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Molume 40 1986 Weather 4

ISSN 0024-0966

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

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

9 October 1986

THE LEPIDOPTERISTS’ SOCIETY

EXECUTIVE COUNCIL

CLIFFORD D. FERRIS, President DouGLas C. FERGUSON,

Don R. Davis, Immediate Past President President-Elect

JERRY A. POWELL, Vice President EDWARD M. PIKE, Vice President RICHARD A. ARNOLD, Secretary ALLAN WATSON, Vice President

ERIC H. METZLER, Treasurer

Members at large:

JOHN M. BuRNS Boyce A. DRUMMOND III MIRNA M. CASAGRANDE FLOYD W. PRESTON JOHN LANE EDWARD C. KNUDSON JACQUELINE Y. MILLER ROBERT K. ROBBINS FREDERICK W. STEHR

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 facil- itate 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 $18.00 Student members—annual dues $12.00 Sustaining members—annual dues $25.00 Life members—single sum $250.00 Institutional subscriptions—annual $25.00

Send remittances, payable to The Lepidopterists’ Society, to: Eric H. Metzler, Treasurer, 1241 Kildale Square North, Columbus, Ohio 48229, U.S.A.; and address changes to: Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A.

Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- ume, and recent issues of the NEWS are available from the Publications Coordinator. The Commemorative Volume, is $6; for back issues, see the NEWS for prices or inquire to Publications Coordinator.

Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A.

Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for $25.00 (institutional subscriptions) and $18.00 (active member rate) by the Lepidopter- ists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Boule- vard, Los Angeles, CA 90007. Second-class postage paid at Los Angeles, CA and addi- tional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, 1900 John St., Manhattan Beach, CA 90266.

Cover illustration: First stage larva of Natada nasoni (Grote) (Limacodidae), from Dyar 1899, J. New York Entomol. Soc. 7:61-67. Suggested by Marc E. Epstein.

JouRNAL OF Tue LeEpPIDOPTERISTS’ SOCIETY

Volume 40 1986 Number 1]

Journal of the Lepidopterists’ Society 40(1), 1986, 1-7

PRESIDENTIAL ADDRESS, 1984: A TRIBUTE TO THE AMATEUR

LEE D. MILLER

Allyn Museum of Entomology of the Florida State Museum, 3621 Bay Shore Road, Sarasota, Florida 33580

First of all, let me state that it has been a great pleasure to serve the Society as its president during the past year. This is a wonderful group and one of the few in this country where the professional and the amateur can speak as equals. But, all too often the statement is made: “He is only an amateur,” usually implying that he doesn’t know what he is talking about, or at least that his opinion is not worth as much as one who makes his living in the field. I do not accept this view, and this presentation is an unabashed tribute to those who do not make their living in the field of lepidopterology.

What is an amateur? The word is derived from the Latin “amator’’ (lover) or the French “amare” (to love), and is defined in the dictionary as “one who cultivates a particular pursuit, study or science from taste, without pursuing it professionally.”” Everyone makes his or her living at something, and I consider an amateur lepidopterist as one who makes his living at something other than lepidopterology. There can, then, be janitors, pipefitters, medical doctors, engineers and mammalian ecologists using lepidopterology as an avocation. There are good am- ateur workers in the field and bad ones, but the variant definition of amateur is not always applicable: “‘one practicing an art without mas- tery of its essentials.’ I am praising that amateur who has gained a certain amount (often a great deal) of expertise in a particular phase of lepidopterology, enough so that he or she is able to impart that knowledge to others.

Amateurs have been the backbone of science since its inception: there were no professional entomologists in Linnaeus’ time, but no one suggests that they did not do the best possible job with the information

2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

available to them. Pieter Cramer was an artist whose avocation was lepidopterology; Dru Drury was a silversmith; Jacob Huebner was a printer; William Chapman Hewitson was a wealthy landowner and otherwise retired; William Henry Edwards was a coal baron; Henry Edwards was an actor; William Barnes was a physician, and Walter Rothschild was a “‘black sheep” who did not fit into the family banking business. From these diverse backgrounds, however, came people who made tremendous contributions to the field of lepidopterology, as great as those of the contemporaneous professionals. Most of the contribu- tions that have been made by amateurs have been in systematics, be- havior, life history, morphology and ecology, subsets of the field that usually require less elaborate equipment; but a number of important contributions also have been made in the field of genetics by non- professionals.

Often the amateur will attack a problem of limited scope and be- come very expert on that topic. The amateur may be expected to gather much pertinent material within a circumscribed group, often more and better material than is readily available to the professional even in the finest of facilities. He studies the material gathered, in- cluding that which is borrowed from museum collections, and gathers together the appropriate literature, just as the professional would. He is at least as likely as the professional to ask opinions from others, and most studies done by amateurs go through a great many more drafts, and hands for comment, than do papers written by those of us in the profession for the simple reason that the amateur is still convinced that he does not know everything there is to know. Most publishing ama- teurs are at least as receptive to new and different ideas as are those of us who do it for a living.

Admittedly, I am talking here of the best amateurs, not those un- guidable persons who nevertheless write on various subjects without the benefit of knowledge or guidance. The “good’’ amateurs are the ones who behave in the field about as we should expect professionals to perform their studies.

The early amateurs worked chiefly in systematics of Lepidoptera. They generally amassed huge (for the day), usually beautifully curated collections of specimens, and spent much of their time writing descrip- tions of new taxa from these collections or from those of their ac- quaintances. Certainly this is what W. C. Hewitson, W. H. Edwards and Henry Edwards, to name a few, did, and their works compare favorably to those of Butler, Walker and other professionals of the day. Once in a while one hears grumblings about the incompleteness by today’s standards of their descriptions, but one also hears this complaint about the descriptions done by professionals of the same time period.

VOLUME 40, NUMBER 1 3

Later, other amateurs like Frederick DuCane Godman and Osbert Salvin, two wealthy English “men of leisure’, began looking at Lepi- doptera as related populations of organisms, rather than as specimens which either varied from other specimens or did not. These amateurs began putting contemporary biological theories into practice and crit- ically examined lepidopteran populations in light of the then new ideas of evolution and biogeography. The result was that they, mostly with Godman’s money, decided to do a total biological inventory of Mexico and Central America for which they would write some parts and enlist experts in other fields to contribute sections. The resulting Biologia Centrali-Americana was published over more than 30 years and ran to nearly fifty sumptuous volumes. This publication still has not been superceded. |

Walter Rothschild, once he decided to collect Lepidoptera seriously, could have been expected to do it with a vengeance. He did, finally amassing something over two and a quarter million specimens, more than the Lepidoptera holdings of most of today’s major museums. He immediately saw the advantages of collecting study series, and for the first time, a private collection had more than a few specimens of any single taxon. At the same time in France, a printer, Charles Oberthuer, began much the same type of accumulation of material, although the scope of his collecting was smaller than Rothschild’s. Oberthuer’s col- lection finally amounted to more than a million specimens. Rothschild was one of the first private collectors to realize that he probably needed professional curatorial help, and he hired Karl Jordan as entomologist at his museum and Ernst Hartert as his ornithological curator. What a team they made! If one goes through the writings of this triumvirate, one can find the first modern concept of geographical subspecies, tri- nominal nomenclature, an elucidation of the biological species concept, and one of the first cladistic analyses of an animal group (it was not so labelled and frequently is not recognized as such). Especially with Walter Rothschild and his curators, modern systematics can be shown to have had its birth. To say that I am a Rothschild fan would be true because of the facts that he (A) collected long series of material for study, (B) had a worldwide bias to his activities, and (C) surrounded himself with those who in conjunction with him developed most of the bases of modern systematics. Their efforts were not always appreciated by their contemporaries, but those must have been halcyon days at Tring!

Strangely enough, another man lived in England at the same time who had an obsession with outdoing Rothschild, thereby setting himself a prodigious task. James J. Joicey was a man of leisure who decided that he would outdo Rothschild in the acquisition of orchids. He tried

4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

this for some years before and during World War I, finally going bankrupt for 30,000 pounds, a very large sum at the time. The judge admonished Joicey, and he agreed that he would not try to build the world’s premier orchid collection, and he held to his promise. He switched instead to Lepidoptera and went broke in the 1930's for over 300,000 pounds, perhaps making a statement about how deeply one can get involved in making a collection if one lets his or her imagi- nation run wild. Joicey did, however, hire curators (perhaps because Rothschild had them), and they produced some excellent work, espe- cially on the Lepidoptera of New Guinea, Hainan Island, and central and eastern Africa. The Joicey, Oberthuer and Rothschild collections are the reason that the British Museum (Natural History) enjoys such numerical superiority over other collections throughout the world.

In the United States, Dr. William Barnes, a physician from Decatur, Illinois, formed a magnificent collection of North American Lepidop- tera. It became readily apparent that many specimens that he was obtaining were undescribed, and with the help of professional curators, he set about to describe them. From this activity came revisionary studies on moth genera which are still standard today. Many of these were published in his own journal, Contributions to the natural history of the Lepidoptera of North America, and were often jointly coau- thored with his curators, Arthur Ward Lindsey, James McDunnough, and Foster H. Benjamin, among others. Many of the finest contribu- tions during the first quarter of the 20th Century came from that journal.

In later years, many amateurs have made contributions to the tax- onomy of Lepidoptera. Without exception, these students have been willing to listen to others and be guided by them, and the resulting papers have been highly informative. Anyone who has ever studied Hesperiidae knows of the legacy left us by Brigadier William Harry Evans, a retired British army officer who served in India during the first third of this century. His Catalogues to the hesperiids are standard works for the professional and the amateur alike. Many people forget that he also wrote the definitive guide to the butterflies of India, which is still being reprinted. Still, he was an amateur who performed like a professional. Another military man turned lepidopterist is John N. Eliot, recently awarded the Karl Jordan Medal, whose studies of the Lycaeni- dae and the Neptis group of nymphalids have earned him a permanent place in our field. He also completely revised Corbet and Pendlebury’s Butterflies of the Malay Peninsula. Our former President, Col. Stanley S. Nicolay, has done some fine work tying together certain of the neotropical hairstreaks, and he is still trying to bring some order out of the chaos that characterizes this group. Stan also works on the hes-

VOLUME 40, NUMBER 1 5

periids, so it might be said that he has selected some very knotty problems to tackle. Surely the taxonomy of North American moths is better because of the activities of Mr. André Blanchard, a retired en- gineer. I have the utmost admiration for one who can tackle these little creatures and make sense of them. The late M. Henri Stempffer was a retired French banker who has added greatly to our knowledge of the taxonomy of African Lycaenidae, a group that has puzzled all workers before him. For his contributions M. Stempffer was elected to receive the first Jordan Medal given by this Society. We can expect other amateurs to receive the award in the future, I am sure. Arthur Rydon studied the Charaxini and has increased our knowledge of this group through his writings. Similarly, the recently deceased Dr. Lionel G. Higgins, an English physician, made a lifetime study of the Melitaei- nae, and his writings are the basis of the classification of this group throughout the world. Our own former President and Honorary Life Member, Dr. F. M. Brown, is a geologist by trade, but he has achieved renown in the taxonomy of both nearctic and neotropical Rhopalocera, and if this is not enough, Brownie has now engaged (at more than 80 years old) in the study of fossil insects, and is writing several compendia on important groups. Cliff Ferris, another amateur subsequently elect- ed President of our Society, is a bioengineer by profession, but his studies on the systematics of Nearctic butterflies are quite professional. Let us not forget the contributions of a former lawyer, Dr. Cyril F. dos Passos, who has worked for several decades on the taxonomy and nomenclature of North American butterflies. The list of amateurs who have contributed to the taxonomy of Lepidoptera is endless, and I apologize to others that I have left out—there simply is not enough space.

A special place must be saved for the late Dick Dominick whose dream of The Moths of North America has been an inspiration to us all. The several volumes that have appeared under the aegis of this series conspire to make the study of moths as popular as that of but- terflies. It was an impressive project, and one can but wish that Dick had lived to see more of it completed.

To finally get to nontaxonomic studies, the life history studies done by Roy and Connie Kendall, their associates, and by those such as Dave Baggett have added the biological information that can turn alpha taxonomic treatises into studies at a higher level. They are providing the building blocks for greater understanding of Lepidoptera, and these studies cannot help but improve the quality of later studies on these insects. Amateur life history work generally does not include the chae- totaxy of the larvae that professionals consider vital, but the workers mentioned here are also preserving egg, larval, and pupal material for

6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

such studies, and this material is readily available to those interested in performing them.

Dr. David Wright is a physician by occupation, still he is embarking on some superb micromorphological studies of various stages of but- terflies, especially the Lycaenidae, using scanning electron microscopy, and he has already published new and innovative ideas on the mor- phology of the larval stages, including showing that some setae have wrongly been attributed to some segments when they belong to adja- cent ones. To the casual observer, these data may seem trivial, but he has found at least one derived (apomorphic) character which seems to define the Lycaeninae, one that was previously unsuspected.

There are other examples of amateurs making contributions in ecol- ogy, physiology, and even genetics of Lepidoptera. The information that these workers are providing surely aids in the understanding of these insects. An example of such a study is Mike Fisher’s rearing of Papilio nitra which unexpectedly turned up the fact that Papilio zel- icaon gothica is the more frequent color morph of nitra. This was an elegant study, done on a low budget, that showed something totally unexpected.

My wife and I have been fortunate enough to work for and with a man with vision much like that of Rothschild. The late Arthur Allyn was the heart of an effort to form a real resource for taxonomic and morphological research on Macrolepidoptera. In the years since we joined him, that collection grew from about 100,000 specimens to its present roughly 550,000 prepared and 250,000 unspread specimens. The decision was made to gather material from throughout the world because we felt that revisionary studies done on groups from only part of their ranges were incomplete, and we wished to encourage a world view of butterflies and large moths. Another goal was to obtain collec- tions in their entirety and not see them broken up as so many had been in the past. The vision of Art Allyn has been an inspiration to us, and we hope that the material he was responsible for gathering will be used for generations to come.

Thus far, almost everyone that I have mentioned in this tribute has been male. Consider the fact that the Rothschilds are and were an amazing lot, true “Renaissance men’. The present Renaissance “man” of the Rothschilds is a woman: Miriam Rothschild has published a quarter of a million words on flea taxonomy, she has worked on plant- insect interactions, on intestinal worms, on mimicry in butterflies, on the behavior of sea gulls and has been involved with worldwide con- servation efforts. Miriam Rothschild was educated basically at home, chiefly by her uncle Walter and his curators at Tring. She earned no university degree, but nonetheless is honored in scientific circles in

VOLUME 40, NUMBER 1 7

England and elsewhere. Her ability and tenacity have made her achievements possible.

The honor role of amateurs is long and distinguished. They have made and are making significant contributions to our chosen field. The next time you hear the statement, ““He’s only an amateur,” realize that this should not be a pejorative; perhaps it is a tribute, since the person in question does not have to be paid to perform. All of us, amateurs and professionals alike, have something to give lepidopterology. The difference between amateur and professional is one of degree, rather than kind. In the final analysis, there is not “amateur science” and “professional science’, there is only good science or poor science. Let us recognize that we all have something worthwhile to say, and we will all benefit from such understanding.

Journal of the Lepidopterists’ Society 40(1), 1986, 7

GENERAL NOTE

TRYON REAKIRT: A SEQUEL

In 1964 I wrote briefly about Tryon Reakirt, a Philadelphia entomologist of note during the 1860’s (Brown 1964, J. Lepid. Soc. 18:211-214). He was a mystery man in his last years. All I knew earlier was that he had fled the country in early 1871. As a result, both his enterprises and his father’s business filed for bankruptcy. I found the answer to his disappearance among newspaper clippings belonging to William Henry Edwards of Coalburgh, West Virginia.

One clipping is from the Philadelphia Inquirer, Wednesday, 8 February 1871. Reakirt had forged notes on large pharmaceutical houses to the tune of more than $110,000! An error in a date caused a bank clerk to go into the matter with the purported issuer. The fat was in the fire! Reakirt left town hurriedly, and ultimately got to Lima, Peru, where, apparently, he died of dysentery in late 1872 or early 1873.

F. MARTIN BROWN, 6715 South Marksheffel Road, Colorado Springs, Colorado 80911.

Journal of the Lepidopterists’ Society 40(1), 1986, 8-19

A NEW SPECIES OF EPIDROMIA (NOCTUIDAE) FROM FLORIDA

M. ALMA SOLIS

Maryland Center for Systematic Entomology, Dept. of Entomology, University of Maryland, College Park, Maryland 20742

ABSTRACT. This preliminary study of the genus Epidromia Guenée (Ophiderinae: Noctuidae: Lepidoptera) describes a new species E. fergusoni and redescribes the type species E. pannosa Guenée.

Twenty species group names are included in the neotropical genus Epidromia Guenée (Ophiderinae). This genus is distributed in north- ern South America and throughout Central America and reaches its greatest diversity in the Antilles. Florida is its northernmost extension.

METHODS

Names applied to wing veins and markings correspond to Forbes (1923). The forewing length is measured from the base of the wing to the apex of the wing. The width is measured from the apex to the anal angle. The depth and width of the cleft of the ostium bursae is mea- sured as shown in Fig. 1. All measurements correspond to the mean value, and the measurements in the parentheses are the range. The numbers in parentheses in the distribution section are USNM (United States National Museum of Natural History) genitalia slide numbers.

RESULTS

Epidromia is being redescribed since the original description is in- complete. The genitalia are described and illustrated for the first time.

Epidromia

Epidromia Guenée, 1852, In Boisduval and Guenée, Hist. Nat. Insectes, Spec. Gén. des Lépid. 7:325. Type: Epidromia pannosa Guenée, 1852, by subsequent designation (Berio, 1966. Annali Museo Civico Storia Naturale Giacomo Doria 76:57).

Penultimate segment of palpi upturned and longer than first and third segments to- gether; ultimate segment ending in blunt point. Abdomen cylindrical, elongated; sternites more hairy than tergites; distal end appearing square-shaped in males and tapered in females (Figs. 3, 6 and 8) distal tergite square-shaped with membranous projections into seventh tergite, projections pointing medially; distal sternite with two sclerotized, long lobes connected by two smaller lobes (Fig. 2). Front legs stout with dense hairs. Wings entire, oblong; underside of wings having beige sheen with silky pubescence; peak of each scallop of adterminal line with immediately adjacent dot. Sacculus simple; uncus simple, widely angled distally; tegumen medially extended to point ventro-laterally; vinculum about as long as wide; vesica distally bilobed without cornuti (Figs. 4, 5 and 9, 10). Genital plate heart-shaped; ostium bursae cleft; no signa on corpus bursae (Figs. 7 and 12).

VOLUME 40, NUMBER 1 9

Fic. 1. Diagrammatic illustration of ostium bursa and measured distances.

Inspection of specimens and representative genitalia slides of genera believed to be closely related revealed that Epidromia is most closely related to Itomia Htibner. These two genera share an uncus that is simple, curved, and with a prominent spine at the distal end, and a costa on the valve that is strongly lobed. Itomia species more closely resemble Epidromia species from South America than those from Cen- tral America and the Antilles.

The type-species of Itomia is Itomia lignaris Hiibner, by monotypy.

1.0 mm

Fic. 2. Last abdominal segment of Epidromia.

10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 3. Epidromia fergusoni, holotype, male.

The holotype is believed to have been destroyed (R. W. Poole, pers. comm.). The original description and illustration of I. lignaris are not adequate for identification of the species. The specimens at the USNM suggest that a complex of species is involved; therefore, a male and a female from the complex have been chosen for the purpose of the genitalia comparison (Female: Mizantlan, Mexico, slide no. 42,618; Male: Mizantlan, Mexico, slide no. 42,619) (Table 1).

Table 2 is simply a list of species group names currently placed in Epidromia. Some names are new combinations. Some of the names have been traditionally recognized as synonyms. Some species de- scribed by Walker from the Antilles are suspected to be synonyms; he described only females, and females are highly variable. Photographs of the Walker and Gueneé types from the British Museum were stud- ied. There are three other new species that will not be described here. A full revisionary treatment is not feasible at this time because of the difficulties of assembling sufficient material and identifying all of the

types.

Epidromia fergusoni Solis, new species

Male holotype. Scales of collar, thorax and abdomen brown with purplish tinge (pur- plish tinge may disappear with age). Third sternite of abdomen without large bristles (n= 7). Ground color of wings brown with purplish tinge. Forewing: Other margin undulated, invaginated at end of R;, forming a peak at apex. Basal band brown (not visible); antemedial line light brown; triangular, chocolate brown area extending distally from base of antemedial line, terminating where postmedial line curves toward base of wing. Reniform spot gray (black, more obvious in older specimens). Postmedial line beige;

VOLUME 40, NUMBER 1

TABLE 1.

Structure

Ultimate labial palp segment

Penultimate labial palp segment

Tergites and ster- nites of abdomen

Distal end of abdo- men

Front legs Underside of wings

Dots at peak of each scallop of adterminal line

Upper side of wings

Sacculus

Tegumen Vinculum Uncus Vesica

Genital plate

Ostium bursae

Epidromia

blunt

four times as long as ultimate palp segment

sternites with more hair than tergites

distal tergite square with membranous projections into 7th tergite bending dorsally; distal sternite with medial sclerotized area but not tubelike

stout, with dense long hairs

beige sheen with silky pubes- cence

immediately adjacent

variable simple

medially extended to a point ventrolaterally

as long as wide

not sharply angled

distally bilobed heart-shaped

with a definite cleft

11

Comparison of Epidromia and Itomia.

Itomia

pointed

two times as long as ultimate palp segment

sternites and tergites equally hairy

distal tergite triangular with membranous projections into 7th tergite bending laterally; distal sternite with medial tubelike sclerotized area

slender, devoid of long hairs

yellow without silky pubes- cence

not immediately adjacent

diagonal lines across the wings

chitinized extension costad and extending distally

simple

longer than wide sharply angled distally

not distally bilobed

rectangular, slightly longer than wide cleft not definite

chocolate brown patch adjacent to postmedial line beginning at M, and extending distally to the apex of forewing. Adterminal line light brown, peak of each scallop with beige dot. Terminal line light brown (beige). Distal underside of wing without gold patch between M, and R,. Forewing length 2.3 cm (2.2—2.4) (n = 25). Length/width ratio 1.5 (1.8-1.7). Hindwing: Outer margin round, area adjacent to margin light brown. Post- medial line same as in forewing; chocolate brown area extending from postmedial line to about halfway to base of wing (Fig. 3). Genitalia: Uncus enlarged at distal end; valve with thumblike process on saccular margin, distal end of valve expands into small, flaplike process; editum on costa round; longest lobe of vesica bifurcate, with short branch round- ed and longer branch rounded; longer branch expanded at base and tapered to blunt point (n = 7) (Figs. 4, 5).

Female allotype. Scales of collar, thorax and abdomen brown. Ground color of wings brown. Forewing: Outer margin same as in male. Basal line (if visible) double, dark brown on inside, yellow on inside; median line dark brown (or absent). Reniform spot same as in male. Postmedial line double, brown on inside and yellow outside. Amount of beige in area between postmedial and adterminal line when present varies. Subter- minal line yellow to brown with dot at peak of each scallop, terminal line brown. Fore- wing length 2.1 cm (1.9-2.2) (n = 25). Length/width ratio 1.5 (1.4-1.6). Hindwing: Outer margin same as in male. Line markings same as forewing, but basal line and antemedial line not visible (Fig. 6). Genitalia: Ostium bursae cleft, approximately 0.45 mm wide at distal end, depth of cleft 0.85 mm (0.28-0.40) (n = 4) (Fig. 7).

12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1.0 mm

1.0 mm

5

Fics. 4 & 5. Male genitalia of Epidromia fergusoni. 4, Egmont, Fla., USNM slide 42,611; 5, aedeagus, Miami, Fla., USNM slide 42,609.

Types. Holotype: Male (Fig. 3), University Reserve, Welaka, Putnam Co., Florida, 6 April 1972, D. C. Ferguson. Allotype: Female (Fig. 6), University Reserve, Welaka, Putnam Co., Florida, 6 April 1972, D. C. Ferguson. Paratypes: Six males, same locality and collector, USNM genitalia slide numbers 42,054, 42,614, 42,607; three females, same locality and collector. All specimens are deposited in the U.S. National Museum of Nat- ural History, Washington, D.C.

Distribution. Other specimens used in this analysis were from the following localities in Florida: Miami, Glenwood, Ft. Myers, Ft. Meade, Dade City, St. Petersburg, Royal Palm State Park, Marcos Island, Lutz, Stemper, Indian River, Egmont, Ft. Lauderdale, Chokoloskee. Specimens with suspect data: Plainsfield, N.Y.; Jemez Springs, N.M.; Cuba.

VOLUME 40, NUMBER 1 13

Fic. 6. Epidromia fergusoni, allotype, female.

Discussion

This is the Florida species that has been known in this country for

nearly a century as Epidromia delinquens (Walker) (=Ophiusa delin- quens Walker, 1858). However, Hampson (1913) referred delinquens to the synonymy of Mocis repanda (F.), a name that Hampson and

TABLE 2. List of species group names in Epidromia. (Names in parentheses are the

original combinations. )

Epidromia pannosa Guenée

POSS

. zetophora Guenée

. xanthogramma Wallengren . zephyritis Schaus

. rotundata Herrich-Schaffer . consperata Dognin

poaphiloides Guenée

profana Walker

flavilineata (Hampson) NEW COMBINATION (Thermesia flavilineata) glaucescens (Walker) NEW COMBINATION (Thermesia glaucescens)

. lenis (Walker) NEW COMBINATION (Thermesia lenis) . antica (Walker) NEW COMBINATION (Ophisma antica)

arenosa (Walker) NEW COMBINATION (Phurys arenosa)

. pedestris (Walker) NEW COMBINATION (Phurys pedestris)

. profecta (Walker) NEW COMBINATION (Poaphila profecta)

. saturatior (Walker) NEW COMBINATION (Remigia saturatior)

. sigillata (Walker) NEW COMBINATION (Thermesia sigillata)

. suffusa (Walker) NEW COMBINATION (Thermesia suffusa)

. tinctifera (Walker) NEW COMBINATION (Thermesia tinctifera) . valida (Walker) NEW COMBINATION (Ophisma valida)

14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1.0 mm

Fic. 7. Female genitalia of Epidromia fergusoni, paratype, Welaka, Fla., USNM slide 42,615.

other authors subsequently (but mistakenly) considered to represent the same species as Mocis latipes (Guenée). The type of delinquens has not been seen, but after reading the description by Walker, it is acknowledged that Hampson was correct in assigning it to the genus Mocis Hubner.

It should be noted that E. fergusoni is not the only species of Epi-

VOLUME 40, NUMBER 1 15

Fic. 8. Epidromia pannosa Guenée, holotype, male.

dromia in Florida. A male specimen from Homestead of one unde- scribed species and a male and a female from Big Pine Key of another undescribed species have also been collected there and are now re- corded from the continental United States for the first time.

E. fergusoni is named after Douglas C. Ferguson who collected the type material.

Epidromia pannosa Guenée

Epidromia pannosa Guenée, 1852, In Boisduval and Guenée, Hist. Nat. Insectes, Spec. Gén. des Lépid. 7:326.

TABLE 8. Comparison of E. pannosa and E. fergusoni.

Overali color Third sternite

Structure pannosa

brown with bristles

fergusoni

purplish without bristles

Outer forewing margin straight undulated

Antemedial and postmedial line double single

Triangular, chocolate brown area extending absent present distally from base of antemedial line

Hindwing outer margin—male angulate rounded

Hindwing outer margin—female rounded rounded

Thumblike process on costa absent present

Short branch of bifurcated lobe of vesica truncated rounded

Cleft of ostium bursae 0.62 mm wide, 0.25 mm wide,

0.43 mm depth 0.35 mm depth On underside of forewing—gold patch be- present absent

tween M, and R,

16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1.0 mm

1.0 mm

10

Fics. 9 & 10. Male genitalia of Epidromia pannosa Guenée. 9, Rio Janeiro, Brazil, USNM slide 42,053; 10, aedeagus, Venezuela, USNM slide 42,065.

Male. Scales of thorax and abdomen brown; head and collar dark brown. Third sternite with bristles (n = 4). Ground color of wings brown. Forewing: Outer margin not undu- lated. Basal band brown; antemedial line double, yellow on inside (may not be evident) and dark brown on outside. Median shade dark brown. Reniform spot outlined in dark brown, filled with gray (black or gray without outline). Postmedial line double, dark brown inside, yellow outside. Subterminal line faint, light brown (chocolate brown or lacking). Dark brown patch between subterminal and adterminal line extending from

VOLUME 40, NUMBER 1 17

Fic. 11. Epidromia pannosa Guenée, female.

M, to anal angle. Adterminal line dark brown, slightly scalloped; peak of each scallop with dark brown dot. Terminal line yellow (brown). (One specimen with vein lines dark brown.) Distal underside of wing with gold patch between M, and R,. Forewing length: 2.3 cm (2.1-2.4). Length/width ratio: 1.6 (1.4-1.8) (n = 10). Hindwing: Outer margin slightly angulate at end of Cu,; edge of wing above Cu, parallel to body. Area from apex to anal angle adjacent to outer margin dark brown (light brown or no shading). Median shade, postmedial and subterminal line same as forewing (Fig. 8). Genitalia: Uncus enlarged at distal end; editum on costa round; vesica with longest lobe bifurcates, short branch truncate, long branch expanded at base, tapering to blunt point (n = 4) (Figs. 9 and 10).

Female. Since Gueneé did not describe the female of this species, the following de- scription of a female believed to represent the same species from Aroa, Venezuela is provided. Three males of this species were collected at Aroa, Venezuela and all four (three males and one female) have the same label information. Scales of head, collar, thorax, and abdomen same color as male. Ground color of wings same as male. Forewing: Median line dark brown. Reniform spot same as male. Postmedial line same as male; subterminal line beige. Length: 2.1 cm. Length/width ratio: 1.7 (n = 1). Hindwing: Outer margin round. Median line, postmedial line and subterminal line same as forewing (Fig. 11). Genitalia: Ostium bursa cleft, approximately 0.62 mm wide at distal end, depth of cleft 0.48 mm (n = 1) (Fig. 12).

Types. The holotype, in the British Museum of Natural History, is a male with no label data other than “‘Bresil.’”” A photograph of the type specimen taken by R. W. Poole was used in this description.

Distribution. Aroa, Venezuela: one female (42,115) and three males (42,065); Edo. Zuela, Venezuela: three males; Rio Janeiro: one male (42,053); Castro, Parana: one male (42,057); Merida, Mexico: one male (42,069); Tamazunchale, Mexico: four males; Ma- zatlan, Mexico: one male (42,056); Poza Rica, Mexico (42,072). All specimens examined are in the collection of the U.S. National Museum of Natural History.

18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1.0 mm

eee Q te Sey

2

Fic. 12. Female genitalia of Epidromia pannosa Guenée, Venezuela, USNM slide 42,115.

Discussion

J. G. Franclemont and E. Todd (1983) mistakenly synonymized pan- nosa and poaphiloides. This synonymy was probably based on notes by Hampson that were never published and transferred from the Brit- ish Museum of Natural History to the U.S. National Museum of Natural History. E. poaphiloides was described from Cayenne, French Guiana by Gueneé in 1852. The type of poaphiloides is believed to have been destroyed or lost (R. W. Poole, pers. comm.). An illustration of the type

VOLUME 40, NUMBER 1 19

is not available. After reading the written description, it is quite ob- vious that poaphiloides is not pannosa. A number of specimens of poaphiloides from French Guiana and British Guiana (Guyana) can be found at the U.S. National Museum of Natural History.

ACKNOWLEDGMENTS

I wish to thank Dr. R. W. Poole, SEL, USDA, for the use of the catalogue of the Noctuidae at the U.S. National Museum of Natural History, for photographing the types at the British Museum of Natural History and for instructing me in dissection techniques. I thank Dr. D. Miller and Dr. D. Ferguson, SEL, USDA for corrections and suggestions on the manuscript. This description was prepared for a course taught by Dr. D. Miller at the University of Maryland at College Park. Mr. A. M. Wilson photographed the specimens. I thank Mrs. Elaine R. S. Hodges, Smithsonian Institution, for instructing me in illustration techniques and for analyzing my illustrations. Scientific Article No. A- 4064, Contribution No. 7049 of the Maryland Agricultural Experiment Station.

LITERATURE CITED

BERIO, E. 1966. Nomi Generici polispecifici di Noctuidae del globo con scelte di specie tipo e observazioni. Annali del Museo Civico di Storia Naturale “Giacomio Doria” 76:57.

FORBES, W. T. M. 1928. Lepidoptera of New York and neighboring states. Part I. Cornell Univ. Agri. Expt. Sta. Mem. 68:19-30.

FRANCLEMONT, J. G. & E. L. Topp. Noctuidae. In Hodges, R. W., et al. 1983. Check list of the Lepidoptera of America north of Mexico. E. W. Classey Ltd. and The Wedge Entomological Research Foundation, London. xxiv + 284 pp.

GUENEE, A. 1852. Noctuelites III. In Boisduval et Guenée, Histoire naturelle des in- sectes, species géneral des Lépidoptéres. 7:215-216, 325-326. Librairie Ency. de Roret, Paris.

Hampson, H. G. 1913. Catalogue of the Noctuidae in the collection of the British Museum 13:84. Board of Trustees. London.

HUBNER, J. 1823. Zutrage zur Sammlung Exotischer Schmettlinge, (etc.), 2. Augsburg. 32 pp. + 8 numbered pages, figures 201-400, 33 plates (1819-1822).

KIMBALL, C. P. 1965. The Lepidoptera of Florida. Arthropods of Florida and neigh- boring land areas, I: v + 363 pages, 26 pl.

NyE, I. W. B. 1975. The generic names of the moths of the World. Vol. 1: Noctuidae, Agaristidae, Nolidae. Trustees of the British Museum, London.

Journal of the Lepidopterists’ Society 40(1), 1986, 20-22

THE LARVA AND PUPA OF LYCOREA PIETERI LAMAS (DANAIDAE)

DAVID KENNETH WETHERBEE

San José 71, Restauracién, Republica Dominicana

ABSTRACT. The larva and pupa of the danaid Lycorea pietri Lamas is described and figured for the first time. It differs most markedly from the larvae of other danaids in having only a pair of tentacles in front and none in the rear. The food plant is Carica papaya (Cariaceae).

During the last quarter of the 18th Century there was a gifted ar- chitect in northern Haiti who was also a naturalist 200 years ahead of his time. His work is unknown to biologists even though he painted from life accurately hundreds of Haitian animals and plants. Most of his subjects did not become known to binomial taxonomy until two generations later. He is unpublished, except that apparently Deshayes pirated some of his work and sent it to Buffon (Wetherbee, in press). Not only did he paint some 50 Lepidoptera, but he reared them from larvae and depicted the early stages and named several of them from their food plants.

He was ““M. de Rabié, marechal de camp, ingenieur en chef de la parties du nord de St. Domingue” and resided at Cap Haitian from at least as early as 1752 to about 1784 and died in Paris in 1785. His first insects were drawn in 1766. Folios of his work are now in the Blacker- Wood Library of McGill University. Through the courtesy of Miss Eleanor MacLean, McGill Librarian, I have been privileged to publish Rabié’s zoological subjects (Wetherbee, 1985a).

Rabié reared Lycorea pieteri Lamas (formerly called L. ceres Cra- mer, 1779) and depicted the larva, pupa, shed “‘robe’’ and imago. He called the larva ‘“‘chenille de papayer (food plant: Carica papaya of the Caricaceae) and the adult “le noeud de ruban’”’ (ribbon-bow). This is L. pieteri cleobaea Godart, 1819, the type-specimen of which was collected, undoubtedly, by Antonio Gonzales of the covert Baudin voy- age to Hispaniola in 1799.

Most of the larvae of Lepidoptera drawn by Rabié were those of moths, especially of Sphingidae (Wetherbee, 1985b). Only four other butterfly species of the 36 species illustrated by Rabié (Wetherbee, 1985a) show early stages: Danaus plexippus, Colobura dirce, Siproeta stelenes and Dione vanillae.

As can be seen from Fig. 1 (the black and white reproduction hardly does justice to the beauty of Rabié’s colored painting), the larva (per- haps fourth instar) is danaid in character, but unlike Danaus, which has pairs of both anterior and posterior fleshy tentacles, and unlike

VOLUME 40, NUMBER 1 x1

C 11 N ata ) . ° a“

\

Le Noeunpr Ruspan.

Fic. 1. Lycorea pieteri Lamas (L. p. cleobaea Godart, 1819) life history as painted by Rabié in Haiti in 1782.

Anetia which has none, Lycorea pieteri has only a pair of anterior tentacles. These are slightly longer than those of Danaus plexippus. The larva is smooth, about the same size as D. plexippus; both the head and posterior segment are black; the thoracic segments are white, followed by seven golden-yellow segments and then one white next to the last segment in front of the black tail-end. The narrow black bands, one at the anterior of each segment, have short, black, single, lateral, oblique, dash-like marks pointing backwards and downwards.

The hanging pupa is shown by Rabié in its lateral aspect only. It is similar in texture, size and shape to D. plexippus (but without the ridge) and is golden yellow with black bump-dots running in two arched lines on the sides, a few on the anterior parts, one prominent “‘occipi- tal” bump-dot and two anal ones. The cremaster is black and contrasts sharply with the white web.

One must pause to consider that this excellent work was accom- plished contemporary with Linnaeus’ 12th edition of the Systema Na- turae in a country which has had essentially no entomological research up to the present time. If we consider that Audubon was somewhat of a pioneer and hero, certainly Rabié was even a greater one.

Since viewing Rabié’s pictures, I have had the good fortune in No-

22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

vember of finding many of the larvae of this species in Restauracion, Republica Dominicana feeding on Carica papaya. The early stages are only slightly tinged with yellow. The fifth instar does not have the posterior segments white as shown by Rabie, but the thoracic segments tend to be whitish. The black tentacles are 15 mm in length, and the larva is about 40 mm. The ten narrow, black bands on the anterior of each segment send a spur toward each black spiracle and sometimes include it. The ultimate yellow segment has only three black dots representing the band. There are a pair of yellow eye-spots on the small black ultimate black segment. Both in Rabié’s illustration and in life, it is easy to mistake the tail for the head in the resting caterpillar.

The pupa is always suspended from the midvein of the green lechosa leaf about midway along the length of the leaf. It is dull waxen-yellow. The black dots run in a mid-dorsal series, a dorso-lateral series, and there is an arc of elongated dots on the wing following the curvature of the wing and an isolated black dot in the middle of the wing. As shown by Rabié, there is a pair of black bumps on the body near the base of the cremaster.

LITERATURE CITED

WETHERBEE, D. K. 1985a. Zoological exploration of Haiti for endemic species. Pub- lished privately, Shelburne, Massachusetts. 556 pp.

1985b. The sphinx-moths (Sphingidae) of Hispaniola and the 18th century moth

paintings of Rabié. Published privately, Shelburne, Massachusetts. 69 pp.

Journal of the Lepidopterists’ Society 40(1), 1986, 23-26

LIFE HISTORY AND HABITS OF EXOTELEIA ANOMALA HODGES, A PONDEROSA PINE NEEDLE MINER IN THE SOUTHWESTERN UNITED STATES (GELECHIIDAE)

ROBERT E. STEVENS!

Rocky Mountain Forest and Range Experiment Station, USDA Forest Service, Fort Collins, Colorado 80526

ABSTRACT. Exoteleia anomala, the larvae of which mine needles of ponderosa pine in Arizona and New Mexico, has a one-year life cycle similar to that of several species of needle-mining Coleotechnites. Each larva requires two needles to complete devel- opment. When numerous, larvae can cause highly visible foliage damage, but outbreaks do not appear to persist.

In summer 1977, I reared an undescribed species of Exoteleia (Ge- lechiidae) from foliage of ponderosa pines, Pinus ponderosa Dougl. ex Laws., near Silver City, Grant Co., New Mexico. In 1978, entomologists with the USDA Forest Service, Southwestern Region, Albuquerque, New Mexico, reported a needle miner infestation in ponderosa pines from an area near Show Low and Pinetop, Navajo Co., Arizona, some 90 km NW of Silver City. No moths were obtained at that time, but a resurgence of the population in 1981 provided material for study; it too was E. anomala. Collections and observations in 1981 and 1982, reported here, have made it possible to outline the species’ life history

and habits.

METHODS

Collections of foliage representing at least two years’ growth were made 21 October 1981, and 4 March, 6 April, 10 May, 22 June, 12 July, and 25 August 1982 for the 1981-82 generation, and a single collection from the 1982-83 generation was obtained on 11 November 1982. Life history events and larval habits were recorded following examination of the foliage, and sufficient numbers of larvae were pre- served to permit determination of instars. Adult voucher specimens are deposited in the U.S. National Museum of Natural History.

DESCRIPTION

The adult is described in detail by Hodges (1985). Briefly, it is a small, fragile moth, forewing length 4-5 mm, with a whitish head, mottled gray-brown to black and white forewings, and mottled brown and whitish abdomen. The hindwings are grayish, and have fringes of long hairs.

‘Present address: Dept. of Entomology, Colorado State University, Ft. Collins, Colorado 80523.

24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

First-stage larvae are tan, with light brown head capsules and pro- thoracic and anal shields. Instars 2, 3, and 4 are brown, with dark brown to black head capsules and anal shields. Prepupal fourth-stage larvae are dark brown to almost black.

Its appearance, coupled with its needle-mining habit, can lead nonspecialists to confuse this species with well known pine needle min- ers in the genus Coleotechnites.

LIFE HISTORY AND HABITS

Exoteleia anomala has a one-year life cycle. The moths fly and oviposit in midsummer, and the larvae overwinter. Pupation takes place in the mined needles. The life history is similar to that of a more northern but potentially sympatric species of needle miner Coleo- technites ponderosae Hodges & Stevens (Stevens 1973, Hodges & Ste- vens 1978), and an undescribed species of Coleotechnites from Pinus jeffreyi Grev. & Balf. in southern California (Luck 1976). However, larval habits of E. anomala are different.

Adults fly in June and July. Eggs were not seen, but they may be laid in old mined needles as with Coleotechnites pine needle miners (Stark 1954, Struble 1972, Stevens 1978), or in other locations near susceptible new foliage.

Larvae readily colonize foliage of the current year’s growth, in con- trast to C. ponderosae larvae, which seem to prefer older foliage (Ste- vens 1973). Examination of shoots from 12 heavily infested trees col- lected 21 October 1981 showed that 72% of the new (1981) needles had been invaded, as had 65% of the 1980 foliage. How much of the damage to 1980 foliage resulted from mining by the 1981 generation was not determined; certainly some was attributable to the 1980 gen- eration. A collection of 14 1981 shoots, made in July 1982 after all larval feeding had ended, showed that only 8% of the needles had escaped infestation.

Larval head capsule measurements indicate four instars (Fig. 1). Each larva utilizes two needles to complete development. First instars enter the first needle in late summer, molt, and remain there as second instars until spring. Most of the larvae in the 4 March 1982 collection appeared to have only recently arrived at the second needle. Larvae enter the first-mined needles in the middle third of the needle. This mine is short, only 1-2 cm; the part of the needle distad of the mine soon dies and fades. Larvae normally enter the second-mined needle within 1 cm of the tip, and more of the second needle is excavated. A set of 28 fully developed mines (from which adults had emerged) averaged 5.7 cm long (SD = 1.2 cm). The larvae cut small holes in the needle surface for disposal of frass; these are covered with silk from

VOLUME 40, NUMBER 1 95

oO

l aks) YV/ Y / h /, —A Wy "| Y; 1; YY rw 6 yy i Wi, ae HHUA om VA VAL

£2428 732 ce ‘444 748. 52° 56 Head capsule widths (mm)

Fic. 1. Head capsule measurements (n = 138) of Exoteleia anomala larvae.

within after they are no longer needed. There is usually only one needle miner per needle, but more than one larva per needle does occur. When this happens, normal entry locations are altered. Larvae located distally to others in a single needle do not complete develop- ment, and may move to another needle.

Pupation, as in the well known Coleotechnites pine needle miners and some species of Exoteleia, takes place within the last mined needle. The larva cuts a hole in the needle surface to allow for adult exit. The hole may be at either end of the mine. Pupae are dark brown to black, cylindrical, and 5.5 to 6.0 mm long. They are usually found 1 cm or more back from the exit hole, head pointing toward it.

POPULATION FLUCTUATIONS AND EFFECTS OF LARVAL FEEDING

Although heavy larval feeding can cause many needles to die, no permanent tree damage has been reported. This may be due to the

26 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

fact that outbreaks of E. anomala appear not to persist. For example, population densities were high enough to attract attention in Arizona in 1978, but no infestation was reported during the next 2 years. Larval numbers in 1981 appeared to decline markedly at the time of transfer from first- to second-mined needles. This may be a time in the insect’s life cycle when it is particularly vulnerable.

ACKNOWLEDGMENTS

I thank J. M. Schmid and J. C. Mitchell for making foliage collections, several of my co-workers for helpful comments on the manuscript, and R. W. Hodges for describing the species.

LITERATURE CITED

HopcEs, R. W. 1985. A new species of Exoteleia reared from ponderosa pine (Lepi- doptera: Gelechiidae). J. Lepid. Soc. 39:139-144.

HonpcEs, R. W. & R. E. STEVENS. 1978. Two new pine-feeding species of Coleotechni- tes (Gelechiidae). J. Lepid. Soc. 32:118-122.

Luck, R. F. 1976. Bionomics and parasites of a needle miner, Coleotechnites sp., infesting Jeffrey pine in Southern California. Env. Entomol. 5:937—942.

STARK, R. W. 1954. Distribution and life history of the lodgepole needle miner (Re- curvaria sp.) (Lepidoptera: Gelechiidae) in Canadian Rocky Mountain Parks. Can. Entomol. 86:]-12.

STEVENS, R. E. 1973. A ponderosa pine needle miner in the Colorado Front Range. USDA For. Serv. Res. Note RM-228, 3 pp. Rocky Mt. For. and Range Exp. Stn., Fort Collins, Colo.

STRUBLE, G. R. 1972. Biology, ecology, and control of the lodgepole needle miner. U.S. Dep. Agric. Tech. Bull. 1458, 38 pp.

Journal of the Lepidopterists’ Society 40(1), 1986, 27-35

BIOLOGY AND IMMATURE STAGES OF HEMILEUCA DIANA AND H. GROTEI (SATURNIIDAE)

PAUL M. TUSKES 7900 Cambridge #111D, Houston, Texas 77054

ABSTRACT. The primary host plant of Hemileuca diana in Arizona is Quercus oblongifolia, Mexican blue oak. Adult flight records extend from August to late Novem- ber, but peak emergence is in October. The primary host plant of H. grotei in central Texas is Quercus fusiformis, live oak. Adult grotei fly from late October to December. Adult Hemileuca grotei from New Mexico are similar to those from Texas. Immature stages and adults of both species are illustrated, as is the holotype of H. diana. Although closely related, hybrid matings between these two species do not produce viable ova.

Hemileuca diana Packard

Hemileuca diana is locally abundant in the mountains of Arizona but frequently difficult to locate or capture. Presently diana is known from Arizona, New Mexico, Colorado, and Sonora, Mexico. Although there are two old Texas records, the data are incomplete and probably in error (Ferguson 1971). Hemileuca diana is associated with the mon- tane oak habitat above 1,100 m (Fig. 1b) and is undoubtedly wide- spread from northern Mexico to Colorado. Because of its similarity to H. grotei Grote & Robinson, the two species have been confused in the literature, making it difficult to accurately determine the extent of either species distribution.

The specimens of diana illustrated by Ferguson (1971) are typical in appearance but slightly smaller than average. The forewing length of males from southern Arizona ranges from 23 to 28 mm, X = 24.3 mm (N = 34); females from 27.6 to 31.5 mm, X = 29.9 mm (N = 14). The fore- and hindwing ground color of the female is black or dark brown. The forewing of the male varies from brown to dark brown, while the hindwing is dark brown. Both sexes have a cream-colored medial line which passes distal to the forewing discal spot (Figs. 2a, b, i, j). The genitalia of a male diana was illustrated by Ferguson (1971) but was accompanied with grotei locality data. Ferguson (pers. comm.) re-examined the specimen and confirmed its identity as diana.

Minor geographical variation has been observed among the males in southern Arizona. This might be expected since many of the mountain ranges diana inhabits are isolated by distance and habitat. Males from the Huachuca Mts. in Santa Cruz and Cochise counties exhibit the most contrast between fore- and hindwing coloration (Fig. 2a). Speci- mens from the Santa Catalina Mts., Pima Co. (Fig. 2b), and Graham Mts., Graham Co., have forewings usually, but not always, darker than those from the Huachuca Mts. Specimens from the Chiricahua Mts.,

28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fics. la-e. Immature stages of Hemileuca diana. a, mature larvae, lateral and dorsal view; b, H. diana habitat, Pima Co., Arizona; ec, egg ring on Quercus oblongifolia; d, ventral and lateral view of male pupa; e, pupal chamber.

Cochise Co., are slightly smaller and have the least contrast between fore- and hindwings. Even though subtle differences are noted, over- lapping phenotypic variation among these populations makes the ap- plication of subspecific names unwarranted.

Adults from central Arizona (Figs. 2c, d) appear intermediate to diana and grotei. The cream colored medial forewing line is thin and disrupted by the discal spot, as in grotei (Figs. 2e, f, g), but the hind-

VOLUME 40, NUMBER 1] 29

f

; ) ; Se Arua atk,

Pham Cucih, Lac

Fics. 2a-m. Adults of Hemileuca diana, H. grotei, and immature stages. a, H. diana 6, Huachuca Mts., Cochise Co., Az; b, H. diana 4, Santa Catalina Mts., Pima Co., Az; e, Hemileuca sp. 6, Oak Creek Canyon, Coconino Co., Az; d, Hemileuca sp.? 4, Sunflower, Maricopa Co., Az; e—h, H. grotei 66, Burnet Co., Tx; i, Holotype °, H. diana; j, H. diana 9, Santa Catalina Mts., Pima Co., Az; k, H. grotei 2, Burnet Co., Tx; 1, mature H. grotei larvae, lateral view; m, mature H. groiei larvae, dorsal view (larvae from Burnet Co., ix),

wing medial line is well developed, and continues to the anal wing margin, as in diana (Figs. 2a, b). The forewing length of the six males examined ranged from 21 to 24 mm. The genitalia are variable but have distinct characteristics. I reared one larva collected on scrub oak

30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

in Oak Creek Canyon, Coconino Co., Arizona. The mature larva was grayish, the intersegmental area was brown, and appeared distinct when compared to diana larvae from southern Arizona, and more similar to grotei larvae from Texas. Females from this population have not been available for examination. The material from central Arizona appears distinct and may represent an undescribed taxon.

An examination of the diana holotype from Plum Creek, Colorado (Fig. 2i) confirms that there is no obvious difference between the type and females from southern Arizona (Fig. 2j). Ferguson also noted that the type agreed well with material from southern Arizona. Therefore, if a new name is proposed it should be applied to the central Arizona phenotype.

Biology

In southern Arizona, the flight period of diana extends from mid- September to late November, with the peak near mid- to late October, but this may vary from one mountain range to the next by as much as three weeks. Adults emerge in the morning and usually mate before 1200 h. Male flight activity usually occurs between 0930 and 1630 h. Females oviposit during the afternoon. The flight of the female is slower, more straight, and higher among the oaks than that of the male, which is rapid, erratic, and usually within 3 m of the ground.

Females deposited 80 to 140 ova in two to four separate egg rings near the tips of the branch close to the leaf clusters (Fig. lc). Field collected egg rings contained 25 to 65 greenish-gray ova. The eggs overwinter, and larvae usually hatch during April or early May.

The larval hostplant in Pima, Santa Cruz, Graham and Cochise coun- ties, is Mexican blue oak, Quercus oblongifolia Torr. Michael J. Smith (pers. comm.) indicates that larvae are occasionally found on Emory oak, Q. emoryi Torr. Early instar larvae are black and feed gregari- ously. During the first two or three instars, larvae prefer developing flower buds and new leaves. If disturbed fourth instar larvae release their grip, fall to the ground, and disperse.

Mature larvae feed singly and do not drop to the ground when disturbed. Some larvae have six rather than five instars but are identical in appearance to those with one less. The mature larva is gray with a distinctly dark gray dorsal area, and purple intersegmental areas (Figs. la, b). Larvae from the Santa Catalina and Huachuca Mts. were reared on five different occasions and mature larvae from the Graham Mts. were examined. Within these three populations, the larval phenotype is uniform. Pupation usually occurs in early June but James S. Mc- Elfresh (pers. comm.) has found mature larvae during early September.

VOLUME 40, NUMBER 1 |

Larvae pupate under leaf litter, where they construct a small chamber of debris tied together with silk (Fig. le).

On four different occasions a total of seven diana females from Arizona attracted and mated with wild male grotei in Texas. Collec- tively, the diana females deposited nearly 900 ova, but none hatched. Dissection of the hybrid ova about one month after pure diana and grotei ova hatched revealed that only a few contained dead, partially developed embryos; most ova appeared to be infertile. The high degree of genetic incompatibility between these two taxa leaves no doubt that they are distinct species. Females of both H. juno (Packard) and H. electra (Hwy. Edwds.) can be used to attract H. diana males (Tuskes 1984). A cross between an electra female and a male diana produced fertile ova, but upon hatching the larvae refused to feed on host plants of either species. Females of electra will also attract and mate with male H. eglanterina (Boisduval) but only infertile ova are produced (Collins & Tuskes 1979).

Larval Description

The larval description is based on 26 larvae reared to maturity from ova collected in 1982 by Mike Smith and the author, at Molino Basin, Santa Catalina Mts., Pima Co., Arizona. Preserved larvae are in the author’s collection.

First instar. Head: Black, diameter 0.7 mm. Body: Length 6 mm, width 1.4 mm. Ground color black. Dorsal area black, lateral and ventral surfaces dark brown to black. Dorsal and dorsolateral scoli forked near apex, one seta on each fork. All scoli black. True legs and prolegs black.

Second instar. Head: Black, diameter 1.5 mm. Body: Length 10-11 mm, width 2.0- 2.2 mm. Similar to first instar except for a small red dot between the dorsolateral and lateral scoli on abdominal (A) segments Al and A7.

Third instar. Head: Black, diameter 2.1-2.3 mm. Body: Length 19-20 mm, width 4.0-4.2 mm. Ground color black. All scoli black and branched. Lateral surface with traces of 3 incomplete lines extending length of larva. Line 1 incomplete, undulating, and white, touching lateral scoli and extending length of larva. Line 2 thin, white dash just anterior of dorsolateral scoli on each abdominal segment. Line 3 thin, white, and broken by black segmental area, passing just ventral to dorsal scoli. Spiracles, prolegs, and true legs black.

Fourth instar. Head: Black with short white secondary setae; diameter 3.5-3.9 mm. Body: Length 32-37 mm, width 9.5-11 mm. Ground color black. Three broken white lateral lines extend length of larva. Line 1 passes between lateral scoli and is well defined only on posterior portion of each segment. Line 2 passes between dorsolateral scoli; prominent on posterior of each segment. Line 8 passes mid-way between dorsolateral and dorsal scoli; broken by intersegmental area. Area between line 1 and ventral surface gray; between lines 1 and 2 (spiracular area) black; between lines 2 and 3, dark gray. Dorsal and mid-dorsal area black. All scoli black with white or hyaline colored spines. Short white secondary setae extend from white or light gray pinacula on lateral and dorsal surfaces. Prominent red-orange dot occurs between lateral and dorsolateral scoli on Al; similar but smaller dot on A7. Spiracles black. All shields black. Inner portion of proleg and ventral surface brown. True legs black.

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Fifth instar. Head: Dark brown with numerous white secondary setae; diameter 5.1- 5.4 mm. Body: Ground color dark gray. Length 47-58 mm, width 7.5-10 mm. Dorso- lateral, lateral and sublateral scoli with black shafts and black and gray spines; base of shaft ringed with gray. Dorsal metathoracic scoli to A8 rosette type with black tips and dark gray base. Pro- and mesothoracic dorsal scoli enlarged. Two prominent light gray lateral lines extend length of larva and divide dorsal and lateral areas. Line 1 undulating, and passes between lateral scoli. Line 2 passes just ventral to dorsolateral scoli and is interrupted by maroon intersegmental area. Lateral segmental area between lines 1 and 2 gray with numerous light gray pinacula which may contain short white secondary setae. Dorsal area black with fewer light gray pinacula than lateral segmental area. Intersegmental area between lateral and dorsolateral scoli black or dark gray. Dorsal intersegmental area maroon. Ventral surface light brown to flesh with light gray pinacula in intersegmental area. Prominent shiny orange spot occurs posterior and near to spiracles on Al and A7. Thoracic shields black. True legs dark brown. Prolegs gray with dark gray shields. Spiracles orange.

Hemileuca grotei Grote & Robinson

Until recently, little information was available on the biology of Hemileuca grotei. Ferguson (1971) illustrated both grotei and diana, and pointed out major differences in adult morphology, wing pattern, and distribution. Kendall and Peigler (1981) provided additional in- formation on the distribution and flight period of grotei in Texas. No mention has been made of the extreme phenotypic variation found in the adults, and only a partial description of the immature stages has been published.

Most males and females have a well defined white medial band on the forewing interrupted by the dark discal spot. On the male hind- wing, the white medial line is usually widest between M1 and M2, then narrows and terminates between Cul and 2A (Figs. 2e, f, g). In females, the medial hindwing line is more developed, seldom strongly tapering, and extends beyond 2A to the anal margin of the wing (Fig. 2k).

Some adults are almost entirely black, with only the white bar in the center of the forewing discal spot present. Others have only a trace of the hindwing medial line (Fig. 2h). Of approximately 400 males examined, 3 to 7% represent the dark phenotype. The occurrence of a dark phenotype is not uncommon in many Hemileuca species (Tuskes 1984). Normally, the ground color of the male is dark brown to nearly black. Sometimes the base of the forewing is dark gray, and the medial and distal portions are light gray (Fig. 2g). The ground color of the female is dark brown to black. Forewing length of males from Inks Lake, Burnet Co.; Texas, ranged from 22.0 to 25.3 mm, X = 24.2 mm (N = 30); females varied from 26.0 to 29.4 mm, X = 28.1 mm (N = 30).

In addition to Texas, Hemileuca grotei occurs in New Mexico. Rich- ard Holland collected a series at Dome Lookout (Sandoval Co., X-11-

VOLUME 40, NUMBER 1 33

84, elev. 2,460 m), and in the northwestern corner of the state at Fort Windgate (McKinley Co., IX-30-1975). The average forewing length of the Dome Lookout males is identical to that of central Texas pop- ulations. There are subtle differences in coloration and the frequency of various phenotypes between central Texas and New Mexico material examined. In 38% of New Mexico males (N = 18), the white medial hindwing line is absent; thus the wing is solid black. Further, there does not appear to be a relationship between the presence or absence of the medial hindwing line and the development of the forewing medial line. In Texas populations, if the hindwing line is reduced or absent, the forewing line also tends to be reduced. One specimen ex- hibits the same grayish scaling on the forewing as illustrated in Figure 2g. The only female examined was identical to those from central Texas. ;

Biology

Both males and females are active day flyers. The flight season in central Texas extends from October to early December, but peak emer- gence is near mid- to late November. Local climatic conditions signif- icantly influence duration and extent of daily adult flight. In late No- vember 1982, 26 males were collected during a one-day trip. The high temperature for the day was 14.4 C, with light rain, strong gusty winds, and 85% cloud cover. The first males were observed at 1020 and the last at 1480 h. During a trip in 1988, conditions at the same location were clear with a high of 25 C and light winds. At that time males were very abundant, and in flight from 0910 to 1800 h. Females were observed and captured in flight from about 1200 to 1800 h.

Newly emerged larvae are black and feed gregariously. The dark coloration may aid thermoregulation and increase activity during the early spring. Like those of H. diana, fourth instar grotei larvae tend to drop from the branch if disturbed. During the late fourth and fifth instars, larvae exhibited almost equal preference for flowers or leaves. The natural larval hostplant is Quercus fusiformis Small. Kendall and Peigler (1981) reported that Q. havardii Rydberg x Q. stellata Wan- genheim, Q. texana Buckley, and Q. marilandica Muenchhausen are also utilized, but to a lesser extent. Mature larvae measure 39 to 48 mm in length, and have a gray ground color. The dorsal surface is darker than the lateral surface and the intersegmental area is reddish brown (Fig. 2], m).

Before pupation the larva appears to darken, shrink in size, and the intersegmental color becomes less prominent and the light yellow pi- nacula become light gray. Larvae wander from one to three days be- fore constructing a loosely woven cocoon in the leaf litter. Seventy-

34 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

seven larvae were reared from ova to maturity; 43 were males and 34 were females. All appeared to have five larval instars, and exhibited little variation. As with H. diana, most grotei pupae produce adults the same year, but both species have the ability to spend two years in the pupal stage.

Larval Description

The description is based on 77 larvae reared to maturity from ova deposited by a female collected by the author at Inks Lake State Park, Burnet Co., Texas.

First instar. Head: Black, diameter 0.8 mm. Body: Length 6 mm, width 1.2 mm. Ground color black to dark brown. Dorsal pro- and mesothoracic scoli forked at tip. Metathoracic scoli forked or spiked, remaining scoli spikelike. All scoli black with light colored setae extending from apex of each shaft. Ventral surface dark brown to black. True legs and prolegs black.

Second instar. Head: Black, diameter 1.4 mm. Body: Length 10 mm, width 1.8 mm. Similar to first instar with one exception: A small orange patch occurs between the dorsolateral and lateral scoli of Al.

Third instar. Head: Black, diameter 1.8-2 mm. Body: Length 18-19 mm, width 3.4 mm. Ground color black. Pro- and metathoracic scoli enlarged; shaft black, spines yellow and black. Dorsal abdominal scoli black; spines yellow with black tips, rosette pattern developing. Dorsolateral, lateral and sublateral scoli black with black spines. Trace of 3 incomplete light yellow lines extend length of larva, lines variable. Line 1 undulating, subspiracular, extending from Al to AQ, and touching base of each lateral scolus. Line 2 broken, consisting of dots passing along base of dorsolateral scoli. Line 3 often well developed, extending from T2 to A8, passing just ventral of dorsal scoli, often disrupted by intersegmental area. Orange dot on Al and smaller dot on A7, both set between dorsolateral and lateral scoli. True legs, prolegs, and spiracles black. Small yellow pinacula occur on all segmental areas, including ventral surface.

Fourth instar. Head: Black, diameter 3.0-3.4 mm. Body: Length 31-33 mm, width 5 mm. Ground color black. Dorsal thoracic scoli T1 and T2 elongated, with black shafts and gold spines. Dorsal abdominal scoli rosette type with black base and gold spines, and light brown to black tips. Dorsolateral, lateral and sublateral scoli with black shaft and gold spines. Three incomplete light yellow lines extend length of larva, some may be poorly developed. Line 1 undulating, subspiracular, connecting base of all abdominal lateral scoli. Line 2 lightly marked, sometimes absent, in line with dorsolateral scoli on segmental area. Line 8 passes just ventral to base of dorsal scoli; interrupted by black intersegmental area. Body covered with small light yellow pinacula with a short hyaline setae extending from each. Mid-ventral surface dull orange-brown. Spiracles orange. Prominent orange dot occurs between lateral and dorsolateral scoli of Al; smaller dot similar in color and location on A7. True legs and prolegs black.

Fifth instar. Head: Black with short white secondary setae, diameter 4.6-5.0 mm. Body: Length 39-48 mm, width 7.0-8.5 mm. Ground color gray. Dorsal prothoracic scoli (T1) elongated; shaft black with black and light yellow to cream colored spines. Dorsal metathoracic scoli similar to dorsal T1 scoli but with light yellow to yellow-gray rosette spines at base. Dorsal abdominal and TS8 scoli rosette type with yellow to yellow- gray spines and black tips. Dorsolateral, lateral, and sublateral scoli with black shaft and light yellow to white spines. Two to three incomplete light yellow to light gray lateral lines extend length of larva; the two dorsalmost may be poorly developed. Line 1 well developed, undulating, subspiracular, extending from A2 to A9, and touching base of each lateral scolus. Line 2 lightly marked, sometimes absent, in line with dorsolateral scoli on segmental area. Line 3 passes just ventral to base of dorsal scoli; interrupted by intersegmental area. Lateral surfaces gray, dorsal area dark grayish-black. Body covered

VOLUME 40, NUMBER 1 35

with small light yellow to cream or light gray pinacula with a short hyaline seta extending from each. Lateral intersegmental are brown to rust. Spiracles light orange. Prominent brown dot occurs between lateral and dorsolateral scoli on Al and A7. Ventral surface light brown. Thoracic shield black. True legs and prolegs dark brown to near black.

Kendall and Peigler (1981) published a partial description of a ma- ture grotei larva but did not give the source of their material. Com- parison of their larval description with larvae from Burnet Co. indi- cates a number of differences. Larvae from Burnet Co. have a shiny black head; reddish brown intersegmental area; scoli of three different configurations and size; and brownish orange spiracles. Kendall and Peigler described grotei larvae as having a rusty brown head with mottled black patches; maroon intersegmental area; almost equally developed scoli; cream colored spiracles; and concluded they were most similar to larvae of H. burnsi (Watson). The larval description of burnsi by Comstock (1937), together with my observations suggest there is little similarity between these two species. In coloration and morphology, groeti larvae from Burnet Co., Texas are most similar to diana and diana-like larvae from central Arizona.

ACKNOWLEDGMENTS

I thank James S. McElfresh, Michael J. Smith, Kenneth C. Hansen, Richard Holland, Douglas Ferguson, and Donald E. Bowman for providing data or material. I also thank Mike Collins and Ann Tuskes for their comments and suggestions on the manuscript.

LITERATURE CITED

COLLINS, M. M. & P. M. TusKEs. 1979. Reproductive isolation in sympatric species of day flying moths (Hemileuca: Saturniidae). Evolution 33:728-733.

ComsTOCK, J. A. & C. M. DAMMERS. 1937. Notes on the early stages of three California moths. Bull. So. Calif. Acad. Sci. 36:68-78.

FERGUSON, D. C. 1971. The Moths of America north of Mexico. Fascicle 20.2a Bom- bycoidea (in part). Classey, London, pp. 1-154.

KENDALL, R. C. & R. S. PEIGLER. 1981. Hemileuca grotei (Saturniidae): Natural his- tory, spatial and temporal distribution. J. Lepid. Soc. 35:41-50.

TusKEs, P. M. 1984. The biology and distribution of California Hemileucinae (Satur- niidae). J. Lepid. Soc. 38:281-309.

Journal of the Lepidopterists’ Society 40(1), 1986, 36-53

NATURAL HISTORY AND ECOLOGICAL CHEMISTRY OF THE NEOTROPICAL BUTTERFLY PAPILIO ANCHISIADES (PAPILIONIDAE)

ALLEN M. YOUNG

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

MURRAY S. BLUM Department of Entomology, University of Georgia, Athens, Georgia 30602

HENRY M. FALES AND Z. BIAN

Laboratory of Chemistry, National Heart, Lung and Blood Institute, Bethesda, Maryland 20014

ABSTRACT. The life cycle, behavior, and chemical ecology of the neotropical but- terfly Papilio anchisiades idaeus Fabricius (Papilioninae) were studied, using larvae from a single cluster of eggs obtained in NE Costa Rica. The butterfly places large clusters of eggs on the ventral surface of older (bluish green) leaves of Citrus. The larvae are cryptically colored and exhibit communal resting, molting, and nocturnal feeding be- havior. Fourth and fifth instars perch on branches and trunk of the host plant. Larvae are parasitized by the braconid wasp Meteorus sp., and the ant Camponotus rectan- gularis attacks and kills pupae located on the host plant. Paper wasps do not attack large larvae or pupae, even though their nests are often abundant in Citrus trees occupied by P. anchisiades. Pupae on substrates away from the host plant may survive ant predation. Larvae readily evert the osmeterium when provoked; a very pungent, disagreeable odor is noticeable (to humans) only in the fifth instar. The principle components of the os- meterial secretions change both qualitatively and quantitatively with the molt to the fifth instar. The major corstituents of the secretions of third and fourth instars are sesquiterpenes including “a-bergamotene’’, a-acoradiene, “‘a-himachalene’’, and isomers of farnesene; the main secretion of the fifth instar is dominated by isobutyric acid and 2-methylbutyric acid with sesquiterpenes, aliphatic hydrocarbons, long-chain alcohols, and carboxylic esters constituting minor constituents. The possible adaptive significance of this shift in the chemistry of the osmeterial defensive secretion is discussed.

The neotropical butterfly Papilio anchisiades idaeus Fabricius (Pa- pilionidae: Papilioninae) is well known in Mexico, Central and South America (Seitz 1908, Ross 1964a, b). It is a large tailless swallowtail with velvety-black wings bearing white patches dorsally on the fore- wings, and deep red to lavender blotches dorsally on the hindwings. This butterfly is commonly seen around clumps or groves of Citrus trees (Rutaceae), the host plant of the caterpillars (Stoll 1781, Carac- ciolo 1981, Dewitz 1878, Moss 1919). The life cycle and early stages have been incompletely described (Caracciolo 1891, Dewitz 1878, Ehr- lich & Ehrlich 1961, Jones 1881, Moss 1919, Oliveira 1977, Ross 1964a, Stoll 1781). In this paper we describe and illustrate the early stages, and present new information on the behavior of immature stages, on parasitism, predation, and egg placement. In addition, we analyzed osmeterial secretions of third, fourth and fifth instar larvae to compare

VOLUME 40, NUMBER 1 37

the chemistry of these defensive secretions with those of other species of Papilio. Since recent investigations demonstrated both qualitative and quantitative changes in the secretions between Papilio fourth and fifth instars of other species (Seligman & Doy 1972, Burger et al. 1978, Honda 1980a, b, 1981), we wondered if this was also the case for P. anchisiades.

MATERIALS AND METHODS

A cluster of 53 eggs was obtained by observing one female P. an- chisiades ovipositing on a 4-m high lemon (Citrus) tree at the edge of a grassy cattle pasture at “Finca La Tirimbina” at 13800 h on 4 March 1982 in NE Costa Rica. This locality is about 10 km E of La Virgen (10°23'N, 84°07’W, 220 m elev.), Heredia Province, and well within the Premontane Tropical Wet Forest region (Holdridge 1967). The eggs were collected by cutting the branch with the leaf bearing them, and placing the cutting in a clean, air-tight, clear plastic bag. The larvae were reared following previously established methods (Young 1972), which included daily observations and periodic changing of leaves and removal of frass and other debris. The duration of each life stage was measured, and feeding and resting behavior noted. The cul- ture of first instars was transported to Milwaukee, Wisconsin, where the rearing continued until adult emergence. During the Wisconsin rearing period, the larvae were fed leaves from a Citrus tree in the greenhouse at the Milwaukee Public Museum.

The rearing period extended from 4 March through 23 April 1982, and during this time osmeterial secretions were collected from all avail- able larvae by instar. These secretions were collected in the standard way: each larva was gently pinched with fine forceps, and the everted osmeterium quickly wiped with a small square of filter paper and dropped immediately into a vial of methylene chloride. Several such “milkings’” were done within an instar between 1400-1500 h, and samples were thus obtained for the third through fifth instars. Milkings from different larvae were pooled at each sampling date as follows: 24 third instars milked 1-2 April; 22 fourth instars milked 5-9 April; 22 fifth instars milked 13-19 April. The apparency of odor associated with everted osmeteria was also noted.

Chemical Analyses

Gas chromatography-mass spectrometry subdivides complex com- pounds into molecular weight fractions. It was done using 15 m x 0.3 mm I.D. OV-17 or SE-30 fused quartz capillary columns (J. and W. Scientific Co., Rancho Cordova, CA) in an LKB 2091 spectrometer, with a splitless injector system (J. and W. Scientific Co.). Confirmation

38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

of the low boiling esters was accomplished on an LKB 900 spectrometer using a 2 m X 2.5 mm I.D. 10% SP-1000 packed column. Synthetic octadecyl and eicosanyl esters were chromatographed on a 2 m X 2.5 mm I.D. 1% OV-17 packed column. Both spectrometers were main- tained during scanning at 70 ev, at a source temperature of 270°, and at 270 amp ionizing current.

Synthesis of Eicosanyl Esters of Valeric and Isovaleric Acids

These compounds were prepared for use as standards by combining an excess (1 g) of the corresponding acid chlorides with 1 g of eicosanol in 8 ml of pyridine. After 1 h the mixtures were poured into water and extracted into ether, washed with dilute sodium bicarbonate, and the ether evaporated. The oils were then chromatographed directly providing only one peak on a 1% SE-54 packed column.

Eicosyl valerate. Retention times, the result of component molecular weights and their percent representation in samples at a specified tem- perature, characterized specific complex compounds with mass spec- trometry. Retention temperature 270°, MS: m/z (rel. intensity) 382(0.9 m*.), 353(0.4), 367(0.060), 340(0.4), 325(2), 280(8), 195(1), 181(1), 167(2), 158(1), 153(2), 139(4), 125(7), 111(14), 103(100, valeric acid + H), 102(17), 97(24), 85(22), 83(23), 82(10), 71(13), 70(9), 69(14), 57(21), 56(7), 43(6).

Eicosyl isovalerate. Retention temperature 265°, MS: m/z (rel. in- tensity) 382(0.8, m*.), 367(0.4), 340(0.2), 325(1.5), 280(7), 252(8), 195(1), 181(1), 167(2), 158(1), 153(8), 189(4), 125(6), 111(10), 103(100, valeric acid + H), 97(15), 85(19), 83(14), 71(7), 70(5), 69(9), 57(11), 48(4).

Preparation of methyl! esters of osmeterial extract. Diazomethane in ether prepared from N-nitro-N-nitrosomethylguanidine (Aldrich Chemical Co., Milwaukee, WI) was added to 20 ul portions of the extract in methylene chloride until a yellow color persisted. Aliquots of this solution were directly injected.

RESULTS Description of Early Stages

Eggs (Fig. 1) spherical, sculptured, about 2 mm diam, with lateral pair of ridges fusing into bilobed knob; honey-colored; not changing in color before hatching; duration of stage: seven days.

First instar cylindrical with fine down and slightly bulbous head; initially about 6 mm long; cuticle translucent amber, darkening to “dirty” greenish brown following first feeding on plant tissues; lateral body profile tapered; no tubercles and no discernible markings on cu- ticle; duration of stadium: seven days.

VOLUME 40, NUMBER 1

‘ g cs ee . wa > ~~ . . : ; Q : ne ‘> ° A a : . ; WAS Bees ae 5 Vea Reo wy: % . : D Fass : hs 2S :- a ‘ Pos a BS « be ‘ a sy Re ye Bs on er” ie a ee

Fic. 1. Papilio anchisiades. (A) position of egg cluster on Citrus leaf (ventral surface); (B) orientation of individual eggs in cluster; (C) surface sculpturing of individual eggs;

(D) second instars.

Second instar (Fig. 1) similar to first but with larger head relative to trunk; more delineation of trunk segments; first three segments and last four dull orange, middle segments greenish; head glossy orange;

attained body length of 10-13 mm in seven to nine days. Third instar (Fig. 2) strikingly different from previous instars; swelled

40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 2. Papilio anchisiades. (A) third instars immediately following molt; (B) fourth instars several days after molting; (C) fifth instar, lateral aspects; (D) aggregative behavior of fifth instars.

thoracic region; coloration a variegated brown and white cuticle with “oily” appearance; head brownish orange and glossy; head hidden and overshadowed by anterior trunk region; attained a length of 20-23 mm in eight days.

VOLUME 40, NUMBER 1 4]

Fourth instar (Fig. 2) similar to third but with decrease in thoracic swelling; more pronounced mottling of rich brown and white blotches on trunk; both third and fourth instars with tubercles further described in Oliveira (1977); attained length of 32-36 mm in ten days.

Fifth instar (Fig. 2) with trunk cuticle “lacework’ pattern of choc- olate-brown background with network of lines and blotches of white; cuticle “warty” due to small tubercles and foldings (further described in Oliveira 1977); prolegs white with brown speckling; head brownish and smaller than anterior trunk; dorsally trunk cuticle bears a series of diamondlike velvety-brown blotches; attained length of 59-62 mm in 20 days. Total larval period: 45 days.

All instars with deep-orange osmeteria, short and stubby in first three instars, long and filamentous in last two. Osmeterium of fifth instar 9- 10 mm long. The prepupa (Fig. 3) contracted in body length and darkened in coloration before the final ecdysis.

Pupa (Fig. 3) 37-39 mm long and 21-23 mm at greatest width; resembles broken twig; color pattern a variable mosaic of brown, gray, green, and white, but usually with large, “lichenlike’’ blotch on pos- terior two-thirds of wing pads extending posteriorly into dorsal area of abdomen; spiracle openings marked in black; duration of stage: 18-22 days. Overall egg-to-adult time: 70-74 days.

Adults eclosed rapidly, and wings were fully expanded (Fig. 3) with- in 25 min, and usually between 0800-0900 h. Sex ratio of 25 pupae: 10 females and 15 males.

Behavior of Stages

Eggs placed in tight rows on the ventral surface of mature Citrus leaf (Fig. 1); even though “‘young” or “‘fresh” (greenish-yellow) leaves available, eggs were placed on older leaf, near the distal end of the branch; butterfly clung to edge of leaf and curled abdomen under while ovipositing for 1 h. When frightened away, it did not return to resume egg laying on the same or several subsequent days. Other ob- servations in Costa Rica indicate that this species oviposits on both mature (greenish-blue) leaves and yellowish-green fresh leaves of Cit- rus in both wet and dry forest regions. All eggs in the cluster touched one another and hatched synchronously taking about 4 h for all larvae to vacate egg shells. Egg shells were immediately devoured by larvae, and larvae remained as one group in the first two instars (Fig. 1). First instars occupied the same leaf as the eggs, and started feeding at the edge of the leaf (Fig. 4). Feeding throughout all instars was synchro- nous and nocturnal. Breakup into two or more subgroups began in the third instar and continued through the fifth (Fig. 2). Fourth instars

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Fic. 8. Papilio anchisiades. (A) prepupa; (B) pupa; (C) aggregation of pupae; (D) freshly eclosed adult.

stayed in small groups on the branch rather than on leaves like previous instars.

In field observations fourth and fifth instars aggregated on the trunk of the host, and pupation occurred on the trunk, on nearby buildings or other substrates near the host.

VOLUME 40, NUMBER 1 43,

Fic. 4. Papilio anchisiades. Feeding pattern of young larvae on a leaf of Citrus.

Larvae of all five instars evert the osmeterium when prodded with forceps, but response is much quicker in the first three instars than in the last two. Eversion of the osmeteria in the first three instars was unaccompanied by odor at close range. A strong, disagreeable odor, best described as ‘‘sweaty socks,’ was apparent when the osmeteria of the last two instars were everted. Growth rates of larvae within a group were mostly similar, and molting was synchronous. Molting required

44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

one to two days. However one to three individuals within the group were smaller, and differed by a full instar. Pupation appeared to occur in two “pulses,” with larger larvae (presumably females) being the last to pupate.

In field observations, more than one cohort of larval P. anchisiades occurred in a single Citrus tree, often in trees with nests of paper wasps (Polistes and various polybiines). In a Citrus tree in a pasture at Palo Verde, Guanacaste Province, Costa Rica, studied between 9- 11 November 1969, separate groups of 40-70 larvae were found, and each group contained 3-8 smaller larvae. Most larvae were fifth instar, and the smaller ones second and third instars. A total of nine pupae were scattered on branches, and all were being attacked by the ant Camponotus rectangularis Emery. An additional 25 pupae, without ants, were found on a nearby weather-beaten tool shed 5 m from the tree and separated from it by tall grasses. A total of 53 paper wasp nests were in the tree and another 74 nests on the shed. Individual wasps often perched near the pupae on the sides of the shed, but did not attack them. Pupae on the tree were not observed to be attacked by wasps. Successful eclosion of four adults was observed. No pupae on the shed were attacked by C. rectangularis during the two days of observation.

Several third instars collected in October 1969 at Naranja, Zaragoza, El Salvador, were parasitized by the braconid Meteorus sp. A second unnamed species of the same genus has been recorded from P. an- chisiades in Venezuela (Paul Marsh, pers. comm.). Together, both rec- ords are new, and represent the first reports of parasitism by Meteorus on the Papilionidae (Paul Marsh, pers. comm.).

A major feature of larval behavior in P. anchisiades in both field and laboratory is the close physical contact among individuals, al- though laboratory individuals sometimes rested and fed solitarily.

Mass Spectral Analysis of Osmeterial Extracts

Extracts of the fifth instar (Fig. 5) showed a poorly resolved series of short-chain acids and esters, followed by traces of sesquiterpenes eluting from 142-170°, and finally (Fig. 6) a series of hexadecyl, oc- tadecyl, and eicosanyl esters of butyric and valeric acids. The early eluting compounds were ethyl isobutyrate, methyl 2-methylbutyrate, ethyl 2-methylbutyrate, isobutyric acid, isovaleric acid and 2-methy]- butyric acid, eluting in that order. Reexamination on a 10% SP-1000 packed column confirmed these assignments and revealed a trace of ethyl 3-hydroxybutyrate eluting just after ethyl 2-methylbutyrate. Also observed were traces of acetic acid and ethyl acetate. All compounds were identified by their mass spectra (Heller & Milne 1976). As found

VOLUME 40, NUMBER 1 45

Ethyl lsobutyrate | | | Isobutyric Acid Isovaleric Acid Ethyl 2-Methylbutyrate

2-Methylbutyric Acid —

RESPONSE

ince Butyrate Octadecyl Valerate

Eicosanyl Butyrate

Eicosanyl Valerate Methyl 2-Methylbutyrate CoH ag Octadecanol CL | \ Va l

249° 228° 205° 170° 142° 80° 50° TIME/TEMP

Fic. 5. Chromatographic analysis of fifth-instar osmeterial secretion in Papilio an- chisiades (SE-30 capillary; 15 m x 0.30 mm I.D.; 10°/min.).

by Honda (1981) for other Papilio species, 2-methylbutyric and iso- butyric acids were major components accompanied by smaller quan- tities of isovaleric acid. Conversion to the methyl esters allowed quan- titation of these acids in the ratio 1:0.75:0.021, respectively, as determined on an SE-54 capillary column.

Expansion of the chromatographic region between 80° and 228° (Fig. 6) allowed two terpenes, a-bergamotene and E-b-farnesene (peaks 1 and 2), to be tentatively identified by comparison of their spectra with published compilations (Heller & Milne 1976). Peak 4 was tentatively identified as a-himachalene by similar comparison, while peak 9, as- sumed to be a sesquiterpene from its mass spectrum (Table 1), was unique to the fifth instar. The mass spectrum of peak 8 was similar

46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Mass spectra of sesquiterpenes identified in the osmeterial secretions of Papilio anchisiades. Retention temperatures (first numbers) and relative amounts ( ) refer to peaks in Fig. 7.

1. a-bergamotene: m/z 204(M*., 2), 161(3), 131(5), 119(53), 93(87), 69(53), 55(40), 43(67), 41(100)

2. A teh isomer: m/z 204(M*., 5), 161(6), 183(14), 119(7), 93(41), 69(100), 55(20), 43(23), 41(99)

3. B-accoradiene: m/z 204(M*., 6), 182(2), 161(6), 183(16), 120(18), 107(8), 93(50), 91(25), 81(25), 79(25), 69(100), 67(25), 57(20), 55(20), 41(90)

4. “a-himachalene”’: m/z 204(M*., 25), 189(7), 161(10), 147(9), 188(17), 121(25), 119(22), 93(100), 79(29), 69(21), 55(31), 48(63), 41(88)

5. A farnesene isomer (same as 2 above, but mixed with a-himachalene)

6. A farnesene isomer m/z 204(M*., 10), 161(18), 1383(20), 120(18), 109(16), 93(49), 69(100), 41(79)

7. “B-selinene”’: m/z 204(Mt., 15), 189(3), 161(3), 121(22), 119(20), 109(21), 98(100), 80(80), 69(25), 41(42)

8. Unknown: m/z 220(M*., 20), 205(5), 177(4), 163(2), 151(6), 149(4), 137(100), 185(53), 110(76), 109(51), 95(46), 82(29), 69(35), 55(40), 43(20), 41(85)

9. Unknown (only in 5th instar): m/z 204(M*., 25), 189(18), 169(31), 183(25), 121(50), 119(44), 105(52), 93(69), 91(50), 79(88), 77(31), 69(25), 55(44), 53(38), 43(81), 41(100)

but not identical to the spectrum of caryophyllene oxide reported by Honda (1981). No evidence was found for monoterpenes or the ele- mene, selinene or germacrenes reported by Honda (1981). Also iden- tified in this sean were C,,-C,, saturated and unsaturated hydrocarbons as well as naphthalene, dichlorobenzene, and phthalates, all of which are regarded as artifacts. |

The acid components of the hexadecyl, octadecyl, and eicosy] esters were expected to be isobutyric and either 2-methylbutyric or isovaleric acids in view of the large quantities of the corresponding free acids that were present (Fig. 5). In fact, comparison of the mass spectra of the last peak with spectra of synthesized samples of eicosyl n-valerate and eicosyl isovalerate reveals that the natural product is the former ester. Thus, the molecular ion of eicosyl isovalerate was slightly less intense relative to high mass peaks, and showed enhanced loss of meth- yl compared to eicosyl valerate. The other three peaks are also esters of the n-butyric and n-valeric acids by the same reasoning. Fig. 6 also shows the presence of the corresponding alcohols, octadecanol and ei- cosanol, easily identified by reference to library spectra.

Gas chromatograms of the third and fourth instars were nearly iden- tical, but presented an entirely different picture (Fig. 7). Both short and long chain acids and esters were missing, and only sesquiterpenes were present. As in extracts from the fifth instar, only a-bergamotene, -acoradiene and three farnesene isomers were identified with confi- dence by comparison with reference spectra. The major component was a compound whose spectrum resembled, but was not identical

VOLUME 40, NUMBER 1] 47

Octadecanol 8 C,H, Dibutylphthalate LW 7) Zz oO Diisobutylphthalate = n-C,,H,, Diethylphthalate cc Eicosanol C_H ly WS ain n-C..H n-C,.H3, cae | Oaeo: : n-C.H ~ | ZA i 22° 46 aS | AAC ee Za)

IL

Hexadecy] butyrate

Hexadecyl valerate |

249° 228° 190° 170° 150° 142° 80° 5Oe

TIME/TEMP

Fic. 6. Mass spectral analysis of fifth-instar osmeterial secretion in Papilio anchisi- ades, highlighting the terpene region of the spectra (SE-30 capillary; 15 m x 0.3 mm I.D.; 10°/min.).

with, caryophyllene oxide as reported by Honda (1981). Peaks 4 and 7 are very similar, and both resemble library spectra (Heller & Milne 1976) of a-himachalene or b-selinene, but neither corresponds to the b-selinene spectrum reported by Honda (1981). Mass spectra of ses- quiterpenes are shown in Table 1.

DISCUSSION

Papilio anchisiades, along with P. cresphontes and P. thoas, and a few others, exploits various Rutaceae as larval food plants (Brower 1958). It differs from other rutaceous-feeding Papilio species by its unique larval aggregative habits, a result of cluster egg placement on the larval host plants. The rutaceous-feeding habit is shared world-

48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

wide by several species of Papilio (Munroe 1960), an association per- haps mediated by the rich profiles of specific alkaloids so characteristic of this plant group (Hegnauer 1963). We propose that the widespread abundance of P. anchisiades from southern Brazil to Mexico and even southern Texas (Ehrlich & Ehrlich 1961), is in part due to the broad distribution of Citrus as an exotic rutaceous host plant coupled with the distribution of other Rutaceae with forest habitats.

Larval host-plant selection in butterflies usually involves a highly structured sequence of visual and olfactory responses (Crane 1955, Swihart & Swihart 1970, Ilse 1937). Vaidya (1969) studied the color preferences of ovipositing P. demoleus L. on Citrus in laboratory stud- ies, and concluded that both color and scent are required for proper egg placement and noted that butterflies preferred blue-green hues of leaves over yellow-green hues. Within the Rutaceae, differences among genera and species for certain substances in the leaves determine pat- terns of host specificity in egg-placement behavior among different papilionids (Ichinose et al. 1981). Age-specific differences in odor and color in Citrus and other Rutaceae may determine patterns of egg placement in P. anchisiades in nature, and the field data for this species in Costa Rica supports a partial preference concept for the older, more bluish-green leaves of host plants. Depending on annual phenological patterns of flushing, such egg-placement substrates may vary in abun- dance at a locality and influence the abundance of the butterfly pop- ulation, or result in oviposition on more yellowish green leaves. In a highly seasonal lowland area such as Guanacaste, such effects might be even more pronounced than in less seasonal Atlantic zone habitats in Costa Rica and elsewhere in southern Central America. Ross (1964b) noted that P. thoas autocles Rothschild & Jordan frequently oviposits on fresh leaves of the larval host plants, including Piper spp. (Pipera- ceae) and Citrus in Mexico. Papilio aristodemus ponceanus oviposits on young shoots of Zanthoxylum fagara (Rutaceae), “Wild Lime,” and first instars readily devour the young leaves without difficulty (Rut- kowski 1971). Tough, thick leaves of Rutaceae used by Papilio species may retard normal growth and development of caterpillars (Watanabe 1982), thereby selecting for egg placement on young, tender leaves.

Several studies reveal that the attraction of parasitoids to their phy- tophagous hosts is often mediated by the aromatic substances emitted by the host plant (Herrebout 1969, Read et al. 1970). We suspect that Papilio species associated with the highly aromatic Rutaceae are sub- ject to such parasitism, and several larvae within an aggregate of P. anchisiades can be killed by the braconid Meteorus sp. Such interac- tions may extend to predatory arthropods such as the ant C. rectan- gularis associated with Citrus in lowland Guanacaste, even though

VOLUME 40, NUMBER 1 49

predation by paper wasps under the same conditions may be minimal or nonexistent.

The size of larval groups of P. anchisiades in Citrus varies, and the group observed in the present study might have been small since the ovipositing butterfly was frightened away. The larvae have been noted to defoliate a tree (Caracciolo 1891), and very large groups of larvae have been found on individual trees (Moss 1919). Pupae are often found on various substrates away from the host tree (Moss 1919), and the present study suggests that mortality from at least one ant species might be less for pupae off the host tree than for those remaining on 1

Although the disagreeable odor from the osmeteria of the older lar- vae is well known (Stoll 1781, Carracciolo 1891, Moss 1919), the func- tional role of the secretion remains unknown, although the components are defensive against ants (Honda 1983). The precise egg-placement behavior of P. anchisiades suggests that the species is a specialist on Rutaceae, a condition that further suggests coevolved associations with parasitoids and predators that cue into the aromatic properties of Cit- rus and other genera within the family. The cryptic appearance and behavior of larvae of all instars, and the cryptic appearance of the pupa, suggest that this species is palatable to visually foraging predators such as lizards and birds (Brower & Brower 1964). When this first-line defense is penetrated by an attacker, the odor defense associated with the osmeterium might be used to thwart attack (Eisner & Meinwald 1965, Honda 1983). All rutaceous-feeding Papilio species appear to have cryptic coloration and habits (Munroe 1960).

We suggest that aggregative behavior in the larval stages of P. an- chisiades enhances visual crypsis to some predators such as birds and lizards. The combined aggregate of several fifth instars on the bark of the host tree creates the image of a mottled blotch of false lichens and bark on the trunk. Similarly, the tightly packed clusters of younger larvae on the ventral surfaces of Citrus leaves resemble dead or dying plant tissue destroyed by a pathogenic microorganism. A large aggre- gation of fifth instars positioned at the junction between the trunk and main branches of a Citrus tree in Trinidad resembled a “‘clot of wet feces” to both L. P. Brower and P. M. Shepard (L. P. Brower, pers. comm.). Aggregative behavior of the larvae, however, may not deter predation by birds. On 28 July 1986 one of us (A.M.Y.) observed an unidentified jay-size bird pluck off a Citrus leaf bearing 50 young third-instar P. anchisiades (at 0530 h) at “Finca La Lola” in Costa Rica. The bird then devoured all the larvae in a few seconds.

The osmeterial secretions from third and fourth instars of P. an- chisiades are similar to those of other Papilio species in being domi-

50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

®

y, B-Farnesene Isomer

®

"‘a-Himachalene”’ Raed

RESPONSE

Farnesene

Isomers \

©) @ a) CL

“B-Selinene”’ ra “‘a-Bergamotene”’

250° 170° = 140° 90°

a-Acoradiene

TIME / TEMP

Fic. 7. Mass spectral analysis of fourth-instar osmeterial secretion in Papilio anchisi- ades, highlighting sesquiterpene region of the spectra.

nated by terpenes. Burger et al. (1978) and Honda (1980a, b, 1981) reported that the secretions of earlier instars of several Papilio species are made up of mono- and sesquiterpenes. While we did not detect monoterpenes in the secretions of P. anchisiades, at least seven ses- quiterpenes fortify the osmeterial exudate (Fig. 7). Honda (1981) pre- viously identified sesquiterpenes in the osmeterial secretions of five species of Papilio, but several of those produced by P. anchisiades appear to be different from those produced by the Japanese species. Earlier investigations established that the osmeterial secretions of a variety of Papilio, Baronia, and Eurytides species were dominated by isobutyric and 2-methylbutyric acids (Eisner & Meinwald 1965, Cross- ley & Waterhouse 1969, Eisner et al. 1970, Burger et al. 1978, Lopez & Quesnel 1970). However, it was subsequently demonstrated that this

VOLUME 40, NUMBER 1 51

acidic duet is characteristic of the osmeterial secretions of the fifth instar. In contrast, the secretions of earlier instars of several Papilio species lack the short-chain acids produced by fifth instars, and a va- riety of terpenes are produced by younger larvae (Burger et al. 1978, Honda 1980a, b, 1981).

The osmeterial secretion of the fifth instar of P. anchisiades contains isobutyric and 2-methylbutyric acids, but, in addition, isovaleric acid, a compound detected as a minor osmeterial constituent in two other Papilio species (Honda 1981). Although isobutyric and 2-methylbutyr- ic acids have been encountered as the acidic moieties of short-chain esters in the osmeterial secretions of P. anchisiades and other species (Burger 1978, Honda 1981), long-chain esters containing butyric and valeric acids (Fig. 5) have not been reported previously from papilionid osmeterial secretions. Thus, fifth-instar larvae of P. anchisiades are distinctive in producing osmeterial secretions containing esters such as hexadecyl! valerate (Fig. 6) and octadecyl butyrate (Fig. 5). It is not clear why the dominant free acids in the secretion—isobutyric and 2-methylbutyric—have not been utilized as the acid moieties of these long-chain esters.

Sesquiterpenes in the osmeterial secretion of the fifth instar is un- usual, since this class of compounds has been identified in the secretions of earlier instar Papilio (Burger et al. 1978, Honda 1980a, b, 1981). However, one sesquiterpene has been identified in the secretion of P. protenor (Honda 1980), and three in that of P. memnon (Honda 1981). Papilio anchisiades is unusual in having almost as many sesquiterpenes (five) in the secretion of the fifth instar as in that of earlier instars (seven).

The secretions of the last instar of P. anchisiades differs from those of any Papilio species similarly analyzed in containing aliphatic hy- drocarbons and long-chain alcohols (Fig. 7). Nine aliphatic hydrocar- bons are present, and these are accompanied by C,, and C,, alcohols. With the presence of esters such as hexadecy] valerate, the distinctive- ness of this osmeterial secretion is further evident.

Previous investigators demonstrated that the secretions of younger larvae were qualitatively richer than those of fifth instars (Burger et al. 1978, Honda 1980a, b, 1981); the opposite is true for the secretions of earlier and fifth instars of P. anchisiades. Whereas the third and fourth instar secretions contain seven sesquiterpenes (Fig. 7), the fifth instar secretion contains more than 30 compounds (Figs. 5, 6) including acids, hydrocarbons, esters, alcohols, and sesquiterpenes. Qualitatively, the fifth-instar secretion of P. anchisiades exceeds that known for any instars of any Papilio species.

If it is assumed that the chemical (osmeterial) defenses of earlier- instar Papilio evolved as deterrents against predators different than

52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

encountered by the fifth instar, then the differences in the chemistry of these instars is explicable. There is little specific evidence on what organisms constitute enemies for larvae of P. anchisiades of any instar. Whatever the selection pressures were for producing the fifth-instar exudate, they have resulted in the most diverse osmeterial secretion encountered in the genus Papilio.

The greater heterogeneity and complexity of the osmeterial secretion of fifth instars in P. anchisiades suggests that the system becomes most functional in this instar. The pungent odor emitted in the fifth instar results from isobutyric and 2-methylbutyric acids, which are lacking in the earlier instars. The occurrence of some components (sesquiter- penes) of earlier-instar osmeterial secretions in the fifth instar indicates that the biochemical pathways underlying the synthesis of these sub- stances are not completely turned off in the fifth instar. Both qualitative and quantitative changes figure in the regulation of secretion in the fifth instar of P. anchisiades.

Our results are largely due to the application of capillary-column gas chromatography, which enabled detecting of minute amounts of specific components in the fifth instar, substances that might have been overlooked otherwise.

ACKNOWLEDGMENTS

Field work in Costa Rica was supported by N.S.F. Grant GB-7805 (D. H. Janzen, principal investigator) in 1969-70; N.S.F. Grant GB 33060, Friends of the Milwaukee Public Museum, and the American Cocoa Research Institute. We thank Thomas Emmel and Michael Salmon for fruitful discussions, and E. O. Wilson for identifying the ant species. Susan Borkin assisted in rearing and milking the larvae. Special thanks to A. Muyshondt for providing the specimens and record of larval parasitism from El Salvador. Paul Marsh identified the braconid wasp and shared unpublished data with us. We thank L. P. Brower for helpful suggestions on the manuscript.

LITERATURE CITED

BROWER, L. P. 1958. Larval foodplant specificity in butterflies of the Papilio glaucus group. Lepid. News 12:103-114.

BROWER, L. P. & J. V. Z. BROWER. 1964. Birds, butterflies, and plant poisons: A study in ecological chemistry. Zoologica 49:137-159.

BurRGER, B. V., M. ROTH, M. LERoux, H. S. C. Spies, V. TRUTER & H. GEERTSMA. 1978. The chemical nature of the defensive larval secretion of the citrus swallowtail, Papilio demodocus. J. Insect Physiol. 24:803-810.

CARRACCIOLO, H. 1891. Notes and news. Entom. News 2:52.

CRANE, J. 1955. Imaginal behavior of a Trinidad butterfly, Heliconius erato hydara Hewitson, with special reference to the social use of color. Zoologica 40:167-196.

CrossLEy, A. C. & D. F. WATERHOUSE. 1969. The ultrastructure of the osmeterium and the nature of its secretion in Papilio larvae (Lepidoptera). Tissue and Cell 1: 525:554.

DewiTz, H. 1878. Entwickelung einiger Venezuelanischer Schmetterlinge nach Beo- bachtungen von Gollmer. Arch. Naturgesch. 44:1-36.

EHRLICH, P. R. & A. H. EHRLICH. 1961. How to know the butterflies. Brown, Dubuque, Iowa. 262 pp.

EISNER, T. & Y. C. MEINWALD. 1965. Defensive secretion of a caterpillar (Papilio). Science 150:1733-1735.

VOLUME 40, NUMBER 1 53

EISNER, T., T. E. PLISKE, M. IKEDA, D. F. OWEN, L. VASQUEX, H. PEREZ, J. G. FRANCLEMONT & J. MEINWALD. 1970. Defensive mechanisms of arthropods— XXVII. Osmeterial secretion of papilionid caterpillars. Ann. Entom. Soc. Am. 63: 914-915.

HEGNAUER, R. 1963. The taxonomic significance of alkaloids, pp. 289-427. In Swain, T. (ed.), Chemical plant taxonomy. Academic Press, London & New York.

HELLER, S. & G. W. A. MILNE, eds. 1976. EPA/NIH mass spectral data base. Publ. NBS 20402 NSRDS-NB563, U.S. Govt. Printing Office, Washington, D.C.

HERREBOUT, W. M. 1969. Some aspects of host selection in Eurarcela rutilla Vill. (Diptera: Tachinidae). Netherlands J. Zool. 19:1-104.

Honpa, K. 1980a. Volatile constituents of larval osmeterial secretions in Papilio pro- tenor demetrius. J. Insect Physiol. 26:39—45.

1980b. Osmeterial secretions of papilionid larvae in the genera Luehdorfa,

Graphium and Atrophaneura. Insect Biochem. 10:583-588.

1981. Larval osmeterial secretions of the swallowtails. J. Chem. Ecol. 7:1089-

11138.

1983. Defensive potential of components of the larval osmeterial secretion of papilionid butterflies against ants. Physiol. Entom. 8:173-179.

ICHINOSE, T., H. ABE & Y. TERAMOTO. 1981. Relationship between host plant ranges of three papilionid butterflies and oviposition-inducing contact chemicals in their host plants. Appl. Entom. Zool. 16:493-496.

ILsE, D. 1937. New observations on responses to colours in egg-laying butterflies. Nature 140:544.

JONES, C. 1882. Metamorphoses of Brazilian Lepidoptera. Proc. Lit. Philos. Soc., Liv- erpool 36:327-377.

Lopez, A. & V. C. QUESNEL. 1970. Defensive secretions of some papilionid caterpillars. Carib. J. Sci. 10:5-7.

Moss, A. M. 1919. The Papilios of Para. Novitat. Zool. 26:259-319.

MUNROE, E. G. 1960. The generic classification of the Papilionidae. Can. Entom., Suppl. 17:1-51.

OLIVEIRA, B. L. DE. 1977. Contribuicaeo ao conhecimento da biologia de Papilio anchisiades capys (Hubner, 1809) (Lepidoptera-Papilionidae). Dusenia 10:89-95.

READ, D. P., P. P. FEENy & R. B. Root. 1970. Habitat selection by the aphid parasite Diaeretiella rapae (Hymenoptera: Braconidae) and hyperparasite Charips brassicae (Hymenoptera: Cynipidae). Can. Entom. 102:1567-1578.

Ross, G. N. 1964a. Life history studies on Mexican butterflies. I. Notes on the early stages of four papilionids from Catemaco, Veracruz. J. Res. Lepid. 3:9-18.

1964b. Life history studies on Mexican butterflies. II]. Nine Rhopalocera (Pa- pilionidae, Nymphalidae, Lycaenidae) from Ocotal, Chico, Veracruz. J. Res. Lepid. 3:207-229.

RUTKOWSKI, F. 1971. Observations on Papilio aristodemus ponceanus (Papilionidae). J. Lepid. Soc. 25:126-136.

SEITZ, A. 1908. Macrolepidoptera of the World. Vol. 5. American Rhopalocera. A. Kernan, Stuttgart.

STOLL, C. 1781. In Pieter Cramer (ed.), De uitlandsche Kapellen voorkomende in de drie Waereld-Deelen Asia, Africa, en America. Suppl. 3, Amsterdam. 384 pp. SELIGMAN, I. M. & F. A. Doy. 1972. -Hydroxy-n-butyric acid in the defensive secretion

of Papilio aegus. Comp. Biochem. Physiol. 41:341-342.

SWIHART, C. A. & S. L. SwIHART. 1970. Colour selection and learned feeding prefer- ences in the butterfly, Heliconius charitonius Linn. Anim. Behav. 18:60-64.

VaIpyA, V. G. 1969. Investigations on the role of visual stimuli in the egg-laying and resting behaviour of Papilio demoleus L. (Papilionidae, Lepidoptera). Anim. Behav. 17:350-355.

WATANABE, M. 1982. Leaf structure of Zanthoxylum ailanthoides Sieb. et Zucc. (Ru- tales: Rutaceae) affecting the mortality of swallowtail butterfly, Papilio xuthus L. (Lepidoptera: Papilionidae). Appl. Entom. Zool. 17:151-159.

YounNG, A. M. 1972. Breeding success and survivorship in some tropical butterflies. Oikos 23:318-326.

Journal of the Lepidopterists’ Society 40(1), 1986, 54

GENERAL NOTE

SATURNIA WALTERORUM (SATURNIIDAE) IN MEXICO: A NEW NATIONAL RECORD

Until now Saturnia walterorum (Hogue & Johnson) had not been taken in Mexico. It had been known to occur only in San Diego, Los Angeles, and Orange counties of California. A previous reference to specimens captured in San Luis Obispo Co. (Tilden 1945, Pan-Pac. Entomol. 21:32-33) is in error, as the cited specimens were examined by Tuskes and Collins (1981, J. Lepid. Soc. 35:1-21) and found not to be typical walterorum.

Both male and female are diurnal, and are not attracted to light. The seasonal flight period appears restricted to a few of the warmest days between late February and early June (Tuskes 1974, J. Lepid. Soc. 28:172-174). The insect is not abundant, and is easily overlooked.

Our experience with this species in coastal San Diego Co. suggests the peak daily flight period for males is mid-morning, diminishing greatly before noon.

On 1 April 1985 at 1130 h, we placed two newly emerged captive-reared females of this species in a screen cage among chaparral near Ensenada, Baja California, Mexico, about 100 km south of the United States border. The small amount of natural vegetation at this site was similar to that of areas near San Diego, California, and included Rhus laurina (Nuttall) and species of Ceanothus, Rhamnus and Adenostoma. Rhus laurina appears to us to be the preferred food plant of Saturnia walterorum in the coastal areas of its range. ;

At 1140 h a single male was attracted to the calling females and was captured. No additional males had appeared by 1300 h, at which time we left the area.

The captured specimen was placed in the San Diego Natural History Museum, San Diego, California.

KirnBY L. WOLFE AND MARVIN D. VALVERDE, Route 5, Box 169-C, Escondido, Cali- fornia 92025.

Journal of the Lepidopterists’ Society 40(1), 1986, 55-58

PYRGUS COMMUNIS AND P. ALBESCENS (HESPERIIDAE) IN NEVADA

GEORGE T. AUSTIN

Nevada State Museum and Historical Society, 700 Twin Lakes Drive, Las Vegas, Nevada 89107

ABSTRACT. Based on more than 500 male genitalia, the Pyrgus communis phe- notype replaces the P. albescens phenotype latitudinally and elevationally in Nevada. Intermediates are known where their distributions meet and overiap.

The status of Pyrgus communis (Grote) and Pyrgus albescens Plotz (Hesperiidae: Pyrginae) has been in question up to the present. They have been treated as separate species, as subspecies, or neither (Tilden 1965). Even the most recent regional and taxonomic treatments vary. They were considered subspecies of P. communis by Stanford (in Fer- ris & Brown 1981) but as full species by Miller and Brown (1981). The two taxa are often segregated by ecology and geography but there are areas of sympatry or near sympatry in southwestern United States and adjacent Mexico. In some latter areas, intermediates are known (Tilden 1965). In others, they are said to occur in close proximity, but no mention is made of intermediates (Ferris 1976, Stanford in Ferris & Brown 1981, Holland 1984); some workers have never seen an inter- mediate (Ferris, H. A. Freeman, pers. comm.). The present paper sum- marizes their status and distribution in Nevada.

More than 500 male adults from Nevada in the Nevada State Mu- seum and in the author’s collection were examined. The left valva of each was classified into one of three configurations, the variations of which are indicated in Fig. 1. These were assigned to P. albescens, P. communis, and intermediate, and their distributions were mapped.

The valvae of individuals assigned to nominate P. communis have a long and recurved dorsal process terminating in two sharply pointed prongs (Fig. 1). The lengths of the dorsal process and the prongs vary. On some individuals, one of the prongs is shorter than the other; on most they are equal. The valvae of individuals assigned to P. albescens have no dorsal process but usually have a single, short prong anterior to the tip (Fig. 1). Intermediates show various degrees of development in the dorsal process and the double prongs (Fig. 1). There was no difference in wing pattern between the genitalic phenotypes; their seasonal variation is likewise identical.

Individuals of the P. communis phenotype occur throughout Ne- vada (Fig. 2); those of the P. albescens and intermediate phenotypes occur in southern Nevada except for one P. albescens from Carson City (Fig. 2). At most stations where P. albescens were taken, inter-

56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

> oe

communis

5D De

intermediates

a/bescens

Fic. 1. Variation in the left valvae of Pyrgus communis in Nevada.

mediates and P. communis were taken also. Individuals with inter- mediate valvae occur only within the range of P. albescens. There is no strict ecological or elevational segregation in southern Nevada, but phenotype proportions do vary. The P. albescens phenotype dominates at lower elevations and latitudes. Intermediates and P. communis be- come more prominent with increase in elevation and latitude (Table 1, Fig. 2). In the Newberry Mountains, Las Vegas Valley, and the lower slopes of the Spring Mountains, P. albescens accounts for more than 60% of the individuals, and P. communis for less than 6%. At moderate elevations of the Spring Mountains, there is an increase in the P. com- munis phenotype and at the higher elevations and in Moapa Valley, intermediates predominate.

The Nevada distribution is compatible with that previously noted

TABLE 1. Proportion of P. albescens, P. communis and intermediate phenotypes from different locations in southern Nevada.

P Inter- P

Location albescens mediate communis N Newberry Mountains (<1,200 m) 60 36 4 25 Las Vegas Valley (600-900 m) 62 33 5 2) Low slopes, Spring Mts. (<1,500 m) 65 29 6 17 Mid elevations, Spring Mts. (1,500- 2,100 m) 57 24 19 84 High elevations, Spring Mts. (>2,100 m) 20 60 20 15

Moapa Valley 34 48 18 91

VOLUME 40, NUMBER 1 57

e@ communis o a/bescens

w intermediates A both

A both and intermediates WV communis and intermediates V a/bescens and intermediates

Fic. 2. Distribution of Pyrgus communis in Nevada.

(Tilden 1965) for Pyrgus communis; the latter is a more northern and higher elevation phenotype, P. albescens, a lower-elevation and more southerly phenotype. Intermediacy, at least in southern Nevada, is greater than previously reported. This indicates that the two pheno- types are closely related, and are probably no more than allopatric subspecies of Pyrgus communis.

58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ACKNOWLEDGMENTS

I thank H. A. Freeman and C. D. Ferris for their comments on these taxa. Thanks are also due two anonymous reviewers for their useful comments on the manuscript.

LITERATURE CITED

FERRIS, C. D. 1976. A checklist of the butterflies of Grant County, New Mexico and vicinity. J. Lepid. Soc. 30:38—49.

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

HOLLAND, R. 1984. Butterflies of two northwest New Mexico mountains. J. Lepid. Soc. 38:220-234.

MILLER, L. D. & F. M. BRown. 1981. A catalogue/checklist of the butterflies of America north of Mexico. Lepid. Soc. Mem., No. 2.

TILDEN, J. W. 1965. A note on Pyrgus communis and Pyrgus albescens (Hesperiidae). J. Lepid. Soc. 19:91-94.

ANNOUNCEMENT

INAUGURATION OF MANUSCRIPT DATING IN THE JOURNAL

Received and accepted dates will appear at the end of all research reports published in the Journal starting with submissions received in 1986. Such dating is practiced by many scholarly journals. It has at least three purposes. First, it encourages editors, re- viewers, and authors to speed manuscript processing. Second, it tells prospective contrib- utors how long manuscript processing might take. Third, it enables more accurate dating of ideas should issues of history or priority arise.

To better serve these purposes, new received dates may be assigned to some revised manuscripts. Examples are those received more than two years after the editor requests them, and those with excessively broadened scopes.

Received and accepted dates should make the Journal more useful to readers and authors alike.

WILLIAM E. MILLER, Editor

Journal of the Lepidopterists’ Society 40(1), 1986, 59-63

FIRST REPORTED MALES, SPECIES STATUS, AND AFFINITIES OF EPARGYREUS SPANNA EVANS (HESPERIIDAE)

KURT JOHNSON

Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024

DAVID MATUSIK

Department of Entomology, Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605

ABSTRACT. Two males of E. spanna, an Hispaniolan endemic formerly reported from only two females, were collected in Pedernales Province, Dominican Republic, in 1985. Male genitalia and other characters support species status of E. spanna but not its long-supposed affinity to E. antaeus Hewitson, endemic to Jamaica, and to E. zestos Geyer of trans-Caribbean distribution. Genitalically, E. spanna resembles E. windi Free- man of Mexico, E. aspina Evans of Colombia and E. tmolis Burmeister of western Argentina. Apparent affinities of E. spanna and some other recently described Antillean endemics do not support the island subspecies view of Caribbean biogeography. These taxa evidence most immediate kinship to particular mainland taxa, not most geograph- ically proximate Antillean congeners.

Epargyreus spanna Evans (1952) has hitherto been known only from the holotype female (labelled “Santo Domingo” [Dominican Republic (DR)]) in the British Museum (Natural History), and a second female reported by Gali and Schwartz (1983) from west of Jayaco, La Vega Province, DR in the Albert Schwartz collection (Miami, Florida).

Brown and Heineman (1972) suggested that E. spanna might rep- resent a subspecies of E. antaeus Hewitson. The latter is endemic to Jamaica, and among Antillean Epargyreus, shares with E. spanna the bright silver-white undersurface stripe on the hindwing. Riley (1975) figured both species, and Gali and Schwartz (1983), on the basis of wing character comparison, considered the two different species, pend- ing examination of E. spanna males. Both Evans and Freeman (1969, 1977) emphasized the importance of male genitalic characters in dif- ferentiating Epargyreus taxa.

In 1985, we collected extensively in the DR and, in Pedernales Prov- ince, collected two female and two male E. spanna. The precise lo- calities of these collections are: (1) a female of 830 mm forewing length (base to apex) taken between 0930 and 1200 h (EDT) on 30 June 1985 at 1,350 m altitude, Aceitillar, 12 km NW of Las Abejas, Pedernales Province, DR, on a mountain path, in mesic broad-leaf deciduous for- est, in sunny weather (in David Matusik Collection [DMC)); (2) a fe- male (not measured) taken at same location at about the same time on 1 July 1985 (in the Museo Nacional de Historia Natural, Santo Domin-

60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 1. “Designated allotype” male of Epargyreus spanna. Left: upper surface; right: undersurface.

go); (3) a male of 28 mm forewing length taken between 0930 and 1200 h (EDT) on 4 July 1985 at 1,250 m altitude, about 5 km NNW of the locality cited in (1) in mesic broad-leaf deciduous forest, in partly sunny weather (DMC); and (4) a male of 29 mm forewing length taken at the same location at about the same time on 8 July 1985. Both females were collected while flying about 13 cm above the ground in zigzag flight, which alternated with periods of alighting on patches of bare ground usually about 4 m apart. Both males were collected while perched on broad-leaf deciduous foliage about 2.4 m above the ground, between leaf-to-leaf flights. The flight of males between leaves was notably slower than that of the females between patches of bare ground. The dates cited above, along with that of Gali and Schwartz [17 Au- gust], indicate the flight period of E. spanna to be at least three weeks.

Considering the apparent rarity of E. spanna, and for reference purposes, we follow Smith (1983) and designate the last-cited male as “designated allotype”. The specimen, marked as such, is in the collec- tion of the American Museum of Natural History (AMNH), and is illustrated here (Fig. 1). The category “designated allotype’” has no status according to the International Commission on Zoological No- menclature Code, but is viewed as having diagnostic utility (Frizzell 1933, Gloyd 1982, Smith 1983).

Collection of the E. spanna male enabled examination of its genitalia (Fig. 2). Comparison of the genitalia with other Epargyreus taxa avail- able to us in AMNH genitalic material of W. H. Evans and H. A.

VOLUME 40, NUMBER 1 61

.

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Fic. 2. Male genitalia of Epargyreus spanna and E. antaeus. Above right: Dorsal view of E. spanna uncus and tegumen. Above left: Lateral view of E. spanna genitalia with aedeagus removed. A, Broad cephalad area; B, Reduced, cephalad-located dorsal process; C, Broadly thickened terminus with wide-based terminal hook. Immediately beneath: Lateral view of E. spanna aedeagus. Bottom: E. antaeus. D, Lateral view of valve; E, Lateral view of uncus; F, Lateral view of saccus; G, Lateral view of aedeagus; H, Dorsal view of tegumen and uncus.

Freeman indicates that E. spanna is specifically distinct from its con- geners. These taxa are spina Evans, aspina Evans, antaeus, orizaba Scudder, exadeus Hiibner, windi, cruza Evans, deleoni Freeman, spi- nosa Evans, clarus Cramer, brodkorbi Freeman, zestos, tenda Evans, plus additional taxa studied from figures in Evans (1952). The genitalia do not reflect the close relation claimed for E. spanna, E. antaeus and E. zestos by Brown and Heineman (1972) and Riley (1975). Although E. spanna is similar to E. antaeus on the wing undersurface, the gen- italia of E. spanna are most like E. windi Freeman (type locality [TL] Ajijic [Jalisco], Mexico) and also similar to E. aspina Evans (TL Bogota, Colombia) and E. tmolis Burmeister (TL Buenos Aires, Argentina). Along with E. spanna, the above three taxa have on the male valvae a broad cephalad area (Fig. 2, A), a reduced and cephalad located

62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

dorsal process (Fig. 2, B), and a broadly thickened terminus with wide- based terminal hook (Fig. 2, C). Epargyreus zestos and E. antaeus share more elongate valvae, with centrad located dorsal process, nar- row terminus, and thin terminal hook (Fig. 2). Epargyreus windi ex- hibits a large, but centrally limited, silver-white patch on the hindwing under-surface, which might further suggest affinity to E. spanna.

The above genitalic similarities do not necessarily override the close sister species relation posited by some for E. spanna, E. antaeus and E. zestos. No one has done a cladistic analysis of Epargyreus, but the apparent affinity of genitalic and wing characters summarized above suggests that Antillean Epargyreus species exhibit character states root- ing them as primitively in phylogenetic diagrams as their mainland congeners. The suggested genitalic affinities of E. spanna are remark- able because they suggest phylogenetic and zoogeographic relations contrary to the common practice of associating Antillean endemics as subspecies of various mainland or Greater Antillean taxa (Riley 1975, Clench 1965). If the latter were true, one would expect the wing char- acter affinities for E. spanna, E. antaeus, and E. zestos to be closely supported by characters of the genitalia.

Johnson and Matusik (1986) noted a similar situation in the genitalic characters of a new, apparently endemic Hispaniolan Tmolus (Ly- caenidae). This species does not resemble its most geographically prox- imate Antillean or mainland congener. Steven Steinhauser [Allyn Mu- seum of Entomology] (pers. comm.) concurs regarding the genitalic characters of Astraptes christyi Sharpe of Hispaniola. Astraptes chris- tyi has long been treated as a subspecies A. xagua Lucas (Riley 1975), even though it differs from that taxon in characters of the wing. Gen- italic examination of A. christyi indicates species status and affinities not simply reducible to sympatric E. xagua. Schwartz and Miller (1985), in describing a new endemic Hispaniolan Strymon, demonstrate other affinities than might be presupposed from the most geographically proximate congener. We recently collected in Hispaniola an unde- scribed species of Nesiostrymon which cannot be regarded as N. celida aibinito Comstock & Huntington of Hispaniola, or of the N. celida Lucas complex now divided into island subspecies. The practice of placing Antillean populations of butterflies as subspecies of other, more common Antillean or mainland species seems to have resulted from certain zoogeographic assumptions. Many authors believed that Antil- lean butterfly distributions represent results of recent (Pleistocene and post-Pleistocene) waif dispersal (Comstock & Huntington 1943, Clench 1965). Such assumptions have been strengthened by limiting compar- isons to wing patterns and interpreting differences in these as “‘varia- tion’”’ without reference to structural characters.

VOLUME 40, NUMBER 1 63

In zoogeography there is currently an increased appreciation of the possible correlation of late Mesozoic and Cretaceous plate tectonic splitting, and allopatric speciation of biological populations now rep- resenting various areas of endemism (Rosen 1975). This view predicts Antillean endemics may exhibit arrays of characters as primitive as any of their mainland counterparts. It is apparent that the affinities of E. spanna do not support the simplistic island subspecies view of Ca- ribbean biogeography. We expect that future cladistic analyses of Epargyreus and other butterfly groups will suggest the early origin of many endemic Antillean taxa.

ACKNOWLEDGMENTS

We thank Albert Schwartz (Miami Dade County Community College, Miami, Florida) and Lee D. and Jacqueline Y. Miller (Allyn Museum of Entomology, Florida State Mu- seum, Sarasota, Florida) for helpful information. Albert Schwartz and H. A. Freeman (Garland, Texas) reviewed the manuscript and made useful suggestions. Frederick H. Rindge (AMNH) kindly facilitated access to the AMNH collections, and also commented on the manuscript.

LITERATURE CITED

BROWN, F. M & B. HEINEMAN. 1972. Jamaica and its butterflies. E. W. Classey, Ltd., London. 478 pp.

CLENCH, H. K. 1965. A synopsis of the West Indian Lycaenidae with remarks on their zoogeography. J. Res. Lepid. 2:247-270.

CoMSTOCK, W. P & E. I. HUNTINGTON. 1943. Lycaenidae of the Antilles (Lepidoptera: Rhopalocera). Ann. New York Acad. Sci. 45:119-180.

Evans, W. H. 1952. A catalogue of the American Hesperiidae. II. Pyrginae. British Museum (Natural History), London. 246 pp.

Gaul, F. & A. SCHWARTZ. 1988. The second specimen of Epargyreus spanna (Hesper- iidae). J. Lepid. Soc. 37:170-171.

GLoyp, L. K. 1982. The original definition and purpose of the term allotype. Syst. Zool. 31:334-336.

FREEMAN, H. A. 1969. Records, new species, and a new genus of Hesperiidae from

- Mexico. J. Lepid. Soc. (suppl. 2). 62 pp.

1977. Six new species of Hesperiidae from Mexico. J. Lepid. Soc. 31:89-99.

FRIZZELL, D. L. 1988. Terminology of types. Am. Midl. Nat. 14:637-668.

JOHNSON, K. & D. Matusik. 1986. A new species of Tmolus (Lycaenidae) from His- paniola. Addendum in A. Schwartz, Butterflies of Hispaniola, Mus. Hist. Nat., Re- publica Dominicana (in press).

RILEY, N. D. 1975. Field guide to the butterflies of the West Indies. Collins, London. 244 pp.

ROSEN, D. E. 1975. A vicariance model of Caribbean biogeography. Syst. Zool. 24:431- 464.

SCHWARTZ, A. & J. Y. MILLER. 1985. A new species of Strymon (Lycaenidae) from Hispaniola. Bull. Allyn Mus. 99:1-6.

SMITH, H. M. 1988. More on allotypes. Syst. Zool. 32:454—455.

GENERAL NOTES

Journal of the Lepidopterists’ Society 40(1), 1986, 64-65

EMERGENCE OF ADULT ECTOMYELOIS MURISCUS (DYAR) (PYRALIDAE) FROM A POD OF THEOBROMA SIMIARUM DONN. SMITH (STERULIACEAE) IN COSTA RICA

The pyralid moth Ectomyelois muriscus (Dyar), a species widely distributed in Central America, northern South America, and the West Indian archipelago, undergoes its life cycle in the pods of Theobroma cacao Linnaeus (Sterculiaceae) and other fruits (Heinrich 1956, American moths of the subfamily Phytcitinae, U.S. Natl. Mus. Bull. No. 207). I have been unable to locate published records of this moth species infesting pods of other Theobroma, a genus represented by several species in tropical America (Cuatrecasas 1964, Cacao and its allies—A taxonomic revision of the genus Theobroma, Contrib. U.S. Natl. Mus. 35:379-614). Given the marked differences in external texture, pubescence, and other morphological features of pod walls among Theobroma species, one might expect some degree of ovipositional selectivity to exist for moth species associated as larvae with pods of these neotropical tree species. Here I report the emergence of 41 adults of Ectomyelois muriscus from one rotted and dried pod of Theobroma simiarum Donn. Smith in Costa Rica, representing a new host record for the moth, and for a Theobroma species with mature pods differing markedly in pod-wall texture from the previously reported host, T. cacao.

One of 12 fallen, mature, and decaying pods of T. simiarum was collected beneath one of four trees of this species in the “Theobroma and Herrania garden” on the grounds of the “Centro Agronomico Tropical de Investigaciones y Ensenanza’’ (CATIE) at Tur- rialba (9°55’'N, 83°41'W; about 600 m elev.), Cartago Province, Costa Rica in mid-August 1984. The 26 x 8 cm brown pod had no external insect emergence holes at the time it was collected. Subsequently the pod was kept on a desk in an office. Following an initial emergence of a few moths, I confined the pod in a plastic bag in the office. All adults were kept, and voucher specimens sent to the U.S. Dep. Agr. Systematic Entomology Laboratory (U.S. National Museum) for determination.

Between 20 October and 7 December 1984, 42 moths emerged from the pod, with an approximately 1:1 sex ratio. From one to four moths emerged on a given day during this period, but there were many days when no moths emerged. Most moths emerged before 0800 h. Several freshly-eclosed moths clung motionless to the pod for several hours, and flew only when disturbed. By the time the last moth emerged, only three exit holes were found on the external surface of the pod. Clearly, several moths used the same exit holes for emergence. Each exit hole had a 10-25 mm long silken tube externally, apparently built by larvae inside the pod wall and pushed out at the time of multiple eclosions. But eclosion behavior was not observed. Nor did I open the pod to determine where larvae were feeding, as the intact fruit was necessary for other research purposes.

The adults exhibited a staggered emergence pattern, because the emergence period lasted about six weeks. The female moth probably deposits clusters of eggs on the external surface of the sand-papery-rough pubescent pod, since adults appeared to emerge in clusters from a few exit holes. Perhaps this particular pod received three different egg batches. Assuming the observed laboratory emergence pattern was similar to that occur- ring in nature, a brood of E. muriscus emerges near the end of the Turrialba rainy season, and before the short, erratic dry season. The availability of decaying pods of T. simiarum varies greatly throughout the year, suggesting a changing pod supply for pod herbivores or pod saprotrophs (whichever the case may be).

Given the previously reported association of E. muriscus with T. cacao in both Central and South America, my discovery of this moth species in a pod of T. simiarum may not be surprising. Theobroma simiarum is one of two species of the genus endemic to Costa Rica. Given the broad geographical range of Ectomyelois muriscus in tropical America, it undoubtedly has other natural hosts, possibly species of Theobroma other

VOLUME 40, NUMBER 1 65

than cacao or simiarum. The dense, thick tomentum (pod wall external surface) may represent a suitable oviposition substrate for Ectomyelois muriscus, but other surface textures must also be suitable given the marked difference in this feature between Theo- broma cacao and T. simiarum. Larvae of Ectomyelois muriscus most likely tunnel through the woody epicarp and softer mesocarp tissues of the pod. Yet they may infest pods once the latter are into advanced stages of decay, perhaps rendering pod-wall tissues more penetrable to larvae.

Near the end of the rainy season at this locality, mature pods of various species of Theobroma are available, in addition to those of T. cacao, the most abundant species due to large commercial plantations. When the dry season arrives near the end of De- cember, dryness may trigger a large moth emergence, a pattern somewhat different than that observed in the office. The very dry conditions of the office may have mimicked the dry season for moth larvae and pupae present inside the T. simiarum pod, leading to a staggered emergence as conditions became increasingly dry.

This research was funded by the American Cocoa Research Institute of The United States of America. I thank D. C. Ferguson for determining the moth and providing the Heinrich reference. The technical assistance of Susan Sullivan Borkin is appreciated.

ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- waukee, Wisconsin 53233.

Journal of the Lepidopterists’ Society 40(1), 1986, 65-66

THE FEMALE OF PAPILIO XANTHOPLEURA GODMAN & SALVIN (PAPILIONIDAE)

Before 1985, literature concerning Papilio xanthopleura Godman & Salvin stated that its female occurs in two forms: a “normal” female resembling the male, and a large yellow one, form diaphora Staudinger (Staudinger 1891, Deut. Entomol. Z. [Iris] Lepid. 4:61-158; Rothschild & Jordan 1906, Novit. Zool. 13:412-752; Jordan 1907, in Seitz, Macrolepidoptera of the World, Vol. 5, Alfred Kernen Verlag, Stuttgart, 592 pp.; Munroe 1961, Can. Entomol. Suppl. 17, 51 pp.; D’Almeida 1965, Catalogo dos Papilionidae Americanos, Soc. Braz. Entomol., Sao Paulo, 366 pp.; D’Abrera, Butterflies of the Neo- tropical Region, Part 1, Papilionidae and Pieridae, Lansdowne Editions, East Melbourne, 172 pp.). None of the literature illustrates a xanthopleura female.

Johnson, Rozycki and Matusik (1985, J. N.Y. Entomol. Soc. 93:99-109), examined the type and other specimens of diaphora, and showed that the type and all known repre- sentatives of diaphora are males, and male genital and wing characters in diaphora indicate it is not conspecific with xanthopleura. As a result, diaphora was accorded species status, it became apparent that females of diaphora are presently unknown in collections, and no “normal” females of xanthopleura were in the following major col- lections: Allyn Museum of Entomology, American Museum of Natural History (AMNH), British Museum (Natural History), Carnegie Museum of Natural History, Collection of David Matusik (Skokie, Illinois), Collection Dep. de Zoologia, Universidade Federal do Parana (Brazil), Collection of Ernesto W. Schmidt-Mumm (Bogota, Colombia), Collection of Rick Rozycki (Chicago, Illinois), Collection Tommasso Racheli (Rome, Italy), Instituto de Zoologia Argricola Maracay (Venezuela), Museu Nacional, Rio de Janeiro (Brazil), Museo de Historia Natural “Javier Prado” (Lima, Peru), National Museum of Natural History (Smithsonian Institution), and the collection of a commercial dealer noted for his holdings in unusual Papilionidae.

Therefore, we borrowed a female of xanthopleura (Fig. 1A, C) from the Staudinger Collection (Zoologisches Museum der Humboldt Universitat, Berlin, German Democratic Republic [ZMH]). The female resembles male xanthopleura on the wing undersurface but, contrary to the above literature, differs markedly from the male on the upper surface of the wings. Males of xanthopleura are black above except for brilliant “powder green’

66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 1. Papilio xanthopleura and P. diaphora, with forewing length (base to apex) in parentheses, B—D, upper surfaces of wings, to same scale, A, undersurface, to different scale. A, P. xanthopleura female (67.0 mm), Iquitos, Peru, ZMH; B, P. diaphora type male (71.0 mm), Manicoré, Brazil, ZMH; C, P. xanthopleura female (of 1A); D, P. xanthopleura male (57.0 mm), Campana [sic], Brazil, AMNH.

in the vein interspaces of the hindwing (Fig. 1D); females are powder green over the entire upper surface of both wings (Fig. 1C). Female xanthopleura are larger than male xanthopleura, but neither exceeds the large size of male diaphora. As noted by Johnson, Rozycki and Matusik, the mean single forewing length (base to apex) of known diaphora males exceeds that of examined xanthopleura males by 12.8 mm and the examined xanthopleura female by 3.0 mm. Thus, wing character differences in the genders of these taxa vary far more than the literature has indicated.

We thank Prof. H. J. Hannemann for loan of the diaphora type and various xantho- pleura specimens. Phillip Ackery, K. S. Brown, O. H. H. Mielke, L. D. Miller, John Rawlins, R. K. Robbins, Tommasso Racheli, and E. W. Schmidt-Mumm aided in survey- ing various collections.

KURT JOHNSON, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024; Rick ROZYCKI, 5830 South McVicker Avenue, Chicago, Illinois 60638; AND DAvID MATUSIK, Department of Entomology, Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605.

VOLUME 40, NUMBER 1 67

Journal of the Lepidopterists’ Society 40(1), 1986, 67

PLACEMENT AND FATE OF MONARCH BUTTERFLY PUPAE IN NORTHERN CALIFORNIA

The placement on a substrate and the subsequent fate of Danaus plexippus (L.) (Dana- idae) pupae were determined on a roadside strip at two sites SE of Davis, Yolo Co., California, in the Sacramento Valley from mid-July to October 1966.

The first site was 3 km south of Davis. It was a strip 4 m wide and about 400 m long containing several clumps of Asclepias fascicularis, other green plants, and dry grass between a paved road and a woven wire fence. The second site was similar and 6 km SSE of Davis. Both were in flat, mostly agricultural land, and near alfalfa, sugar beets, and grains. The only trees were around a dwelling and an old church and cemetery.

Each site was thoroughly searched on weekends and occasionally during the week from 14 July to 30 October 1966. Each pupa was marked with a paper tag located nearby with a serial number and date of discovery.

Vertical distributions of pupae were from ground level to 1 m. One pupa was found 4 m high on an old creosoted pole.

Choices of substrate for 409 pupae were: A. fascicularis, 17.7%; other green plants, 6.2%; dry grass, 47.7%; wire fence, 25.5%; and wooden fence posts, 2.9%. Only 4% of the pupae on the A. fascicularis were placed after 31 August. On the wire fence, 6% were placed in July, 25% were placed in August, 53% in September, and 16% in October. On dry grass, 24% were placed in August, 53% in September, and 23% in October. Pupae are rarely placed on Asclepias spp. in southeastern Canada (Urquhart 1960, The Monarch Butterfly, Univ. Toronto Press, 361 pp.).

Following initial observations, several categories of pupal fate were defined but only three are reported (Table 1). The latter are: discolored and dead, empty shell left after eclosion, and disappeared leaving no evidence of fate (because pupae were marked with white tags, passersby may have taken some, but predation seems more likely).

TABLE 1. Danaus plexippus pupal fate by substrate, Davis, California, 1966.

Number of pupae

Asclepias Other green Fate Dry grass fascicularis plants Wire fence Total Died 83 25 9 21 138 Eclosed 74 32 14 35 155 Disappeared 54 18 4 23 99 Total 211 75 27 79 392

Without regard to substrate’ 39.4% eclosed, 35.1% died, and 25.5% disappeared (Table 1). Of the 99 that disappeared, 55% were on dry grass, 28% on the wire fence, and 24% on green plants. The data were analyzed in a standard contingency table; x? = 7.18, df = 6, .500 > P > .250. The sample may be too small, but tentatively, pupae on green plants appear to be safest from predators. A higher percentage of pupae eclosed on green plants than on dry grass or the wire fence.

I thank A. M. Shapiro of the University of California, Davis, for reviewing the manu- script and doing the statistical test.

LESLIE V. SMITH, 7589 Twin Oaks Ave., Citrus Heights, California 95610.

68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Journal of the Lepidopterists’ Society 40(1), 1986, 68

ABERRANT POSTMEDIAL SPOTS IN ICARICIA ICARIOIDES LYCEA AND I. ACMON (LYCAENIDAE)

Several nearctic lycaenids tend to abnormal expression of ventral postmedial spots. This generically takes the form of some or all spots being elongated. Illustrations recently published include a multiple occurrence in Satyrium calanus falacer (Godart) (Wright 1981, J. Lepid. Soc. 35:158-159). Single occurrences have been shown for Satyriwm acadica acadica (W. H. Edwards) (Leeuw 1979, J. Lepid. Soc. 33:204—205), Glaucopsyche lygdamus couperi Grote (Neil, 1983, J. Lepid. Soc. 37:258), Euphilotes rita rita (Barnes & McDunnough), Icaricia acmon texana Goodpasture (Holland 1980, J. Res. Lepid. 19: 88-95) and Icaricia acmon lutzi (dos Passos) (Cannon, J. Lepid. Soc. 39:329-330). The similarity of the two aberrant acmon specimens is striking, especially since the acmon lutzi is from Idaho and the acmon texana is from New Mexico.

To this list of aberrant lycaenids, I now add Icaricia icarioides lycea (W. H. Edwards). Figure 1 shows an aberrant, an intermediate and a typical specimen. About 5% of 100 specimens were found to have noticeably enlarged postmedial spots. The two atypical examples illustrated here represent the most extreme.

I call attention to a well illustrated and useful article on the mechanics of color pattern formation and malformation in butterflies and moths (Nijhaut 1981, Sci. Am. 245(5): 140-151). This work seems not to have received the recognition it should among lepi- dopterists.

RICHARD HOLLAND, 1625 Roma NE, Albuquerque, New Mexico 87106.

Fic. 1. Icaricia icarioides lycea ventral surface. Left, aberrant male, Corral Tank, Canyon de Los Corrales, NE slope, Jemez Mts., Rio Arriba Co., NM, 27.V.84, 2,438 m; Middle, intermediate male, 1.6 km N of Poncha Pass, Chaffee Co., Colo., 5. VII.63, 2,743 m; Right, normal male, 20.9 km W of Espanola on road to Santa Clara Peak, E slope, Jemez Mts., Rio Arriba Co., NM, 18.V1.83, 2,499 m; all leg. R. Holland.

VOLUME 40, NUMBER 1 69

Journal of the Lepidopterists’ Society 40(1), 1986, 69-71

NOTES ON A COSTA RICAN “MONKEY SLUG” (LIMACODIDAE)

Little is known about the life cycles, larval food plants and other aspects of natural history of neotropical limacodids (Dyar 1924, Limacodidae, in Macrolepidoptera of the World, Vol. 6. American Heterocera [A. Seitz, ed.], A. Kernan Verlag, Stuttgart). Herein I describe the final instar caterpillar, some aspects of caterpillar behavior, and one larval food plant for Phobetron hipparchia Cramer in northeastern Costa Rica. Dyar mentions that P. hipparchia caterpillars feed on “different forest trees” and that this species occurs in Mexico, Panama, Ecuador, Colombia, Venezuela, Guiana, Brazil, and Argentina.

On 27 February 1985, two late-instar caterpillars of P. hipparchia were collected from one 6 m tall Gliricidia sepium (Jacq.) Steud. (Papilionoideae: Galegeae, Robiniinae) supporting vanilla vines at “Finca La Tirimbina,” near La Virgen, Sarapiqui District, Heredia Province (10°23’N, 84°07’W;; 220 m elev.). The tree was one of several thousand G. sepium planted there for vanilla production (Allen & Allen 1981, The Leguminosae: A source book of characteristics, uses and nodulation, Univ. of Wisconsin Press, Madison, Wisconsin, 812 pp.). The caterpillars were placed in a clear-plastic bag, along with cuttings of G. sepium, for rearing to adulthood.

A thorough search of the G. sepium having the caterpillars revealed no other individ- uals of P. hipparchia, as was also the case for an additional eight trees of this species examined in the same area. Both caterpillars were discovered on the dorsal (upper) exposed surfaces of old, tough leaves (Fig. 1), and about 1 m apart at eye level (about 1.8 m above the ground). At the time of discovery, the majority of G. sepium trees at the site were without flushes of new (fresh) leaves. From a distance of about 1 m, the caterpillars resembled curled, dry leaves (Fig. 1).

Within three days after collection, both caterpillars molted to the final instar, exhibiting little change in overall appearance from the previous instar. The final instar lasted only a few days; both caterpillars formed loose silken cocoons in the leaves by 7 March. Each cocoon (Fig. 1) consisted of a thin sheet of silken mat across several leaves, and dorsally mostly the larval tubercles shed during spinning. The pupal stage lasted about one month (under laboratory conditions of 65—70°C and 30-40% RH). Both adults emerged between 1500-1600 h. One was female, the other a male (Fig. 1).

At the time of discovery, the caterpillars were both about 24 mm long and 20 mm wide, including the greatest expanse of the laterally positioned “horns” (Fig. 1). When the caterpillar was motionless on a leaf, the horns were held laterally against the leaf surface (Fig. 1). When moving, the caterpillar appeared to be rocking back and forth, with a slight rotation of the horns. The squarish, angular body profile of a motionless caterpillar on a leaf (Fig. 1) became spherelike when the caterpillar was disturbed: the caterpillar curled itself into a ball, and partially tucked in and interlocked some of the lateral horns. Several sustained prods with a forceps were needed to ellicit this curling behavior.

The overall color of the caterpillar consisted of a patchwork of brown shades. Cramer (1791, Uitlandsche Rupsen, Supplement) reported the larva (Plate XVIII) of P. hippar- chia to be light-brown in color. As in all Limacodidae, the head capsule (about 4 mm diam) was small and hidden at all times. Both the head capsule and thoracic legs were glossy orange. The first two thoracic segments were almost translucent, and without prominent tufts of setae or lateral extensions of the cuticle. The third thoracic segment had a ringlet of six bulbous, orange tufts of hairs. The lateral horns were present on the first three abdominal segments (one pair per segment), and dorsally were darker brown than below. The horns of the third segment were dorsally more markedly brown than those of the previous two segments. Those of the fifth abdominal segment were dark brown dorsally, while those of segments 7-9 were light brown (tan) dorsally. No lateral horns were present on abdominal segments 4 and 6. The lateral horns appeared to be extensions of the cuticle, and were covered with short setae (see Dyar 1896, J. New York Entomol. Soc. 4:167-190). Dorsally, each abdominal segment had a dark brown rectan-

70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fic. 1. Phobetron hipparchia. Left top and bottom: Final-instar caterpillars on leaves of Gliricidia sepium. Right top: Cocoon. Right bottom: Reared male (above) and female (below).

gular patch, with a light spot in the center and even darker borders (Fig. 1). A dorsal- medial raised line of tan-colored setae ran lengthwise on both thoracic and abdominal regions. The body of the caterpillar was 8 m wide at the thickest point within a day of cocoon formation. Following cocoon formation there was no noticeable change in the colors of the larval cuticle, even though it became part of the cocoon.

VOLUME 40, NUMBER 1 ral

Phobetron hipparchia, the single species of the genus represented in the neotropical region, apparently gets the name “monkey slug” from the curiously shaped caterpillar stage. One of the two North American species of the genus, P. pithecium (J. E. Sm.) is the “hag moth,” and its caterpillars feed on a broad range of trees, none of which apparently is within the Leguminosae (Papilionoideae) (Covell 1984, A field guide to the moths of eastern North America, Houghton Mifflin Co., Boston, 496 pp.). Dyar (1896, op. cit.) noted that P. pithecium larvae perch on the undersides of leaves until the last instar, and that Phobetron larvae in general cryptically resemble dead leaves. Even though perched on the upper sides of leaves, the final-instar larva of P. hipparchia appears cryptic, resembling the yellow, brown, and green blotch pattern of older G. sepium leaves. The observed pattern of cocoon construction, in which cast-off tubercles are added to the silk during spinning, is considered typical by Dyar for New World Phobetron, enhancing crypsis of the pupal stage.

Of the several genera and species of North American Limacodidae discussed in Covell, none apparently utilize legumes as larval food plants. Yet an outstanding feature of these moths in general appears to be their highly polyphagous food habits as caterpillars (Dyar 1924, op. cit.; Covell, op. cit.). Several genera and species of Limacodidae feed on legumes in Australia.

McFarland (1979, J. Lepid. Soc. 33: Supplement, 72 pp.) reports that Australian li- macoids are associated with Acacia and other legume genera, and that caterpillars of some species invariably occur on old, tough leaves and stems of food plants as in the present observations. Within the neotropical region, the limacodid Sibine apicalis (Dyar), or “gusano montura,’ sometimes defoliates banana (Musa spp.) trees (Jaramillo & Ji- menez 1974, Turrialba 24:106-107). Thus both dicotyledenous and monocotyledenous larval food plants for the Limacodidae are known from the neotropical region. Cramer reported P. hipparchia on Granadilla (Passifloraceae). Eucalyptus spp. (Myrtaceae) are major larval food plants for the Limacodidae in Australia and Africa (McFarland; Se- vastopulo 1983, J. Lepid. Soc. 37:91, respectively). But in East Africa some limacodids feed on indigenous legumes (Acacia spp.), but not introduced species of the family. Gliricidia is endemic to the neotropical region, and given the great evolutionary diver- sification of the papilionoid legumes in tropical America (Richards 1964, The Tropical Rain Forest, The Univ. Press, Cambridge, England, 437 pp.), one might expect to dis- cover several other legume larval food plants for P. hipparchia. Gliricidia is widely distributed throughout the American tropics, both in natural habitats, as a result of its extensive use as a shade tree for cacao and coffee, and as a support for vanilla vines.

In spite of several years of casual observation during both dry and rainy seasons, I did not notice other P. hipparchia caterpillars on the trees.

This research was a by-product of a grant from The American Cocoa Research Institute of The United States of America. I thank J. Robert Hunter for allowing me access to Finca La Tirimbina. Adult moths reared in this study have been deposited in the collec- tions of the Milwaukee Public Museum. Detailed comments of the reviewers were most helpful, and one reviewer provided me with the Cramer (1791) and Dyar (1896) refer- ences, for which I am grateful.

ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- waukee, Wisconsin 53238.

Journal of the Lepidopterists’ Society 40(1), 1986, 72-73

BOOK REVIEWS

MILKWEED BUTTERFLIES, by P. R. Ackery and R. I. Vane-Wright. 1984. British Museum (Natural History), Cromwell Road, London SW7 5BD, England. v-ix + 425 pp., includ- ing 261 figures, 12 colored and 73 halftone plates, quarto size. Price £ 50. Also published in U.S. with Cornell Univ. Press, Ithaca, New York.

The full title of this book, “Milkweed butterflies: their cladistics and biology, being an account of the natural history of the Danainae, a subfamily of the Lepidoptera, Nym- phalidae” states the goals of Ackery and Vane-Wright’s study. They succeed admirably in this difficult task. While the major thrust of the book is a taxonomic revision of the danaines, there is a wealth of well documented biological information about distribution, behavior, life history, chemical attraction and defense, genetics, ecology, mimicry and faunistics. This makes the book not only valuable to the systematist, but also to the worker in other biological disciplines.

The fact that the revisionary section is a cladistic study probably will bother phenet- icists and at least some evolutionary taxonomists, but this does not concern the authors. It is reassuring to see a study that is done strictly on characters, and draws no systematic conclusions not supported by those data; the authors freely admit that certain relations are not shown by the characteristics they used, and suggest that these problems may be solved later by the addition of new characters. What is speculation by the authors is clearly so labeled, and definitive statements are not made without reference to underlying reasons. The revision is, therefore, highly scientific; and it is a pleasure to have the authors lead one through the reasoning to their conclusions without the appearance of the occult that one is so often left with in systematic revisions where conclusions are “correct” simply because authors state they are.

The cladistic section of the book involves discussions of the characters treated and the classification scheme derived from them. Each character is discussed, numbered in the text and, perhaps most importantly, is illustrated clearly in the figures, along with alter- nate character states. Again, one can follow the reasoning through a logical progression to the classification adopted. There are some valuable, but unconventional, thoughts on classification intertwined with the data, such as those on clades and polytypic species (pp. 20-22), including the concepts of “‘cladospecies” and “paraspecies”’, and those im- mediately following on larval versus adult characters. Their suggestions for further sys- tematic research on the danaines (pp. 61-66), most of which the authors promise to tackle later are valuable and thought-provoking. The book would have been even more useful had it considered the myriad subspecies, but an attempt to analyze cladistically some 2,000 names would have been humanly impossible, and the result would not have been economically feasible to print. Nevertheless, the handling of the subject is reasonable enough to make a confirmed cladist of the reader: if only people of that taxonomic school were always so rigorously tied to logic!

The authors anticipated that the revision might be controversial because it upsets prevailing nomenclature. Ackery and Vane-Wright comment about the genus “Danaus” of authors. “Danaus” is at best a grade taxon and is probably polyphyletic, perhaps paraphyletic. A number of species are placed in genera far removed from their “con- ventional” placement, but the authors recognize the difficulty that others might have accepting these new assignments, stating (p. 8), “. . . because of the still overriding influence of ‘Seitz’ and the acceptance of ‘Danaus’, we can be sure that a dual nomen- clature, with ‘Danaus’ sita for Parantica sita, ‘Danaus’ similis for Ideopsis similis, ‘Danaus’ hamata for Tirumala hamata and so on, will unfortunately continue in exis- tence for a considerable length of time—probably until about A.D. 2179 judging by past performances!” Such a statement might be made about any revision that accepts a different nomenclature than that in a “standard” work, but Ackery and Vane-Wright present a large and impressive body of data, and it is up to their detractors (if any) to examine such data in detail and show where they think the authors are wrong. I hope

Ackery and Vane-Wright are incorrect on the period of time until their nomenclature is accepted.

VOLUME 40, NUMBER 1 73

Ackery and Vane-Wright have gathered an impressive array of biological data on the danaines which are summarized in Part 2: Biology (pp. 67-102). They have also gener- ated a large body of data on co-mimicry and faunistics (pp. 1038-158). Both of these sections should be of great interest to ecologists, evolutionists and other nontaxonomists, as well as to systematists in the broadest sense. Part 4 consists of identification keys, often utilizing novel characteristics not stressed in the typical key; and all data are summarized in Part 5, the specific taxonomic and biological catalogue (pp. 173-245), an impressive compendium of information that should convince even the most skeptical. There are some new synonyms, combinations and taxa, but finding them requires some searching because they are buried throughout the text. Short of a section summarizing changes, there is no other way the data could have been presented conveniently. A short adden- dum follows, and precedes an exhaustive bibliography of 86 pages.

The work is remarkably free of typographical errors—the one I noticed was the ren- dering of Japan as “Japen” on p. 21. It is an attractive book and easy to follow. The illustrations are uniformly of high quality, including line drawings, colored plates, half- tone plates, and 394 additional figures illustrating all aspects of danaines and their biol- ogy. Colored plates depict some danaine mimetic associations in the Philippine and Indonesian Islands. It is too bad that illustrations are not cross-referenced to the pages on which descriptions occur (the pages with descriptions do have references to plates), but this is a minor complaint; also each species is mentioned several times in the text, and to which page would the plate reference refer? The comprehensive Index clearly leads the reader to any place a taxon is discussed.

I must consider this work to be one of the major taxonomic revisions of this century, and it is a good book for the reader who is not taxonomically inclined. The authors have attacked a problem, solved much of it, and have honestly admitted those parts that have resisted solution so far. To say that I am impressed with this work is an understatement; it is the kind of work that one always hopes to be able to do. The book, even though expensive, is well worth the cost; I would recommend it to anyone interested in well explained and defended cladistic analyses, in systematics of Lepidoptera, or in the Danai- nae.

LEE D. MILLER, Allyn Museum of Entomology of the Florida State Museum, 3621 Bay Shore Road, Sarasota, Florida 33580.

Journal of the Lepidopterists’ Society 40(1), 1986, 74

A MONOGRAPH OF THE BIRDWING BUTTERFLIES. Volume I, parts 1-8, the subgenera Aetheoptera, Ornithoptera, Schoenbergia; Volume II, parts 1-2, the genera Trogonop- tera, Ripponia, Troides (partim), by J. Haugum and A. M. Low. Vol. I. 308 pp., 12 plates. Vol. II. 240 pp., 12 plates. Scandinavian Science Press, Ltd., Klampenborg, Den- mark. 1978-1984.

No other group of butterflies has attracted as much attention or interest as the birdwing butterflies. In this two-volume set, the authors provide the most detailed analysis of these showy butterflies to date. The first part of Vol. I, for example, on Aetheoptera, deals with only two species, yet numbers 84 pages; and Vol. II, part 1, which describes three species, numbers 104 pages. The text of this work will complement the beautiful illus- trations in the recently published Birdwing Butterflies of the World, by B. D’Abrera.

Volume I covers the genus Ornithoptera, Volume I, Trogonoptera, Ripponia, and Troides. The genera and species are introduced with descriptions and notes on biology, followed by keys. However, no keys are given for most subspecies, which means that most (as with so many subspecies of butterflies) must be determined by geographic locality. Each species and subspecies is described in detail, and notes are given on the phylogenies of each taxon. Several illustrations, including distribution, accompany the description of each taxon. The authors tend to recognize almost every described taxon, and include descriptions of several “forms” or aberrations. Even though the authors recognize that formal names given to such infraspecific forms are not nomenclatorally valid, I find it annoying to see several new names applied to these forms.

A group as well known and as popular as the birdwing butterflies is bound to generate controversy in the literature, and such is the case here. I found the authors to be partic- ularly critical of D’Abrera, often disagreeing with what is said in his volume. For ex- ample, D’Abrera recognizes only two subspecies of Ornithoptera goliath, Haugum and Low, five. D’Abrera considers O. richmondia and O. urvillianus separate species from O. priamus, not so Haugum and Low. Ripponia is used for hypolitus, but D’Abrera considers it a Troides.

For all the detailed analysis given for each subspecies, I find it disappointing that no quantitative data were used to back up the authors’ assertions that these taxa are, indeed, taxonomically distinct. Most of the diagnostic comments are qualitative, as two examples will show: Males of O. tithonus “waigeuensis may be recognized by having a narrow HW [hind wing] with a notable reduction of the apical area; the wing is even further modified and less angular at the apex than in subspecies misresiana and tithonus ...,” and “The male HW [of T. amphrysus ruficollis f. loc. euthydemus] tends to be more rounded than in ruficollis on average, also the FW [fore wing] may be broader ... ,” which means to the reader that geographic locality will still be the only way to identify subspecies.

Puzzling also is the uneven treatment of “Material examined.” For some species, a detailed listing is given, for others, the data are incomplete, as for O. goliath atlas: “photographs of a further series of E. Weyland Mts. imagines” [how many?]. The loca- tions of types are, unfortunately, not addressed for all taxa. No cladistic analysis was carried out on this group, although I hope the authors may consider this in their final volume.

The books are handsomely bound, the printing, layout, and illustrations are good, but, alas, errors in typography and syntax abound. The color photos of Vol. I are excellent, but those of Vol. II have a white halo within the marginal black areas of the wings figured.

Despite their shortcomings, these volumes represent the best compilation of data for these magnificent butterflies. As one who is fascinated by these gorgeous insects, I heartily recommend this work for every lepidopterist interested in Old World Papilionidae.

ROSSER W. GARRISON, 1030 Fondale St., Azusa, California 91702.

Journal of the Lepidopterists’ Society 40(1), 1986, 75-76

OBITUARIES

ARTHUR C. ALLYN (1913-1985)

Dr. Arthur Cecil Allyn, life member of the Lepidopterists’ Society and Director Emer- itus of the Allyn Museum of Entomology, died on 22 March 1985, after a lingering illness. He will be remembered by the lepidopterological community for his generosity, dedication, and service to the science.

Dr. Allyn was born in Evanston, Illinois on 24 December 1918, completed primary and secondary schooling there, and attended Dartmouth and Beloit Colleges. He received a D.Sc. from the University of Florida in 1981 in recognition of his accomplishments in and service to entomology. He is survived by his wife, Dorothy D. Allyn, and three children, David D. Allyn, William N. Allyn and Dorothy A. Lavick, as well as eight grandchildren.

Dr. Allyn had a successful and diversified business career with international interests in oil, maufacturing, farming and sports in many countries, including Australia, Indo- nesia, Canada, South Africa and countries in Latin America. He was a philanthropist of note, being responsible for a wing at Chicago's Mercy Hospital and the Convention Center and Robarts Sports Arena in Sarasota. He was interested in the arts, especially theater, and he and his wife coproduced a number of plays at the Asolo State Theater in Sarasota. But, his abiding interest was in lepidopterology, and he amassed a huge collection of these insects. Finally he decided that the chore was too much for one man part-time; thus began our association with him in 1968.

Arthur Allyn’s interest in Lepidoptera and his desire to establish a quality institution for their study led to the formation of the present-day Allyn Museum of Entomology. During the early days of the Museum, Dr. Allyn continually purchased material that was needed to enhance the Museum collections, often obtaining entire collections or entire season's catches from people throughout the world. Later, collections or individual

76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

specimens were donated to the museum, and less material was purchased, but the growth of the collection continued to its present size of about 850,000 specimens from throughout the world. Dr. Allyn insisted that the collection be worldwide and accepted the necessity of long series of specimens to show individual variation within taxa.

In 1973 the Museum moved to its present home, and it became apparent that it would outgrow the new collection range rapidly if something were not done. Dr. Allyn had been impressed by the compactor housing the collections at the Missouri Botanical Gar- dens, and we immediately decided that the museum needed such housing. Thus was installed the first entomological compactor in the United States, a system that served as the model for similar systems elsewhere in this hemisphere. It was an excellent choice.

Concern about the backlog of papers in many lepidopterological journals led to the founding of the Bulletin of the Allyn Museum late in 1971. This publication, patterned after American Museum Novitates, has gone through 105 numbers as of this writing, with indices after every 20 numbers. The Bulletin rapidly evolved into a refereed pub- lication, but final responsibility for the content of numbers rests with individual authors.

Dr. Allyn became an accomplished scanning electron microscopist largely through his own efforts. One of his photographs was featured on the cover of an issue of Annals of the Entomological Society of America. After studying a number of entomological papers with which he disagreed, based on his knowledge of the physical sciences, he began to publish papers on structures of Lepidoptera not only with Museum staff, but also with such authors as Dr. John C. Downey, Dr. Miriam Rothschild and Professor David Spencer Smith. Other papers were in varying degrees of completion at his death, some of which may appear in future Bulletin numbers. Those papers that were published are standards for the field, and it is a tribute to Arthur Allyn and his coauthors that they are frequently cited not only in lepidopterological, but also in sources oriented toward scanning electron microscopy. Dr. Allyn also cooperated with various researchers in other fields, and his photographs have appeared in publications on optic, muscular and nervous systems of both vertebrates and invertebrates. A complete listing of Dr. Allyn’s publications is given in the Bulletin of the Allyn Museum, number 97.

Eventually, Dr. Allyn began to search for an orderly transition from the independent nature of the Allyn Museum of Entomology to a more structured, but secure status to assure its permanence. After examining many options, he presented the Board of Direc- tors of the Allyn Museum with a proposal from the University of Florida Foundation which was accepted, and the Museum became a part of the Florida State Museum in 1981. Eventually the present facility will be moved to Gainesville, and the Museum’s metamorphosis will be complete.

Arthur Allyn was a benefactor of the Lepidopterists’ Society in numerous ways. In the late 1960's, the Society faced severe financial difficulties from which Dr. Allyn rescued it in return for financial accountability from the officers. Through a series of excellent treasurers, the Society has managed to remain a viable entity ever since. Equally impor- tant was the establishment of the Karl Jordan Medal for papers of lepidopterological excellence. There have been ten Jordan Medal awards since 1973, and the award winners have been truly international. Not only is the United States represented by Jordan Medal laureates, but also France, Canada and England.

Those of us who were close to Arthur Allyn will miss him for his generosity, excellent judgment and common sense. He would not want this, however, to become an overriding emotion: he would demand that we continue as before. Lepidopterology, along with many other pursuits, is better for its association with him.

LEE D. MILLER AND JACQUELINE Y. MILLER, Allyn Museum of Entomology of the Florida State Museum, 3621 Bay Shore Road, Sarasota, Florida 33580.

Journal of the Lepidopterists’ Society 40(1), 1986, 77-78

LIONEL GEORGE HIGGINS (1891-1985)

With the recent death of Lionel Higgins, at the age of 94, one of our few remaining links with pre-War entomology has been severed. Perhaps reflecting those more leisurely times, in common with many of his generation he was a ‘generalist’ of distinction, equally at home in art and music as in his chosen profession of medicine, or indeed entomology. Following rheumatic fever in childhood, Lionel was pronounced too delicate for a formal school education. As a result, the interests that so enlivened his lifetime were probably kindled. Taking a medical degree at Clare College, Cambridge, he qualified at St. Thomas’ Hospital before serving in the 1914-18 War as a surgeon-lieutenant. Spe- cializing in gynaecology and obstetrics, he practiced from 1922 onwards in Woking, Surrey, where he continued to live after his retirement.

In the field of Lepidoptera, he is, of course, best known for his collaborative book with the late N. D. Riley, “A Field Guide to the Butterflies of Britain and Europe” (1970), a standard work translated into at least nine languages, with world-wide sales approaching 200,000; and its companion volume “The Classification of European Butterflies’ (1975). These works were the culmination of fifty years of serious study. While this is not the place to include a full bibliography, C. R. Smith (Type specimens of the taxa described by L. G. Higgins in the British Museum (Natural History), in preparation) lists more than 70 titles, which truly reflects his contribution since 1924, when the first work ap- peared. Probably he would have regarded the Melitaeinae as “his” group. Major contri- butions in 1941, 1950, 1955, 1960, 1978 and 1981 provided a firm foundation for future work. Tangible recognition of his contributions to natural history include the Stamford Raffles Award (Zoological Society of London) and the H. H. Bloomer Award (Linnean Society).

As well as being a prolific author, he was an indefatigable collector, both of butterflies and books. Accompanied by his wife, Nesta, he collected extensively in the holarctic

78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

region. As recently as 1978, shortly before his wife’s sad and unexpected death, they were to be found wielding their nets in the mountains of Kashmir! The British Museum (Natural History) is the direct beneficiary—his incomparable collection in excess of 30,000 specimens was generously bequeathed to this Institution. As for his library, much passed to the Hope Entomological Collection, Oxford; but over the years, many volumes were presented to the BM(NH), including several rare works by Jacob Htibner.

Despite Lionel Higgins’ eminence, it must be acknowledged that some of his ideas attracted criticism. Aware of this, he maintained a dignified, but never dogmatic, con- fidence in his convictions. In methodology, he was a man of his times (few of us could claim to be anything else!). While the numerical and cladistic revolutions passed him by, he remained true to his basic principles—overall similarity and the equivalence of species- groups and genera. It is impossible to predict how the future will judge his work, but personal experience suggests that the results of “traditional” methods should be treated with the utmost respect.

How can the loss of such a man be measured? A glance at his correspondence file gives some indication. There can hardly be a lepidopterist of note who has not com- municated with him over some problem. All were given the benefit of his lifetime’s experience. The personal touches perhaps give most indication of the esteem and affec- tion in which he was held: photographs sent by correspondents, showing themselves and sometimes their families, at ease in their homes. For me, the abiding memory will be of a battered briefcase, a cork postal-box containing the latest treasure from the Pamirs or Urals, and above all the half serious admonition, always delivered with a twinkle in the eye, that I really should learn something about palaearctic butterflies.

LITERATURE CITED

Hiccins, L. G. 1941. An illustrated catalogue of the Palearctic Melitaea (Lep. Rhopa- locera). Trans. Roy. Entomol. Soc. London 91:175-365.

1950. A descriptive catalogue of the Palaearctic Euphydryas (Lepidoptera:

Rhopalocera). Trans. Roy. Entomol. Soc. London 101:435-—489.

1955. A descriptive catalogue of the genus Mellicta Billberg (Lepidoptera:

Nymphalidae) and its species, with supplementary notes on the genera Melitaea and

Euphydryas. Trans. Rey. Entomol. Soc. London 106:1-131.

1960. A revision of the melitaeine genus Chlosyne and allied species (Lepi-

doptera: Nymphalinae). Trans. Roy. Entomol. Soc. London 112:381-467.

1975. The classification of European butterflies. Collins, London.

1978. A revision of the genus Euphydryas Scudder (Lepidoptera: Nymphali-

dae). Entomol. Gaz. 29:109-115.

1981. A revision of Phyciodes Hiibner and related genera, with a review of the classification of the Melitaeinae. Bull. Brit. Mus. (Nat. Hist.) (Entomol.) 43:77- 243.

Hiccins, L. G. & N. D. Ritey. 1970. A field guide to the butterflies of Britain and Europe. Collins, London.

PHILLIP R. ACKERY, Butterfly Section, British Museum (Natural History), Cromwell Road, London SW7 5BD, England.

Date of Issue (Vol. 40, No. 1): 9 October 1986

EDITORIAL STAFF OF THE JOURNAL WILLIAM E. MILLER, Editor

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

Associate Editors: Boyce A. DRUMMOND III, DOUGLAS C. FERGUSON, THEODORE D. SARGENT

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study of Lepidoptera. Short manuscripts concerning new state records, current events, and notices should be sent to the News, June Preston, Editor, 832 Sunset Drive, Lawrence, KS 66044 U.S.A. Journal contributors should prepare manuscripts according to the following in- structions.

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Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch paper. Titles should be explicit and descriptive of the article’s content, including the family name of the subject, but must be kept as short as possible. The first mention of a plant or animal in the text should include the full scientific name with author. Mea- surements should be given in metric units; times should be given in terms of the 24- hour clock (0930 h, not 9:30 AM). Underline only where italics are intended. References to footnotes should be numbered consecutively, and the footnotes typed on a separate sheet.

Literature Cited: References in the text of articles should be given as Sheppard (1959) or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the heading LITERATURE CITED, in the following format:

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, references should be given in the text as Sheppard (1961, Adv. Genet. 10:165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1:23-30).

Illustrations: Only half of symmetrical objects such as adults with wings spread should be illustrated, unless whole illustration is crucial. Photographs and drawings should be mounted on stiff, white backing, arranged in the desired format, allowing (with particular regard to lettering) for reduction to their final width (usually 4% inches). Illustrations larger than 8% x 11 inches are not acceptable and should be reduced pho- tographically to that size or smaller. The author’s name, figure numbers as cited in the text, and an indication of the article’s title should be printed on the back of each mounted plate. Figures, both line drawings and halftones (photographs), should be numbered consecutively in Arabic numerals. The term “plate” should not be employed. Figure legends must be typewritten, double-spaced, on a separate sheet (not attached to the illustrations), headed EXPLANATION OF FIGURES, with a separate paragraph devoted to each page of illustrations.

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CONTENTS

PRESIDENTIAL ADDRESS, 1984: A TRIBUTE TO THE AMATEUR. Lee D. Miller) jo A NEw SPECIES OF EPIDROMIA (NOCTUIDAE) FROM FLORIDA. M. Alma Solis 2.00 i eC

THE LARVA AND PUPA OF LYCOREA PIETERI LAMAS (DANAIDAE). David Kenneth Wetherbee 00

LIFE HISTORY AND HABITS OF EXOTELEIA ANOMALA HODGES, A PONDEROSA PINE NEEDLE MINER IN THE SOUTHWESTERN UNITED STATES (GELECHIIDAE). Robert E. Stevens ................

BIOLOGY AND IMMATURE STAGES OF HEMILEUCA DIANA AND H., GROTEI (SATURNIIDAE). Paul M. Tuskes ae

NATURAL HISTORY AND ECOLOGICAL CHEMISTRY OF THE NEO- TROPICAL BUTTERFLY PAPILIO ANCHISIADES (PAPILIONIDAE). Allen M. Young, Murray S. Blum, Henry M. Fales & Z. Bian oe a

PYRGUS COMMUNIS AND P. ALBESCENS (HESPERIIDAE) IN NEVADA. George. T. Austin: 0

First REPORTED MALES, SPECIES STATUS, AND AFFINITIES OF EPARGYREUS SPANNA EVANS (HESPERIIDAE). Kurt Johnson dx David Matusik 00 ee

GENERAL NOTES Tryon Reakirt: A Sequel: F. Martin Brown 1...

Saturnia walterorum (Saturniidae) in Mexico: A New National Record. Kirby L. Wolfe & Marvin 'D.: Valverde ue

Emergence of Adult Ectomyelois muriscus (Dyar) (Pyralidae) from a Pod of Theobroma simiarum Donn. Smith (Steruliaceae) in Costa Rica. Allen M. Young te i OO CAA ee

The Female of Papilio xanthopleura Godman & Salvin (Papilionidae). Kurt Johnson, Rick Rozycki\<> David, Matusik) 0000)

Placement and Fate of Monarch Butterfly Pupae in Northern California. Leslie V. Smith, so oats

Aberrant Postmedial Spots in Icaricia icarioides lycea and I. acmon (Lycaen- idae), Richard Holland i

Notes on a Costa Rican “Monkey Slug’ (Limacodidae). Allen M. Young ... Book REVIEWS OBITUARIES

TARR Ren nee e nee nen ene en nnn e eee naan nen n anne Rane nnn en aeenenaeanennnennanenananenaeeensanaaennnann sana aanenestenaeeeennsenaneneneeansanceneneneannaee

ANNOUNCEMENT Inauguration of Manuscript Dating in the Jourrral cc ccsesssseeevecssuceeneresneeie

Volume 40 1986 Number 2

ISSN 0024-0966

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

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

23 October 1986

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Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the Lepidopterists’ Society, % Richard A. Arnold, Secretary, 50 Cleaveland Rd. #3, Pleasant Hill, CA 94523. Second-class postage paid at Los Angeles, CA and additional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, Julian P. Donahue, Assistant Secretary, Dept. of Entomology, Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007 U.S.A.

Cover illustration: First stage larva of Natada nasoni (Grote) (Limacodidae), from Dyar 1899, J. New York Entomol. Soc. 7: 61-67. Suggested by Marc E. Epstein.

JouRNAL OF Tue LeEpIpoPTERISTS’ SOCIETY

Volume 40 1986 Number 2

Journal of the Lepidopterists’ Society 40(2), 1986, 79-92

WHY PIERIS RAPAE IS A BETTER NAME THAN ARTOGEIA RAPAE (PIERIDAE)

ROBERT K. ROBBINS

Department of Entomology, MRC NHB 127, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560

PAMELA M. HENSON

Smithsonian Archives, A & I 2135, Smithsonian Institution, Washington, D.C. 20560

ABSTRACT. We show that there is no phylogenetic justification for changing the name of Pieris rapae to Artogeia rapae. We “define” Pieris by the presence of andro- conial basal lobes, and suggest that this grouping, which includes P. rapae, P. brassicae, and P. napi, is monophyletic. Female genital characters indicate that Perrhybris, Ita- ballia, and Ganyra are the closest relatives of Pieris. We discuss criteria for choosing generic nomenclature, and suggest that the following guidelines will best promote no- menclatural stability. If a genus is monophyletic, do not change the name. If a genus is not monophyletic, choose the combination of monophyletic generic groupings that will create the fewest name changes. If another option causes more name changes now but will be more stable in the future because of better evidence for monophyly, then present the reasons and evidence for that choice.

Pieris rapae Linnaeus is one of the best known and commonly en- countered temperate area butterflies. Although native to the Palaearc- tic, it is now nearly ubiquitous in suitable disturbed habitats in North America (Howe 1975), New Zealand (Gibbs 1980), and Australia (Common & Waterhouse 1981). Because P. rapae is widely distributed, easily reared, and a pest on cultivated crucifers, it has been extensively studied in the agricultural, ecological, and physiological literature (Harcourt 1966, Dempster 1969, Aplin et al. 1975, Slansky & Feeny 1977, Blau et al. 1978, Kobayashi & Takano 1978, Yamamoto & Ohtani 1979, Wolfson 1980, Chew 1981, Jones et al. 1982, Gilbert 1984, Ma- guire 1984).

The generic placement of P. rapae has recently been changed from Pieris to Artogeia Verity. Schrank (1801) placed rapae in Pieris when he originally described the genus, and Klots (1933) retained this generic

80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

placement in his systematic treatment of world Pieridae. Verity (1947) proposed Artogeia as a subgenus including rapae, and Kudrna (1974) and Higgins (1975) elevated it to generic rank, an action that has been followed in some general works (Pyle 1981, Miller & Brown 1981) but not others (Kawazoé & Wakabayashi 1976, Opler & Krizek 1984). This situation was further complicated when Kudrna later treated Artogeia as a subgenus (Blab & Kudrna 1982), and Feltwell and Vane-Wright (1982) suggested that Artogeia might not be monophyletic.

In this paper we assess the evidence for switching rapae from Pieris to Artogeia. We took up this project because of repeated inquiries from scientists in a variety of biological disciplines to the National Museum of Natural History concerning the proper generic name for rapae. We address the following questions: ““What is the evidence for the change in generic nomenclature and is it compelling?’”’, ““What is the ‘best definition’ for Pieris?’’, “What are the closest relatives of Pieris?’’, and “What criteria will promote stability of generic nomenclature?”

It is not our intent to produce a definitive work on Pieris systematics. Besides reporting the results of a few representative female genitalic dissections, we discuss published information only, all of which was available to Kudrna and Higgins, with the exception of two recent papers on isozymes. We discuss characters sequentially, note their states and distributions, and generally limit our discussion to those species for which we have information. Many species level decisions, partic- ularly in the P. napi group, are controversial (Warren 1961, Bowden 1972, Eitschberger 1983, Geiger & Scholl 1985); we avoid entering the fray because it is largely irrelevant to our purpose. Finally, we show that treatment of all Pieris species would not alter our conclusions.

PIERIS RAPAE OR ARTOGEIA RAPAE

In this section, we ask whether rapae is more closely related to napi Linnaeus—the type of Artogeia—or to brassicae Linnaeus—the type of Pieris. The classification of Kudrna (1974) and Higgins (1975) im- plies that the former is correct, while others (Mariani 1937, Geiger 1981, Geiger & Scholl 1985) suggest the opposite. These two possibil- ities are represented by alternative phylogenies (Figs. 1 & 2).

To determine primitive character states among these species (the state at point A in the phylogenies), we used two sets of outgroup species. The first set is Pontia daplidice Linnaeus and Synchloe callid- ice Hubner. Pierid specialists (Klots 1933, Bernardi 1947) considered them to be closely related to the brassicae-napi complex, and some- times included them in Pieris. Kudrna (1974) placed them next to Pieris and Artogeia. They are the immediate outgroups of the brassi- cae-napi lineage in dendrograms constructed from isozyme data (Gei-

VOLUME 40, NUMBER 2 81

of of > & > Kj Yr eg ¢ Yr Oa $ ¥ * $ & v C c B B A A

1 2

Fics. 1 & 2. Phylogenies showing cladogenesis among Pieris brassicae, P. rapae, and P. napi. The letters designate ancestral species in the branching sequence.

ger 1981, Geiger & Scholl 1985). The second outgroup set is Ganyra Billberg, Itaballia Kaye, and Perrhybris Hiibner (sensu Klots 1938). We discovered that they share female genitalic characters with the brassicae-napi complex (detailed below), and may be more closely related to them than has been previously realized.

The first character that Kudrna (1974) and Higgins (1975) used in their taxonomic analysis was androconial structure (illustrations in Dix- ey 1910, 19382, Bernardi 1947, Warren 1961). There are four major shapes in the “Pieris group” with slight quantitative interspecific vari- ation within each type: P. brassicae has one type of androconium (Fig. 3), rapae and A. napi a second (Figs. 4 & 5), outgroups P. daplidice, Perrhybris, and Itaballia a third (Figs. 6 & 7), outgroup Ganyra a fourth (Fig. 8), and outgroup S. callidice lacks androconia. On either phylogeny this distribution of character states can be explained, no matter which outgroup state occurred at point A, by the rapae-napi androconium evolving at point B and the brassicae androconium evolv- ing at point C. Although there are other equally parsimonious possi- bilities, either phylogeny could produce the distribution of character states simply—each androconium type evolved once. Thus, although rapae and A. napi share a similar androconial structure, this distribu- tion provides no evidence for choosing between the phylogenies in Figs. 1 and 2.

Kudrna (1974) and Higgins (1975) also used male genitalic charac-

82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

W

4 4 3 cig 5 ] 8

Fics. 3-8. Outlines of androconia, redrawn from Dixey (1932). Arrows in first three figures point to the right basal lobe. 3, Pieris brassicae; 4, P. rapae; 5, P. napi; 6, Pontia daplidice; 7, Itaballia demophile; 8, Ganyra josepha.

oo

ters for defining Artogeia. The male genitalia of A. napi and rapae are similar to each other, and differ from P. brassicae (illustrated in Klots 1933, Bernardi 1947, Kudrna 1974, Higgins 1975). The penis of P. brassicae has a dorsal hump and each valva has a distal pointed process while rapae and A. napi lack the dorsal hump and the process. The outgroups, like rapae and A. napi, lack a penial dorsal hump and process on the valva (except for Ganyra, which has a differently shaped valva process). Thus, the P. brassicae penis and valva morphology is derived, defines only P. brassicae (evolved at point C on either phy- logeny), and provides no information about the phylogenetic position of rapae.

Higgins (1975) also used haploid chromosome numbers to differen- tiate Pieris from Artogeia. Reported haploid chromosome numbers (De

VOLUME 40, NUMBER 2 83

Fics. 9-14. Right dorsolateral view of the corpus bursae and anterior portion of the ductus bursae (except for Pontia callidice, which is a dorsal view with an additional lateral aspect of the cervix). 9, Pieris napi; 10, P. rapae; 11, P. brassicae; 12, Perrhybris pyrrha; 13, Ganyra josepha; 14, Pontia callidice.

84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Lesse 1967, 1970, De Lesse & Brown 1971, Robinson 1971) are P. brassicae—15, rapae—25-26, A. napi—25-28, and for the outgroups P. daplidice—26, S. callidice—26, Itaballia—25-26, and Perrhybris— 27-29. We infer that the lower haploid chromosome number of P. brassicae is a derived character state that defines only P. brassicae—it evolved at point C in the phylogenies—and that provides no infor- mation on the systematic position of rapae.

Since the characters used by Kudrna (1974) and Higgins (1975) pro- vide no evidence for choosing between the phylogenies in Figs. 1 and 2, the placement of rapae in Artogeia was phylogenetically unjustified. We now ask whether other published characters provide information on the generic placement of rapae.

Mariani (1937) and Bernardi (1947) examined “Pieris” female gen- italia, and reported interspecific variation in morphology of the single signum (often called “lamina dentata” in the pierid literature) on the corpus bursae. The signum of A. napi has a long posterior process (“flagello” of Mariani, “‘tail’’ of Chang [1963]) that is lacking in rapae and P. brassicae and in all outgroup species (Figs. 9-14, figures in Mariani and Bernardi). Thus, the posterior process is a derived char- acter state that apparently evolved once on the lineage leading to A. napi, and does not give us information with which to distinguish the phylogenies in Figs. 1 and 2.

Geiger (1981) and Geiger and Scholl (1985) electrophoresed enzymes from species of Pieris, Artogeia, Pontia, and Synchloe, but not for the other outgroups. They obtained a dendrogram of relationships by using an unweighted pair-group average clustering method on genetic sim- ilarities. They found that rapae is more similar to P. brassicae than to A. napi and that all three are more similar to each other than to Pontia and Synchloe. This result supports the phylogeny depicted in Fig. 1. There are numerous methods for coding and analyzing electrophoretic data, and Mickevich and Mitter (1981) propose criteria for judging different methods. Before uncritically accepting their dendrograms, we would want to know if other methods of coding and analysis corrob- orate their results.

In summary, analysis of published characters indicates that the use of Artogeia as a genus or subgenus including rapae is phylogenetically unjustified. Although the male genitalia, androconia, and haploid chro- mosome numbers of rapae are more similar to A. napi than to P. brassicae, the opposite relationship is true with regard to the female genitalia and isozymes. Further, these similarities are based on primi- tive character states, as Feltwell and Vane-Wright (1982) had predict- ed, and do not provide the information necessary to choose between the phylogenies in Figs. 1 and 2. Characters of “Pieris” immature

VOLUME 40, NUMBER 2 85

stages may provide the information necessary to decide this point, but have not been used in Pieris revisions.

THE GENUS PIERIS

Since rapae is such a widely known species and since Kudrna and Higgins’ concept of Pieris and Artogeia leave rapae without certain generic placement, we ask in this section whether there are other, more reasonable definitions for Pieris. Klots (1933) revised the world pierid fauna. Although he narrowed the definition of Pieris—it previously had been a catchall genus for many questionably related pierines— subsequent authors have split the genus further. We ask whether any of these groupings are monophyletic. For outgroup comparisons, we use those genera that Klots considered to be most closely related to Pieris: Leptophobia Butler, Itaballia, Perrhybris, Ascia Scopoli (in- cluding subgenus Ganyra), Tatochila Butler, Phulia Herrich-Schaffer, and Baltia Moore.

Three different concepts of Pieris besides that of Kudrna and Hig- gins have been used since 1933. For ease of communication, we list representative species for each grouping, and refer the reader to the original work for a complete list. Klots (1933) placed brassicae, rapae, napi, callidice, daplidice, and pylotis in Pieris. Mariani (1937) and Bernardi (1947) put the first four of these species in Pieris, while Hig- gins and Riley (1970) restricted Pieris to the first three. (Note that Higgins [1975] later narrowed the genus further, like Kudrna, to in- clude only brassicae and close relatives.)

Klots (1933) defined Pieris with a paragraph of character states. For the most part, however, they are too ambiguous to code accurately. For example, how does one code “antenna long, with abrupt club” (Pieris), “antenna long, with usually somewhat abrupt club” (Ascia), and “antenna long, with flattened abrupt club” (Tatochila)? Further, each of Klots’ generic character states is shared with at least one out- group genus. Because of character state ambiguity and the lack of unique, potentially defining character states, we found no evidence in Klots’ work to indicate that his concept of Pieris is monophyletic.

Mariani (1937) and Bernardi (1947) apparently ignored pylotis (a neotropical species that does not “look” like other Pieris species) and moved daplidice to Pontia. Pontia daplidice has forewing veins R; and R,,; fused while they are separate in the other Pieris species. Outgroups Phulia and Perrhybris have the fused veins while the other outgroups have separate veins (Klots 1933). Because both character states are found in the outgroups, the primitive character state is ambiguous. Phylogenetic interpretation of this character is thus equivocal.

Higgins and Riley (1970) joined callidice with daplidice in Pontia,

86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

so that their Pieris grouping consisted of brassicae, rapae, napi, and close relatives. The androconial lateral edges of these species and their close relatives protrude basally to form lobes (Dixey 1932, Bernardi 1947, Warren 1961) (Figs. 83-5) while the androconial basal edge is flat in daplidice and relatives (Bernardi 1947) (Fig. 6), pylotis (Dixey 1932), and all outgroup genera (Dixey 1932) (Figs. 7 & 8). Since daplidice and pylotis share the primitive state—that which occurs in the out- group genera—the androconial basal lobes would appear to be a de- rived, defining character for the brassicae-napi complex. This situation contrasts with the one in the previous section, in which rapae and napi share an androconium type that does not reflect phylogenetic relat- edness because brassicae does not possess the primitive character state.

We “define” Pieris by the androconial basal lobes. Although we are reluctant to define a genus on the basis of one character state, there are no alternatives in this case. From published illustrations of andro- conia (Dixey 1932, Bernardi, 1947), we place the following specific taxa—listed in Bernardi—in Pieris: brassicae, deota de Niceville, bras- sicoides Guerin, krueperi Staudinger, tadjika Groum-Grshimailo, ca- nidia Sparrman, manni Mayer, rapae, dubernardi Oberthur, extensa Poujade, stoetzneri Draeseke, napi, virginiensis Edwards, ochsenhei- meri Staudinger, ergane Geyer, melete, and davidis Oberthur. Al- though there are other taxa, particularly in the napi group, that are given specific rank by some authors, we leave species level decisions to others.

We believe that this grouping is the most reasonable and stable one for Pieris. There is an enormous biological literature on Pieris brassi- cae, Pieris rapae, and Pieris napi, and the name Pieris is widely rec- ognized by nontaxonomists in connection with these species. Our grouping will preserve this association, and because it is based on the best available evidence for monophyly, it is most likely to be stable in the future.

There are three morphologically distinct groups within Pieris. The P. brassicae group (brassicae, deota, brassicoides) has the androconial and male genital structures of brassicae, and is probably a monophy- letic lineage defined by these structures. The P. napi group (napi, virginiensis, ochsenheimeri, ergane, melete, davidis, and presumably stoetzneri, extensa, and dubernardi—Bernardi [1947]) has a posterior process on the signum, which probably defines this group as a mono- phyletic lineage. The P. rapae group (krueperi, tadjika, canidia, man- ni, rapae) lacks derived character states. There is no evidence to in- dicate whether it is monophyletic or whether it is phylogenetically more closely related to the P. brassicae or P. napi groups. Thus, even if we had examined all Pieris species in the previous section, it would

VOLUME 40, NUMBER 2 87

not have provided us with evidence on the phylogenetic position of P. rapae. Interestingly, the same three groups result when isozyme data are analyzed phenetically (Geiger & Scholl 1985).

THE RELATIVES OF PIERIS

In this section we ask which genus or genera are most closely related to Pieris. From the work of Mariani (1937) and Bernardi (1947), it appeared that the bursa copulatrix, particularly signum location and shape, had states that might provide information on the phylogenetic position of Pieris. Because this character was promising, but unrecord- ed for many of the outgroups, we dissected the female genitalia of species in Pieris and related genera.

We recorded three character states of the bursa copulatrix. In the first, the signum is a narrow transverse band located at the posterior end of the corpus bursae just around the entrance to the ductus bursae (Fig. 14). We recorded this state in Pontia (daplidice, protodice), Synchloe (callidice), Leptophobia (eleone Hewitson, aripa Boisduval), and Ascia (monuste Linnaeus). It also occurs in Tatochila, Phulia, Baltia, and close relatives (Field 1958, Herrera & Field 1959, Field & Herrera 1977), in the pierine Aporia Htibner (Mariani 1937) and the coliadines Colias Fabricius (Mariani 1937) and Eurema Hubner (Field 1950).

In the second character state, the signum is located on the right dorsolateral side of the corpus bursae well anterior to the entrance of the ductus bursae (Figs. 9-13). Signum shape varies, particularly in how far it extends posteriorly and in the amount of sclerotization of the median line. We recorded this character state in Pieris (brassicae, rapae, napi, melete), Ganyra (josepha Godman & Salvin, limona Schaus), Itaballia (demophile Linnaeus, viardi Boisduval, pisonis Hew- itson), and Perrhybris (pyrrha Fabricius, pamela Cramer [=lypera Kol- lar], lorena Hewitson). Mariani (1937) noted its occurrence in all 12 Pieris species that he examined.

A third character state is limited to Glennia pylotis. There is no signum. The corpus bursae and ductus bursae are greatly modified into a long tube that occupies the length of the abdomen. This tube grad- ually increases in diameter anteriorly, and the usual abrupt change in size that distinguishes the corpus from the ductus is absent.

The closest relatives of Pieris appear to be Perrhybris, Itaballia, and Ganyra. The position of the signum on the right dorsolateral side of the corpus bursae is an unusual character state that is apparently re- stricted to these four genera. The other genera that Klots (1933) placed near Pieris have the signum at the posterior end of the corpus bursae, which is probably the primitive state for the pierines because it is also

88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

found in the coliadines Eurema and Colias. A definitive survey of the distributions of female genital structures is obviously desirable.

Those holarctic species that Klots (19383) put in Pieris, but which have recently been placed in Pontia (Higgins 1975, Miller & Brown 1981) are often considered to be close relatives of Pieris. However, we know of no evidence that this group is more closely related to Pieris than to other genera, such as Tatochila, Phulia, and relatives (Shapiro 1979). Further, we have found no. published characters to determine whether this group is monophyletic. In short, there is a glaring need for a worldwide treatment of the pierines.

The placement of Glennia pylotis remains a problem. Although Klots (1933) treated Glennia as a subgenus of Pieris, it lacks the an- droconial basal lobes and signum of Pieris. There is currently no evi- dence to decide whether the divergent female genitalia of Glennia evolved from the Pieris type or from the Pontia type.

STABILITY AND GENERIC NOMENCLATURE

In this section we use the confusion over the generic nomenclature of P. rapae as an example to discuss the relationship between taxonom- ic method and nomenclatural stability.

We suggest that butterfly generic nomenclature can be more objec- tively chosen than in the past by using the criteria of “stability” and “monophyly”. The Preamble to the International Code of Zoological Nomenclature (Int. Comm. on Zool. Nomenclature 1985) states that “.. the object of the Code is to promote stability and universality in the scientific names of animals ....’’ Ehrlich and Murphy (1982) dis- cuss the widespread support for a stable generic nomenclature.

By monophyly, we refer to taxa defined by derived characters. As Jordan (1898) noted, “... we have here an instructive illustration of the fact—so very often entirely disregarded in classificatory work— that the presence of the same character in two different [taxa] .. . is, evidence of closer relationship only, if the character is a specialisation and not of the ancestral type.”’ Jordan’s logic is simple, but has been largely ignored by butterfly systematists.

The application of stability and monophyly to groups with an estab- lished generic nomenclature, such as the bulk of the butterflies, is straightforward. If a genus is monophyletic, do not change the name. If a genus is not monophyletic, choose the combination of monophy- letic generic groupings that will create the fewest name changes. If another option causes more name changes now but will be more stable in the future because of better evidence for monophyly, then present the reasons and evidence for that choice.

Ehrlich and Murphy (1982) suggested that the concept of balance

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(the equivalence of categorical rank in related taxa, sensu Mayr 1969) also be used to decide generic nomenclature. Despite Mayr’s discussion of how balance might be applied, this method is subjective, particularly since it is unclear exactly what the method is supposed to estimate. Although objectivity is not itself justification for using a criterion, we believe that an obviously subjective one, such as balance, will promote instability of butterfly generic nomenclature.

Kudrna (1974) and Higgins (1975) used the criterion of “‘similarities and differences” to justify their recognition of Artogeia, and did not mention stability and monophyly. For example, Higgins (1975) stated, “Their [Artogeia] genitalia, androconial scales and chromosome num- bers differ from those of P. brassicae and it is not satisfactory to include them in the same genus.” Neither worker suggested that Pieris, as used by Klots (1938) or Bernardi (1947), was polyphyletic. Neither discussed the possible confusion that would result from changing the generic nomenclature of P. rapae and P. napi.

There are many problems with the criterion of similarities and dif- ferences. (1) The similarities and differences used by Kudrna and Hig- gins do not provide information on the phylogenetic position of P. rapae. This example is a clear illustration that similarities and differ- ences alone are insufficient to establish monophyly. (2) If Kudrna and Higgins had examined female genitalia and isozymes (as opposed to male genitalia, androconia, and chromosome numbers), they would have put rapae in Pieris. When taxonomic conclusions depend upon the character set used, the result is instability. (8) We are certain that Kudrna and Higgins believed that Artogeia should be split from Pieris because it is “sufficiently different.’’ However, if one “‘authority”’ states that a difference is sufficient to split a genus, but another disagrees, then how can these conflicting views be resolved? It is evident that the criterion of similarities and differences promotes instability, and should not be used.

Kudrna (1974) and Higgins (1975) assumed that the divergent mor- phology of P. brassicae is the result of phylogenetic distance, but did not consider that it might be the result of rapid evolution. We hypoth- esize that rearrangement of genes caused by extensive chromosomal fusion—haploid chromosome number decreased from about 26 to 15 at point C in Figs. 1 and 2 (discussion in White 1973)—affected gene expression during development (the “position effect”; Dobzhansky 1957, White 1973), and is causally related to the divergent male genital, androconial, and larval (D. Weisman, pers. comm.) morphology of P. brassicae. Chromosomal rearrangements would not be expected to af- fect the protein products of structural genes, however—an expectation consistent with isozyme data (Geiger 1981, Geiger & Scholl 1985). Our

90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

hypothesis generates the testable prediction that morphology and chro- mosome numbers are perfectly correlated; all species in the P. brassicae group should have reduced haploid chromosome numbers (about 15) while none in the P. rapae and P. napi groups should have the reduced numbers.

J. H. Comstock (1893) wrote: “Here I believe lies the work of the systematist of the future. The description of a new species, genus, family or order, will be considered incomplete until its phylogeny has been determined so far as is possible with the data at hand.’’ Com- stock’s vision of the holarctic butterfly “systematist of the future’ is, by and large, still just a vision. Until we have reasonable phylogenies, generic nomenclature is bound to be unstable. In the meantime, sug- gested changes in generic nomenclature will hopefully be based on evidence of monophyly, and proposed with due regard for stability.

ACKNOWLEDGMENTS

We conducted this project under the auspices of the Maryland Center for Systematic Entomology, a consortium of the Department of Entomology, Smithsonian Institution, Department of Entomology, University of Maryland, and the Systematic Entomology Laboratory (USDA). We thank C. W. Mitter of the Center for his support. We thank L. D. Miller of the Allyn Museum of Entomology for the loan of a specimen. For providing thoughtful criticisms of various manuscript drafts, we thank P. R. Ackery, A. Aiello, S. R. Bowden, M. D. Bowers, J. M. Burns, C. J. Callaghan, F. S. Chew, S. P. Courtney, C. V. Covell, R. de Jong, J. C. Downey, U. Eitschberger, J. N. Eliot, L. F. Gall, J. S. Glassberg, R. W. Hodges, G. Lamas, C. W. Mitter, S. S. Nicolay, P. A. Opler, R. W. Poole, J. E. Rawlins, R. L. Rutowski, A. M. Shapiro, G. B. Small, R. I. Vane-Wright, and B. A. Venables. We thank G. Venable for professional illustrative aid.

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‘Journal of the Lepidopterists’ Society 40(2), 1986, 938-96

RESTINGA BUTTERFLIES: BIOLOGY OF SYNARGIS BRENNUS (STICHEL) (RIODINIDAE)

CurRTIs J. CALLAGHAN

Rua yeddo Fiuza 595, Petropolis, Rio de Janeiro, Brazil, and “Pesquisador Associado” of the Museu Nacional, Rio de Janeiro

ABSTRACT. Synargis brennus (Stichel) isa myrmecophilous riodinine butterfly that inhabits the restinga, a low forest on the Brazilian coast. The larval food plant Dalbergia ecastophylla has nectaries attractive to ants of the genera Camponotus and Azteca. Ants attend the larvae, drumming on them to stimulate a secretion which they subsequently eat from two glands located on the eighth abdominal segment.

This paper continues the series of studies on restinga butterflies start- ed with the biology of Menander felsina (Callaghan 1977). The ur- gency of the study of this habitat is underlined by the fact that the site of the latter study near Rio de Janeiro has been destroyed by a housing development.

Synargis brennus (Stichel) (Fig. 1), a myrmecophilous riodinine but- terfly, is not an endemic restinga species, but is found mostly in this habitat, and forms an important element of its fauna. The butterfly ranges from the coast in southeast Brazil across the Planalto to the Amazon basin, where it intergrades with S. calyce (Felder). The habitat where the observations were made is called “restinga’’, and consists of low, scrubby, dense, woody vegetation growing along the coast. The vegetation and physical characteristics are summarized in Callaghan (1977). The observations in the present study were made over three months during visits to Buzios, a very dry section of coastline 170 km E of Rio de Janeiro (Fig. 2).

Observations were made in the field and in the laboratory. The letters T and A followed by a number refer to thoracic and abdominal segments, respectively.

DESCRIPTION OF IMMATURE STAGES

Egg (Fig. 3). Rounded laterally, flattened dorsally and ventrally, giving the appearance of a fat tire. Color shiny bronze, with a network of small ribs forming a hexagonal pattern smaller around the micropyle and ventrally; top of ribs irregular, with a small tubercle at each intersection. Micropyle circular, depressed, with numerous small openings. Du- ration 9 days (N = 5).

First instar. Length 1.8 mm, head width 0.2 mm. Form rounded dorsally, flat ven- trally. Head black, numerous small setae on face. Thorax light brown with black dots, pair of reddish, broken lines dorsally; T1 bilobed, with