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7 ISSN 0024-0966 a 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
Ma 3 23 February 1994
THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE COUNCIL
PAUL A. OPLER, President JORGE E. LLORENTE-BOSQUETS, Ray E. STANFORD, Immediate Past Vice President
President FREDERICK W. STEHR, CHEN-SHING LIN, Vice President Vice President JULIAN P. DONAHUE, Acting Secretary ROBERT J. BORTH, Treasurer Members at large: Charles V. Covell, Jr. Eric H. Metzler John V. Calhoun Linda S. Fink Robert K. Robbins Robert C. Lederhouse Scott E. Miller J. Benjamin Ziegler William E. Miller
EDITORIAL BOARD
PAUL A. OPLER (Chairman), FREDERICK W. STEHR (Member at large) JOHN W. BROWN (Journal), WILLIAM E. MILLER (Memoirs) STEPHANIE S. MCKOWN (News)
HONORARY LIFE MEMBERS OF THE SOCIETY
CHARLES L. REMINGTON (1966), E. G. MUNROE (1973), ZDRAVKO LORKOVIC (1980), IAN F. B. COMMON (1987), JOHN G. FRANCLEMONT (1988), LINCOLN P. BROWER (1990), DOUGLAS C. FERGUSON (1990), HON. MIRIAM ROTHSCHILD (1991), CLAUDE LEMAIRE (1992)
The object of the Lepidopterists’ Society, which was formed in May 1947 and for- mally constituted in December 1950, is “to promote the science of lepidopterology in all its branches, .... to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards these aims.
Membership in the Society is open to all persons interested in the study of Lepi- doptera. All members receive the Journal and the News of the Lepidopterists Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. There are four numbers in each volume of the Journal, scheduled for February, May, August and November, and six numbers of the News each year. |
Active members—annual dues $25.00 Student members—annual dues $15.00 Sustaining members—annual dues $35.00 Life members—single sum $500.00 Institutional subscriptions—annual $40.00
Send remittances, payable to The Lepidopterists’ Society, to: Robert J. Borth, Treasurer, 6926 North Belmont Lane, Fox Point, WI 53217, U.S.A.; and address changes to: Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007- 4057 U.S.A. For information about the Society, contact: Julian P. Donahue. To order back issues of the Journal, News, and Memoirs, write for availability and prices to the Publications Manager: Ronald Leuschner, 1900 John St., Manhattan Beach, CA 90266- 2608, U.S.A.
Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for $40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Second-class postage paid at Los Angeles, California and ad- ditional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. If you have used a Society address many years after its publication date, and your mail is returned as undeliverable, please redirect it to the Natural History Museum address provided above.
Cover illustration: The black swallowtail, Papilio polyxenes asterius (Stoll), is a wide- spread species of the eastern United States. The larvae feed on a variety of umbelliferous plants (Apiaceae). Original drawing by Erik Russell Wild, Museum of Natural History, University of Kansas, Lawrence, Kansas 66045-2454.
JOURNAL OF
Tue LEPIDOPTERISTS’ SOCIETY
Volume 48 1994 Number 1
Journal of the Lepidopterists’ Society 48(1), 1994, 1-7
PRESIDENTIAL ADDRESS 1993: ON THE COMPARATIVE DISTRIBUTIONS OF LEPIDOPTERA AND LEPIDOPTERISTS
RAY E. STANFORD 720 Fairfax Street, Denver, Colorado 80220, USA
Both Lepidoptera and lepidopterists, each part of a much larger group of similar organisms, represent but fleeting moments in time. But the Lepidoptera have flitted a great deal longer than we have as their students, by four orders of magnitude. I have been interested in the spatial and temporal distribution of butterflies for nearly 50 years, beginning in the neighborhood of my house and expanding incremen- tally to all of western North America (Stanford & Opler 1993), but began to notice a consistent artifact on the dot maps: butterflies are shown to be more common in cities, along paved roads, and in beautiful mountain meadows than in deserts, on farms, or in abandoned mine- fields. This presentation is an attempt to put these biases into perspec- tive, based on several years’ attempts by me and others to find out what species do occur (and even thrive) in the less-well-known habitats of our large region. Before summarizing my own research in this realm, it is necessary to give some historical perspective both on bugs and buggers.
Although some insect orders are known from fossils dating from early Cretaceous time, the first Lepidoptera appeared along with flowering plants about 120 million years before the present (mybp), and probably all families of Lepidoptera that exist today had differentiated by 66 mybp (Emmel et al. 1992). Most fossil butterflies date from the late Eocene to early Oligocene epochs, about 48-34 mybp, or later. Exten- sive movements of species occurred during and following the Pleisto- cene glaciations, as has been determined both from examination of fossils and study of today’s distributions taking geologic evidence into consideration. Humans undertook equally extensive movements at the same time, extending into much of North and South America from
2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Eurasia perhaps as long ago as 35 thousand ybp. Lepidoptera appear in pictographs and caves dating from then, but it would be a stretch to refer to the artists as lepidopterists! The first drawings that are fairly easily determined to species are from Egyptian and Sumerian tombs from 5000 to 3000 ybp; Danaus chrysippus can be dated from drawings in Luxor created about 3500 ybp (Larsen 1990). The first surviving descriptions of Lepidoptera which may be considered scientific were by the Greek philosopher /scientist Aristotle, who lived and wrote in the 4th century BC. His accounts are sufficiently detailed that several species of butterflies and moths which still occur in his country could be considered described by him, but he failed to assign them Latin names or designate type localities! Common names (in Greek for chry- sippus) are entirely appropriate for common species (Miller 1992), but are a conundrum for uncommon ones (Scott 1993) for which every author seems to make up a different common name. For example, what should be ‘““Edwards’ skipper’? He described 51 species of them (as presently classified) from the western United States alone. I shall not attempt to review the history of lepidopterists since the time of Linnaeus (1753), Cramer (1775), or Fabricius (1807), but the late F. Martin Brown wrote many historical papers in addition to his monumental series on the W. H. Edwards taxa. Brown influenced many of us over several decades, and I find it appropriate to dedicate my remarks here to his memory since he was long a guiding force behind my research.
Many factors influence the distribution of butterflies in space and time, including climate, host plants and other biological requirements, and the effects of human activity. In order for range maps to show the actual distribution of a species for any given interval of time, these factors and the potential biases and artifacts mentioned at the outset must be considered carefully. Also, the mapper must beware of intro- ducing errors by the very process of making maps (Monmonier 1991), and the changes in a species’ range over time require either several maps or different symbols denoting different time periods on a single map (e.g., Heath 1970). The British Atlas (Heath 1970) also shows different intensities of observation/collecting among the thousands of 10 km grid squares, so that the presence or absence of a species in a certain area may be evaluated in terms of observation density as well as other factors.
First off, I shall address the issue of errors which originate from the mapping process itself. All maps tell little white lies of necessity. Most of us grew up seeing Mercator-projection maps on classroom walls, where Greenland appears larger than the United States, and Antarctica is as long as the equator, but we learned quickly to adapt to these “lies.” My well-worn Colorado highway map shows a prominent north-south
VOLUME 48, NUMBER 1 5 °
ribbon about 2 mm wide running from Cheyenne, Wyoming, to Raton, New Mexico, Interstate Highway 25. If that width were to scale, a DC- 10 could easily land crosswise and never see or hit an automobile! Similar types of misinterpretation are possible on dot maps of small scale, where only a single dot in the center of each county (or other unit) will fit. For example, a common species such as Vanessa cardui, known from all counties of both Kansas and Nevada, appears to be very much more common in Kansas (105 tiny counties) than in Nevada (18 gigantic ones and 4 tiny ones) simply because the dots are nearly confluent in Kansas. Also, Boloria acrocnema shows in 4 large Colorado counties, with a total area of thousands of square miles, but the insect actually occurs in only a few several-acre colonies above treeline on the San Juan massif. Of course, in a scientific paper addressing either of these issues, the maps would be prepared in different formats, with equal size grid squares for the painted lady in Kansas and Nevada, and a large scale map to show the specific localities for the Uncompahgre fritillary in Colorado. In a work with over 1000 identical-format maps, these matters must be summarized by a simple caveat in the introduction: Beware of [implied] lies! Most range maps, whether showing discrete dots or shading, are based on county records in the United States because the county of the locality on the specimen label is usually easy to determine given the other label data and ready access to historical maps; also there is no standard grid system used among disciplines, although longitude/ latitude could be used. However, using counties as the basis requires considerable care to avoid plotting errors. Counties sometimes come or go, or change boundaries, or even names, and the names of towns correlate poorly with county names. Here are some examples. Before Colorado became a State in 1876, Denver was in Montana! That is, Montana Co., Kansas Territory. Several butterfly species were described from “Denver” which do not occur anywhere near the city and certainly didn’t then either, so a dot must not be placed there without better information. Grand Co., Utah Territory, included what are now Grand Co., Colorado, Grand Co., Utah, and nearly everything in between. In more recent years, Washabaugh Co., South Dakota, vanished into Jack- son Co., South Dakota (combining the map dots was quite easily ac- complished), while Yuma Co., Arizona, and Valencia Co., New Mexico split into 2 counties each (we had to go back to ground zero for many records), and Denver Co., Colorado gobbled up a lot of real estate in Adams Co. (requiring transfer of a few county dots). Bullfrog Co., Nevada, was created and then abolished so quickly that no action was necessary on our part. Several of the Sierra foothill counties in California changed their boundaries almost weekly in the days of the Gold Rush and afterward, but have fortunately been stable during most of col-
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
a
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LEPIDOPTERISTS, 1993 (Season Summaries
1993
CABBAGE WHITE (P. RAPAE),
VOLUME 48, NUMBER 1 )
lecting efforts there since the time of Lorquin. The Sonora blue, de- scribed from “Sonora,” was from near the California gold camps, not Mexico! Then, beware of ambiguous and misleading names: Bent, Col- orado, is in Las Animas Co., while Las Animas is the seat of Bent Co.: also Cheyenne Co. is in Nebraska, while Cheyenne, Wyoming is in Laramie Co., and Laramie, Wyoming is in Albany Co. (NOT New York)!
Secondly, maps reflect the habitats of lepidopterists at least as much as those of the Lepidoptera they study, and also the goals and biases of the students. I mentioned earlier that populated, easy-access, and beautiful places tend to be better known than unpopulated, remote, and barren ones, but another principle has been called Powell’s Law for its perpetrator J. A. Powell: distant places are more thoroughly studied than close ones, or “No field biologist does any significant work closer than 1000 miles from his home!’ A case in point could be the Chiricahua Mtns of Arizona, or the Galapagos Islands, but if one looks at the evolving knowledge of common species’ distributions, the exact opposite seems to be true. I have chosen the cabbage white, Pieris rapae, to illustrate this point. Panel A of Fig. 1 shows the range of this introduced Eurasian species in 1800, several years before it first ap- peared on our continent, side-by-side with the range of serious lepi- dopterists in the same year in the western United States: both zero! Panel B shows the known locations for each group in 1956, again nearly the same, in cities and towns only. Panel C shows where they are documented to exist in 1993—again identical, but this time nearly everywhere. So my corollary to Powell’s Law is the converse: “No one notices cabbage whites except in his/her own back yard!’ Except that a certain map-dotter finally picked one up on the west side of Loveland Pass, Colorado, 12,000’ above sea level, to complete the dots for all 63 counties and prove Powell correct after all. The distribution of lepi- dopterists also is shown by the fact that 110 butterfly species are known from Scott Co., Kansas, compared with 30 to 45 in surrounding and equally depauperate counties, because Virgil Calkins lived and recorded species in his back yard in Scott City for several decades this century. In his honor I have therefore formulated Calkins’ Law: If one studies a habitat for long enough, more than 100 butterfly species will be found there. Anywhere! Another artifact is caused by the interests of the observers. Swallowtails and fritillaries are much better known (but probably NOT more widespread) than skippers or noctuid moths, be-
—_—
Fic. 1. County records for the cabbage white (Pieris rapae) (left column) and the distribution of lepidopterists (right column). A) 1800; B) 1956; C) 1993.
6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
cause they are more popular. Methods of observers affect the results also; like authors who accept literature records or sightings show more polka dots than those who accept only captures with voucher specimens and extremely detailed label data (frequently only their own). Iam not faulting careful work, and am sometimes annoyed at papers that show everything from soup to nuts thereby conferring a measure of credibility on frank errors that should be expunged, but I support a middle ground. Maps that are revised frequently, or are on computer files, can be updated to show deletions and corrections as well as additions and format changes.
Finally, I pose a somewhat rhetorical question: When do range maps reach their maximal utility? An important point is reached when all common species are known from all or most grid squares or counties, because it can be inferred then that the ranges of rarer species are well- known also since most biologists are more interested in them, and that a scattergram stray species (such as Eurema mexicana) has not been encountered very frequently but can pop up nearly anywhere. We are nearing that inflection point for western butterflies now, and a map showing E. mexicana for all counties from Canada to Mexico in a few centuries would tell a different sort of lie, unless the relentless march of Eurasian weeds and fungi into the region—replacing most native plant species—should allow this particular species to flourish continent- wide. Will the range maps for cabbage whites in the year 2098 in western North American look very much like today’s, with a few more squares filled in, or will they look like those in 1800? In either case, the comparative distributions of common Lepidoptera and common people will finally be identical, but where will the uncommon ones be? Hopefully the uncommon lepidopterists will still be pursuing the un- common insects, as Rindge urged in his 1965 presidential address, which I had the privilege to hear in person. Collect NOW, he urged, because it soon would be too late. His message is still true, and even more urgent than nearly 30 years ago. Much collecting is possible with a good telephoto camera, but museums and universities still need material, especially of yet unknown or poorly known species, so the process of obtaining permits to collect specimens is well worth the effort. When all that is left is cabbage whites, a permit will probably not be required. If so, the butterflies may have to issue it.
LITERATURE CITED
EMMEL, T. C., M. C. MINNO & B. A. DRUMMOND. 1992. Florissant butterflies: A guide to the fossil and present-day species of central Colorado. Stanford University Press, California. x plus 118 pp., 9 color plates.
HEATH, J. 1970. Provisional atlas of the insects of the British Isles, Part 1: Lepidoptera (Rhopalocera, Butterflies). Biological Records Centre, Monks Wood Experimental
~I
VOLUME 48, NUMBER 1
Station, Abbots Ripton, Huntingdon, England, front matter plus index plus 57 full page maps.
LARSEN, T. B. 1990. The butterflies of Egypt. American University in Cairo Press. 112 pp., 8 color plates.
MILLER, J. Y. (ed.). 1992. The common names of North American butterflies. Smith- sonian Institution Press, Washington, D.C. ix plus 177 pp.
MONMONIER, M. 1991. How to lie with maps. University of Chicago Press. xi plus 176 pp.
RINDGE, F. H. 1965. Presidential Address 1965: The importance of collecting—Now. J. Lepid. Soc. 19(4):193-195.
ScoTT, J. A. 1993. The common names of North American butterflies (book review). J. Lepid. Soc. 47(2):170-171.
STANFORD, R. E. & P. A. OPLER. 1993. Atlas of Western U.S.A. butterflies, including adjacent parts of Canada and Mexico. Privately published. x plus 275 pp.
Received and accepted for publication 20 October 1993.
Journal of the Lepidopterists’ Society 48(1), 1994, 8-23
A NEW GENUS OF WINTER MOTHS (GEOMETRIDAE) FROM EASTERN CALIFORNIA AND WESTERN NEVADA
JERRY A. POWELL Essig Museum of Entomology, University of California, Berkeley, California 94720, USA
AND
DOUGLAS C. FERGUSON
Systematic Entomology Laboratory, ARS, USDA, Washington, District of Columbia 20560, USA
ABSTRACT. Tescalsia, a new genus of Geometridae, is described and assigned to the subfamily Larentiinae. It is represented by two new species: Tescalsia giulianiata Ferguson, known from 3 localities in and adjacent to the Owens Valley, Inyo Co., California, and T. minata Ferguson, from Mineral Co., Nevada. The female of T. giu- lianiata has linear, straplike forewings, vestigial hindwings, and long, slender legs that enable agile climbing in shrubs; the female of T. minata is unknown. Both sexes lack the proboscis and tympana. Adults of T. giulianiata are active at sundown and nocturnally in November and December, despite near freezing temperatures.
Additional key words: brachypterous, flightless, Larentiinae, tympana.
In 1976 Mr. Derham Giuliani, a keen naturalist of Big Pine, Cali- fornia, brought a brachypterous moth to Berkeley that he had collected in Deep Spring Valley, east of Big Pine, in early December 1978. So bizarre was the specimen, with peculiarly bristled, straplike forewings and vestigial hindwings and mouthparts, that Powell could not identify it to family. The taxonomic placement remained a mystery after Fer- guson and other lepidopterists at the National Museum of Natural History (NMNH) examined the specimen in 1977.
Two additional females were taken in pitfall traps at Deep Spring Valley in December 1978, one of which laid eggs that produced first instar geometrid larvae. After additional visits by Giuliani, Powell, and others during November and December 1978-82, we had assembled 9 winged males and 8 females, a sufficient sample to provide con- vincing circumstantial evidence for the association of the sexes. The males indicate that the mystery moth is an undescribed species best assigned to the Larentiinae (Geometridae).
Later, males of a congeneric species, from Mina, Mineral Co., Ne- vada, that had been collected in 1914, were discovered by Ferguson in unidentified material at the NMNH. Although descriptions of the new genus and species were written several years ago, we delayed publi- cation, anticipating that the larva and host plant might be discovered; but that hope has not been realized. Hence, we present the descriptions, and characterize the egg, together with observations on the habitats and adult behavior.
VOLUME 48, NUMBER 1 9
MATERIALS AND METHODS
Trapping method. The first female was observed on the sand, and a few males were netted near sundown; but most of the series was taken in pitfall traps. This technique is widely used for sampling ground- dwelling insects in various habitats and is particularly effective for nocturnal insects such as many beetles. Cups may be deployed empty, baited with truly disgusting materials, or partially filled with anti-freeze (ethylene glycol), which allows long-term sampling. If deployed empty, traps need to be checked frequently because many predators, spiders, scorpions, carabid beetles etc., are trapped.
We used 9 oz. squat plastic tumblers (7 cm deep and 9 cm diameter at the rim); they are inexpensive and nest in compact packages for transport. We deployed them in transect lines of 50-100, about 2 m apart, usually situated at the bases of shrubs. We trapped with empty cups during single nights (when two of three females were killed by a predaceous mite and spider) or with anti-freeze over a several day period. Specimens taken from the latter were washed in water, then transferred to alcohol, and later dried for pinning. Understandably, lepidopterists rarely are familiar with the method, but it is an effective one for brachypterous forms. Winged males are also sometimes trapped, and most of our male Tescalsia were collected this way. The holotype was taken in early morning in an empty trap about 50 m from the nearest trapped female.
SYSTEMATICS Tescalsia Ferguson, new genus
Type species: Tescalsia giulianiata Ferguson, new species.
Diagnosis. Characterized by the following combination of unusual characters: 1) in male, veins Sc and Rs fused for half length of hindwing, forking just before end of the very long cell, and beyond that point Rs and M, stalked together for “4 or % of distance from end of cell to outer margin; 2) female with greatly reduced forewing, linear and straplike, with numerous setae longer than width of wing along entire length of both fore and hind margins; 3) female with hindwing present only as a vestige concealed beneath base of forewing; 4) tympanic cavities in base of abdomen, characteristic of nearly all Geometridae, missing in both sexes; 5) proboscis missing in both sexes, as is true of some other winter moths.
Description. MALE: Length of forewing: 12.0-17.5 mm. Head: An- tenna bipectinate, extremely delicate, with slender shaft and long, wide- ly spaced, setose branches and large scape 3-4 times thickness of shaft;
10 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
shaft scaled dorsally. Labial palpus short, hardly extending beyond front. Eye not very large, its vertical length almost equal to width of front at narrowest point, but strongly protuberant in the type-species, less so in the other. Ocellus absent. Front protuberant, rounded. Chae- tosema normal, moderately to well developed, the two chaetosemata not extended transversely to meet behind head as in many Larentiinae. Tongue absent. Vestiture untufted. Legs slender, normal except that foretibia is extremely reduced and bears a large, conspicuous, double claw (Fig. 9), and foretarsus much more elongated than mid- or hind- tarsus; hindtibia with one or two pairs of spurs.
Fore- and hindwing somewhat elongated, produced especially toward apex; tornus of both wings rounded; forewing length/width ratio about 2.4/1.0, narrower than the 2.0/1.0 ratio of Operophtera (length = base to apex; width = length of line through anal angle meeting costa at 90°); wing pattern mostly diffuse and indistinct, variable between and within species; hindwing maculation differing from that of forewing in its reduced pattern consisting of discal spot only or discal spot and diffuse postmedial band only; fringes of both wings unusually long; wings covered with wide, tulip-shaped to almost round scales, dentate with 3-5 short points distally or simple. Venation as described and figured (Fig. 8); Sc of forewing free from radials; discal cells long, that of forewing more than half and of hindwing about half length of wing, closed off almost straight transversely at ends; forewing with two large accessory cells; hindwing with discal cell unusually wide because of long fusion of Sc and Rs, with M, a fully developed, tubular vein, and with lst A a well-developed fold, 2nd A a well-developed vein, and 3rd A entirely lost. Frenulum well developed and very long.
Male genitalia (Figs. 10, 11). Valva and uncus simple; uncus ap- pearing to be fused to tegumen, forming a solid unit; gnathos absent; transtilla distinct, forming slender, complete bridge; manica spinulate, heavily so in one species, flanked by pair of knoblike, setose processes, the derivation of which is unclear; each knoblike process with delicate, sclerotized connections both to juxta and base of costa of valva; juxta appearing to have a large, pointed, conical or thornlike medial process adjoining its posterior margin, seemingly apposed to end of aedeagus, and derivation of this structure also unclear. Vesica with clumps of small cornuti.
FEMALE (Fig. 14, type species only): Head: Antenna simple, slen- der, sparsely setose, with scape much smaller than that of male. Labial palpus small, not exceeding front. Eye nearly as large as that of male. Ocellus absent. Tongue absent. Front, chaetosema, and legs similar to those of male. Brachypterous; forewing 6-7 mm long, % to % length of body, narrow, straplike, tapering to a pointed end, roughly clothed
VOLUME 48, NUMBER 1 1
with scales narrower than those of male and with both margins fringed with long, straight, bristlelike setae slightly longer than width of wing; hindwing present as small vestige beneath base of forewing and bearing long setae distally. Venation (Fig. 12) very reduced but bearing three longitudinal elements that are probably the stems of Sc, R, and Cu.
Female genitalia (Fig. 13). Simple and without very significant fea- tures except an extremely long ostial cavity, with a pair of lateral sclerotized supports at base or in what may be a funnel-like posterior section of the ductus bursae, that on right side the larger; corpus bursae ovoid, membranous, delicate, without signum; ductus seminalis arising from bursa near ductus bursae. Sclerotized parts darkly pigmented.
Distribution. Known only from arid habitats of eastern California, and western Nevada.
Early stages. Eggs and first instar larvae were obtained, but larvae would not feed. Food plant unknown. Young larva typically geometroid, with no indication of a third pair of prolegs such as might indicate a relationship to the Alsophila group.
Remarks. Sattler (1991) has reviewed wing reduction in Lepidoptera and analyzed implications of flightlessness. Examples of brachyptery in female winter moths in the Northern Hemisphere are known in several families; they are particularly numerous in Geometridae (e.g., Alsophila, Oenochrominae; Phigalia, Ennominae; and Operophtera, Larentiinae). The taxonomic relationships of Tescalsia proved difficult to determine and are still not clear. A combination of such features as the double accessory cell, free subcosta, extremely long fusion of Sc + Rs in the hindwing, and condition of the anal veins, points to a probable connection with the Larentiinae. Because of the elongate discal cells, loss of the proboscis, and reduced tympanic cavities, the possibility of relationship to the Alsophila group (currently in the Oenochrominae but probably misplaced) was also considered. All species of Alsophila, as well as the similar Phthorarcha primigera Staudinger (Central Asia) and Inurois tenuis Butler (Japan), were examined. These agree with one another in venation, especially with respect to the anal veins of the hindwing consisting of a weak Ist anal fold and well-developed 2nd and 3rd anals. Tescalsia clearly differs in having a strong Ist anal fold, well-developed 2nd anal, but no 3rd anal. This would seem to relate Tescalsia to the Larentiinae, in which the Hydriomenini and Operoph- terini have anal veins of this type. Surprisingly, Phthorarcha has wide, rounded wing scales almost exactly like those of Tescalsia, although other members of the Alsophila group and Larentiinae examined do not.
The venation agrees best with that of the Larentiinae, although it is of an exaggerated type with two large accessory cells, elongated discal
12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
cells, and elongated fusion of Sc and Rs in the hindwing, and the almost straight, transverse closure of the ends of the cells is unusual. The genitalia, both male and female, seem closer to those of Operophtera than anything else, the male genitalia are quite like those of Operophte- ra in general form, in the shape of the valvae, and in the apparent homology of all components of the anellus. In female Operophtera, the beginnings of the same kind of large ostial cavity and short, sclerotized ductus bursae are apparent, and the simple, membranous corpus bursae, lacking a signum, agrees exactly. Operophtera has fairly well-developed tympanic cavities in the base of the abdomen, but their loss, together with the loss of the tongue, extremely wide pectinate male antennae, and curiously specialized female wings are adaptations to an extreme habitat. The large foretibial claw appears here and there in many groups of geometrids and noctuids, especially of desert habitats, and it has no important systematic significance. A palearctic geometrid that occurs in winter in Turkmenia and Kazakhstan, Cheimoptera pennigera Danil. (Danilevskiy 1969) shows many of the same features, including the foretibial claw and loss of the tympanic cavities, although it is unrelated to Tescalsia and believed to belong to the Ennominae. Tescalsia should, for the present, be assigned to the tribe Operophterini of the Laren- tiinae, although there are notable differences. For example, other species of Operophterini possess a reduced proboscis, large chaetosemata that meet in the middle in some instances, coarsely ciliate rather than bi- pectinate antennae in the male, lack the foretibial claw, and have either one accessory cell in the forewing (Operopthtera), or two (Epirrita) as in Tescalsia.
The superficially similar, gray, long-winged Chesiadodes morosata Hulst (Ennominae) flies in the same area near Lone Pine, Inyo County, California, in December, and also has a foretibial claw, although of different shape. It differs in having a proboscis. The two are not closely related, and the female of Chesiadodes has fully developed wings.
Tescalsia giulianiata Ferguson, new species
(Figs. 1, 2, 5-9, 11-14)
Diagnosis. Males large, forewing length 16.0-17.5 mm; wings gray with blackish markings; hindtibia with two pairs of spurs. Female as described for genus and illustrated.
Description. MALE: Head: labial palpus small, not surpassing front; eye of about same dimensions as that of T. minata but more protuberant, its form exceeding that of half a sphere; front bulging, roundly convex, with large, broad, gray-brown or whitish scales, tending to be oriented toward middle of front; eye rimmed anteriorly and ventrally with
VOLUME 48, NUMBER 1 13
Fics. 1-7. 1, Tescalsia Pliaihata Eur dteoe holstene male. 2, T. Balaniata para- type female sare data as holotype). 3, T. minata Ferguson, felon ee male; 4, T. minata, paratype male. 5-7, T. giulianiata, female (Deep Spring Valley, CA, XII-15-78): 5, perched on sand; 6, in repose balanced on wingtips; 7, climbing on Chenopodium branch.
contrastingly pale border of radiating whitish scales. Thorax beneath without long, hairlike scales; legs similar to those of T. minata except that femora lack long, hairlike scales, and hindtibia has two pairs of spurs. Wings whitish, dusted with blackish scales and thus appearing gray, although forewing sometimes so heavily suffused that markings are obscured; forewing normally with diffuse, slightly dentate or sinuous
14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
dark antemedial and postmedial bands, the former slightly convex and subparallel to postmedial, which is nearly parallel to outer margin; subterminal shade, if present, indistinct, similarly parallel to outer mar- gin; some veins, especially in medial area, faintly outlined with dark scales; black discal spot present; fringe white, checkered with dark brown and preceded by weak terminal line of diffuse dark spots. Hind- wing paler gray, almost unmarked except for small discal spot and diffuse but complete transverse band crossing just beyond middle of wing; fringe whitish, unmarked, preceded by faint, broken terminal line in some specimens. Underside with fore- and hindwing nearly alike, light gray with discal spots, diffuse postmedial bands, and variable, black, interrupted, terminal lines. Length of forewing: holotype, 17.0 mm; other 6, 16.0-17.5 mm. Genitalia (Fig. 11), differing from those of T. minata mainly in their wider, blunt or obtusely pointed medial juxtal process, less heavily spined manica, and smaller, paired, setose processes attached to bases of costal sclerite. Eighth sternite rectangular, not triangular, and eighth tergum without a posterior border of per- sistent scales.
FEMALE: As described for genus. Antenna simple, covered with whitish scales. Hindtibia with two pairs of spurs. Body gray, variably dusted with black scales; in paler specimens a pair of dark subdorsal spots toward posterior margins of abdominal segments 2-5. Legs gray brown with each tarsal segment pale-tipped. Forewing with light and dark scales almost evenly mixed and no other markings. Length of forewing: 5-6 mm. Genitalia (Fig. 18) as illustrated and described for genus.
Types. Holotype male and allotype female: CALIFORNIA, Inyo Co., Alabama Hills, 4 miles [5 airline km] southwest of Lone Pine, 4550’ [1390 m elev.], 6/7-XII-1982, in pitfall traps (J. De Benedictis & J. Powell); deposited in Essig Museum of Entomology, U. California, Berkeley. Paratypes (15), all Inyo Co., CA: 44, 3 2, same data as holotype, 7 /11-XII-1982 (D. Giuliani, De Benedictis, Powell); 1 6, Alabama Hills, N. fork Lubkin Cr., 4 mi. S, 1 mi. W of Lone Pine, 4800’, 11-XII-1982 (Giuliani); 1 2, Deep Spring Valley, 8-XII-1978, sand dunes (Giuliani); 2 2, same locality, 15/16-XII-1978, pitfall traps (Powell); 1 6, same locality, 17-XI-1980, flying at sundown (Giuliani); 2 6, same data, 17- XI-1982; 1 2, Owens Lake, mid XI to mid XII-1978, ethylene glycol pit trap, Atriplex-Franseria assoc. (Giuliani, F. Andrews, D. Hardy); paratypes deposited in California Dept. Food & Agric., Sacramento, Essig Museum, and U.S. National Museum of Natural History, Wash- ington, D.C.
Habitats. Tescalsia giulianiata is known from three sites in the Owens Valley region, California, which are similar in general aspects of veg-
VOLUME 48, NUMBER 1 15
sl mee
8
Fic. 8. Tescalsia giulianiata Ferguson, male, wing venation.
etation architecture, with a low scrub of scattered shrubs interspersed with open patches of sand, yet they differ in dominant plant species.
1) Deep Spring Valley is a closed basin at 1500-1600 m elevation, situated southeast of the White Mountains and surrounded by arid
16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
mountain ridges. To the west the valley is separated from the Owens Valley by Westgard Pass (2225 m) and to the east from the valleys of Nevada by Gilbert Summit (1950 m). From the highway along the north edge of the valley, the terrain slopes southward towards Deep Spring Lake, which is dry except following winter storms. Fine aeolian sand from the dry lakebed has been deposited to form low ridges and dunes that are stabilized by low scrub dominated by Atriplex confer- tifolia (Chenopodiaceae), Thamnosma montana (Rutaceae) and scat- tered patches of Chrysothamnus nauseosus (Asteraceae). Female moths were taken from open sand and in pitfall traps placed at the base of Atriplex and Thamnosma, about 1 km north of the lakebed.
The valley is about 300 m above the floor of Owens Valley, and it evidently acts as a basin for cold air drainage, with snow patches per- sisting much longer than at comparable elevations on the hills around Owens Valley. During our December visits, daytime temperatures of 10-18°C fell rapidly, to 4.5-5.0°C towards sundown and 1.6-4.5° by dusk.
2) The Alabama Hills are remnants of an ancient uplift that has eroded to low hills of decomposed granitic, alluvial sand subtending weathered, granite outcrops. The collecting site is situated at 1890 m elevation about 5 airline km southwest of Lone Pine, off Indian Springs Road. This area is characterized by gently sloping expanses of coarse, granitic sand stabilized by a low scrub consisting mainly of Tetradymia glabrata (Asteraceae), a low spiny Atriplex, and scattered Chrysotham- nus nauseosus. Most of the type series was taken at this site, December 7/11, 1982, in pitfall traps deployed December 6. Although the locality is only about 100 m lower than Deep Spring Valley, it isa much warmer habitat during winter.
3) The former Owens Lake was drained by the Los Angeles water district, beginning in the 1920’s, and has long been a dry lakebed. Aeolian sand is deposited in low ridges along its east margin. One female of T. giulianiata was taken here in a pitfall trap. Giuliani and F. G. Andrews had monitored the area by pitfall trapping, with 12 traps in each of 7 vegetation types for one year. The site that yielded the Tescalsia is about 3 km northwest of Keeler (1100 m). The traps were set east of a few low sandhills at the margin of the lakebed, in a low area that acts as a catch basin for the sparse rainfall runoff. It is char- acterized by shadscale scrub, alkaline tolerant species of Chenopodia- ceae, including Atriplex hymenelytra, the spiny Atriplex of the Ala- bama Hills, Allenrolfia occidentalis, and Sarcobatus vermiculatus, as well as Tetradymia glabrata. Thus the dominant vegetation is more similar to the Alabama Hills site than to Deep Spring Valley. We did not deploy additional pitfall traps at this locality after. 1978.
VOLUME 48, NUMBER 1 17
Diel rhythm. Temperatures in December in the Owens Valley area typically drop to 2-10°C below freezing at night but rise well above freezing during the day, often to 10-15°C. Nonetheless, our observations suggest that Tescalsia giulianiata is not diurnal and begins activity near sundown. Its period of activity may vary daily with a low temperature threshold and may exclude some evenings. Males were observed flying only twice, by Giuliani on November 17, 1980 and Nov. 17, 1982, near sundown. One female was found on open sand in late afternoon, and one was trapped in a pitfall between 1530-1615 PST (ca. 8—10°C; sunset at 1610), and another between 1630 and 1030 PST the following day, in temperatures below 5°C. Other individuals were trapped during longer intervals, including one male and a female at the Alabama Hills between 1230 and 0930 the following day. Mating was not observed.
On each of the four dates in December, 1977-82, we made continuous observations for 2-8 h, on days when temperatures rose from 0.5—2.7°C at 0930-1030 PST to 15°C in mid afternoon, dropping to 4.5-5.0°C by sundown. No males were seen, and just one female was trapped during these intervals, which included extensive pitfall trapping, beating of shrubs, sifting and net brushing of the sand. T. giulianiata evidently is not adapted to diurnal flight when temperatures are highest. We ran a blacklight trap just once; the temperature was 4.5°C at dusk, and not one insect was trapped.
One captive female survived more than 4 days, with lab temperatures at 10-14°C at night to 21°C diurnally. Activity periods were sporadic, but in absence of disturbance, she seemed most active during evening, 1800-2100 h, moved only slowly when viewed by flashlight at 0500- 0700, and rarely during morning hours.
During periods of activity, the female held the antennae and fore- wings at about a 45° angle to the plane of the body (Figs. 5-7). When quiescent, she positioned the antennae back along the body, and the wings were curled downward. On the sand surface, she seemed very awkward, scarcely able to ambulate forward or walk evenly. Walking on the sand, she dragged her abdomen, leaving linear tracks, but this did not seem to be a part of oviposition behavior. By contrast, when debris or the branchlet was encountered, she climbed quickly with agile movement from twig to twig and often hanging by one or two legs, reminiscent of a miniature orangutan. She frequently perched on the highest reach of the branchlet and moved quickly to maintain that position if the branch was moved. It seems that the species is adapted to life in shrubs rather than on the sand, which fails to explain why females were trapped in pitfalls but none could be beaten from shrubs. In fact, there may be strong selection against life on the sand at that time of year because predaceous mites and lycosid spiders appeared in
18 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics. 9-11. 9, Tescalsia giulianiata Ferguson, female, right prothoracic leg; small process arising near middle of double clawed tibia is a vestigial epiphysis. 10, T. minata Ferguson, male genitalia; a, aedeagus. 11, T. giulianiata, male genitalia; a, aedeagus.
VOLUME 48, NUMBER 1 19
33-50% of our pitfall traps. Two of the female T. giulianiata fell victim to these predators overnight, when we used dry cups.
Whether on sand or on twigs, the fore tarsi were extended, so that the tibial hooks were well back from the substrate; evidently they are not employed in adult locomotion. Upon disturbance, the female con- sistently feigned death, falling on her side with all legs retracted, and remained so for 10—20 seconds. Also, at times the female assumed a repose stance, balancing on her wingtips and curled abdomen, with the legs partially folded (Fig. 6). This may have been an abnormal behavior in confinement, but it was repeated several times for lengthy periods.
Oviposition. The single female retained alive was housed in a plastic box with sand and a dry branch of Atriplex (JAP 78M1). A cotton wick was provided, but there was no indication that the female ever imbibed moisture or touched the cotton with her vestigial mouthparts. She de- posited 10 eggs in captivity, none during the first 48 h, 7 in the 8rd 24-h period, and 3 the following day. Eggs were deposited between 0940-1000 PST (1), 1055-1125 (1), 1400-1700 (6), and one later. Most were placed on loose sand, but the female was not observed to probe the sand with her abdomen. Two eggs were deposited on the damp cotton wick after the female had been coaxed onto it, and one egg was affixed to a dry Atriplex leaf.
Cuming (1961) reported the activity periods of the related larentiine winter moth, Operophtera brumata (L.), to be nocturnal. Caged moths in outdoor conditions mated between 1730-2305 at temperatures of —0.8 to +12.2°C, and females oviposited between 1515-0020 at —1.6 to +12.2°C. T. giulianiata displays a comparable diel rhythm and temperature tolerance.
Eggs. The eggs (n = 10), which were deposited singly, were oblong, slightly variable in shape, 0.80-0.90 mm long x 0.60 mm wide x 0.48 mm thick, somewhat flattened on the side away from the substrate. The chorion was opaque, white, very weakly rugose, appearing smooth under low magnification. When first deposited the eggs were pale greenish, similar in color and shape to new leaves of Atriplex confer- tifolia.
The eggs were confined in individual gelatin capsules; five were placed in a tightly covered plastic container in refrigeration each night (ca. 8:16 h, 1.6°: 15-20°C), while the remainder were retained at lab temperatures (10—20°C), for 30 days. During development, eggs dark- ened and most showed a depressed area away from the micropylar end as though collapsing. Larvae began eclosing in the refrigerated sublot January 21, after 34 days; the others were refrigerated 8:16 h from day 36 to 46, and first instar larvae emerged February 2-4, after 46-48 days.
20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Yyf
S
Md
Fics. 12-18. 12, Tescalsia giulianiata Ferguson, female, forewing showing setation of margins and rudimentary venation. 13, T. giulianiata, female genitalia.
First instar larvae. The eclosing larva ate a ragged, round hole at the micropylar end but did not continue to feed on the chorion. Each was placed in a small cup with synthetic diet (modified Shorey Diet used for Choristoneura) and a sprig of fresh Atriplex (but not A. con- fertifolia) from the U. C. Botanic Garden. None fed. After two days,
VOLUME 48, NUMBER 1 | WAI
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YR
aS aS = aw
SQ Sant A i
=e ww SS
Fic. 14. Adult female of Tescalsia giulianiata.
some larvae also were offered a sprig of Prunus or Salix, but by that time they likely were too weak to feed. All died by the 3rd day after eclosion.
If failure to accept synthetic diet and the plants provided indicates a narrow host specificity, we cannot suggest a probably food plant. In retrospect, after observing the three habitats from which the flightless females were taken (just the one site was known at the time the eggs were obtained), Atriplex is the most plausible of the dominant, woody shrubs, with one or more species occurring in all three habitats. Chrys- othamnus, which was present only in scattered patches, or some her- baceous perennial are possible alternative candidates.
Tescalsia minata, Ferguson, new species
(Figs. 3, 4, 10)
Diagnosis. Males small, forewing length 12-15 mm; wings brown with mostly indistinct, darker brown markings; hindtibia with one pair of spurs. Female unknown.
22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Description. MALE: Head: labial palpus slender and delicate, but with its terminal scales clearly surpassing front; eye about as in the other species but much less protuberant, its shape distinctly less than that of half a sphere; front somewhat protuberant but flattened rather than roundly convex, with large, broad scales tending to be oriented toward middle of front; eye rimmed anteriorly and ventrally with brown scales concolorous with those of front, not contrasting. Thorax beneath and tegulae above sparsely clothed with long, brown, hairlike scales; legs similar in the two species except that femora of T. minata have long, hairlike scales, and the hindtibia only one pair of spurs. Wings gray brown with variable, darker brown markings; forewing with antemedial band often strongly convex and enclosing a paler area toward base; faint basal band also may be present; postmedial not always distinct but, if present, not parallel to outer margin, more curved, concave in posterior half of wing and convex in costal half; a vague, pale, subterminal band may be present; dark-brown discal spot present but weak; fringe brown, concolorous with wing, unmarked; terminal line wanting. Hindwing brown, hardly paler than forewing, unmarked except for weak discal spot; fringes concolorous. Underside much like upperside except that a faint, diffuse, convex, transverse band may cross just beyond discal spot, and veins on underside of hindwing may in part be faintly outlined with darker brown scales. Length of forewing: holotype, 18 mm; other 6, 12-15 mm. Genitalia (Fig. 10), similar to those of T. giulianiata; the most obvious difference is in the shape of the large medial process of juxta. In T. minata it has an abruptly acuminate, thornlike shape, with a sharp-pointed apex. Spines on man- ica more numerous and larger, and paired, setose processes flanking manica and adjoining base of costal sclerite of valva are larger. Eighth sternite small and triangular rather than quadrate, and eighth tergum with wide, dense tuft of short, persistent scales along its posterior mar- gin, not easily removed in dissection.
FEMALE: Unknown. Types. Holotype 6, Mina [Mineral County], Nevada, November 17, 1914, A. Wetmore. Paratypes: 4 4, same data. Type series in collection
of U.S. National Museum of Natural History, Washington, D.C.
ACKNOWLEDGMENTS
The discovery of this genus and subsequent important collections were made by Derham Giuliani, of Big Pine, California, who has contributed considerably to our knowledge of the insects of the interior deserts in California. F. G. Andrews, California State Department of Food & Agriculture, Sacramento, provided support for pitfall trap sampling by Giuliani. J. A. De Benedictis, now at University of California, Davis, J. T. Doyen, J. K. Liebherr, now at Cornell University, Ithaca, New York, endured winter field trips while at Berkeley. The genitalia and wing venation drawings were done by Linda H. Lawrence, Staff
VOLUME 48, NUMBER 1 23
Illustrator, Systematic Entomology Laboratory; the habitus drawing by Tina Jordan, U. California, Berkeley. The photographs of specimens were made by Ferguson and those of living moths by Powell.
LITERATURE CITED
CUMING, F. G. 1961. The distribution, life history, and economic importance of the winter moth, Operophtera brumata (L.) (Lepidoptera, Geometridae) in Nova Scotia. Canad. Entomol. 93:1385-142.
DANILEVSKI, A. S. 1969. Two remarkable new species of winter moths from the deserts of Soviet Central Asia: Dasyethmia hiemalis, gen. et sp. n. (Lepidoptera, Ethmiidae), and Cheimoptera pennigera, gen. et sp. n. (Lepidoptera, Geometridae). Entomol. Rev., Washington, D.C. 48:176-191.
SATTLER, K. 1991. A review of wing reduction in Lepidoptera. Bull. Brit. Mus. Nat. Hist. (Entomol.) 60:243-288.
Received for publication 5 June 1993; accepted 31 July 1993.
Journal of the Lepidopterists’ Society 48(1), 1994, 24-45
SPLIT SKIPPERS: MEXICAN GENUS POANOPSIS GOES IN THE ORIGENES GROUP—AND YVRETTA FORMS THE RHESUS GROUP—OF POLITES (HESPERIIDAE)
JOHN M. BuRNS
Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, District of Columbia 20560, USA
ABSTRACT. Genitalia show that the montane Mexican genus Poanopsis Godman fits snugly within the origenes group of Polites Scudder and that Yoretta Hemming constitutes a new species group of Polites. Relationships have been masked in part by evolutionary reduction and loss of various characters. The origenes group, with the northern superspecies Polites mystic (P. mystic (Edwards) + P. sonora (Scudder)), the intermediate linking species P. origenes (Fabricius), and the southern sister species P. puxillius (Mabille) new combination and P. pupillus (Plétz) new combination, extends from the width of southern Canada to southern Mexico. The rhesus group of Polites, with P. rhesus (Edwards) and the sister species P. carus (Edwards) and P. subreticulata (Pl6tz) (new combinations, all), extends from southern central Canada to Panama. Geo- graphic distributions of species within these groups (and in the themistocles and vibex groups of Polites, as well) still strongly reflect allopatric speciation.
Additional key words: genitalia (male and female), variation, generic limits, character reduction and loss, speciation (allopatric).
In dealing with biodiversity, systematists divide and conquer. For- mally described divisions assume a life of their own. The longer they live and the more we use them, the sounder they seem. Sometimes we get so accustomed to overly fine divisions that we cannot see the forest for the trees. The American genus Polites Scudder is one such forest, a good bit larger and denser than we thought.
Once upon a time, Godman (1900) created Poanopsis for a small, brown, high montane, Mexican skipper, Pamphila puxillius Mabille, which he made the type of his monotypic new genus. He observed that puxillius, with its short, broad wings, looks like massasoit Scudder, the type of the genus Poanes Scudder (hence the name Poanopsis), but differs conspicuously in having an oblique stigma on the forewing of the male from the origin of vein 3 to vein 1. Godman (1900:pl. 98, fig. 40—my Fig. 1) illustrated the male genitalia of Poanopsis puxillius without comment. In mid century, Evans (1955) added a similar Mex- ican skipper, pupillus Plotz (=catahorma Dyar), to Poanopsis, carica- tured the male genitalia of both species, and set Poanopsis next to Poanes, eight genera removed from Polites. However, genitalia show that both species of Poanopsis belong in one of the species groups of Polites.
Polites is a familiar, widespread, New World genus, stretching from Canada to central Argentina, from Atlantic to Pacific coasts in both Americas, and through much of the West Indies. On the basis of genitalic
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Fic. 1. Male genitalia of Polites puxillius (minus left valva) in left lateral view (ex Godman 1900:pl. 93, fig. 40). This figure presents the inner surface of the right valva whereas those that follow present the outer surface of the left valva. The specimen is “from the high tablelands of MEXICO.”
and stigmal characters, MacNeill (1993) recognized four subdivisions: the themistocles group, the origenes group, the vibex group, and Polites baracoa (Lucas). He pointed out that, within groups (the themistocles group in particular), genitalia may be so conservative and so individ- ually variable that differences between species blur but that, paradox- ically, superficial color pattern will distinguish some of the genitalic look-alikes.
THE ORIGENES GROUP OF POLITES
The origenes group contains the eastern and central North American Polites origenes (Fabricius) plus the continent-spanning and largely allopatric sisters P. mystic (Edwards) and P. sonora (Scudder), all of which show some internal differentiation. From an evolutionary per- spective, P. mystic and P. sonora can be considered a superspecies (see Stanford & Opler 1993:74, 75 for maps that more or less reflect the mutual geographic replacement of these two similar species, and see Burns 1964, 1983 for detailed analyses of superspecies in other American skippers). The origenes group ranges from Canada to the southern United States and, disjunctly, to higher elevations in the Sierra San Pedro Martir of Baja California Norte, Mexico. Now, with Polites pux- illius new combination and Polites pupillus new combination, it jumps from three to five obvious species and from the southern United States to southern mainland Mexico. Though these five species vary greatly in facies, they are genitalically close, especially in males.
To be sure, male genitalia are grossly similar throughout the genus Polites (see figures in Scudder 1889, Skinner & Williams 1924 [or Lind- sey et al. 1931], MacNeill 1993). Of all parts, the aedeagus is most generally useful in clustering species. Its accessories at once stamp the origenes group: the paired, toothed titillators are large, boldly dentate plates, something like semicircular saws (Figs. 1-5), extending down- ward from the lower part of the flared, distal end of the aedeagus (Figs. 1, 3, 5); and the lone cornutus is a rolled scroll bearing a crown of thorns (Figs. 2-5). In all other Polites the paired, toothed titillators assume
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Fics. 2,3. Male genitalia of Polites puxillius from 28 mi (45 km) E EI Salto, 8400 ft (2560 m), Durango, MEXICO, 6-7 August 1972, Viers & MacNeill [genitalia dissection number X-2652] (USNM). Scale = 1.0 mm. 2, Tegumen, uncus, tip of gnathos, both valvae, both titillators (each with four teeth), and the single cornutus (like a rolled scroll bearing a crown of thorns)—all in posterior view; 3, Complete genitalia (minus right valva, right titillator, and juxta) in left lateral view, with vesica everted.
—
Fics. 4,5. Male genitalia of Polites pupillus from Sierra de Guerrero, MEXICO, July 1913, R. Miller [X-2699] (USNM) (type of Amblyscirtes catahorma Dyar). Scale = 1.0 mm. 4, Tegumen, uncus, tip of gnathos, both valvae, both titillators (left with 10 teeth, right with 8), and the single cornutus (like a rolled scroll bearing a crown of thorns)— _ all in posterior view; 5, Complete genitalia (minus right valva, right titillator, and juxta) in left lateral view, with vesica everted.
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very different shapes, come from the upper (rather than lower) part of the distal end of the aedeagus, and, if they extend anywhere, extend backward (not downward); decorated cornuti number two or three (instead of one) and suggest thorny scrolls only in the vibex group (where the aedeagus sports a unique, midventral, caudally-directed prong). The distal end of the uncus, which is divided, is bent farther dorsad in the origenes group (Figs. 1, 3, 5) than in any other group of Polites. Female genitalia in Polites present less of a generic gestalt, varying more both within and between groups—and, to some degree, in par- allel—so that group characterization is not as simple. In the origenes group (also in the vibex group, but in no others) the apophyses anteriores join the lamella postvaginalis via continuously sclerotized bands (Figs. 6-9). In the origenes group (also in P. baracoa) sclerotization of the lamella postvaginalis is intermediate (P. puxillius [Figs. 6, 7], P. pupillus [Figs. 8, 9], and P. origenes) to extensive (superspecies P. mystic) rather than limited (as it is in the themistocles group [MacNeill 1993] and in the vibex group [where the lamella postvaginalis carries a unique, mid- ventral, short, sclerotized, finely spined keel]). In the origenes group sclerotization of the ductus bursae is partial and posterior; but it may be ventral only (superspecies P. mystic), ventral and lateral (P. pupillus [Figs. 8, 9]), or ventral, lateral, and, in a very narrow band, dorsal (P. puxillius [Figs. 6, 7], P. origenes, and, at times, vaguely, P. pupillus)— always with a longitudinal, midventral groove (Figs. 6-9) or break in the sclerotization. (Sclerotization of the ductus bursae is partial, pos- terior, but 360° and broadly ringlike [though midventrally weak] in the vibex group; partial, lengthy, mostly ventral but also somewhat lateral [resembling a long, essentially ungrooved, scoop] in P. baracoa; and complete, extending more or less the entire length of the ductus bursae and running all the way around, usually with a strong middorsal groove and a ventral pouch, in the themistocles group [MacNeill 1998].) While the northern major differentiates of the origenes group of Polites (P. origenes and superspecies P. mystic) are widely known, repeatedly shown in the rising tide of North American butterfly books, and readily separable superficially, the southern differentiates (P. pux- illius and P. pupillus) are not and must be treated here. Whereas in both sexes of mystic and sonora and in males of origenes extensive yellow to yellow-orange brightens the dorsal wing surface, in females of origenes and in both sexes of puvxillius (Figs. 23, 24) and pupillus (Figs. 25, 26) the wings above look basically brown (with a variable set of small, pale spots [yellowish in origenes, white in puxillius and pupil- lus]). Wingshape is sexually dimorphic in the northern three species but not in the southern two: in mystic, sonora, and origenes the wings of males are narrower and more pointed, those of females, broader and
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rounder; but in puxillius and pupillus the wings are broad and round in both sexes (Figs. 23-26). So, with respect to both color and wingshape, puxillius and pupillus tend to resemble females of origenes.
In the two Mexican species, wing spots, which are white, are expressed better by pupillus (Figs. 25, 26) than by puvxillius (Figs. 23, 24). Fore- wing spots of pupillus look whiter and brighter partly because, in some to most of them, some scales stand up from the surface of the wing so as to let light through. In puvxillius all the white scales lie flat against the wing so that all spots are opaque, none hyaline. Usually, pupillus develops a full set of forewing spots—in spaces 1b, 2, 3, 4, 5, 6, 7, 8, and the cell (Figs. 25, 26)—but puxillius rarely does, almost always dropping the one or two spots in space 1b (Figs. 23, 24), often skipping those in spaces 4 and 5, as well as that in 8 (Fig. 23), and, in one male examined, losing all but spots 3 and 6. At least some hindwing spots (which are opaque) appear dorsally in pupillus (usually spots 2 to 6) (Figs. 25, 26) while none really do in puxillius (Fig. 23) (at most, there may be a suggestion of a spot in space 3—see Fig. 24). Ventrally, where spot development is better, up to a full set may surface in pupillus— in spaces lc, 2, 3, 4, 5, 6, 7 (rare) and the cell (Figs. 25, 26); in puxillius ventral expression runs the gamut from all except the rare spot 7 (Fig. 24) to nothing whatsoever.
As for real interspecific differences in the conservative but individ- ually variable male genitalia, titillator teeth are fewer in puxillius (Figs. 1-3), varying from 4 to 6 (usually 4) on each of the paired titillators, and commoner in pupillus (Figs. 4, 5), where they vary from 6 to 10 (usually 7 or 8). The valva of puxillius, in lateral view (Fig. 3), is more tapered distally along its ventral margin and hence not as broad (from top to bottom) at its distal end as it is in pupillus (Fig. 5). (Evans does not mention these differences; yet once you know what to look for, you can detect both of them in his [1955:pl. 78, M.21.1, M.21.2] genitalic cartoons.) In posterior view, the lower part of the divided distal end of the valva usually forms more of a U in puxillius (Fig. 2) than it does in pupillus (Fig. 4). Though conspicuous in the specimens figured, this difference almost vanishes when certain variants are compared. The greater valval height of pupillus, evident in lateral view, usually shows too in posterior view as a ventral, vertical extension below the U (com- pare Fig. 4 with Fig. 2).
One interspecific difference in the highly variable female genitalia crept into the foregoing group comparisons: the incomplete scleroti- zation of the ductus bursae (mainly ventral and lateral) continues dor- sally to form a narrow but unmistakable sclerotized ring around the ductus in puxillius (Figs. 6, 7) but not—or, at best, very weakly—in pupillus (Figs. 8, 9). In addition, the sides of the midventral groove in
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Fics. 6, 7. Female genitalia of Polites puxillius from 25 mi (40 km) W Durango, Durango, MEXICO, 5 August 1972, MacNeill & Viers [X-2653] (USNM). Scale = 1.0 mm. 6, Ovipositor lobes (retracted), eighth tergite with apophyses anteriores (sclerotically connected to the lamella postvaginalis of the sterigma), sterigma, and bursa copulatrix in ventral view; 7, The same, plus the right apophysis posterioris and part of the ductus seminalis, in right lateral view.
the ductal sclerotization are more divergent anteriorly in puvxillius (Fig. 6) than in pupillus (Fig. 8), and the sclerotization itself is more fluted in puxillius (Figs. 6, 7) than it is in pupillus (Figs. 8, 9). The lateral part of the ductal sclerotization (which is more extensive on the left side than on the right—see Figs. 7, 9) is more extensive on both sides in pupillus than it is in puxillius; so it better hides the roof of the
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Fics. 8,9. Female genitalia of Polites pupillus from 34 mi (55 km) SE Acatlan, 6000 ft (1830 m), Puebla, MEXICO, 9 July 1952, E. E. Gilbert, C. D. MacNeill [X-3198] (MacNeill collection). Scale = 1.0 mm. 8, Ovipositor lobes (exserted) with apophyses posteriores, eighth tergite with apophyses anteriores (sclerotically connected to the lamella postvaginalis of the sterigma), sterigma, and bursa copulatrix in ventral view; 9, The same, plus part of the ductus seminalis, in right lateral view.
32 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
midventral groove in lateral view in pupillus (Fig. 9) than in puxillius (Fig. 7). Again in lateral view, the roof of the midventral groove is convex in puxillius (Fig. 7) but straight in pupillus (Fig. 9). All these interspecific differences are rather subtle.
Polites pupillus is larger than P. puxillius. In each species the fore- wing of the female averages about one mm longer than that of the male, and in each sex the forewing of pupillus averages about one mm longer than that of puxillius:
SEX mean range n puxillius 3 13.6 12.6-14.4 9 2 14.9 14.0-15.6 13 pupillus 3 14.8 13.5-16.2 16 2 ony 15.0-16.4 OD
The number of segments in the nudum of the antenna ranges from 11 to 13 in puxillius (mean = 12.2, n = 21) and from 11 to 14 in pupillus (mean = 12.8, n = 20). In both species nudum segments are evenly split between the body of the club and the apiculus.
Although both species are montane, P. puxillius is higher, altitudi- nally tighter (recorded from 8000 to 8500 ft [2440-2590 m]), and less widely distributed (Durango [Fig. 10]—to as far south as Guerrero according to Godman 1900, Hoffmann 1941). Polites pupillus extends from 4200 to 7500 ft (1280-2285 m) and from Sonora and Sinaloa to Colima, Puebla, Guerrero, and Oaxaca (Fig. 10). So far as known, flight times are similar: dates on specimens of puxillius examined run from 18 July to 11 August; on pupillus, from 3 July to 8 August.
THE RHESUS GROUP OF POLITES
The rhesus group of Polites comprises at least three species (usually treated as two) that collectively range from the grasslands of southern Saskatchewan and Alberta, Canada, through the western Great Plains, southern Rocky Mountains, and southwestern United States, through Mexico and Central America, to the middle of Panama. Although God- man (1900:474) noted that “the structure of the genitalia of the males is very similar in the two species’ when he put them in his new genus Chaerephon, he did not see the great similarity between their genitalia (Godman 1900:pl. 93, figs. 4 and 7—my Figs. 15, 16) and those of what he called Thymelicus vibex (pl. 93, fig. 14) and Poanopsis puxillius (pl. 93, fig. 40—my Fig. 1), which were on the same plate and which now are both in Polites. Likewise, Skinner and Williams (1923) figured the
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\ i) i | \
1 a, —™. Vay Sn ~L,
400 MILES
300 500 KILOMETERS
Fic. 10. Geographic distribution (based on plottable material examined) of Mexican sister species Polites puxillius (triangles) and P. pupillus (dots).
male genitalia of the two US. species of Chaerephon and then (1924) those of nine U.S. species of Polites without seeing how remarkably close they are. Enamored of names, Hemming (1935) saw that Chae- rephon Godman is a junior homonym and replaced it with Yvretta. Commenting that its “genitalia [are] very like those of Polites,’ Evans (1955:315) still set Yuretta next to Hylephila, six genera removed from Polites (and 21 genera from Poanopsis). MacNeill (1993:177) observed that “male genitalic similarities suggest that the nearest relatives of Polites are the genera Yuretta Hemming, Hylephila Billberg, and Wal- lengrenia Berg.” I am going much further by reducing Yoretta to a new species group of Polites, with the species Polites rhesus (Edwards), P. carus (Edwards), and P. subreticulata (Pl6tz) (new combinations).
Like all other elements of the male genitalia, the aedeagus in the rhesus group has that Polites look: anteriorly narrow, it at least doubles in size (in lateral view) with the entry of the ductus ejaculatorius and more or less flares at the elaborate distal end, which flaunts a pair of
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Fics. 11-18. Male genitalia of Polites subreticulata from Coatepec, Veracruz, MEX- ICO, October 1910, R. Miller [X-3499] (USNM). Scale = 1.0 mm. 11, Tegumen and uncus (gnathos hidden) in dorsal view; 12, Complete genitalia (minus right valva and juxta) in left lateral view (left titillator more dorsal and more tightly dentate than right), with vesica everted showing all three cornuti (two dentate [upper one curved, lower one straight] and one without teeth); this individual somewhat malformed anteriorly: saccus curved sharply upward, aedeagus bent to right, and bottom of tegumen not curved ventrad near juncture with vinculum (compare Fig. 14); 13, Aedeagus in dorsal view (left titillator more tightly dentate than right), with vesica oieted (only the two dentate cornuti visible).
toothed titillators as well as toothed cornuti in the vesica (Figs. kj 3s0% 12-16, and figs. 11-19 in MacNeill 1993).
In the rhesus group, the paired, toothed titillators come from the upper part of the distal end of the aedeagus, extend backward with
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Fic. 14. Male genitalia (digital image) of Polites rhesus from Sapillo Creek Valley, 5800 ft (1770 m), 12 mi (19 km) N Pinos Altos, Grant Co., New Mexico, USA, 28 May 1959, J. M. & S. N. Burns [X-2712] (USNM). Complete genitalia in left lateral view (right titillator out of focus and toothless cornutus hidden), with vesica everted showing both dentate cornuti (upper one curved, lower one straight).
one twist along their narrow length, and then expand distally into the dentate portion, with 3 to 7 teeth (usually 5) on the left titillator and 4 to 7 teeth (usually 4, 5, or 6) on the right (Figs. 12-16). The teeth are closer together on the left titillator than they are on the right one (Figs. 12, 13). In another asymmetric touch, the left titillator is more dorsal than the right one (Fig. 12). Like the number of teeth, the length of the titillators varies individually instead of interspecifically or geo- graphically: in each species of the rhesus group, the titillators may be equal (Figs. 12, 13, 15), or the left one may be a little to a lot shorter than the right. Two of the three cornuti are conspicuous and tentlike— one essentially straight, the other slightly to very curved, each with 3 to 6 teeth, which are closer together on the straight cornutus than they are on the curved one (Figs. 12-14). The third cornutus is very incon- spicuous, amounting to nothing but a bit of lightly sclerotized vesica (Fig. 12).
Right down to the titillators and cornuti, the aedeagus of the rhesus group recalls that of the themistocles group (compare Figs. 12-16 with figs. 11-19 in MacNeill 1993 and see MacNeill’s diagnosis of the themis- tocles group). Some variation is less rampant in the themistocles group, even though that group is larger: in P. themistocles (Latreille) itself, the two titillators are always the same length (fig. 19 in MacNeill 1993); and in what I consider the peckius subgroup comprising P. peckius (Kirby), P. sabuleti (Boisduval), P. norae MacNeill, P. draco (Edwards), and P. mardon (Edwards), the left titillator is always much shorter
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Fics. 15, 16. Male genitalia of rhesus group Polites (minus left valva) in left lateral view (ex Godman 1900:pl. 93, figs. 4, 7). 15, Polites subreticulata from MEXICO or GUATEMALA (aedeagus artificially rotated so that the left titillator is lower than the right whereas the reverse is really true); 16, Polites rhesus from Milpas, 5900 ft (1800 m), Durango, MEXICO (tegumen plus uncus artificially twisted so that the underlying gnathos and the gap between it and the uncus fail to show).
than the right (figs. 11-18 in MacNeill 1993). Throughout the themis- tocles group, the third, lightly sclerotized cornutus (inconspicuous in the rhesus group) is elongate and suggestive of a scouring pad (with a surface that MacNeill [1998] calls “minutely scobinate’’). Otherwise, the basic forms and arrangements of aedeagal accessories are closely similar in the rhesus and themistocles groups (small differences will be evident in the comparison of figures—but a few figures cannot adequately convey individual variation, which tends to weaken such differences). In contrast, forms and arrangements of aedeagal acces- sories differ sharply (and in different ways) in the origenes group (Figs. 1-5), the vibex group, and P. baracoa.
What aspects of the male genitalia set the rhesus group apart? In Polites the distal end of the uncus is divided: each of the two uncal tips terminates in a two-layered comb, with extremely close-set. tines curving backward and downward in each layer. These ““uncal combs” (called “‘pectines’” by MacNeill 1993) are slightly enlarged in the rhesus group (Figs. 11, 12, 14, 16), intermediate in the themistocles and ori- genes (Figs. 2-5) groups, reduced in the vibex group, and absent in P. baracoa. There is a sizable gap between the distal end of the uncus and the underlying divided gnathos in the rhesus (Figs. 12, 14, 15), origenes (Figs. 1, 8, 5), and vibex groups, and in P. themistocles (fig. 10 in MacNeill 1993)—but not in other members of the themistocles group (i.e., the peckius subgroup), where the gnathos is up against the bottom of the uncus (figs. 2-9 in MacNeill 1993). There is no gap in P. baracoa, either, but for a totally different reason: this species has lost the un- derlying divided gnathos.
In Polites the dorsodistal corner of the valva is split by a small notch (Figs. 3, 5, 12, 14-16, and figs. 2-10 in MacNeill 1993): the dorsal margin of the valva immediately anterior to this notch supports a dense field of more or less dorsally-directed bristles; and the posterior margin of the valva immediately posterior to this notch displays (in lateral view) at least one sizable, dorsally- or dorsocaudally-directed tooth
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which, in turn, joins one or more teeth (readily visible in posterior view [Figs. 2, 4]) that extend mediad beneath the more or less mediaily- expanded posterior edge of the dense field of bristles. Medial expansion of the bristled area is least in the vibex and themistocles groups, strong in the origenes group, and greatest in P. baracoa and the rhesus group. In lateral view this expansion produces a slight hump (in the origenes group, Figs. 3, 5) to a very perceptible hump (in P. baracoa and the rhesus group, Figs. 12, 14) on the dorsal margin of the valva anterior to the valval notch. The posterior edge of the dense field of bristles is without teeth in the rhesus group, finely dentate in the themistocles group, more coarsely dentate in P. baracoa and the origenes group (Figs. 2, 4), and most coarsely dentate in the vibex group.
About halfway down the posterior margin of the valva (in lateral view) a massive projection extends backward in P. baracoa and one or more sizable teeth point backward in the rhesus group (Figs. 12, 14- 16). This portion of the posterior margin is finely dentate (Figs. 3, 5) to smooth in the origenes group and essentially smooth in the vibex and themistocles groups (figs. 2-10 in MacNeill 1998).
The one or more teeth—visible in posterior view (Figs. 2, 4)—that extend mediad beneath the posterior edge of the dense field of bristles, are conspicuously multiple in the origenes group (Figs. 2, 4), in the vibex group, in P. baracoa, and in P. themistocles but are single to inconspicuously multiple especially in the peckius subgroup but also, somewhat less consistently, in the rhesus group.
The posterior margin of the valva (in lateral view) has more of a “chin” —so looks more squared off—in the rhesus group (Figs. 12, 14- 16) than in any other group of Polites. Certain members of the origenes group (P. mystic, P. sonora, P. pupillus [Fig. 5], and some individuals of P. origenes) come closest to this well-chinned look. Farthest from it, in a sense, is the vibex group because the posterior half of the ventral margin of the valva is narrowly excised.
Female genitalia in the rhesus group (Figs. 17-22) are broadly rem- iniscent of those of the origenes group—particularly P. origenes, P. puxillius, and P. pupillus (Figs. 6-9)—except that the apophyses an- teriores are not sclerotically connected to the lamella postvaginalis. Sclerotization of the lamella postvaginalis is intermediate, as in P. or- igenes, P. puxillius, and P. pupillus. Sclerotization of the ductus bursae is partial—mostly ventral and lateral, with a narrow, dorsal extension anteriorly—in P. rhesus (Figs. 17, 18) and P. carus (Figs. 19, 20), much as in P. origenes, P. puxillius (Figs. 6, 7), and P. pupillus (Figs. 8, 9); but it is virtually complete in P. subreticulata (Figs. 21, 22). In all three species of the rhesus group, as in all members of the origenes group, the ductus bursae has a ventral, longitudinal groove or break. This
38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fics. 17, 18. Female genitalia of Polites rhesus from Denver, Colorado, USA [X- 3490] (USNM). Scale = 1.0 mm. 17, Ovipositor lobes (exserted) with apophyses posteriores, eighth tergite with apophyses anteriores (not sclerotically connected to the lamella post- vaginalis of the sterigma), sterigma, and bursa copulatrix in ventral view; 18, The same, plus part of the ductus seminalis, in right lateral view.
groove is to the right of center in the rhesus group (Figs. 17, 19, 21) instead of more or less midventral as it is in the origenes group (Figs.
6, 8). |
Owing to their overall conservatism and their individual variation,
VOLUME 48, NUMBER 1 39
the male genitalia of the rhesus group are not diagnostic at the specific level. But here, as in the themistocles group, external color pattern separates species: the underside of the hindwing instantly sets P. rhesus (Figs. 27, 28) apart from P. carus (Figs. 29, 30) and P. subreticulata (Figs. 31, 32). These last two species are much more similar in pattern (so much so that they are mistakenly regarded as subspecies). Color may help in determining unworn specimens: the spots of the upperside are usually creamy to pale yellow in P. carus and light to medium yellow-orange in P. subreticulata. Although the undersides tend to be creamier in carus, yellower in subreticulata, they overlap considerably.
It is the female genitalia that best distinguish these species. The ductus bursae is only about half sclerotized (ventrally and laterally) in P. carus (Figs. 19, 20) but almost fully sclerotized in P. subreticulata (Figs. 21, 22). In both, the sclerotized ductus bursae usually looks rather globular in ventral view (Figs. 19, 21); but in P. subreticulata it tends—dorso- laterally—to extend backward slightly (and usually to flare slightly) at the ostium bursae (Fig. 21). Genitalic differences between P. carus and the superficially distinct P. rhesus are much more subtle because in rhesus, as in carus, the ductus bursae is only about half sclerotized (ventrally and laterally) (Figs. 17, 18). However, the sclerotized ductus bursae looks a little more elongate in P. rhesus, especially in ventral view (Fig. 17); and, where it approaches the corpus bursae, the anterior edge of the sclerotization is more irregular in rhesus (Fig. 18) than it is in P. carus (Fig. 20) (and, for that matter, P. swbreticulata [Fig. 22)).
Polites rhesus ranges from southern Canada (Saskatchewan and AI- berta), in a fairly narrow strip through the western Great Plains and southern Rocky Mountains of the United States (Stanford & Opler 1993), to high mountains of central Mexico (10,000 ft [3050 m] in the state of Mexico); P. carus, from the southwestern United States (western Texas to southeastern California—see Stanford & Opler 1993) to central Mex- ico (Distrito Federal); and P. subreticulata, from central Mexico (Sina- loa, Jalisco, Colima, Michoacan, Distrito Federal, Morelos, and Vera- cruz), through Central America, to Panama (as far, at least, as the Canal).
DISCUSSION Generic Limits and Vanishing Traits
Extending generic limits this way calls for more discussion of variation in characters—especially their reduction and loss. The form of the antennal club and the length of its reflexed apiculus, which are widely used in skipper classification, are undeniably valuable. However, though they tend to be conservative at the generic level, they are hardly im-
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Fics. 19, 20. Female genitalia of Polites carus from Sunny Glen Ranch, 5000-7000 ft (1525-2135 m), near Alpine, Texas, USA, 1-15 May 1926 [X-3488] (USNM). Scale = 1.0 mm. 19, Ovipositor lobes (exserted) with apophyses posteriores, eighth tergite with apophyses anteriores (not sclerotically connected to the lamella postvaginalis of the ste- rigma), sterigma, and bursa copulatrix in ventral view; 20, The same, plus part of the ductus seminalis, in right lateral view.
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Fics. 21,22. Female genitalia of Polites subreticulata from Mazatlan, Sinaloa, MEX- ICO, J. A. Kusche [X-3494] (USNM). Scale = 1.0 mm. 21, Ovipositor lobes (exserted) with apophyses posteriores, eighth tergite with apophyses anteriores (not sclerotically connected to the lamella postvaginalis of the sterigma), sterigma, and bursa copulatrix in ventral view; 22, The same, plus part of the ductus seminalis, in right lateral view.
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Fics. 23-32. Adults of species moved to Polites (all x1) (in USNM unless otherwise indicated); in each figure, upperside on left, underside on right. 23, puxillius 6, 28 mi (45 km) E El Salto, 8400 ft (2560 m), Durango, MEXICO, 6-7 August 1972, Veirs & MacNeill (MacNeill collection); 24, puxillius 2, 25 mi (40 km) W Durango, 8100 ft (2470 m), Durango, MEXICO, 20 July 1964, J. A. Chemsak [X-3196] (Univ. Calif. Berkeley collection); 25, pupillus 6, Sierra de Guerrero, MEXICO, July 1913, R. Miller [X-2699] (type of Amblyscirtes catahorma Dyar); 26, pupillus 2, 2 mi (3 km) SW Potrerillos, 4200 ft (1280 m), Sinaloa, MEXICO, 7-8 August 1986, J. Brown & Powell [X-3203] (Univ. Calif. Berkeley collection); 27, rhesus 6, Sapillo Creek Valley, 5800 ft (1770 m), 12 mi (19 km) N Pinos Altos, Grant Co., New Mexico, USA, 28 May 1959, J. M. & S. N. Burns [X-2712]; 28, rhesus 2, Cedar Creek Canyon, 6900 ft (2100 m), Ruidoso, Lincoln Co., New Mexico, USA, 24 May 1959, J. M. & S. N. Burns [X-3504],; 29, carus 6, Portal, Chiricahua Mountains, 4800 ft (1465 m), Cochise Co., Arizona, USA, 18 July 1974, J. M. & S. N. Burns [X-2708]; 30, carus 2, 6.5 mi (10.5 km) NE Sawtooth Mountain, Davis Mountains, 5600 ft (1705 m), Jeff Davis Co., Texas, USA, 28 April 1959, J. M. & S. N. Burns; 31, subreticulata 6, Mexico City, MEXICO, 7 June 1897, O. W. Barrett [X-3498] (forewing with a long tear in space 1b); 32, swbreticulata 2, Mexico City, MEXICO [X- 3491], 7
VOLUME 48, NUMBER 1 43
mutable. For example, Amblyscirtes alternata (Grote & Robinson), which is an Amblyscirtes in the narrowest sense, is unique among all those species in having the sizable, delicate, sharp apiculus suddenly reduced to a short, blunt fraction of itself (Burns 1990). Lumping Yvretta with Polites may bother those who overweigh antennal clubs because the very short but definite apiculus of the rest of Polites seems to be lacking in the rhesus group. (Its blunt antennal club is one of the main reasons why Yvretta was created in the first place and kept well removed from Polites.) But in most specimens of each of the species of the rhesus group the blunt club actually ends in a slightly reflexed nubbin (comprising a few segments) that looks like a variably vestigial apiculus. A supposedly important difference between the rhesus group and the rest of Polites becomes relatively trivial.
Differences in stigmal expression are demonstrably insignificant. Al- though males of P. carus and P. subreticulata of the rhesus group have a fairly well developed, Polites-type stigma, males of P. rhesus do not: the stigma is, at best, variably vestigial, and is usually missing altogether. In apparent contrast, “‘real’’ Polites males “always” develop a good stigma—or they did until MacNeill (1993) described P. norae, which produces nothing at all. (Polites norae is near P. sabuleti in the stigmally well endowed themistocles group.) Stigmal expression has been shown to vary ina similar, unpredictable manner within other genera. Despite a respectable male stigma in most species of Atrytonopsis, the lunus group entirely lacks one (Burns 1982), and A. deva (Edwards) “runs a gamut from no stigma, through many and various vestigial and reduced expressions, all the way to the complete three-part structure’ (Burns 1982:551).
On the genitalic front, no importance can be attached to the fact that the uncal combs of the rhesus group are bigger than those of other Polites. They are only about as much larger than those of the themisto- cles and origenes groups as those of the vibex group are smaller. At the farthest extreme, Polites includes baracoa, whose uncal combs have vanished without a trace. Moreover, baracoa has lost its gnathos. And, in a different kind of unique development, its paired, toothed, caudally- extending titillators have broadly joined each other, near their anterior ends, across the underside of the aedeagus. Altogether, in its genitalic morphology, the rhesus group is considerably closer to the Polites main- stream than is P. baracoa.
Some may be jolted by the broad, round, female-like wings of males of P. puxillius (Fig. 23) and P. pupillus (Fig. 25) within the origenes group, where males of all other species have narrower, more pointed wings than do females. But sexual dimorphism in wingshape (which is typical and extremely widespread in skippers) has abruptly disappeared
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elsewhere within the genus Polites—in P. mardon within the peckius subgroup of the themistocles group.
Distribution and Speciation
In the origenes group, P. origenes (mainly from the eastern and central United States) morphologically and geographically links the northern, transcontinental superspecies P. mystic (P. mystic and P. sonora) and the Mexican sister species P. pupillus and P. puxillius. These montane sisters appear to be closely allopatric, in part through different altitudinal preferences; but data are too few to say for sure (see Fig. 10). Phylogenetically closer to the Mexican sisters than to superspecies P. mystic, P. origenes is out of touch with them and broadly sympatric with the superspecies—mostly with P. mystic (see maps in Opler & Malikul 1992, Stanford & Opler 1993). However, across all five species of the group, sympatry is limited. Their spatial distribution strongly reflects a set of allopatric speciation events.
In the rhesus group, the superficially distinctive and more northward ranging P. rhesus extensively overlaps P. carus in the southwestern United States and northern Mexico, whereas P. carus and P. subreticu- lata, which are sister species, seem (from meager distributional data) almost to replace each other geographically.
Again, the four strictly western members of the peckius subgroup of the themistocles group—P. draco, P. sabuleti, P. norae, and P. mar- don—are essentially allopatric (MacNeill 1993). So are several members of the vibex group, which Evans (1955) erroneously treated as a single, very widely distributed polytypic species.
All four species groups of Polites overlap in distribution, ranging in broadly repetitious—though different—patterns from the neotropics to the nearctic: the vibex group from Argentina, Paraguay, Brazil, Bolivia, and Peru to Mexico, the West Indies, and (primarily) the southeastern United States; the rhesus group from Panama to southern central Can- ada; the origenes group from southern Mexico to the width of southern Canada; and the themistocles group from central Mexico to the width of southern Canada, plus Yukon Territory. But all four species groups of Polites still give clear distributional evidence of allopatric speciation within themselves. This is independent evidence that the morpholog- ically defined species groups are correct.
ACKNOWLEDGMENTS
Frederick H. Rindge at the American Museum of Natural History, New York, New York, furnished some important material; Jerry A. Powell, Essig Museum of Entomology, University of California, Berkeley, John E. Rawlins, Section of Invertebrate Zoology, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, C. Don MacNeill, and Doug Mullins loaned some; and MacNeill, John Kemner, and H. A. Freeman donated
VOLUME 48, NUMBER 1 45
some. Don Harvey helped with the map (Fig. 10). Elizabeth Klafter and the late Adrienne Venables dissected genitalia (126 KOH-preparations of Polites, about evenly split between the sexes). Daniel Otte at The Academy of Natural Sciences, Philadelphia, Pennsylvania, made a digital image of one dissection (Fig. 14), Victor E. Krantz photographed adults (Figs. 23-32) as well as the Godman genitalic illustrations (Figs. 1, 15, 16), and Young Sohn drew eight dissections (Figs. 2-9, 11-18, 17-22) and then mounted all figures. Sarah Burns aided in many ways. George T. Austin and C. Don MacNeill perused the manuscript. My thanks to everyone.
LITERATURE CITED
BurRNS, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif. Publ. Entomol. 37:1-217.
1982. Lychnuchoides frappenda from central Mexico joins lunus and zweifeli
in a lunus group of Atrytonopsis (Lepidoptera: Hesperiidae: Hesperiinae). Proc.
Entomol. Soc. Wash. 84:547-567.
1988. Superspecies Atrytonopsis ovinia (A. ovinia plus A. edwardsi) and the
nonadaptive nature of interspecific genitalic differences (Lepidoptera: Hesperiidae).
Proc. Entomol. Soc. Wash. 85:335-358.
1990. Amblyscirtes: Problems with species, species groups, the limits of the genus, and genus groups beyond—A look at what is wrong with the skipper classi- fication of Evans (Hesperiidae). J. Lepid. Soc. 44:11-27.
Evans, W. H. 1955. A catalogue of the American Hesperiidae indicating the classifi- cation and nomenclature adopted in the British Museum (Natural History). Part IV. Hesperiinae and Megathyminae. British Museum, London. 499 pp., pls. 54-88.
GODMAN, F. D. 1900. In Godman, F. D. & O. Salvin. 1879-1901. Biologia Centrali- Americana; Insecta; Lepidoptera-Rhopalocera. Vol. 2, 782 pp.; Vol. 3, 113 pls.
HEMMING, F. 1935. Notes on seventeen genera of Rhopalocera. Stylops 4:1-3.
HOFFMANN, C. C. 1941. Catalogo sistematico y zoogeografico de los Lepidopteros Mexicanos. Segunda parte. Hesperioidea. Anales Inst. Biol. [Mexico] 12:237-294.
LINDsEY, A. W., E. L. BELL & R. C. WILLIAMS JR. 1931. The Hesperioidea of North America. Denison Univ. Bull., J. Sci. Lab. 26:1-142.
MACNEILL, C. D. 1993. Comments on the genus Polites, with the description of a new species of the themistocles group from Mexico (Hesperiidae: Hesperiinae). J. Lepid. Soc. 47:177-198.
OPLER, P. A. & V. MALIKUL. 1992. A field guide to eastern butterflies. Houghton Mifflin Co., Boston, New York, London. xvii + 396 pp., 48 pls.
SCUDDER, S. H. 1889. The butterflies of the eastern United States and Canada with special reference to New England. Publ. by the author, Cambridge, Massachusetts. Vol. 3, pp. vii + 1775-1958, pls. 1-89, 3 maps.
SKINNER, H. & R. C. WILLIAMS JR. 1923. On the male genitalia of the Hesperiidae of North America. Paper III. Trans. Am. Entomol. Soc. 49:129-158.
1924. On the male genitalia of the Hesperiidae of North America. Paper V. Trans. Am. Entomol. Soc. 50:141-156.
STANFORD, R. E. & P. A. OPLER. 1993. Atlas of western USA butterflies, including adjacent parts of Canada and Mexico. Publ. by the authors, Denver and Fort Collins, Colorado. xi + 275 pp.
Received for publication 6 August 1993; accepted 12 September 1998.
Journal of the Lepidopterists’ Society 48(1), 1994, 46-50
A NEW SPECIES OF ACROLEPIOPSIS AND THE DESCRIPTION OF THE FEMALE OF A. CALIFORNICA (ACROLEPIIDAE)
REINHARD GAEDIKE
Deutsches Entomologisches Institut (DEI), Schicklerstrasse 5, D 16202 Eberswalde, Germany
ABSTRACT. The family Acrolepiidae, of the superfamily Yponomeutoidea, includes three genera: Digitivalua Gaedike, Acrolepiopsis Gaedike, and Acrolepia Curtis. Acro- lepiopsis liliitvora, new species, from California and Oregon, is described herein; the male and female genitalia are illustrated. Adults have been reared from the bulbs of Lilium washingtonianum. The female of A. californica, previously unknown, also is described, and the genitalia are illustrated. The larvae are reported to feed on Disporum hookeri.
Additional key words: Acrolepiopsis liliivora, genitalia (male and female), Lilium washingtonianum, Disporum hookeri.
Acrolepiidae is a family in the superfamily Yponomeutoidea. It is represented in all zoogeographical regions and includes 84 described species in three genera: Digitivalvua Gaedike (40 species), Acrolepiopsis Gaedike (35 species), and Acrolepia Curtis (9 species). In the Nearctic and Neotropical regions there are 15 described species, three in Digi- tivalva, ten in Acrolepiopsis, and two in Acrolepia (Gaedike 1984a, 1984b). Undoubtedly, more detailed study of each zoogeographical region will increase the number of known species.
The known life histories indicate that acrolepiid larvae are leaf- skeletonizers or miners of leaves, fruit, or bulbs of Asteraceae (Digiti- valva), Dioscoriaceae and Liliaceae (Acrolepiopsis), and Solanaceae (Acrolepia). While examining acrolepiid from the western United States, I discovered a new species of Acrolepiopsis and the previously unde- scribed female of A. californica. These are described below.
Acrolepiopsis liliivora Gaedike, new species
Wingspan 15-16 mm. Head dark brown, except vertex light brown; labial palpi dark brown. Thorax dark brown. Forewing dark brown, with white triangular spot, sometimes overlaid with dark scales, ex- tending obliquely forward from margin of dorsum at % distance from base, with minute white dot on dorsum at base of cilia, and with individual light scales scattered throughout the dark area; cilia beneath apex pale distally. Males somewhat lighter brown than females. Male genitalia (Fig. 1): saccus elongate, narrow, apically rounded; valva with
—
Fics. 1-4. Genitalia of Acrolepiopsis liliivora Gaedike, new species. 1, Male genitalia; 2, Female genitalia (sterigma); 3, Female genitalia (signa); 4, Female genitalia (signa).
AT
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base broad, narrowest medially and expanded apically, costal margin concave; aedeagus more than 2.5 times valva length, broad basally, tapering to apex, with minute sclerotizations in vesica. Female genitalia (Figs. 2-4): Eighth segment with a pair of relatively narrow, somewhat clublike structures bearing setae on the broadly rounded base; ostium with rectangular sclerotization, the lower half more strongly sclerotized; ductus bursae strongly sclerotized over most of its length; corpus bursae with two long signa; signa slightly bent with dentate inner surface, the appearance variable depending upon the preparation.
Type locality: California: Auto Rest. (I have been unable to determine the exact location.)
Types: Holotype male: Auto Rest, Cal [ifornia] 18. 8.[19]18, on Lilium washingtonianum, Coll. David Griffiths (genital slide R. Gaedike No. 2421). Paratypes: 1 male, 3 females, same data as holotype; 1 female Oregon, Santiam Natl. For., reared from bulb of Lilium washingto- nianum, emerged 9. X. 1931. The holotype and three paratypes are deposited in the U.S. National Museum of Natural History, Smithsonian Institution, Washington, D.C.; two paratypes are deposited in the Deutsches Entomologisches Institut (DEI) Eberswalde.
Biology: Larva in bulb of Lilium washingtonianum (Liliaceae), pupa in a net-like cocoon.
Remarks: All specimens are in poor condition. One female lacks forewings; another lacks a hindwing. The new species is closely related to A. californica. It differs in the coloration of the cilia below the apex, in having somewhat broader valvae and longer signa than californica.
Acrolepiopsis californica Gaedike, 1984 Entomol. Abh. Staatl. Mus. Tierk. Dresden 47(1983)10:183-184, Fig. 20.
In material sent to me by J. Powell there were four specimens of this species, two of which were females. Because the female was previously unknown, I describe the female genitalia below.
Female genitalia (Fig. 5): The pair of ventrolateral clublike structures bearing setae on the broadly rounded base, tapering to a point; ostium with a somewhat cup-shaped sclerotization, the lower half more strongly sclerotized; ductus bursae strongly sclerotized; signa short, with dentate inner surface.
Material examined: Two males, two females, as follow: One male: Calif.: El Dorado Co., Blodgett Forest, 13 mi E Georgetown, 4000- 4500’, 27/28.V.1978, leg. J. Powell. One male: Calif.: Siskyou Co., McCloud River at Ash Creek Rgr. Sta., 9.VI.1974, leg. J. Powell. One female: Calif.: Myers Flat, Humboldt Co., 14.VIII.1963, leg. J. Powell. One female: Calif.: Big Creek Reserve (UCNLWR), Monterey Co., 26./
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Fic. 5. Female genitalia of Acrolepiopsis californica.
28.V.1987, leg. J. Powell. These specimens are the first records of this species since its description.
Biology: J. Powell (in litt.) gives some remarks on the biology of this species: “... In late April [1990] I made additional collections of the larvae of Acrolepiopsis at Big Creek on Disporum hookeri and suc- ceeded in rearing a few adults. This confirms the hostplant for the species represented by one specimen that I sent you from 1987. The larvae sometimes start to eating the inflorescence but sometimes do not; later they skeletonize the leaves, feeding on either upper or lower surfaces.”
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ACKNOWLEDGMENTS
I am much obliged to J. B. Heppner and an anonymous reviewer for their help with the manuscript. I thank Heppner for the loan of material from the Smithsonian Institution, Washington, D.C., and J. A. Powell for the loan of material from the University of California, Berkeley.
LITERATURE CITED
GAEDIKE, R. 1984a. Revision der nearktischen und neotropischen Acrolepiidae (Lepi- doptera). Ent. Abh. Staatl. Mus. Tierk. Dresden 47(1983)10:179-194.
1984b. Die Epermeniidae und Acrolepiidae der danischen Sidamerika-Expe-
ditionen 1978/79 und 1981/82 (Lepidoptera). Reichenbachia Staatl. Mus. Tierk.
Dresden 22(16):125-181.
Received for publication 17 December 1998; revised and accepted 27 July 1998.
Journal of the Lepidopterists’ Society 48(1), 1994, 51-57
A NEW SPECIES OF LAPARA (SPHINGIDAE) FROM SOUTHEASTERN UNITED STATES
VERNON ANTOINE BROU JR. 74320 Jack Loyd Road, Abita Springs, Louisiana 70420
ABSTRACT. Lapara phaeobrachycerous, new species, is described and illustrated. The new species presently is known to occur in extreme eastern Louisiana and the state of Mississippi in the southeastern United States. The species can be distinguished from Lapara coniferarum (J. E. Smith) by its slightly smaller size, darker color, the presence of a single postcellular dash, narrower wings, shorter antennae, and different flight period.
Additional key words: endemism, hawkmoths, Louisiana, voltinism.
Hodges (1971) recorded two species of Lapara in North America north of Mexico: Lapara bombycoides Walker and Lapara coniferarum (J. E. Smith). He treated Lapara halicarnie (Strecker) as a synonym of L. coniferarum, and Lapara pinea Lintner as a synonym of L. bom- bycoides. In his review of Lapara, Riotte (1972) recognized L. halicarnie as distinct on the basis of features of the labial palpus, pretarsus struc- ture, color, size, maculation, genitalia, and larvae. Riotte also questioned the synonymy of L. pinea with L. bombycoides. Previous authors (e.g., Holland 1908, Clark 1919), also noted or discussed the problems asso- ciated with the “hypertrophied” type specimen of L. halicarnie.
At the time of Riotte’s (1972) review, no Lapara species were known from west of the Mississippi River in the United States, and none was known to occur in Louisiana. Riotte examined 24] examples of L. coniferarum from 18 states, mostly bordering the east and Gulf coast, from New York and Rhode Island southward to Florida and westward to Mississippi. He also examined 649 examples of L. bombycoides from a much greater range, 48 examples of L. halicarnie from four south- eastern states, and the type of L. pinea from New York.
Since 1972, 8422 specimens of Lapara from Louisiana have been collected by the author using ultraviolet light traps. Most of the better quality specimens are pinned, spread, and labeled, and are in the au- thor’s collection. From 1972 through 1985, 1946 specimens were col- lected and recorded as L. coniferarum. From 1986 through 1992, spec- imens were segregated into two phenotypes. During these seven years, 6473 Lapara specimens were collected: 2247 (85%) are L. phaeobra- chycerous, new species, and 4226 (65%) are L. coniferarum.
Lapara phaeobrachycerous Brou, new species (Figs. lA & B) Male. Head: Dark charcoal gray to occassional brownish gray in color, scales on front and vertex form a bluntly rounded area between an-
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Fic. 1. The Lapara of Louisiana: L. phaeobrachycerous, A. male holotype, B. female allotype. L. coniferarum, C. male, D. female. Specimens pictured were collected at Louisiana, St. Tammany Parish, 6.8 km NE Abita Springs.
tennae. Antennae laminate, biciliate, weakly hooked tip, length x = 10.3 mm (9.0-12.0; n = 40). Thorax: Color above as described for head, below a slightly lighter shade. Forewing: Dorsal color charcoal gray, whitish scales mostly limited to area basad along curving postmedial line, usually one prominent postcellular dash, though a second dash to varying degrees of prominence is not uncommon. Ventral color usually lighter shade than above, unremarkable without bands or maculation, length x = 27.35 mm (25-81.5; n = 40). Hindwing: Unicolorous charcoal gray dorsally without whitish scales or maculation. Ventral surface as described for forewing. Genitalia: (n = 12) Valve generally oval, process of sacculus variable, narrow to broad projection, either acuminate, dentate, or combination of both, uncus apex mildly hooked (Fig. 2A).
Female. Head: As described for male. Antennae simple, length x =
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sew chi . is Sig |
Fic. 2. Genitalia of L. phaeobrachycerous, Louisiana, St. Tammany Parish, A. male, B. female.
10.6 mm (10.0-12.0; n = 18). Thorax: As described for male. Forewing: Dorsal and ventral color as described for male, length x = 31.6 mm (29.5-35.5; n = 14). Hindwing: Dorsal and ventral color as described for male. Genitalia: (n = 4) Sclerotized genital plate, rounded center convexity along distal edge. Entire structure unremarkable otherwise (Fig. 2B).
Types. Holotype ¢ (Fig. 1A), USA, Louisiana, St. Tammany Parish, 4.2 miles (6.8 km) NE Abita Springs, sec. 24T6SR12E, 9 Sept. 1991. Allotype ¢ (Fig. 1B), same locality, 19 Sept. 1991. Paratypes: 879 6 and 18 2, same locality, April 1 to Oct. 28, 1983-92. Holotype and allotype deposited in U.S. National Museum of Natural History. Paratypes de- posited in Florida State Collection of Arthropods, Gainesville, Louisiana State University, Baton Rouge, and the author’s collection.
Diagnosis. In contrast to L. phaeobrachycerous, scales on the front and vertex of the head of L. coniferarum form an acute distal projection between the antennae and lower margin of the frons, among Louisiana and Mississippi specimens.
Maculation is variable within populations, especially among different broods. In Louisiana, individuals of the first brood of L. coniferarum are larger than those of subsequent broods, a trait seen in other Louisiana Sphingidae such as Isoparce cupressi (Bdv.). The first brood of L. coni- ferarum also exhibits the greatest variation in color and shade. Speci- mens can be very dark, or have a predominance of brown scales es- pecially on the forewings. It is not uncommon for darker specimens to exhibit a range of very dark gray to off-white or even near white hindwings.
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In Louisiana, both Lapara species exhibit varying shades of gray or occasionally brownish gray. An area along the forewing inner margin between the transverse postmedial line and the base is chestnut brown in both species. In fresh specimens of coniferarum, the dorsal forewing ground color is usually ash gray. Unlike phaeobrachycerous, the entire dorsal surface of the forewing of coniferarum is suffused with white scales. In both species, these scales are especially concentrated basad of the along the postmedial line. The upper surface of the hindwing of coniferarum is noticeably lighter than the forewing, owing to white scales, which are more numerous near the base. This pattern occurs to a lesser degree in phaeobrachycerous but is absent in some specimens, the moths appearing unicolorous gray. Some coniferarum, usually in- dividuals of the first brood, also have unicolorous gray hindwings.
Among the sparse forewing maculation of Larara are bold to faint black streaks occurring as a furcating transverse postmedial line, es- pecially on the veins between the postmedial line and the outer margin. These black vein lines flare out as they approach and include the fringe near each vein. The fringe between each vein is usually white in conifer- arum and white to off-white in phaeobrachycerous.
The forewing maculation of both species includes a faint to bold median line, which arises from the chestnut-colored area along the inner margin and proceeds towards the apex. The upper one-third of this line abruptly turns inwardly and intersects the costal margin at varying angles, acutely to near perpendicular. In phaeobrachycerous, this line often intersects the costal margin more basad than seen in coniferarum. The sometimes faint antemedial line roughly parallels the median line. |
Riotte (1972) noted that female L. coniferarum have less accentuated markings than males. Markings of both Louisiana species similarly exhibit less accentuated markings. The forewing postcellular dashes vary in number in both Louisiana species. In coniferarum there are usually two dashes, though one or three occur in varying degrees of prominence and with less frequency. Visually, males of the two species are easy to distinguish. Male phaeobrachycerous appear as darker, slightly smaller, short and narrow-winged specimens with short antennae. The antennae of male coniferarum in Louisiana average 29% longer, x = 13.3 mm (11.5-14.5; n = 40), than in phaeobrachycerous. Differences in both wing length and antennae length of coniferarum and phaeo- brachycerous are statistically significant as defined by t-test (highest level a = 0.0005). In males of coniferarum, the antennae cilia are twice the length of those on phaeobrachycerous, while the shaft diameter is roughly equivalent in both species.
In Louisiana, the average forewing lengths for coniferauam are:
Ol Or
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males x = 29.25 mm (27.5-31.5; n = 40), females x = 31.1 mm (29.5- 35.0; n = 23). Riotte (1972) listed measurements for coniferarum as males 24-30 mm (n = 40), females 27-36 mm (n = 8), with no averages given. For L. halicarnie, Riotte listed forewing lengths as males 29-36 mm (n = 18), females 32-40 mm (n = 6).
The genitalia of L. coniferarum have been illustrated by Rothschild and Jordan (1903), Hodges (1971), and Riotte (1972). Both Riotte and Hodges discuss the variability of certain structures of the male genitalia, and Riotte illustrates the variation in the sacculus. Comparing female specimens from Louisiana, the distal edge of the lamella postvaginalis of L. coniferarum is gently rounded, without the degree of convex protrusion noted in L. phaeobrachycerous. This genital plate attribute does vary slightly over L. coniferarum’s vast range.
Distribution. Despite extensive collecting throughout Louisiana, L. coniferarum has been taken only in six upper, southeastern parishes, all east of the Mississippi River: West Feliciana, East Feliciana, East Baton Rouge, Ascension, Tangipahoa, and St. Tammany. This region of the state is known as the Florida Parishes. :
I have studied hundreds of Lapara specimens from Mississippi to Florida, and upwards along the east coast states to Pennsylvania. From Georgia and South Carolina, specimens exist along with coniferarum which appear assignable to neither coniferarum nor bombycoides. They appear as small, narrow-winged specimens, like phaeobrachycerous, and may account for prior literature records indicating bombycoides occurring south to Florida. Due to limited material, it is unclear if these specimens have a relationship to phaeobrachycerous, though I suspect they are indeed phaeobrachycerous. All specimens examined from the area around Clemson, South Carolina, were melanic (very dark charcoal gray to near black) narrow-winged small specimens with short antennae.
The verified range of L. phaeobrachycerous includes only the states of Louisiana and Mississippi at present. Besides the type locality, spec- imens have been collected at Fluker, Tangipahoa Parish, Louisiana, and from Harrison, Pike, and Lee counties in Mississippi.
Etymology. The specific epithet is derived from the Greek phaeo, meaning dark, dusky, and brachycerous, meaning short horns or an- tennae.
DISCUSSION
Lapara is a difficult genus for several reasons. Most field collected specimens are worn because the moths have a tendency to flutter along the ground through brush and leaf litter for considerable distances, especially when they approach light traps operating within a few feet of ground level. This problem appears to be reduced by operating traps
06 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
at a greater height above ground, for example, above five meters. Females comprised less than one percent of the total Lapara specimens collected. Apparently, females are not attracted to ultraviolet light to the degree males are.
In Louisiana, flight periods of L. coniferarum are different from those of L. phaeobrachycerous, though both species have five annual broods. Based on dates of capture over a seven-year period (1986-92), L. coniferarum has its initial annual brood peaking mid-April, with specimens beginning to appear in mid-March, and on occasion even as early as late February. The initial brood peak is separated from the second brood peak by an approximately two-month interval. Brood peaks two through four begin around mid-June and occur at 30-day intervals, In contrast, L. phaeobrachycerous has its first brood peak about mid-May, though the numbers of individuals of this brood are usually quite low compared to subsequent broods. All five broods occur at 30-day intervals.
Koebele (1881) stated that L. coniferarum has at least two annual broods in the southern United States. Riotte (1972) also stated conife- rarum has two distinct flight periods in the southeastern coastal states. Those authors apparently did not recognize that their second brood/ flight period involved multiple sequentially occurring broods.
Based on specimens collected at ultraviolet light traps, the nightly flight time for male coniferarum in Louisiana begins about midnight and lasts for approximately three to four hours, while females have appeared only at dusk. No flight time information is known about phaeobrachycerous. :
Nothing is known about the immature stages of phaeobrachycerous, but I suspect that the larvae may feed on Pinus species. The area in which this new species occurs in Louisiana is classified as a longleaf pine region.
Specimens fade quite rapidly. Pinned specimens several years old appear brown, quite different in color than freshly collected specimens. This is due in part to ultraviolet degradation as well as the effects of chlorine gas from vaporizing insect pest fumigants in storage cases (Brou 1991). Descriptions of colors and other attributes are representative of those occurring on freshly collected and dried Louisiana specimens, unless stated otherwise. Clark (1919) commented that the type of L. halicarnie in Strecker’s collection appears faded.
ACKNOWLEDGMENTS
I thank the following individuals who supplied specimens, records, or aided in this project: Rick Kergosien, the late Charles Kimball, Bryant Mather, Richard S. Peigler, Eric L. Quinter, and J. C. E. Riotte.
VOLUME 48, NUMBER 1 o7
LITERATURE CITED
Brou, V. A. 1991. Health related information on chemicals commonly used by ento- mologists and adjunct methods of pest control in stored entomological collections. South. Lepid. News 13:1-3.
CLARK, B. P. 1919. Some undescribed Sphingidae. Proc. New England Zool. Cl. 6: 67-68.
Hopces, R. W. 1971. The moths of America north of Mexico, Fasc. 21 Sphingoidea. E. W. Classey Ltd. and R.B.D. Publications. 158 pp.
HopcEs, 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.
HOLLAND, W. J. 1903. The moth book. Doubleday, Page and Co., New York. xxiv + 479 pp. 48 plates.
KOEBELE, A. 1881. Descriptions of and notes upon various larvae. Bull. Brooklyn En- tomol. Soc. 4:20-22.
RIOTTE, J. C. E. 1972. A review of the North American hawk moth genus Lapara (Lepidoptera: Sphingidae). Life Sci. Contr., Royal Ont. Mus. No. 79. 40 pp.
Received for publication 10 December 1992; revised and accepted 18 September 1993.
PROFILE
Journal of the Lepidopterists’ Society 48(1), 1994, 58-67
WILLIAM WITTFELD: THE FLORIDA CONNECTION
JOHN V. CALHOUN! 1731 San Mateo Drive, Dunedin, Florida 34698
ABSTRACT. William Wittfeld (1828-1913) collected numerous natural history spec- imens, including Lepidoptera, at “Indian River, Florida,” in the vicinity of Georgiana, Brevard County. Many of his Lepidoptera specimens were sent to William H. Edwards, Henry Edwards, and Berthold Neumoegen. No fewer than thirty-three taxa of Lepidop- tera were described from material collected by Wittfeld. He contributed to the life history descriptions of at least nineteen species. Three patronyms of Lepidoptera honor Wittfeld. His daughter, Annie M. Wittfeld (1865-1887), also reared numerous species.
Additional key words: Lepidoptera, “Indian River, Florida,” type locality, life his- tory, Annie M. Wittfeld.
During the late nineteenth century, Florida was still largely a vast, undeveloped frontier. New railroad systems were allowing pioneers to establish homesteads in formerly remote Indian territory. In 1880, the entire population of Florida totalled less than 270,000 (Tebeau 1980). In March of that year, William Wittfeld (1828-1913) began collecting Lepidoptera at his home on “Indian River, Florida.” Wittfeld’s spec- imens and observations of Florida species became invaluable to the prominent Lepidopterists of the period. For more than a decade, “In- dian River, Florida” was a renowned source of unique and undescribed species of Lepidoptera.
William Wittfeld (Fig. 1) was born in Germany on 3 April 1828. In 1853, when he was 25 years old, Wittfeld immigrated to the United States, joining thousands of others seeking to escape political unrest in Germany. He left behind a large family, including sisters Hermine and Marie and brothers Herrmann, Hoeinreich, and Peter (Peter immi- grated to the United States in 1869). William lived in Philadelphia where he worked as a bookkeeper. Here, he married Rosalie Gottlieb in 1863, with whom he had two children: Annie, born in 1865, and Harry, born in 1867. During Wittfeld’s residency in Philadelphia, the city experienced “the coldest winter on record” (Weigley 1982). From that time forward, Wittfeld probably yearned for a milder climate.
Lured by the prospects of comfortable temperatures and available land, the Wittfelds moved in 1869 to Merritt Island, Florida where they
' Research Associate, Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Gainesville, Florida.
VOLUME 48, NUMBER 1 09
ints Hae ; sv ne © 5 ae .* Be. Sd mn a fis Fic. 1. William and Rosalie Wittfeld at “Fairyland” (ca. 1890).
were among the first settlers (Hellier 1965). In May 1870, Wittfeld applied for a federal land patent to secure 376 hectares (152 acres) of fertile land on what is now the southern end of Merritt Island, just south of the town of Georgiana in Brevard County. The property rose in a long slope from the Indian River (an inshore marine habitat, now part of the Intracoastal Waterway), eastward across a narrow peninsula to the shores of the Banana River (a marine lagoon). Wittfeld could view both bodies of water from the top of this slope, which he called Hon- eymoon Hill. Into the side of the hill, Wittfeld dug a hurricane shelter nearly large enough to stand in. Forests of live oak (Quercus virginiana Mill.) and cabbage palm (Sabal palmetto (Walt.) Lodd ex Schultes) dominated the hill overlooking a clear lake that Wittfeld appropriately named Honeymoon Lake.
Wittfeld cultivated tropical fruits such as pineapples, bananas, and citrus. He also grew a variety of tropical plants that he generously distributed to his neighbors. His home was built next to a large Indian midden and burial mound and for many years was considered a “‘show- place” on the island (McAleenan 1991). Wittfeld started a school for his children next door to their home, bringing in a teacher who lived in a room adjoining the classroom. Over the years, Wittfeld developed his property into what he called “Fairyland” (or “Fairy Land’’), which
60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
2 \ W Merritt Island
~f Wi Rockledge
Banana River
"Fairyland"
Je
Fic. 2. Location of “Fairyland” in Brevard County, Florida.
became a popular tourist attraction during the late 1880’s (Fig. 2). Tourists rented rooms within five cottages built adjacent to Wittfeld’s home and strolled along trails that meandered through the local forest and around Honeymoon Lake. Although Wittfeld preserved much of the original landscape, developmental pressures began to have an effect on surrounding properties as early as the 1890’s. Holland (1898) and Skinner (1907) both mentioned the destruction of a natural area near Wittfeld’s home due to expanding cultivation.
At least seven different spellings of Wittfeld’s name have been em- ployed. William H. Edwards consistently misspelled his name as ““Wiss- feld’’ in letters to Henry Edwards during 1880 and 1881. This spelling was published at least three times (W. H. Edwards 1881a, 1881b, Neu- moegen 1881) and was the result of Wittfeld’s own signature which closely resembled “‘Wissfeld.”” Additional spelling variations include Whitfeld (McAleenan 1991), Whitfeldt (Langlais 1984), Whittfeld (Anonymous 1918), Whitfield (Kjerulff 1972), Wittfield, and Whittfield (W. H. Edwards 1882d, H. Edwards 1883b, U.S. Census Bureau 1900, Hellier 1965). Although he was usually referred to as “Dr. Wittfeld” and W. H. Edwards (1881f) stated that Wittfeld was a “physician,” it is unclear whether he ever practiced medicine or the title was granted out of respect for his horticultural expertise (e.g., “herb doctor’’). Witt- feld was congenial, educated, spoke eloquent English, and had a pre- dilection for smoking cigars.
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Wittfeld’s botanical interests eventually led to a fascination with insects, especially Lepidoptera. In March 1880, Wittfeld contacted W. H. Edwards about his desire to collect insects. Edwards persuaded his sister and her husband, who resided at Lake Jessop, Florida, to visit the Wittfelds during a trip to the Indian River region. She convinced Witt- feld to collect Lepidoptera specimens for Edwards who generously forwarded the necessary equipment including nets, poison, and insect pins. It appears that Wittfeld also agreed to collect a limited number of Coleoptera specimens at the request of W. H. Edwards, but the disposition of these specimens is unknown.
Shipments of specimens from Wittfeld were eagerly accepted by Edwards who retained only the butterflies, offering the moths (and duplicate butterflies) to Henry Edwards and Berthold Neumoegen. As an incentive to Wittfeld, W. H. Edwards suggested that H. Edwards and Neumoegen begin contributing money to Wittfeld for the speci- mens they received (“it will be well and encouraging’’). To facilitate this endeavor, W. H. Edwards proposed a fixed price of 10 cents each for the desirable specimens. Edwards collected these payments and forwarded them directly to Wittfeld. Over the next twelve years, Witt- feld and W. H. Edwards developed a close relationship.
During 1880 and 1881, Wittfeld supplied thousands of specimens to W. H. Edwards, H. Edwards, and Neumoegen. Individual shipments consisted of 500 or more specimens. In a letter dated 10 December 1880, W. H. Edwards proposed to H. Edwards that Wittfeld could supply them four thousand specimens during 1881 for a fee of two hundred dollars. W. H. Edwards continued to instruct Wittfeld in proper collecting techniques and sent additional equipment especially suitable for collecting moths.
As with many nineteenth century naturalists, Wittfeld’s passion to collect was not limited to one group of organisms. In February 1881, Wittfeld contacted Spencer F. Baird of the Smithsonian Institution and offered to collect snake specimens for the National Museum. He ex- plained that collecting Lepidoptera and Coleoptera “does not occupy all my time.” Wittfeld ultimately supplied Baird with numbers of small mammals, snakes, turtles, frogs, lizards, fish, shells, and a few insects (Coleoptera and Orthoptera; to be forwarded to W. H. Edwards if unwanted). All these specimens were shipped in jars of alcohol which Baird supplied. Wittfeld also procured some Indian artifacts for the museum. Baird paid Wittfeld for this material and based compensation on rarity.
It is apparent that Wittfeld’s inexhaustible collecting activities oc- casionally exceeded the demand for his specimens, especially of ““com- mon” species. In late 1881, Henry Edwards complained about having
62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
to pay Wittfeld for all the specimens he had received. In response, W. H. Edwards wrote Wittfeld and asked him to reduce the number of shipments and to “collect only the rarest in such as he conveniently can and not give all his time to it as he has been doing.” S. F. Baird a'so advised Wittfeld (letter dated 27 March 1882) that many of the specimens he was sending were of little value to the Smithsonian col- lections. Despite the consistency of Wittfeld’s shipments, payments were sometimes painfully tardy. Delinquent payments were further exacer- bated by uncontrollable postal delays. W. H. Edwards lamented that Wittfeld’s letters were “sometimes 10 or 11 days in getting to me... and mine to him just the same.” Hardships associated with life in Florida also contributed to delays in shipments and payments.
In 1881, Wittfeld bemoaned the hordes of mosquitoes he encountered whenever he attempted to collect, resulting in a reduction in the number of specimens he was able to secure. It is notable that one of the highest densities of mosquitoes ever recorded in Florida was near Wittfeld’s home in Brevard County where early residents of Merritt Island sup- posedly escaped mosquito bites by stuffing their clothing with crumpled newspaper (Myers and Ewel 1990). In September 1880, the Wittfelds were nearly devastated by a hurricane. W. H. Edwards was greatly concerned about their welfare and wrote (letter to H. Edwards dated 11 September 1880) that he feared that Wittfeld “suffered by the late cyclone on that coast. Since that occurred I have had no letter from him. He has not missed a mail for months.’ Edwards’ fears were par- tially realized when Wittfeld confirmed that the hurricane “nearly ruined them.” The damage was severe. The storm “blew all the leaves off trees, tore down fences and buildings (but not his house), broke off all ripe oranges, broke flat the bananas” and “pulled pine apples from the roots.”’ As a result of the hurricane, Wittfeld temporarily suspended shipments of specimens to Edwards. The storm had damaged the mail boat, causing the mail to run so irregularly that he could not trust boxes to it. Surprisingly, Wittfeld resumed shipments to Edwards by early October of that year.
Wittfeld soon became concerned that the increasing popularity of his captures would result in unwanted requests for specimens. In April 1880, he asked W. H. Edwards to keep his name as confidential as possible. Only W. H. Edwards, H. Edwards, Neumoegen, and E. T. Cresson, Sr. were initially aware of Wittfeld’s identity. In January 1881, W. H. Edwards reminded H. Edwards not to publish Wittfeld’s name “for fear that it will lead to everybody writing him for butterflies.” Four months later, W. H. Edwards inexplicably ignored his own advice and published Wittfeld’s name (as ““Wissfeld’’) (W. H. Edwards 1881a).
In early 1882, Wittfeld sought to provide specimens of Papilio to the
VOLUME 48, NUMBER 1 63
German entomologist, Otto Staudinger, while reserving the right to continue supplying material to W. H. Edwards and Neumoegen. After 1888, shipments from Wittfeld became increasingly irregular. Very few shipments were sent during the late 1880’s. The correspondence be- tween Wittfeld and W. H. Edwards appears to have ceased in 1892. The Smithsonian Institution continued to receive small mammals and reptiles from Wittfeld until at least 1894.
No fewer than 383 taxa of Lepidoptera were described from Wittfeld material and possess type localities of “Indian River, Florida.’ They are currently included in the families Tineidae (Beutenmueller 1887), Yponomeutidae (H. Edwards 1881la), Sessiidae (H. Edwards 1883b), Hesperiidae (W. H. Edwards 1880a, 188le, Lintner 1881, Skinner 1896), Pieridae (Skinner 1894, Miller and Brown 1981), Lycaenidae (W. H. Edwards 1883), Nymphalidae (W. H. Edwards 1880b), Limacodidae (H. Edwards 1886), Pyralidae (Hulst 1886, Kimball 1965), Geometridae (Grote 1882, Kimball 1965), Arctiidae (H. Edwards 1882, 1883a, Neu- moegen 1881), Apatelodidae (H. Edwards 1886), Sphingidae (Butler 1881), and Noctuidae (Grote 1884, H. Edwards 1881b, 1886, Kimball 1965). Wittfeld also discovered a new species of muskrat (Rodentia: Cricetidae) (True 1884).
Wittfeld was the first to confirm the occurrence of several species of Lepidoptera in North America, including Battus polydamas (L.) (W. H. Edwards 1882a, 1882d), Hypolimnus misippus (L.) (W. H. Edwards 1881d, 1888b), and Siproeta stelenes (L.) (W. H. Edwards 1885). Ed- wards was constantly impressed by the species that Wittfeld found, telling H. Edwards (letter dated 17 July 1881) ‘“‘one Cuban or Mexican species after another turns up there.” In addition to supplying dried adult specimens, Wittfeld also reared many species and sent W. H. Edwards live and preserved examples of ova, larvae, and pupae. Many of the preserved early stages of moths were forwarded to H. Edwards.
W. H. Edwards personally reared many species of Lepidoptera from ova and larvae supplied by Wittfeld. Edwards had a particular fondness for Heliconius charitonius (L.). With the assistance of Wittfeld, Ed- wards reared this species numerous times and published a description of its early stages (W. H. Edwards 1881c). On 27 May 1880, Edwards wrote “I turned a 6 Heliconia Charitonia loose (from chrysalis) in my parlor yesterday and it was pretty to see it course about just as I remember adult Heliconias on the Amazon.” The behaviors of adult H. charitonius were carefully documented by Wittfeld and published by W. H. Edwards (1881f). Wittfeld’s specimens and detailed obser- vations contributed to the life history descriptions of at least nineteen species of Lepidoptera (see W. H. Edwards 188 1a, 1881b, 1882b, 1882c, 1887-97, H. Edwards 1887, Scudder 1889). W. H. Edwards (1888)
64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
wrote that Wittfeld “has done more to elucidate the biological history of the Lepidoptera of Florida than any one who has preceded him.”
Additional references to Wittfeld, “Indian River’ or Georgiana, Flor- ida can be gleaned from over one hundred years of literature on Lep- idoptera. Such sources include W. H. Edwards (1881g, 1884b), French (1885), Maynard (1891), Skinner (1911, 1921), Grossbeck (1917), Skin- ner and Williams (1924, 1925), Burns (1964) and Kimball (1965). Hol- land (1898, 1908, 1931) photographed several of Wittfeld’s specimens for his plates. Wittfeld’s specimens are also discussed in detail by Brown (1967, 1970), Brown and Miller (1975, 1977, 1980, 1987), and Calhoun (1993). Most major North American institutional collections possess Wittfeld Lepidoptera specimens.
Annie M. Wittfeld shared her father’s interest in Lepidoptera. In September 1884, Annie documented the strange effects that a lightning strike had on the larvae of Limenitis archippus floridensis Strecker (W. H. Edwards 1884a). She was a talented artist and occasionally supplied drawings of new captures and early stages of Lepidoptera to W H. Edwards. Some of the life history information attributed to William Wittfeld is actually referable to Annie. In November 1887, Annie ob- served H. misippus ovipositing on purslane (Portulaca sp.) and sub- sequently reared the species on this plant (W. H. Edwards 1888b, dos Passos 1951). This remains one of only two known records of H. mis- ippus reproducing in North America. W. H. Edwards (1888b) char- acterized Annie as “intelligent and painstaking.’ Unfortunately, on 10 April 1887, Annie died of rheumatic fever at the age of 23 (W. H. Edwards 1888a). Fifteen months earlier, on 10 January 1887, the Witt- felds’ only son, Harry, died suddenly of meningitis at the age of 20 after becoming ill on a boat trip down the Indian River. William and Rosalie were devastated by these losses. It is likely that these tragedies contributed to William’s decreasing contact with W. H. Edwards and the Smithsonian Institution during the late 1880's.
The final blow to William occurred with the death of his wife on 16 December 1906. She was interred with Annie and Harry under a large oak tree near their home at “Fairyland.” In 1911, their remains were relocated to the Georgiana Cemetery. William’s health gradually failed. During the closing years of his life, he was cared for at the home of his friends, Mr. and Mrs. John Frye, of Rockledge, Florida. It was here that William Wittfeld died of edema on 13 July 1918 at the age of 86. We was interred beside his wife and two children at Georgiana Cem- etery.
Today, there is little evidence of “Fairyland” or the habitats that Wittfeld frequented. Honeymoon Lake is nearly surrounded by housing
VOLUME 48, NUMBER | 65
developments. Nevertheless, neighboring properties still support trop- ical plants that grew from Wittfeld’s cuttings.
Patronyms in Lepidoptera named in honor of William Wittfeld:
wittfeldii, W. H. Edwards, Thecla, 1883, Canadian Entomol. 15:136-137. wittfeldii, H. Edwards, Alypia, 1883, Papilio 3:34. wittfeldii, H. Edwards, Pyrohotaenia, 1883, Papilio 3:156.
ACKNOWLEDGMENTS
I thank the late Theresa Bier, Nancy C. Sieck, Lucy Tomdale, Bill Wittfeldt, Rose Wooley, and Vera Zimmerman. Without their invaluable assistance, this biography could not have been possible. Lucy Tomdale generously supplied the photograph of William and Rosalie Wittfeld. Lee D. Miller and Jacquiline Y. Miller of the Allyn Museum of Entomology, Florida Museum of Natural History, kindly granted me access to copies of letters written by William H. Edwards. The West Virginia State Archives supplied copies of the two remaining Wittfeld letters in the W. H. Edwards files. Mark Epstein and William Cox provided copies of correspondence between Wittfeld and officials of the Smithsonian Institution which are deposited in the Smithsonian Institution Archives. I also express thanks to Alice Sanders, Librarian at the Division of Plant Industry, Florida Department of Agriculture and Consumer Services, who patiently helped me obtain numerous literature references. Finally, thanks are extended to Marc Epstein and William D. Winter for critically reviewing the manuscript.
LITERATURE CITED
ANONYMOUS. 1913. [Obituary of Dr. William Wittfeld]. East Coast Advocate (8 Aug.):5.
BEUTENMUELLER, W. 1887. Descriptions of new species of North American Tineidae. Entomol. Am. 3:139-140.
BROWN, F. M. 1967. The types of the nymphalid butterflies described by William Henry Edwards—Part III, Nymphalinae, Limenitidinae, Apaturinae and Charaxinae. Trans. Am. Entomol. Soc. 93:319-393.
1970. The types of the lycaenid butterflies described by William Henry Edwards. Part II—Theclinae and Strymoninae. With four neotypes established by Paul A. Opler. Trans. Am. Entomol. Soc. 96:19-77.
Brown, F. M. & L. D. MILLER. 1975. The types of the hesperiid butterflies named by William Henry Edwards. Part I, Hesperiidae; Pyrginae. Trans. Am. Entomol. Soc. 101:597-649.
1977. The types of the hesperiid butterflies named by William Henry Edwards.
Part II, Hesperiidae: Hesperiinae, Section I. Trans. Am. Entomol. Soc. 103:259-302.
1980. The types of the hesperiid butterflies named by William Henry Edwards
Part II, Hesperiidae: Hesperiinae, Section IJ. Trans. Am. Entomol. Soc. 106:43-88.
1987. The types of the hesperiid butterflies described by William Henry Ed- wards. Part II—Hesperiidae: Hesperiinae, Section III and Megathymidae. Trans. Am. Entomol. Soc. 113:29-71.
Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif. Publ. Entomol. 37:1-216.
BUTLER, A. G. 1881. Notes on some North American Lepidoptera. Papilio 1:103-106.
CALHOUN, J. V. 1993. Designation of a lectotype of Nisoniades somnus and notes on the occurrence of Erynnis icelus in Florida (Hesperiidae). J. Lepid. Soc. 47:49-54.
DOs Passos, C. F. 1951. The entomological reminiscences of William Henry Edwards. J. New York Entomol. Soc. 59:129-186.
EDWARDS, H. 188la. A new genus and some new forms of North American Zygaenidae. Papilio 1:80-81.
66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
1881b. Descriptions of some new species of Heterocera. Papilio 1:115-121. 1882. New specie of Heterocera. Papilio 2:9-15.
1883a. New forms of the genus Alypia. Papilio 3: 33-34.
1883b. New species of Aegeriadae. Papilio 3:155-157.
1886. Notes on North American Zygaenidae and Bombycidae with descriptions of new forms. Entomol. Am. 2:8-15.
1887. Early stages of some North American Lepidoptera. Entomol. Am. 3:161-
Wl
EDWARDS, W.H. 1880a. Description of a new species of Pamphila from Florida. Canad. Entomol. 12:224-225.
1880b. Description of a new species of Limenitis. Canad. Entomol. 12:246-
291.
188la. Description of the preparatory stages of Apatura flora, Edw. Canad.
Entomol. 13:81-85.
1881b. Description of the preparatory stages of Papilio palamedes, Drury.
(Calchas, Fab.). Canad. Entomol. 13:119-123.
1881lc. Description of preparatory stages of Heliconia Charitonia, Linn. Canad.
Entomol. 13:158-162.
1881d. Capture of Diadema bolina, Linn, in Florida. Papilio 1:30.
188le. Description of a new hesperian from Florida. Papilio 1:78-79.
1881f. On certain habits of Heliconia Charitonia, Linn., a species of butterfly
found in Florida. Papilio 1:209-215.
1881g. Notes on the species of Callidryas found within the United States. Trans.
Am. Entomol. Soc. 9:9-14.
1882a. Note on Papilio polydamas, Linn. Canad. Entomol. 14:120.
1882b. Description of the preparatory stages of Neonympa areolatus, Smith-
Abbot. Canad. Entomol. 14:163-166.
1882c. Description of the preparatory stages of Grapta interrogationis, Fab.
Canad. Entomol. 14:201-207.
1882d. Papilio polydamas in Florida. Papilio 2:122.
1883. Description of a new species of Thecla from Florida. Canad. Entomol.
15:1386-137.
1884a. [Letter from Annie M. Wittfeld]. Canad. Entomol. 16:180. _
1884b. Revised catalogue of the diurnal Lepidoptera of America north of Mexico.
Trans. Am. Entomol. Soc. 11:245-337.
1885. Miscellaneous notes on butterflies, habits of larvae, etc. oer Entomol.
17:108-114.
1887-1897. The butterflies of North America. Vol. II]. Am. Entomol. Soc,
Philadelphia, Pennsylvania. [432] pp.
1888a. Obituary of Annie M. Wittfeld. Canad. Entomol. 20:140.
1888b. On Diadema misippus, Linn, in Florida. Canad. Entomol. 20:128.
FRENCH, G. H. 1885. The butterflies of the eastern United States. J. B. Lippincott Co., Philadelphia, Pennsylvania. 402 pp.
GROSSBECK, J. A. 1917. In Watson, F. E. (ed.), Insects of Florida IV. Lepidoptera. Bull. Am. Mus. Nat. His. 37: (Article 1):1—147.
GROTE, A. R. 1882. Notes on certain Geometridae, with a new Byssodes from Florida. Papilio 2:100-101.
1884. A new species of Nystalea. Papilio 4:7.
HELLIER, W. R. 1965. Indian River, Florida’s treasure coast. Hurricane House, Pub- lishers, Inc., Coconut Grove, Florida. 128 pp.
HOLLAND, W. J. 1898. The butterfly book. Doubleday, Page & Co., New York, New York. 382 pp.
1903. The moth book. Doubleday, Page & Co., New York, New York. 479 pp.
1931. The butterfly book, new and thoroughly revised edition. Doubleday and Co., Inc., Garden City, New York. 424 pp.
HuLsT, G. D. 1886. Descriptions of new Pyralidae. Trans. Am. Entomol. Soc. 13:145- 168.
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KIMBALL, C. P. 1965. Arthropods of Florida and neighboring land areas. Vol. 1. Lep- idoptera of Florida. Div. of Plant Industry, Gainesville, Florida. 363 pp.
KJERULFF, G. G. 1972. Tales of old Brevard. The South Brevard Hist. Soc., Melbourne, Florida. 121 pp.
LANGLAIS, V. G. 1984. Cemetery census of Brevard County Florida, including Sebastian (Indian River County). Dist. by the author, Titusville, Florida. 155 pp.
LINTNER, J. A. 1881. On some species of Nisoniades. Papilio 1:69-74.
MAYNARD, C. J. 1891. A manual of North American butterflies. De Wolf, Fiske and Co., Boston, Massachusetts. 226 pp.
MCALEENAN, J. 1991. Romantic horticulturist named M_[erritt] I.[sland] landmarks. Florida Today (19 Oct.):2A.
MILLER, L. D. & F.M. BRown. 1981. A catalogue/checklist of the butterflies of America north of Mexico. Lepid. Soc. Memoir No. 2. 280 pp.
Myers, R. L. & J. J. EWEL (eds.). 1990. Ecosystems of Florida. Univ. of Central Florida Press, Orlando, Florida. 765 pp.
NEUMOEGEN, B. 1881. On a new species of Arctia from Florida. Papilio 1:9-10.
SCUDDER, S. H. 1889. The butterflies of the eastern United States and Canada with special reference to New England. 3 Vols. Publ. by the author, Cambridge, Massa- chusetts. 1958 pp.
SKINNER, H. 1894. Tachyris ilaire n. var. neumoegenii. Entomol. News 5:110.
1896. Two new hesperids. Canad. Entomol. 28:187-188.
1907. Rare butterflies. Entomol. News 18:22-24.
1911. The larger boreal American Hesperidae, including Eudamus, Erycides,
Pyrrhopyge and Megathymus. Trans. Am. Entomol. Soc. 37:169-209.
1921. Atrytone kumskaka Scudder (Lep., Rhop.). Entomol. News 32:276-277.
SKINNER, H. & R. C. WILLIAMS JR. 1924. On the male genitalia of the Hesperiidae of North America. Paper III. Trans. Am. Entomol. Soc. 49:129-153.
1925. On the male genitalia of the Hesperiidae of North America. Papers [V- VI. Trans. Am. Entomol. Soc. 50:57-73, 141-156, 177-208.
TEBEAU, C. W. 1980. A history of Florida. Univ. of Miami Press, Coral Gables, Florida. 527 pp.
TRUE, F. W. 1884. A muskrat with a round tail. Science 4:34.
U.S. CENSUS BUREAU. 1900. Twelfth census of the United States, Brevard County, Florida, Precinct 13:184.
WEIGLEY, R. F. (ed.). 1982. Philadelphia, a 300-year history. W. W. Norton & Co., New York, New York. 842 pp.
Received for publication 1 May 1998; revised and accepted 9 August 1993.
GENERAL NOTES
Journal of the Lepidopterists’ Society 48(1), 1994, 68-69
MUSINEON TENUIFOLIUM (APIACEAE): NEW HOST OF FOUR NEBRASKA PAPILIO (PAPILIONIDAE)
Additional key words: Artemisia dracunculus, Papilio machaon-complex, swallow- tail.
The recorded larval host plants of four Papilio species, P. zelicaon nitra W. H. Edwards, P. indra Reakirt, P. polyxenes asterius Fabricius, and P. machaon bairdii (W. H. Ed- wards), encompass a variety of genera in three plant families, Asteraceae, Apiaceae, and Rutaceae. Here we report narrow-leaved musineon, Musineon tenuifolium Nutt. (Api- aceae), as a new larval host plant for these four swallowtails.
Musineon tenuifolium is endemic to the west-central Great Plains, ranging from the Black Hills of South Dakota and Wyoming, south through eastern Wyoming and the Nebraska panhandle (Barkley 1977). Farrar (1990) characterizes M. tenuifolium as a native perennial favoring dry, rocky outcrops, growing from a caudex atop a thickened taproot. It possesses stemless, dissected leaves rising from a root crown to form a compact tuft of foliage. Small, cream to bright yellow flowers form compound umbels, approxi- mately 2.5 cm across, atop a leafless stalk normally less than 15 cm tall. In Nebraska, M. tenuifolium usually stays green the entire season except under severe drought conditions. It is an extremely local umbel, but often common in the appropriate habitat.
M. tenuifolium was first brought to our attention in 1983 while collecting in the canyons of northwestern Sioux County, Nebraska. A search of M. tenuifolium foliage yielded larvae of both P. z. nitra and P. indra. M. tenuifolium is believed to be the exclusive host plant for these swallowtails in Nebraska, and supports a second brood of P. indra in our area. We also have observed larvae of P. z. nitra and P. indra feeding on M. tenuifolium in the Black Hills of South Dakota.
On 20 June 1991, a collecting trip to Bull Canyon, Banner County, Nebraska resulted in about 40 swallowtail larvae collected on M. tenuifolium. Of these larvae, approximately one-half emerged as adults from mid-July through mid-August. Surprisingly, only three adults were P. z. nitra; others were P. p. asterius.
While collecting in southern Scotts Bluff County, Nebraska on 25 May 1992, five 5th instar swallowtails were discovered on M. tenuifolium by the first author and Nick D. Theis. The larvae were removed and fed cut M. tenuifolium until pupation. Starting 15 June 1992, pupae began to eclose. Emergent adults were P. m. bairdii, a species thought to have fed exclusively on silky wormwood, Artemisia dracunculus L. (Asteraceae), in Nebraska.
These observations raise some intriguing questions regarding possible range extensions and the use of alternative host plants by members of the P. machaon-complex. It is likely that continued efforts of naturalists and collectors within and outside the state will lead to insights concerning these and subsequent questions.
We thank Brett C. Ratcliffe and J. Ackland Jones for review of the manuscript. We are grateful to Neil E. Dankert and Jeff C. Germer, who assisted in collecting larvae. This is paper number 10264 of the journal series of the Nebraska Agricultural Research Division, University of Nebraska. This work was supported by the University of Nebraska Agricultural Experiment Station Project 17-055.
LITERATURE CITED
BARKLEY, T. E. 1977. Atlas of the flora of the Great Plains. Iowa State Univerin Press, Ames. 600 pp.
FARRAR, JON. 1990. Field guide to wildflowers of Nebraska and the Great Plains. Nebraska Game and Parks Commission, Lincoln. 216 pp.
VOLUME 48, NUMBER 1 69
J. M. REISER, Department of Horticulture, University of Nebraska, Lincoln, Nebraska 68583-0724, AND S. M. SPOMER, Department of Entomology, University of Nebraska, Lincoln, Nebraska 68583-0816.
Received for publication 25 February 1998; revised and accepted 29 August 1998.
Journal of the Lepidopterists’ Society 48(1), 1994, 69-71
TWO NEW SYNONYMIES IN NEARCTIC EUCOSMA (TORTRICIDAE: OLETHREUTINAE)
Additional key words: Eucosma wandana, E. uta, E. ustulatana, Kentucky, genitalia.
In the course of identifying Eucosma specimens captured recently in Kentucky, we reached the conclusion that E. wandana Kearfott, E. uta Clarke, and E. ustulatana Blanchard & Knudson are one and the same morphospecies. Males captured at the same Kentucky locality on the same date match male holotypes of all three taxa. All specimens we examined that had been or could be assigned to these taxa have similar underlying forewing patterns varying in the proportion of dark brown to light orange brown. On average, males (n = 12) are darker brown than females (n = 44).
Eucosma wandana was described from one male by W. D. Kearfott in 1907, but we also have seen a female with a “type” label not mentioned in the original description; E. uta was described from two males and one female by J. F. G. Clarke in 1953; and E. ustulatana was described from two males by A. Blanchard and E. C. Knudson in 1983. None of these short type series reveals the full range of color variation we observed in longer series. Moreover, Clarke seems to have overlooked E. wandana when he described E. uta; for differentiation, he used E. atomosana (Walsingham), a western species (Hein- rich 1928). In describing E. ustulatana, Blanchard and Knudson used E. wandana (as E. eumaea Meyrick) for comparison, but the diagnostic differences cited by them—smoother, more evenly convex anterior valval margin in E. ustulatana—can be ascribed to individual variation. No differences are discernible among holotype genitalia of E. wandana (as E. eumaea) (Heinrich 1923), E. uta (Clarke 1953), or E. ustulatana (Blanchard & Knudson 1983).
We summarize as follows:
Eucosma wandana Kearfott (Figs. 1-3)
Eucosma wandana Kearfott (1907) (holotype: male, Cincinnati, Ohio, 25 July [not August as in original description] 1904, A. F. Braun, in American Museum of Natural History [AMNH], New York, N.Y., wings illustrated in Fig. 1 here); Klots (1942).
E. eumaea E. Meyrick (1912); Heinrich (1923) (holotype genitalia photo-illustrated); Klots (1942) (invalid replacement name).
E. uta Clarke (1953) (holotype: male, Putnam Co., Illinois, 10 July 1939, M. O. Glenn, in National Museum of Natural History [NMNH], Washington, D.C.; male and female genitalia sketch-illustrated, wings of paratype photo-illustrated); Godfrey, et al. (1987) wing of female photo-illustrated. New Synonymy.
E. ustulatana Blanchard & Knudson (1983) (holotype: male, Brenham, Washington Co., Texas, 4 June 1979, E. C. Knudson, genitalia prep. WEM 36924, in NMNH, wings photo-illustrated, male genitalia of paratype photo-illustrated). New Synonymy.
Eucosma wandana is probably multivoltine. Capture dates range from 4 June to 15 September (n = 53). The known distribution is Kentucky, Ohio, Illinois, Missouri, Arkansas, Kansas, Georgia, Florida, and Texas. Forewing length of males ranges from 7.5 to 8.5 mm (n = 12); of females, 7.0 to 9.25 mm (n = 44). Eucosma wandana is one of at least 11 eastern species of Eucosma having ventral extensions of female papillae anales (Fig.
70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
3
Fics. 1-3. Eucosma wandana. 1, Wings of holotype. 2, Genitalia of male. 3, Genitalia of female. Z
3), an accessory whose function is not yet known (Miller 1987). The larval food plant is unknown.
We thank F. H. Rindge, G. L. Godfrey, K. R. Methven, J. R. Heitzman, and D. Profant for specimen loans.
Material examined. We examined specimens as follows, labeling all except primary types “Voucher, Gibson & Miller 1992”: ARKANSAS: Devil’s Den St. Pk., Washington Co., one male, 6.VII.66; one female, 6.VII.66 (genit, prep. USNM 70317); one female, 26. VI.66; one female, 3. VII.66 (genit. prep. USNM 70316) all R. W. Hodges; Washington Co., “type”, female. KANSAS: Pittsburg, E. L. Todd, one female, 28.VI.54; one female, 2.VII.54. GEORGIA: Atlanta, P. W. Fattig, one male, 31.VII.41. TEXAS: E. ustulatana holotype; Dallas, “714”, Fernald collection, one female (all foregoing in NMNH). OHIO: E. wandana holotype (in AMNH). KENTUCKY: Campbell Co., Jolly Co. Pk., three females, 15. VII.82 (female genit. prep. LDG 134) (Fig. 3); Gallatin Co., Markland Dam, two females, two males, 3. VIII.88 (male genit. prep. LDG 048) (Fig. 2); Boone Co., Big Bone Lick St. Pk., one female, 7.VII.81; one female, 27. VII.89; Owsley Co., near Boone-
VOLUME 48, NUMBER 1 7a
ville, two females, two males, 22.VI.84 (male genit. prep. LDG 047); Bullitt Co., near Shepherdsville, one female, 8.VII.88 (all L. D. Gibson, in L. D. Gibson collection). MIS- SOURI: Cape Girardeau, one female, 25. VII.78; Independence, one male, 30. VI.76; Clay Co., Coolie Lk., one female, 18.VII.68; one female, 22. VII.72 (all J. R. Heitzman, in J. R. Heitzman collection). ILLINOIS: E. uta holotype; Oconee, two females, 8-15. VII; one female 1—7.VII (female genit. prep. USNM 70670); Decatur, one female, 8-15.VII; Put- nam Co., M. O. Glenn, one female, 30. VII.74; one male, 18. VII.73; one female, 3. VIII.65; one male, 25.VII.74 (male genit. prep. WEM 176923); one female, 13.VIII.43 (female genit. prep. USNM 70320) (E. uta paratype); (all in NMNH); Putnam Co., M. O. Glenn, one male, 23.VI.56, “5057”; one male, date missing, “5157”; one female, 5.VII.53; one female, 24. VII.68; one female, 11.VIII.74; one female, 15.VII.64; one female, 5. VIII.61 (abdomen missing); one female, 28.VII.64; one female, 2.VII.56; one female, 15.1X.54; one female, 28. VII.56; one female, 21. VII.56; one female, 23. VII.73; one female, 10. VII.39; one female, [illegible]. VII.65; one female, 17.VII.64; one female, 29.V1I.64; one female, 5. VIII.47 (E. uta paratype) (all in Illinois Natural History Survey Collection). FLORIDA: Liberty Co., Torreya St. Pk., one female, 7.VII.88 (female genit. prep. LDG 131) (D. Profant, in D. Profant collection).
LITERATURE CITED
BLANCHARD, A. & E. C. KNUDSON. 1983. New North American species of Eucosmini (Lepidoptera: Tortricidae). Proc. Entomol. Soc. Wash. 85:845-852.
CLARKE, J. F. G. 1953. New species of Olethreutidae from Illinois (Lepidoptera). J. Wash. Acad. Sci. 43:226-231.
GopFREY, G. L., E. D. CaAsHaTT & M. O. GLENN. 1987. Microlepidoptera from the Sandy Creek and Illinois River region: An annotated checklist of the suborders Dacnonypha, Monotrysia, and Ditrysia (in part) (Insecta). Il]. Nat. Hist. Surv. Spec. Pub. 7. 44 pp.
HEINRICH, C. 1923. Revision of the North American moths of the subfamily Eucosminae of the family Olethreutidae. U.S. Natl. Mus. Bull. 123. 298 pp.
KEARFOTT, W. D. 1907. New North American Tortricidae. Trans. Am. Entomol. Soc. 33:1-98.
Kiots, A. B. 1942. Type material of North American microlepidoptera other than Aegeriidae in the American Museum of Natural History. Bull. Am. Mus. Nat. Hist. 79:391-424.
MEYRICK, E. 1912. On some impossible specific names in micro-Lepidoptera. Entomol. Mon. Mag. 23:32-36.
MILLER, W. E. 1987. Guide to the olethreutine moths of Midland North American (Tortricidae). U.S. Dept. Agr. Handb. 660. 104 pp.
LORAN D. GIBSON, 8496 Pheasant Drive, Florence, Kentucky 41042, AND WILLIAM E. MILLER, Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108.
RECEIVED FOR PUBLICATION 28 JANUARY 1993; REVISED AND ACCEPTED 27 JULY 1993.
Journal of the Lepidopterists’ Society 48(1), 1994, 71-78
ADDITIONAL DATA ON THE GEOGRAPHICAL DISTRIBUTION AND ADULT ACTIVITY OF THE DIURNAL, MIMETIC PLUME MOTH, OIDAEMATOPHORUS CHAMELAI (PTEROPHORIDAE)
Additional key words: Costa Rica, Mexico, Croton, Cordia.
An extraordinary, black pterophorid with enlarged scale tufts on the hind legs was described from Mexico by Gielis (1992), but neither its strikingly wasp-like appearance
2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
nor its diurnal behavior, unusual for the family, were emphasized. In addition to the type series of seven specimens, most of which were collected by Chemsak in 1985-86, we have taken a larger number in more recent years, documenting a more extensive flight period. Moreover, two specimens from Costa Rica were erroneously recorded from Mexico by Gielis.
Geographical distribution. Oidaematophorus chamelai has been recorded along the immediate Pacific coast of Middle America, in Mexico near Elota in central Sinaloa, and Chamela, Jalisco, and in northwestern Costa Rica in Guanacaste Province. An inland occurrence in Mexico, indicated on the mapped distribution by Gielis in the vicinity of Guadalajara, evidently was based on his misinterpretation of the label on specimens from Costa Rica (“C.R.: Playa Tamarindo, Guan. Prov.’’) as the State of Guanajuato, Mexico. A specimen from south of Elota, Sinaloa, was illustrated and cited but not mapped.
Seasonal occurrence. In Mexico we have taken adults of O. chamelai during each month that we have visited the Estacion de Biologia Chamela: July, August, September, October, December; while the Costa Rica specimens were collected in June. Whether or not the species flies during the dry season (December—June in Mexico, January—April in Costa Rica) cannot be determined from our sampling.
Mimetic appearance and diurnal activity. Oidaematophorus chamelai and two closely related South American species, O. ochracealis (Walker) and O. paraochracealis Gielis, differ markedly from typical members of the genus in having the wings relatively broad and uniformly black with purplish iridescence or “brown-black”’ (Gielis 1992). Oidae- matophorus chamelai is particularly wasp-like, an image enhanced by broad, cream- colored, dorsolateral spots on abdominal segments 2 + 3 that render the appearance of the constricted petiole of many ichneumonids and sphecids, and by seemingly aposematic red tegulae and large yellowish “eyespots” of the metathorax. Another pair of cream colored dorsal spots at the base of segment 8, which were lacking on the Sinaloa female Gielis illustrated, are variable from a trace to conspicuous in both sexes. The male has long, white, eversible hair brushes flanking the genitalia that were not mentioned by Gielis. The enlarged scale tufts of the hind tibiae (misinterpreted as the first tarsal segment in the original description) and tarsi in the male and tarsi only in the female are reminiscent of many ctenuchine Arctiidae, such as Macrocneme, that are believed to be members of mimetic assemblages. D. L. Janzen has reared one species of Macrocneme and two of the closely related Poliopastea from Mesechites trifida (Apocynaceae), a plant family well known for its cardiac glycocides. The diurnal, aposematic larvae and adults of these two genera are superficially quite similar and presumably are distasteful to vertebrate predators (Janzen in litt., Dietz 1994).
While nearly all Nearctic and Neotropical pterophorids that we have encountered are nocturnal, O. chamelai (and presumably the other species of the O. ochracealis complex) are primarily diurnal. We have observed adults of O. chamelai at flowers of three shrub and tree species of Croton (Euphorbiaceae) at the Estacion de Biologia Chamela. Their time of activity corresponded with that of other flower visitors such as cerambycid and scarabaeid beetles, wasps, bees, and hesperiids. In Costa Rica, males were collected by sweeping inflorescences of an arboreal Cordia (Boraginaceae), 5-6 m above the ground, employing an extensible tropical net. In addition, however, about 7 individuals of 17 taken at Chamela were attracted nocturnally to ultraviolet or white lights.
Location of specimens. The holotype is housed in the Essig Museum of Entomology, University of California, Berkeley (UCB), along with later collections. Specimens also are deposited in the Instituto de Biodiversidad, Santa Domino de Heredia, Costa Rica (INBio); Instituto de Biologia, Universidad Nacional Autonoma de México, Mexico City (UNAM); Los Angeles County Museum of Natural History (LACM); U. California, Riverside (UCR); and the U. S. National Museum of Natural History, Washington, D.C. (NMNH). The type series was erroneously cited as LACM; the specimens had been on loan to J. P. Donahue from UCB and UCR and were so labelled when forwarded to Gielis, but their ownership was not communicated.
Material examined. COSTA RICA: Playa Tamarindo, Guan. Prov., 2 6, VI-14-86, “arboreal Croton fls. 15-18 ft. above ground, tropical net’’ (Chemsak, H: Katsura, A. & M. Michelbacher). MEXICO: Estacion de Biologia Chamela, Jalisco, ¢ Holotype, ? VII-
VOLUME 48, NUMBER 1 73
8/16-85 (Chemsak, Katsura, Michelbacher); 2 4, 2 X-13/23-86 (Chemsak); 2 VII-12/15- 87 (Chemsak & E. G. Linsley); 2 2 X-16/19-87, 6 X-21/22-87, blacklight (Chemsak & Powell); 6 [X-27/28-88, blacklight (P. A. Opler); 2 XII-1-88 (Chemsak); 2 6, 2 VII-23-90, 1 6 at light (Chemsak); ° VII-13-92, 6 VII-20-92, at lights (Chemsak); 1 6, VIJ-15-93, at Croton flowers (B. Eya).
We thank D. H. Janzen for unpublished observations on ctenuchid biology in Costa Rica and J. P. Donahue for review and comments on the manuscript.
LITERATURE CITED
Dietz, R. E. Ill. 1994. Systematics and biology of the genus Macroeneme Hiibner (Lepidoptera: Ctenuchidae). Univ. Calif. Publ. Entomol. In press.
GIELIS, C. 1992. Neotropical Pterophoridae 6. The Oidaematophorus ochracealis com- plex (Lepidoptera). Phegea 20:81—94.
J. A. POWELL AND J. A. CHEMSAK, Essig Museum of Entomology, University of California, Berkeley, California 94720.
Received for publication 6 April 1993; revised and accepted 4 September 1993.
Journal of the Lepidopterists’ Society 48(1), 1994, 74-76
BOOK REVIEWS
KEYS TO THE INSECTS OF THE EUROPEAN PART OF THE USSR (G. S. Medvedev, chief editor). VOLUME IV (LEPIDOPTERA), PART II, by M. I. Falkovitsh (ed.) et al. 1990. E. J. Brill, Leiden. (translation of: OPREDELITEL NASEKOMYKH EVROPEISKOI CHASTI SSSR, TOM IV, CHESHUEKRYLYE, VTORAIA CHAST. Nauka Publishers, Leningrad, 1981—+trans- lator: B. R. Sharma). x + 1092 pp., 675 figs. Hard cover, 16 x 24 cm, ISBN 90-04-08926- 8. $160.00 U.S. Available from E. J. Brill (U.S.A.) Inc., 24 Hudson Street, Kinderhook, New York 12106.
This remarkable, bulky handbook is the second part of a work devoted to the Lepi- doptera of western Russia. The English version of Part One was issued in 1987 (Amerind Publishing Co. Pvt. Ltd, New Delhi—edition supervised by the U.S.D.A., Washington, D.C.), and it covered the non-ditrysian families (with only a superficial treatment of the Nepticulidae), and eight families among the lower Ditrysia, namely the Psychidae and all members of the Zygaenoidea, Cossoidea, Sesioidea and Tortricoidea. Part Two deals with a larger number of families (29 if one accepts the classification that I proposed in 1991: see Entomol. Scand. 22:90-91). These are the Eriocottidae and Tineidae (including the “Euplocamidae”’ and “Hieroxestidae’’), all members of the Gracillarioidea, Ypono- meutoidea, Choreutoidea, Urodoidea (Wockia Heinemann: p. 508, as a “plutellid” genus), Schreckensteinioidea, and Epermenioidea, and all the gelechioid families with the ex- ception of most Coleophoridae (only the Amphisbatinae being treated: pp. 792 and 801, among the Oecophoridae sensu auct.). Parts One and Two thus provide keys to the “Micromoths” of nearly all the families known to occur in the European part of Russia.
The present book is firmly bound in boards, nicely presented, and has a good quality paper, i.e., opaque and whiter than that of the Russian edition. Compared with the latter, the book is thicker (approximately 7 cm) and of a different format: 15.5 by 24 cm instead of 17.5 by 27 cm. Most figures, however, have not been reduced and are as good as those of the original publication. Only a few figures are less satisfactorily printed, such as Figures 191 and 193. The page numbers of the Russian original are mentioned in the lefthand margin, and thus often precede the numbers of the figures (which may be a bit confusing). The arrangement of the figures is usually rather practical, although it should have been slightly different in a few cases: for example, Figures 11 and 12 (Euplocamus Latreille) are found on page 16, in the “Key to suborders and families,” whereas they correspond to the “Euplocamidae”’ text (pp. 24-25); Figures 499 to 505 (Schreckensteinia Hubner and Heliodines Stainton) should have been placed between the Schreckenstei- niidae (pp. 697-698) and Heliodinidae (pp. 699-700), rather than at the end of the Stathmopodidae chapter; etc.
On the whole, the English translation appears quite faithful, and no changes were introduced, either nomenclatural or taxonomic. That is definitely a reasonable option, although typographical errors might have been rectified for certain Latin names. For instance, Bucculatrix gnaphaliellea (p. 193), B. frangulella (p. 195), Caloptilia populo- torum (p. 252), Milliereia (p. 414), Mompha propinguella (p. 704), Pseudatemella (p. 792), and Oliaria (p. 1001), are all incorrect spellings for, respectively, Bucculatrix gna- phaliella (Treitschke), B. frangutella (Goeze), Caloptilia populetorum (Zeller), Millieria Ragonot, Mompha propinquella (Stainton), Pseudatemelia Rebel, and Uliaria Dumont.
Nine authors contributed to Part Two, namely: A. S. Danilevsky (Choreutidae), M. I. Falkovitsh (Heliodinidae, Scythridae, Stathmopodidae, Schreckensteiniidae, Epermeni- idae), Z. S. Gershenzon (Yponomeutidae), V. I. Kuznetzov (Gracillariidae, Glyphipteri- gidae, Choreutidae), A. L. Lvovsky (Oecophoridae sensu auct.), V. I. Piskunov (Sym- mocidae, Lecithoceridae, Blastobasidae, Gelechiidae), S. V. Seksajeva (Bucculatrigidae, Gracillariidae, Phyllocnistinae, Lyonetiidae), S. Yu. Sinev (Elachistidae, Batrachedridae, Momphidae, Cosmopterigidae), and A. K. Zagulajev (Tineoidea, Douglasiidae, Roesler- stammiidae, Urodidae, and several yponomeutoid and gelechioid families). Most of these entomologists are well known specialists of the families in question, so that the proposed keys are usually quite reliable. Sometimes they could not examine a number of foreign (European) species, their work in such cases being largely based on that of other specialists
VOLUME 48, NUMBER 1 To
(e.g., R. Gaedike’s papers on Douglasiidae and Epermeniidae). Much in the same way, the drawings were made by several artists or even, sometimes, by certain of the above- mentioned lepidopterists. These illustrations range from fair to excellent, and are often original, though explicitly taken from other works in several cases. I would like to em- phasize the abundance of the figures provided throughout the book: nearly 530 drawings represent adults or their wing pattern; nearly 240, wing venation schemata; about 1700 and nearly 470, male and female genital structures, respectively; and so on. As in Part One, photography was not used, probably because diagnostic features may be more easily distinguishable with line drawings.
After a short abstract (p. v), a preface (pp. vii-viii) and a table of contents (pp. ix—x), the handbook starts with a key to suborders and families (pp. 1-23). Elaborated by M. I. Falkovitsh and A. K. Zagulajev, this key is almost identical with that published in Part One, except for minor changes: the translation is better, abbreviations have been avoided, and illustrations are slightly more numerous. Various parts of the key are interesting and original, but a few remarks have to be made:
* many of the characters selected for a given family cannot be generalized on a
worldwide scale
according to the seventh alternative, tibial spurs would be absent from the Hepialidae, but they are actually present in genus Gazoryctra Hiibner, one species of which is treated in Part One (as a member of Korscheltellus Borner)
* alternatives 13 and 22 are chiefly based on a slight difference in the “breadth” of the head, not on a clear-cut demarcation between two traits
several inaccuracies must be corrected: ocelli are present in many Eriocottidae (con- trary to alternative 16 statement), the maxillary palpi may be well developed in Adelidae since Nematopogon Zeller definitely belongs to this family (90), the Sesiidae possess a long frenulum (118), tympana are distinctly present in the Thaumetopoeinae (Notodontidae) (182), ete.
* “Endromidae’” has been omitted after “In forewings R with 4 branches (R2-R5) stalked together” (alternative 190); in the description of the Nolidae (alternative 202), “ocelli” obviously represents a slip, and should be replaced by “compound eyes
the nomenclature is wrong or outmoded for certain groups: in particular, Brachodidae must be substituted for “Atychiidae,’’ Saturniidae for “Attacidae,” and Thyatirinae (Drepanidae) for “Tetheidae”
several “families” have to be downgraded to subfamily rank (or even to tribe rank): “Phyllocnistidae,” “Ethmiidae,”’ ““Galleriidae,” “Phycitidae,” “Pyraustidae,”’ “Lith- osiidae,’ “Ctenuchidae,” and so on.
*
The following section deals with about 30 families, for which are provided keys for the identification of genera and species (pp. 24-1024). For each family, one can find a general diagnosis and at least one illustrated key to species, based on external characters and genitalia. Frequently, there are two separate keys to species (one being based on external characters; the second, on male genitalia), and there may also be a third one, based on female genitalia. In addition, there is a key to genera (or to subfamilies and genera), unless the family contains only one genus in western Russia. By and large, all these keys are more accurate than the one to suborders and families. Although the book is not intended as a taxonomic revision, it provides useful data for each species (e.g., distribution and larval food plants), and includes some interesting proposals or suggestions: Eretmocera Zeller, often misplaced in recent literature, is rightly placed within the Scythridae (“Scythrididae’’); in the Oecophoridae sensu auct., Anchinia Hibner follows immediately Hypercallia Stephens, probably because A. L. Lvovsky regards these genera as close relatives, a point of view in agreement with my recent reappraisal of the gelechioid classification (I have transferred both genera to the Elachistidae Hypertrophinae: Alexanor 16:239-255); Uliaria Dumont is rightly considered a member of the Gelechiidae Di- chomerinae (the genus has been occasionally allocated to the “Autostichinae’’); etc. Of course, the adopted classification and nomenclature would also require a number of amendments: Odites Walsingham is definitely not a member of the Xyloryctidae [see,
76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
e.g., Hodges 1978: Moths Am. N. of Mexico 6(1):8-9]; Millieria Ragonot belongs to the Choreutidae, not to the Glyphipterigidae (see, e.g., Heppner 1982: Smithson. Contr. Zool. 370:1-27); Phyllonorycter Hiibner must be substituted for Lithocolletis Hiibner, Ypso- lopha Latreille for Ypsolophus Fabricius, Lecithocera nigrana (Duponchel) for Lecithoce- ra luticornella (Zeller), and so on.
A list of references (pp. 1025-1041) and an index of Latin names (pp. 1061-1092) conclude the book, along with a list of abbreviations of authors (pp. 1042-1043), and a useful list of botanical names, which gives the correspondence between common and Latin denominations (pp. 1044-1060). The present handbook is highly valuable because it includes a large portion of the European fauna of “Micromoths,” for which no equiv- alent, recent work has been published up to now. For instance, it takes into account a high percentage of the species known to occur in France: nearly 80% for Bucculatrix Zeller, more than 90% for Caloptilia Hubner and Ypsolopha Latreille, 90% for Ethmia Hubner, about 60% for the Scythridae, 93% for the Momphidae, 100% for the Choreutidae, etc.
In conclusion, this work should be included in the private library of any microlepi- dopterist interested in the Palaearctic fauna. It is to be wished that a similar translation will soon be available for Part Three (dealing chiefly with Pyrales and Plume Moths).
JoEL MINET, Muséum national d Histoire naturelle, Entomologie, 45 rue Buffon, F-75005, Paris, France.
Journal of the Lepidopterists’ Society 48(1), 1994, 76-77
BUTTERFLY GARDENING IN THE SOUTH: CULTIVATING PLANTS THAT ATTRACT BUTTER- FLIES, by Geyata Ajilvsgi. 1990. Taylor Publishing Company, Dallas, Texas. xi + 348 pp., 196 color photographs, 2 line drawings and 7 diagrams. Hard cover, 20.3 em x 28 cm, ISBN 0-87833-738-5. $34.95.
Novice and master gardeners in the southern U.S., particularly in the Rio Grande Valley, Texas, have an extraordinary treat in store with this volume. Ms. Ajilvsgi inves- tigates every aspect of a butterfly garden, from her brief introduction on the significance of butterflies and their means of survival to appropriate commercial and private sources for obtaining seed and other garden supplies. The brief forward by Chess Ezzell McKinney, Chairman, Preservation of Butterflies, National State Garden Clubs, sets the stage for this treasury of information, and, although the title focuses on butterfly gardening, the em- phasis is on gardening with a capital “G.”
There are special sections of the book devoted to creating a personal butterfly garden and to methods of attracting butterflies, highlighted by personal observations on such topics as the important characteristics of floral nectaries (color, shape and fragrance) and how to choose the appropriate plants, with one of my favorites—adopt a weed. Two other chapters detail the actual planning and planting of a butterfly garden. For the novice, or for butterfly watchers with limited space, there is the “instant” butterfly garden. For example, a fence, porch, or wall can accommodate hanging baskets or vines, and even the edges of driveways or window boxes can function as butterfly gardens. Special attention is given to the selection of plants, maintenance of the garden, and even to the introduction of caterpillars and chrysalids. For those lepidopterists with a grander vision in mind, a series of diagrams (pp. 78-83) provides garden plans for almost any geographic setting in the south. This thorough chapter on planning includes discussions of soils, preparation of the flower beds, and selection of appropriate plants, including native species, in addition to some forethought about flight patterns through the garden and appropriate areas for water and puddling stations. A chapter on “butterfly-friendly pest controls” emphasizes biological and physical controls, companion and repellent plants, and natural insecticides. Butterflies of south Texas, and especially of the Rio Grande
VOLUME 48, NUMBER 1 Tt
Valley, are featured in the special south Texas garden, which lists local butterflies and their associated larval hostplants based on the current literature and on the author’s own field observations.
Detailed analyses for 50 butterfly species that are easily visible in the field or backyard are included and organized according to the common name followed by the scientific name. Each diagnosis generally includes a color illustration, size, range, flight period, number of broods, overwintering stage, and brief descriptions of the adult, egg, and larva, and of the hostplant and parts of the plant consumed. Curiously, descriptions of the chrysalis are omitted. Favorite nectar sources, their bloom period, height class, cultivation, and other notes are also listed. Descriptions of adult behavior, especially favorite perches and interaction with nectar sources, are interspersed throughout this section.
Separate chapters treat larval hostplants and nectar resources, with the plants subdivided according to type (trees, vines, and herbs). These chapters feature both cultivated and native species with a special perspective on south Texas butterflies and with descriptions and accompanying color illustrations for 40 plants in each category. The chapter on larval hostplants provides information on their cultivation, associated larval taxa, and other notes. A longer list of all potential host plants gives the common name of their associated butterflies, the normal plant range, and the geographic area in which these plants are located in Texas (pp. 209-242). Similarly, the chapter on nectar sources lists the most appropriate yet common plants, their habitat association, and identification as a larval hostplant, if appropriate. An addendum list of nectar plants (pp. 281-311), including both cultivated and native naturalized species, is subdivided annuals versus perennials (herb), with special annotation indicating height, color and bloom period.
A wealth of other information can be found in this book, including basic advice and tips on the photography of butterflies. Other features include maps of U.S. hardiness zones and of the regional subdivisions of Texas considered. A compendium of appendices lists the major butterfly and plant organizations and societies and their associated publications, butterfly gardens and special events, and garden magazines and newsletters with articles on butterfly gardening. Other lists provide sources for additional information on habitat preservation, garden seeds, and supplies. The diverse bibliography supplies references on butterflies and other insects; garden, plant and wildflower guides; and photography. The butterfly and plant index is consistent in format, listing species by common name followed by the associated scientific name.
This book is exceedingly well researched and delightfully written. It is refreshing to read the author's personal observations on gardening and butterflies, especially those on behavior. For the taxonomist, the organization of the butterfly diagnoses is a little dis- concerting, with, for example, the Janais Patch (Chlosyne janais) and the Theona Check- erspot (Thessalia theona) interspersed among representatives of the Hesperiidae. Similar problems occur in the Lycaenidae.
Although the use of common names as the main reference point may be attractive to introduce the uninitiated gardener to butterflies, such names vary in use from one section of the country to another, leading to confusion. Thus, the Gray Calico and Gray-skirted Calico refer to the same beast, Hamadryas februa. One further minor hurdle concerns the type size of the index (6 pt.), which is generally in vogue for curatorial staffs but may be a little small for the average reader.
These few problems notwithstanding, this book is beautifully illustrated and provides a treasury of vital information on butterfly gardening. For anyone even remotely interested in southern gardening and butterflies, this book is absolutely essential reading.
JACQUELINE Y. MILLER, Allyn Museum of Entomology, Florida Museum of Natural History, 3621 Bay Shore Road, Sarasota, Florida 34284.
Journal of the Lepidopterists’ Society 48(1), 1994, 78-79
BUTTERFLY GARDENING: CREATING SUMMER MAGIC IN YOUR GARDEN, created by the Xerces Society in association with the Smithsonian Institution. 1990. Sierra Club Books, San Francisco, and National Wildlife Federation, Washington, D.C. xv + 192 pp., 118 color photographs, 4 diagrams. Softcover, 18.5 x 23.5 cm, ISBN 0-87156-615-X. $18.95.
The cooperative spirit of this volume is reflected not only in the educational and conservation organizations involved in its production, but also by the wealth of individ- uals—authors, naturalists, and photographers—who so generously donated their time and efforts. Butterfly Gardening is dedicated to the broad scale conservation of butterflies and their native food plants. It also celebrates the diversity of nature and encourages all lepidopterists to observe ecology in action. With the loss of native habitats, there also has been a noticeable disappearance of butterfly species, which serve as indicators of major changes in ecosystems.
A brief Introduction by Dennis Murphy focuses on some of the problems in butterfly conservation, suggests possible solutions, and sketches the contributions of the other nine authors of the book. Such a conservation approach to butterfly gardening would be incomplete without a contribution from the Honorable Miriam Rothschild, the eminent entomologist who has devoted so much time to the study of butterflies and other inver- tebrates. Dr. Rothschild’s keen insight has made naturalists more aware of conservation and of the significant role of wildflowers in ecosystems. Dr. Rothschild wrote three chapters for this book. The first describes the joys and challenges of Gardening with Butterflies; the second explores The Visual Perception of Lepidoptera and the importance of pro- viding appropriate nectar sources in butterfly gardens; the third describes The Life Cycle of the Large White Butterfly. Some life history aspects crucial to the survival of butterflies and moths in nature—metamorphosis, mate recognition, migration, temperature regu- lation, natural predators, and defense through camouflage and cryptic coloration and mutualism—are discussed by Dave Winter in his erudite chapter (The Struggle to Sur- vive). From a more personal perspective, Jo Brewer recounts her experiences as a butterfly gardener—providing descriptions of some of her more spectacular triumphs, sprinkled with the inevitable problems she encountered (Notes from a Butterfly Gardener). She also evaluates the propriety of introducing butterfly species into new habitats and discusses the use of native plant species versus exotic species in butterfly gardens.
Mary Booth, a landscape architect, with supporting text by Melody Mackey Allen, provides butterfly garden designs with emphasis on color. Four basic arrangements from simple to more sophisticated plans are provided and suggestions for flowering plants are included. Some do’s and don'ts with regard to conservation and to introduction of plant species that may be difficult to control are also considered. This chapter (Butterfly Garden Design) concludes with a master plant list of 30 flowering plants that are common nectar sources and, in some cases, larval hostplants. This annotated list provides a description of the type (shrub versus hardy perennial), height, color, bloom season, exposure, and soil requirements for each species. The food requirements for moths are not neglected owing to the thoughtful inclusion of Dave Winter’s chapter on nocturnal nectar sources (Moths and the Garden at Night).
Conservation issues concernign habitat are addressed in the chapter Wildflowers in the Planned Landscape by David Northington. As the Executive Director of the National Wildflower Research Center, he discusses the disappearance of plants and the resulting ecological consequences for animal species. The poignant discussion by Stanwyn Shetler in Butterfly Gardening and Conservation addresses these subjects from a naturalist’s viewpoint and delves further into the importance of plant/insect interactions. Dr. Shetler actively supports increased public education and awareness of conservation efforts by promoting gardening to teach preservation, rehabilitation, and restoration of diverse natural habitats.
The final chapters offer suggestions for enhancing personal enjoyment of butterfly
VOLUME 48, NUMBER 1 79
gardens, with excellent tips on close-up photography and butterfly observations by the incomparable Edward Ross (Enriching Your Personal Landscape and Butterfly Photog- raphy) and additional intriguing ideas for butterfly watching by Robert Michael Pyle. Following a short Afterword by Pyle are two appendices: a list of nectar plants for North American butterflies and moths and an annotated list of the most familiar North American butterflies and their larval food plants. Resource lists for obtaining garden plants and entomological equipment, a lengthy bibliography, and an index to both scientific and common names complete this useful volume.
An attractive glossy cover adds an exquisite touch to this extraordinarily illustrated volume. Superbly written, this book offers an excellent mixture of conservation philosophy and biological common sense with a focus on butterflies and moths. With such a com- pilation of authors, the book is anecdotal, and if there is any fault, it is that parts of some chapters may appear redundant. However, such duplicate treatment is refreshing, inas- much as different perspectives reflect the diversity of thinking on various subjects, such as appropriate nectar sources and problems encountered in rearing species. The Xerces Society and Smithsonian Institution are indeed to be commended for producing this volume. Through conservation, restoration, and management of native habitats initiated at the backyard level, we can increase public awareness of how complex yet fragile nature is and make a new beginning at protecting Lepidoptera.
JACQUELINE Y. MILLER, Allyn Museum of Entomology, Florida Museum of Natural History, 3621 Bay Shore Road, Sarasota, Florida 34234.
Journal of the Lepidopterists’ Society 48(1), 1994, 79-80
THE ONTARIO BUTTERFLY ATLAS, by Anthony M. Holmes, Quimby F. Hess, Ronald R. Tasker, and Alan J. Hanks. 1992. Toronto Entomologists’ Association, 34 Seaton Drive, Aurora, Ontario L4G 2K1, Canada. 167 pp., 13 color plates. Soft cover, 20.5 x 25.5 cm, ISBN 0-921631-11-1. Available from the Association for $29 Canadian (including taxes & p/h) or $26 U.S. (including p/h) (no Master-Card or Visa).
This attractively covered paper-back culminates the efforts of the Toronto Entomol- ogists Association (TEA) by recording the skippers and butterflies found in Ontario. The title may be somewhat misleading, however, as this publication is more of an annotated checklist of Ontario Rhophalocera.
The preface, brief author backgrounds, and table of contents are followed by an
introduction that describes the purposes of the Atlas: “to summarize ... the distribution and some characteristics of Ontario butterflies .. . , to encourage . . . others to explore the distribution ..., and as a reference for planning efforts to conserve ... rare species. ...”
This introduction also includes information on TEA, nomenclature, timetables, habitat, status (employing symbols of occurrence used by the Nature Conservancy of Canada), records, figures showing counties and districts, physiographic features (Hudson Bay low- lands to deciduous forest region in southwestern Ontario), life zones (Hudsonian, Cana- dian, Transition and Carolinian), and problem species. The latter category includes several butterflies treated as species by some authors and as subspecies by others; also, because some species of Erynnis and Satyrium are difficult to identify, some records may be inaccurate.
The main portion of the Atlas treats 138 species, including Basilarchia arthemis ar- themis and B. a. astyanax as separate entities. The authors make no attempt to differentiate subspecies except in a few cases. Each family is introduced with beautiful photographs of adults in natural settings or poses, with the exception of the Hackberry Butterfly on page 111 that suspiciously resembles a pinned specimen!
The authors devote a full page to each species, with brief notes on timetable, including broods and “hibernates,”’ and occurrence, including habitat, food plant, distribution, and
80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
status. Also included on the species page is an occurrence calendar-graph for each stage of the species and a map of Ontario showing distribution by dots placed in each county and district of occurrence. I wish the authors had included more biological information and “characteristics” of each species instead of devoting more than half a page to each distribution map. The 12 color plates include 2 habitat photographs and 27 individual photographs that show 7 larvae and pupa and 20 adults, representing 22 species.
Following the main species section is information on 19 stray species that have been vouchered by a single specimen (e.g., Erynnis zarucco) or rarely recorded (e.g., Speyeria idalia). It appears to me that these strays should have been included in the main species section, which already includes such strays as Pyrgus communis, Hylephila phyleus, Battus philenor, to name a few. Next is a section listing five unconfirmed species that may range into Ontario based on records from adjacent provinces and Michigan. This section might better have been called a hypothetical species list. The final species section has information on five doubtful species that have been erroneously referred to Ontario as a result of misidentification, improper labelling, or lack of a voucher specimen.
The last four pages of the Atlas comprise a bibliography, including check-lists, TEA publications, and general works, and a provincial ranking of status indicating number of occurrences within the province.
This publication is a valuable addition to the literature on Lepidoptera of the region from Hudson Bay to the Great Lakes, although there are a few questionable assertions. For example, the authors state that Lycaeides melissa samuelis hibernates “as a larva,” but according to James Scott (The Butterflies of North America, 1986, Stanford University Press, CA), “eggs hibernate in ssp. samuelis...” And for Danaus plexippus, the authors mention “positive evidence for a cycle of abundance peaking about every 11 years.” In both cases, the authors fail to include literature citations or to otherwise identify the source of this information.
This Atlas would have been more valuable if specific rearing experiences and personal food plant observations had been included. Furthermore, there is no discussion under L. m. samuelis or other rare or threatened species as to the authors’ recommendations of proposed management methods and techniques to preserve or enhance existing popula- tions. This shortcoming appears to be inconsistent with the authors stated purpose “to provide a reference for planning efforts to conserve our rare species. ...' Also, I found it interesting that the authors assert that the food plants of Erynnis persius are “willows, poplars and aspens.” In Michigan, persius feeds on lupine and most recent authoritative references state that lupine is the preferred larval food.
The Ontario Butterfly Atlas, with an 8 x 10” format, is printed on high quality paper with easy to read type and excellent photographs. I found no typographical errors or improper use of nomenclature. I recommend this publication to all students interested in the butterflies and skippers of Ontario and the Great Lakes region.
MOGENS C. NIELSEN, Adjunct Curator-Lepidoptera, Department of Entomology, Michigan State University, East Lansing, Michigan 48824.
Journal of the Lepidopterists Society 48(1), 1994, 80-82
OUT OF AFRICA
THE BUTTERFLIES OF KENYA AND THEIR NATURAL History, by Torben B. Larsen. 1992. Oxford University Press, Oxford, New York and Tokyo. xxii + 490 pp., 19 + A4 figures, 64 color plates. Hard cover, 18 x 25 cm, ISBN 0-19-854011-6. $195.00.
BUTTERFLIES OF TANZANIA, by Jan Kielland. 1990. Hill House, Melbourne and London. 363 pp., 3 maps, 179 text figures, 68 color plates. Hard cover, 22.5 x 30 cm, no ISBN. $120.00.
VOLUME 48, NUMBER 1 81
These are two rather comparable volumes devoted to the butterfly fauna of adjacent East African countries. Both are authoritative, even though Kielland’s volume represents many more years of sampling the Tanzanian fauna than does Larsen’s. Despite this shortcoming, however, Larsen has managed to produce a very readable and useful volume because he has consulted with many authorities and resident Kenyan collectors and has organized the field notes of the latter into an excellent compendium. Kielland’s book is no less well-documented and both volumes contain much useful information for either the collector or the butterfly-watcher.
The plates, of course, are the highlight of any butterfly book, and these are of top quality. The specimens illustrated are the best ones available to the authors: those in the Kenya book are presumably the best available at The Natural History Museum, London, whereas the specimens in the Tanzanian book are largely those available in Mr. Kielland’s collection, sometimes to the detriment of the Tanzanian work. The larger format of Kielland’s book makes possible the life-size illustrations of all taxa. Larsen’s book illustrates only half of each spread specimen for larger species and has more reduced figures, but where applicable, this fact is stated on each caption page.
The Kenya volume appears to have been proofread somewhat more stringently than does the Tanzanian one. A number of annoying “typos” in the latter distract the reader. For example, when one seeks the illustration for Bicyclus kiellandi Condamin, the plate on which it is figured is stated in the text (page 81) to be Plate 18; the butterfly is actually illustrated in Plate 17! Occasional inappropriate capitalizations of some, but not all, patronymial specific epithets occur throughout the text.
Both books use species citations that do not strictly conform to the Rules of Zoological Nomenclature because they do not place the authors’ names in parentheses when a species or subspecies was described in a genus other than the one in which it is presently contained. This habit, while maddening to the purist, seems to be gaining acceptance among authors, and perhaps it will no longer be required in future Rules.
Larsen is a superb writer who manages to use humor to its greatest advantage. He does not always write humorously, but rather interjects it occasionally. For example, on page 34, during a discussion of urine being attractive to butterflies, he provides this aside: “(When collecting in the tropics I often take a couple of Johnnie Walker bottles full of urine with me; whoever once stole two such bottles from my car must have been in for a surprise when he reached home).”’ I suspect that there is more useful information about habits of butterflies in the Kenya book, but both provide much useful data.
I have only two major criticisms, neither perhaps important in itself, but both of which are an annoyance. Both books contain a number of descriptions of new taxa, especially in the Kielland book, which make the books indispensable to the taxonomist. There seems to be a “conspiracy on the part of some authors to include such descriptions in books to enhance sales, but perhaps this not intentional. Steve Collins and Larsen do describe their new species in an appendix (pp. 438-445) almost apologetically, which seems to mitigate my objection mentioned above. Kielland does not so segregate his new names (as well as those of other authors), which are included throughout the text. Type specimens are illustrated in both books, thereby further increasing their utility to the specialist.
The second criticism applies only to the Kielland book, and it is not the author’s fault: the type faces employed throughout the book are so similar to one another (family-group names are in slightly larger type) that it is extremely difficult to locate information quickly or determine where one discussion ends and the next begins. In Larsen’s book, the main headings are much better accentuated, and one can tell at a glance what constitutes any species discussion. In addition, genera are well separated in that text, whereas in the Tanzanian volume there is no difference in typography between genera, species or sub- species.
These are really minor complaints, however, and both books are significant additions to the African butterfly literature. They are well-written and superbly illustrated, and though both volumes are somewhat expensive, they are well worth the money. If the reader has to choose only one of the volumes (they do cover roughly the same fauna of nearly 900 butterfly species), I would have to recommend Larsen’s volume over Kielland’s based chiefly on the clarity of the former’s presentation. Hopefully, no one will have to
82 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
choose and those interested in east African butterflies can have both volumes—the authors are to be congratulated for their accomplishments.
LEE D. MILLER, Allyn Museum of Entomology of the Florida Museum of Natural History, 3621 Bay Shore Road, Sarasota, Florida 34234.
Journal of the Lepidopterists’ Society 48(1), 1994, 82-83
A PRACTICAL GUIDE TO BUTTERFLIES AND MOTHS IN SOUTHERN AFRICA, by S. E. Wood- hall (co-ordinating editor) et al. 1992. Lepidopterists’ Society of Southern Africa, P.O. Box 470, Florida Hills, 1710, Transvaal, Republic of South Africa [Sponsored by Monsanto South Africa (Pyt) Ltd., Agricultural Group]. 223 pp., 48 color photographs on 8 plates, numerous black and white photographs and text figures. Softcover, glossy paper, 14.5 x 21 cm, ISBN 0-620-16774-2. Available from the Lep. Soc. So. Africa for $25 U.S. plus postage ($1 for surface mail or $20 for Airmail).
This techniques guide was compiled through the efforts of nineteen of the top lepi- dopterists in southern Africa. It is an easily totable resource (approximately 6” x 8” x %2") for almost every aspect of studying Lepidoptera, meant for use by beginners as well as by others who desire to learn more about techniques used in other countries. It is easy reading, even amusing at times, and the techniques appropriate for southern Africa are certainly usable here. Although the price is high for a paperback, the book is absolutely stuffed with information!
Chapter topics include the history of lepidopterists and lepidopterology in southern Africa, the lepidopteran life cycle, morphology and terminology, binomial nomenclature, species theory, evolution of Lepidoptera, a discussion on “lumpers vs. splitters, ’ taxonomic classification, and the rules that govern scientific nomenclature (ICZN). As expected in a work meant to address beginners, the book is amply illustrated with black and white photographs, line drawings, cartoons, and eight pages of color photographs depicting 48 live specimens of butterflies, moths (even some micros), and larvae which accompany the systematic classification of Lepidoptera and a brief overview of Lepidoptera.
A chapter devoted to Lepidoptera conservation in southern Africa presents the various habitat changes and impacts that man’s activities have had on insect populations and includes discussions of monitoring methods (conducting mark-recapture studies, hostplant monitoring, habitat monitoring). Also in this chapter is mention of the South African Red Data Book—Butterflies, a publication documenting 102 species considered to be exposed to some level of threat. This guide provides a table of 190 or so rare, endangered or vulnerable butterfly species cross tabulated with habitat, habits, distribution, taxonomy, food, and reproduction. Checkmarks in the columns denote a lack of information about a specific aspect of the particular butterfly. This table draws attention to those aspects unknown, with the idea that special efforts should be made to remedy these “unknowns.”
Collecting and field techniques for butterflies are detailed and are accompanied by photographs, cartoons, and line drawings. Included are instructions on the construction of equipment and the proper use of nets, traps, decoys and baits, killing methods, and specimen storage in the field, along with suggestions for observing hilltopping, territo- riality, and egg laying behavior. Especially interesting are the instructions for preparing rotten shrimp bait, various fruit baits, and different combinations of scat and urine to attract specimens. Apparently, the fresher the dung, the better (baitwise). However, a cautionary note is included to remind the lepidopterist to keep a sharp lookout for the originator of the dung; e.g., lions, elephants, baboons, etc. Discussion of moth collecting includes traps (many diagrams and photos), sugaring, and the collecting of immatures.
Preservation and mounting techniques are covered thoroughly. Relaxing techniques (chambers, injection, cutting wing muscles), selecting pin sizes, pinning specimens, con-
VOLUME 48, NUMBER 1 | §3
struction of various spreading boards, use of various materials as “setting strips,” and setting pins are all discussed. The actual spreading technique for butterflies and macros is described in detail, with many cautions included for the inexperienced. There are also instructions for baking (=drying) specimens while on the spreading board. Although the description of techniques for pinning micros is brief, detailed instructions are given for preserving and mounting early stages, and for dissection and preservation of genitalia, palpi, legs, androconia, and larval head shields. Degreasing, dust removal, mold removal, repairing damaged specimens, housing the collection, prevention of infestation, proper labelling, and hints for successful mailing of specimens are also included in this chapter.
Other topics covered in this guide include rearing and breeding Lepidoptera (cages, sleeving, foodplants, artificial diets, and larval diseases) and photography in the field and in the studio (backdrops and composition, as well as proper storage of photographs). There are chapters on good scientific practice, the British Butterfly Conservation Society Code of Practice, several appendices (providing addresses of equipment dealers, names and addresses of journals worldwide that accept Lepidoptera papers, reviews of threatened species in southern Africa), a glossary of terms, a bibliography, and an index.
Throughout the book the authors encourage all lepidopterists to expand and further the science of lepidopterology. Emphasis on observations in the field and the elucidation of life histories are recurring themes in this handbook that espouses a common-sense approach to the study of Lepidoptera. Amateur lepidopterists are hailed for tirelessly providing valuable information, continually taking on the onerous task of breeding and recording life histories, and contributing much of what is known about Lepidoptera. The debate over collecting vs. no-collecting is broached. Admitting that it is an emotionally charged issue, the authors agree that general bans on collecting are detrimental to the conservation of Lepidoptera and of nature as a whole. Conservation is promoted and responsible collecting is encouraged. The point is made that the reproductive capabilities of insects far outweigh the impact of the small numbers collected by lepidopterists. A reminder to all is that the younger generation should be encouraged to study invertebrates; that first-hand collecting and field experiences, besides being enjoyable, entice children to become more aware of nature and more able at a later date to make informed judgements on conservation matters; and that if “... no more children are starting at the basics, one day there will be no lepidopterists to be consulted on conservation.”
I found the book to be an inspiring and complete guide to handling Lepidoptera, written in an easy to understand, often entertaining manner. It was rewarding to read a techniques book that continually encouraged and lauded the efforts of avocational lepi- dopterists. It sets an excellent example for our own future Lepidopterists’ Society Tech- niques Manual.
STEPHANIE McKown, 650 Cotterell Drive, Boise, Idaho 83709.
Journal of the Lepidopterists’ Society 48(1), 1994, 84
MANUSCRIPT REVIEWERS, 1993
The merit of a scientific journal depends on the quality of its reviewers as well as of its authors, but the former are usually unknown to readers. The Journal relied on the expertise of 68 reviewers last year to provide 89 evaluations of manuscripts. It is with much gratitude that the Journal acknowledges the services of the people listed below from whom manuscript reviews were received in 1993.
Phil R. Ackery, London, England Annette Aiello, Panama
Paul H. Arnaud, Jr., San Francisco, CA *George T. Austin, Las Vegas, NV
Susan S. Borkin, Milwaukee, WI *M. Deane Bowers, Boulder, CO
Andrew V. Z. Brower, Ithaca, NY
Lincoln P. Brower, Gainesville, FL
*Keith S. Brown, Sao Paulo, Brazil
*Richard L. Brown, Mississippi State, MS
*John M. Burns, Washington, DC
John V. Calhoun, Dunedin, FL
Reginald Chapman, Tempe, AZ
Frances S. Chew, Medford, MA
Ian F. B. Common, Toowoomba, Queensland, Australia
Charles V. Covell, Jr., Louisville, KY
P. T. Dang, Ottawa, Ontario, Canada
Don R. Davis, Washington, DC
John J. Dayton, Santa Cruz, CA
*John A. De Benedictis, Davis, CA
*Philip J. DeVries, Austin, TX
Julian P. Donahue, Los Angeles, CA
*Boyce A. Drummond, Woodland Park, CO
Thomas D. Eichlin, Sacramento, CA *John F. Emmel, Hemet, CA Marc Epstein, Washington, DC
David K. Faulkner, San Diego, CA Douglas C. Ferguson, Washington, DC Clifford D. Ferris, Laramie, WY
Lawrence F. Gall, New Haven, CT *Glenn A. Gorelick, Glendora, CA
David L. Hancock, Brisbane, Queensland, Australia
John B. Heppner, Gainesville, FL
Ronald W. Hodges, Washington, DC
* Reviewed two or more manuscripts.
Daniel Janzen, Philadelphia, PA Kurt Johnson, New York, NY
Niels P. Kristensen, Copenhagen, Denmark
Gerardo Lamas Muller, Lima, Peru Ron Leuschner, Manhattan Beach, CA
*C. Don MacNeill, San Francisco, CA Deborah L. Matthews, Gainesville, FL Tim L. McCabe, Albany, NY Jacqueline Y. Miller, Sarasota, FL William E. Miller, St. Paul, MN
Raymond W. Neck, Houston, TX Mogens C. Nielsen, East Lansing, MI
Robert W. Poole, Washington, DC *Jerry A. Powell, Berkeley, CA
Frederick H. Rindge, New York, NY *Robert K. Robbins, Washington, DC
Walter H. Sakai, Santa Monica, CA David A. Schooley, Reno, NV *James A. Scott, Lakewood, CO Arthur M. Shapiro, Davis, CA Oakley Shields, Mariposa, CA
John A. Shuey, Traverse City, MI Stephen M. Spomer, Lincoln, NE Stephen R. Steinhauser, Sarasota, FL Ray E. Stanford, Denver, CO *Stephen Stone, Lakewood, CO
Thomas Turner, Clearwater, FL Paul M. Tuskes, San Diego, CA
R. I. Vane-Wright, London, England Adrienne Venables, Washington, DC
*David L. Wagner, Storrs, CT
Warren H. Wagner, Jr., Ann Arbor, MI
David A. West, Blacksburg, VA
William D. Winter, Dedham, MA
Date of Issue (Vol. 48, No. 1): 23 February 1994
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Associate Editors: M. DEANE BOwERS (USA), BoYCE A. DRUMMOND (USA), LAWRENCE F. GALL (USA), GERARDO LAMAS (Peru), ROBERT C. LEDERHOUSE (USA), ROBERT K. ROBBINS (USA), CHRISTER WIKLUND (Sweden)
NOTICE TO CONTRIBUTORS
Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Technical Comments, Book Reviews, Obituaries, Feature Photographs, and Cover Illustrations. Reviews should treat books published within the past two years. Obituaries must be authorized by the President of the Society. Re- quirements for Feature Photographs and Cover Illustrations are stated on page 111 in Volume 44(2). Journal submissions should be sent to the editor at the above address. Short manuscripts concerning new state records, current events, and notices should be sent to the News, Stephanie McKown, Editor, 650 Cotterell Drive, Boise, Idaho 83709 U.S.A. For information regarding book reviews, contact Boyce A. Drummond, Natural Perspectives, P.O. Box 9061, Woodland Park, CO 80866. Journal contributors should submit manuscripts in triplicate, typewritten, entirely double-spaced, with wide margins, on one side only of white, letter-sized paper. Prepare manuscripts according to the following instructions, and submit them flat, not folded.
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SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 209 pp.
196la. Some contributions to population genetics resulting from the study of
the Lepidoptera. Adv. Genet. 10:165-216.
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CONTENTS
PRESIDENTIAL ADDRESS 19938: ON THE COMPARATIVE DISTRIBU- TIONS OF LEPIDOPTERA AND LEPIDOPTERISTS. Ray E. Stan- : fot tc ee Reh Sa RS Nb i iin eC eg ]
A NEW GENUS OF WINTER MOTHS (GEOMETRIDAE) FROM EASTERN CALIFORNIA AND WESTERN NEVADA. Jerry A. Powell and Douglas C: Ferguson: 22) 600s a 8
SPLIT SKIPPERS: MEXICAN GENUS POANOPSIS GOES IN THE ORIG- ENES GROUP—AND YVRETTA FORMS THE RHESUS GROUP— OF POLITES (HESPERIIDAE). John M, Burns ..3) 02a 24
A NEW SPECIES OF ACROLEPIOPSIS AND THE DESCRIPTION OF THE FEMALE OF A. CALIFORNICA (ACROLEPIIDAE). Reinhard
Gaedike: (00 co RS OO SS A6 A NEW SPECIES OF LAPARA (SPHINGIDAE) FROM SOUTHEASTERN UNITED STATES. Vernon Antoine Brou Jr. 2) ol PROFILE William Wittfeld: The Florida connection. John V. Calhoun 58 GENERAL NOTES Musineon tenuifolium (Apiaceae): New host of four Nebraska Papilio (Pa- pilionidae). “J. M. Reiser and S. M. Spomer, 1... oo) ee 68 Two new synonymies in Nearctic Eucosma (Tortricidae: Olethreutinae). Loran D. Gibson and William E. Miller 69 Additional data on the geographical distribution and adult activity of the diurnal, mimetic plume moth, Oidaematophorus chamelai (Pterophor- idae).\J. A. Powell and J. AsChemsak: 2 00 2) 3 ee 71 Book REVIEWS Keys to the insects of the European part of the USSR. Volume IV (Lepidoptera), Part T1.. \Joel Mimet fo 2G00h oo Von Ss Oh IY AeA AS a 74 Butterfly gardening in the south: Cultivating plants that attract butterflies. Jacqueline Y. Millere isco s0 285 0 VE 76 Butterfly gardening: Creating summer magic in your garden. Jacqueline Y. Miller: eee yk MN ete CUD SC 78 The Ontario’ butterfly atlas» “Mogens, C. Nielsen (00... eee 79 The butterflies of Kenya and their natural history. Butterflies of Tanza- nial ee DD. Miller: 3000 Val 80 A practical guide to butterflies and moths in southern Africa. Stephanie ME CK Otay ces S88 aN eh nl UES Saal eee lad ce POEL Fo MANUSCRIPT REVIEWERS, 199800 oO 84
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
q Volume 48 1994 | Number 2
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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
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17 May 1994
THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE COUNCIL
PAUL A. OPLER, President JORGE E. LLORENTE-BOSQUETS, Ray E. STANFORD, Immediate Past Vice President
President FREDERICK W. STEHR, CHEN-SHING LIN, Vice President Vice President MICHAEL J. SMITH, Secretary ROBERT J. BORTH, Treasurer Members at large: Charles V. Covell, Jr. Eric H. Metzler John V. Calhoun Linda S. Fink Robert K. Robbins Robert C. Lederhouse Scott E. Miller J. Benjamin Ziegler William E. Miller
EDITORIAL BOARD
PAUL A. OPLER (Chairman), FREDERICK W. STEHR (Member at large) JOHN W. BROWN (Journal), WILLIAM E. MILLER (Memoirs) STEPHANIE S. MCKOWN (News)
HONORARY LIFE MEMBERS OF THE SOCIETY
CHARLES L. REMINGTON (1966), E. G. MUNROE (1978), ZDRAVKO LORKOVIC (1980), IAN F. B. COMMON (1987), JOHN G. FRANCLEMONT (1988), LINCOLN P. BROWER (1990), DOUGLAS C. FERGUSON (1990), HON. MIRIAM ROTHSCHILD (1991), CLAUDE LEMAIRE (1992)
The object of the Lepidopterists’ Society, which was formed in May 1947 and for- mally constituted in December 1950, is “to promote the science of lepidopterology in all its branches, .... to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures’ directed towards these aims.
Membership in the Society is open to all persons interested in the study of Lepi- doptera. All members receive the Journal and the News of the Lepidopterists’ Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. There are four numbers in each volume of the Journal, scheduled for February, May, August and November, and six numbers of the News each year.
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Send remittances, payable to The Lepidopterists’ Society, to: Robert J. Borth, Treasurer, 6926 North Belmont Lane, Fox Point, WI 53217, U.S.A.; and address changes to: Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007- 4057 U.S.A. For information about the Society, contact: Michael J. Smith, 7428 Holworthy Way, Sacramento, CA 95842-4165. To order back issues of the Journal, News, and Memoirs, write for availability and prices to the Publications Manager: Ronald Leuschner, 1900 John St., Manhattan Beach, CA 90266-2608, U.S.A.
Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for $40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Second-class postage paid at Los Angeles, California and ad- ditional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. If you have used a Society address many years after its publication date, and your mail is returned as undeliverable, please redirect it to the Natural History Museum address provided above.
Cover illustration: The buckeye butterfly (Junonia coenia) is a common and widespread species of North America. Original drawing by Marianne C. Filbert, P.O. Box 155, Logan, Utah 843821.
JOURNAL OF
Tue LeEerPIDOPTERISTS’ SOCIETY
Volume 48 1994 Number 2
Journal of the Lepidopterists’ Society 48(2), 1994, 85-105
SAMPLING STRATEGIES FOR ESTIMATING MOTH SPECIES DIVERSITY USING A LIGHT TRAP IN A NORTHEASTERN SOFTWOOD FOREST
A. W. THOMAS
Canadian Forest Service, Natural Resources Canada, P.O. Box 4000, Fredericton, New Brunswick E3B 5P7, Canada
AND
G. M. THOMAS University of New Brunswick, Fredericton, New Brunswick E8B 6C2, Canada
ABSTRACT. A 22-watt black-light trap was operated for 29 nights within a forest canopy in the Maritime Lowlands Ecoregion of the Acadian Forest. The species-abun- dance frequency distribution (pattern of species abundance) was a good fit to the log series model and this model was used for subsequent data analysis. No single-night sample adequately estimated the log series alpha index of diversity based on the total catch; some sampling effort was required each night. Each night’s catch was separated into 16, 30- minute samples. The alpha index of diversity for the summed catch for each time-period was compared with the overall alpha based on the total catch. A strategy that involved operating the trap for just a 1-hour period each night had no effect on the pattern of species abundance and gave a value for alpha equal to that obtained by operating the trap for an 8-hour period each night. This strategy reduced the catch from 6088 to 971 moths and the number of species from 255 to 161. Processing costs associated with the larger sample and any possible negative effect on the moth population caused by removal trapping were greatly reduced. This new sampling strategy is thus useful for comparing indices of species diversity between several sites when data are collected simultaneously, but is of limited use for species-inventory studies.
Additional key words: species-abundance distribution, 30-minute samples, log-series model, partial-night sampling.
In recent years, the challenge to maintain biodiversity on this planet has become a major public concern. Most attention focuses on Neo- tropical ecosystems (Mares 1992). However, the importance of main- taining Canada’s biodiversity was addressed in Environment Canada’s Green Plan (Hyslop & Brunton 1991), and the launching, in 1991, of “Canadian Biodiversity’’ produced by the Canadian Centre of Biodi- versity at the Canadian Museum of Nature lends credence to the recent
86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
national interest in biodiversity. The values of maintaining current biodiversity have been stated by many authors, and summarized by Ehrlich (1990) into ethical, aesthetic, economic, and “ecosystem ser- vices.’ Salwasser (1990) added the legal obligation for conserving bio- logical diversity. Intimately linked with the concept of maintaining biodiversity, and especially protection of areas rich in species, is the need for a “quick and dirty survey to chart biodiversity of the planet” (Roberts 1988), a view reiterated by Ehrlich (1992). The ‘quick and dirty’ approach does not advocate poor science; rather, it recognizes that the scope of diversity from individual gene systems through pop- ulations of species, communities, ecosystems, and ultimately all life in the biosphere (Wilson 1988) cannot be addressed in the short-term. It suggests that studies should be focused on certain taxonomic groups over an extensive area. The hope is that areas with many species or high endemism in the selected groups will reflect similarly high values for other groups (Roberts 1988). Because of logistic and knowledge constraints, the number of species within a community can be deter- mined for only a limited number of taxonomic groups.
This study addresses just one segment of biodiversity, i.e., the diversity of moths in a single ecosystem. Diversity is used here to mean the number of species and their relative abundance (Magurran 1988), and to prevent ambiguity we will always use ‘species diversity’ where appropriate. Relative abundance is considered in the form of species-abundance frequency distributions, which show the relationship between the abun- dance of individuals and the number of species possessing that abun- dance (May 1975); abbreviated in this paper as the pattern of species abundance. The ecosystem studied is one locality in the Maritime Low- lands Ecoregion of the Acadian Forest (Loucks 1962).
The use of the moth community, in the 15 families used in this study (see Appendix), as an exemplar of the species diversity of this ecosystem has advantages that include the relative ease of identification at the species level, the somewhat standardized sampling methodology (Wil- liams 1951, Williams et al. 1955, Taylor & French 1974, Bowden 1982), and the high correlation of insects, in general, with the spatial, archi- tectural, and taxonomic diversity of plants (Southwood et al. 1979).
No community consists of species of equal abundance (Magurran 1988). It is normally the case that the majority of species are rare while a number are moderately common with the remaining few species being very abundant (Williams 1964, May 1975, Pielou 1975, South- wood 1978, Magurran 1988). Within this general distributional form, communities have characteristically different patterns of species abun- dance which remain stable despite changes in species composition (Pie- lou 1975, May 1976, Kempton 1979). The pattern of species abundances
VOLUME 48, NUMBER 2 87
at a site allows for comparison with similar sites that have different mixes of species, and a change in the pattern of abundance at one site has been shown to be a useful indicator of environmental disturbance (Kempton & Taylor 1974, Taylor et al. 1978, Kempton 1979).
Four main species-abundance models (the geometric series, the log- arithmic series, the log normal distribution, and MacArthur’s broken stick model) have been developed to describe species diversity in terms of an ‘index parameter’ as well as the pattern of species abundance. In addition there are several non-parametric indices based on the pro- portional abundances of species (May 1975, Southwood 1978, Magurran 1988).
The log series model was the first to describe the pattern of species abundance (Fisher 1943). Since then it has been found to have a wide application for catches of many invertebrates, e.g., moths in light traps (Williams 1948, 1945, 1964, Taylor & Brown 1972, Taylor & French 1974, Kempton & Taylor 1974, Taylor et al. 1976, 1978, Taylor 1986), Ichneumonidae (Owen & Chanter 1970), cockroaches (Wolda 1983), Psocoptera (Broadhead & Wolda 1985), Hymenoptera (Noyes 1989), and the community of phytophagous arthropods on apple (Brown & Adler 1989). Its wide applicability is because it is based on the abun- dances of the species with medium abundance rather than the very abundant and very rare species (Taylor et al. 1976, Kempton 1979, Brown & Adler 1989).
The log series is a simple two-parameter model, with two defining multispecies population parameters, chi and alpha. Chi is devoted to sample characteristics and varies with sample size as it is a function of the mean number of individuals per species. Alpha is independent of sample size and characterizes the required population quality (Kempton & Taylor 1974). Fisher’s (1948) initial suggestion was that alpha might ‘be useful as a measure of ‘species richness’ when comparing samples. Williams (1943) suggested that the parameter alpha be known as a community’s ‘index of diversity.’ Later he recognized that this term was applicable to other functions having the same properties and re- ferred to Fisher’s alpha as ‘diversity calculated on the basis of the logarithmic series’ (Williams 1964). The log series model can be derived from two statistics, S, the total number of species, and N, the total number of moths. It is a discontinuous frequency series with an infinite number of terms:
Me GK 2 Sp IK Aas oe, where n, is the number of species with 1 individual and successive
terms with 2, 3, 4, etc. individuals, and x (chi) is a constant <1 (Williams 1947).