymphalinae), perching on the author’s left calf.
Brazil: Rondonia, 60 km S of Ariquemes; 11 November 1989.

—
PLATE 1. Above: Prepona demophon L. (Nymphalinae), feeding at rotting banana.

Brazil: Rondonia, 60 km S of Ariquemes; 3 November 1989. Below: Temenis pulchra
Hewitson (Nymphalinae), upperside of basking male. Brazil: Rondonia, 60 km S of
Ariquemes; 11 November 1989.

VOLUME 44, NUMBER | 59

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

PLATE 2. Above: Euselasia (eutychus species group) (Riodinidae), pair in copula.
Brazil: Rondonia, 60 km S of Ariquemes; 9 November 1989. Below: Amarynthis meneria
Cramer (Riodinidae). Brazil: Rondonia, 60 km S of Ariquemes; 5 November 1989.

VOLUME 44, NUMBER | 61

bananas to make several bait stations on the ground and periodically visit them in sequence
to see what has arrived.

To photograph metalmarks or skippers perched beneath leaves, I must lie on my back
and inch slowly under the plant. Of course, just as Iam in the perfect position and ready
to “capture” my quarry on film, it may flit off to another leaf. Once, in Costa Rica, I
repeated my back inching approach to a perched Eurybia lycisca (Riodinidae) ten times
before I was able to focus in and with trembling hands takes several photographs (Fig. 2).

Nymphalids may land on sweat-drenched clothing or arms instead of the intended
bait, and, rather than just shoo them away, I try to photograph them. This can be especially
difficult because they are so close that the minimum lens to subject distance is just barely
possible, and because I have to hold the lens, camera, and flash in one hand while the
butterfly perches on the other (Fig. 3). Sometimes a butterfly may perch on my pants in
a position awkward for photography (Fig. 4). Despite occasional difficulties and discomfort
resulting from biting insects, high heat and humidity, fogged lenses, and occasional
dehydration, patience and persistence will be rewarded with crisp, close-up photographs
of some remarkable butterflies (Plates 1 & 2).

In summary, photographing butterflies in the tropics can be more rewarding than just
trying to capture and kill each butterfly as quickly as you discover it. In the course of
my photography I have learned much about the behavior of tropical butterflies. It is a
pleasure to share this knowledge through my photographs.

All photographs were taken with a Nikon FA camera, fitted with a Micro-NIKKOR
200mm 1:4 lens, and, when appropriate, Nikon extension rings PN-11 and PK-13. A
Nikon SB-15 electronic flash, attached to the hot shoe on the camera, was used in con-
junction with Ektachrome 200 slide film. The Nikon FA camera has a TTL (through the
lens) metering system that automatically admits the necessary amount of light from the
Nikon SB-15 flash in TTL mode. Exposures were taken at 45, sec at apertures from £16-
£32.

GEORGE O. KRIZEK, 2111 Bancroft Place NW, Washington, D.C. 20008.

Journal of the Lepidopterists’ Society
44(1), 1990, 62

MANUSCRIPT REVIEWERS, 1989

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 relied on the
expertise of 61 reviewers last year to provide 87 evaluations of manuscripts. It is with
much gratitude that the Journal acknowledges the services of the people listed below
from whom manuscript reviews were received in 1989.

Richard A. Arnold, Pleasant Hill, CA
George T. Austin, Las Vegas, NV

Howard D. Baggett, Palatka, FL

*Carlos R. Beutelspacher, Mexico, DF,
Mexico

*Keith S. Brown, Jr., Campinas, Sao Paulo,
Brazil

*Richard L. Brown, Mississippi State, MS

Gary R. Buckingham, Gainesville, FL

*Johns M. Burns, Washington, DC

Timothy M. Casey, New Brunswick, NJ

*Everett D. Cashett, Springfield, IL

I. F. B. Common, Toowoomba, Queens-
land, Australia

*Charles V. Covell, Jr., Louisville, KY

P. T. Dang, Ottawa, Ontario, Canada
P. J. DeVries, Austin, TX
Julian P. Donahue, Los Angeles, CA

John N. Eliot, Taunton, Somerset, England
John F. Emmel, Hemet, CA

*Thomas C. Emmel, Gainesville, FL
David L. Evans, Watertown, MA

*Douglas C. Ferguson, Washington, DC
Clifford D. Ferris, Laramie, WY

*Lawrence F. Gall, New Haven, CT

David L. Gibo, Mississauga, Ontario, Can-
ada

George L. Godfrey, Champaign, IL

L. Paul Grey, Lincoln, ME

Jane Leslie Hayes, Stoneville, MS
Bernd Heinrich, Burlington, VT
John B. Heppner, Gainesville, FL

David C. Iftner, Worthington, OH

* Reviewed two or more manuscripts.

Kurt Johnson, New York, NY

Roy O. Kendall, San Antonio, TX
Ian J. Kitching, London, England

Bernard Landry, Ottawa, Ontario, Cana-
da

W. Harry Lange, Jr., Davis, CA

Robert C. Lederhouse, East Lansing, MI

Rosemary Leen, Albany, CA

Don C. MacNeill, Oakland, CA
Lee D. Miller, Sarasota, FL
*William E. Miller, St. Paul, MN
James R. Mori, Modesto, CA

Stanley S. Nicolay, Virginia Beach, VA
Mogens C. Nielsen, Lansing, MI
Frederick H. Nijhout, Durham, NC

*Paul A. Opler, Fort Collins, CO

Dorothy P. Pashley, Baton Rouge, LA
John W. Peacock, Marion, OH
*Richard S. Peigler, Denver, CO
Kenelm W. Philip, Fairbanks, AL
*Jerry A. Powell, Berkeley, CA

John E. Rawlins, Pittsburgh, PA
*Robert K. Robbins, Washington, DC

Dietrich Schneider, Seewiesen, Federal
Republic of Germany

*James A. Scott, Lakewood, CO

Oakley Shields, Mariposa, CA

Frank Slansky, Jr., Gainesville, FL

Maria Alma Solis, Washington, DC

Ray E. Stanford, Denver, CO

Steve Stone, Lakewood, CO

*Paul M. Tuskes, San Diego, CA
James P. Tuttle, Troy, MI

G. A. Wood, Atherton, Queensland, Aus-
tralia

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),
GERARDO LAMAS (PERU), 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
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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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Literature Cited: References in the text of Articles should be given as Sheppard (1959)
or (Sheppard 1959, 1961a, 1961b) and listed alphabetically under the heading LITERATURE
CITED, in the following format without underlining:

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

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

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

In General Notes and Technical Comments, references should be shortened and given
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CONTENTS

PRESIDENTIAL ADDRESS, 1989: WHY CAN’T LEPIDOPTERISTS BE
MORE LIKE BOTANISTS? Julian P. Donahue UR!

AMBLYSCIRTES: PROBLEMS WITH SPECIES, SPECIES GROUPS, THE
LIMITS OF THE GENUS, AND GENUS GROUPS BEYOND—-A LOOK
AT WHAT IS WRONG WITH THE SKIPPER CLASSIFICATION OF
EVANS (HESPERIIDAE). John’M. Burns 2 ee eee Ty

A NEW SPECIES OF PIRUNA FROM MEXICO (HESPERIIDAE). Hugh
Avery Freeman (00 oe 28

GENERAL NOTES

Predation of five species of Noctuidae at ultraviolet light by the western
yellowjacket (Hymenoptera: Vespidae). Andrew D. Warren 82

Lycaena hyllus (Cramer) (Lycaenidae): New host and mating behavior in a
drought-induced population explosion. David B. MacLean & Bonnie
Ke Maclean: 223 ae es 33

Seasonal variation of occurrence of deformed cocoons of the tasar silk moths
Antheraea mylitta (Drury) and Antheraea paphia (L.) (Saturniidae) in
India. A. K. Dash: + B.iKs Nayak, 0.50) Sa ot ee 34

Book REVIEWS
CIE Guides to Insects of Importance to Man. 1. Lepidoptera. Ronald W.

heres | 2 a IS EU SO RSA A NO 87
Nordeuropas Prydvinger (The Oecophoridae of Northern Europe). Ronald

W Hodges) 25. 8). \ 0) N08 Ly Tee SENET SRS 0 J 38
Saturniidae: Ecological and Behavioral Observations of Select Attaci-

ni. Richard 'S) \Peiglen 2 222 Ne a 89
The Butterflies of Manitoba. Clifford Dy Ferris: cece 40
Mariposas*de' Venezuela.) Angel. Vilona) 2.2 ee 4]
Atlas of the Japanese Butterflies. Thomas C. Emmel Paneer! 42
Butterflies of Laos. Thomas C. Emmel YU ES 43

OBITUARY

James Wilson Tilden (1904-1988). Arthur C. Smith 45

FEATURE PHOTOGRAPHS
Butterfly Photography in the Tropics. George O. Krizek 56

MANUSCRIPT: REVIEWERS, 1989 oe 62

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