Volume 2

1 January 2000

Number 1

77 ?<? Taxonomic Report ^0

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

A NEW ELFIN BUTTERFLY (LYCAENIDAE: EUMAEINI)

FROM NORTHERN CHINA

WITH COMMENTS ON THE NOMENCLATURE OF PALAEARCTIC ELFINS

KURT JOHNSON

Environmental Department, The Ethical Culture Society,

53 Prospect Park West, Brooklyn, New York 11215

ABSTRACT. Ahlbergia hsui, new species, is described from two specimens recently collected in China’s Nan

Shan area of endemism located along the boundary of northern China with Mongolia. In discussing the new species, the

historical literature concerning Palaearctic elfin butterflies is reviewed and compared to nomenclatorial usages in D’Abrera’s

widely disseminated 1993 treatment of Palaearctic butterflies; 25 errors or omissions in the latter treatment are corrected.

Additional key words. Species group placement.

INTRODUCTION

In July of 1999 Dr. Yu-Feng Hsu collected two specimens of a tailless elfin butterfly (Lycaenidae;

Eumaeini) of the genus Ahlbergia Bryk (Figs. 1-2) in the Gansu Province of north-central China. In the

context of my monograph of Palaearctic elfin butterflies (Johnson 1992), these individuals clearly represent

a previously unknown species.

Dr. Hsu's specimens are from the northern “Nan Shan” area of endemism (Johnson 1992) located

along the boundary of northern China with Mongolia (Figs. 5-6). Of the five areas of endemism noted for

Palaearctic elfin butterflies in my 1992 monograph, the Nan Shan region is the least explored. Previously,

the Nan Shan was known to have only three endemic elfin species: one from the diverse genus Ahlbergia ,

and two from the more primitive elfin genus Cissatsuma Johnson. Dr. Hsu’s discovery adds a fourth

endemic and second Ahlbergia to the elfin fauna of this region.

This new species is typical of previously unknown elfins from isolated areas of China with poorly

known butterfly faunas in that its wing color and pattern, and genitalic features are unique and usefully

comparable (as summarized in the description below) only to disparate elfin species from elsewhere in the

eastern Palaearctic Region. Terminology follows Johnson (1992) as reviewed herein in Figure 3.

Ahlbergia hsui Johnson, new species

Diagnosis. Currently known from only the female. Wings (Figs. 1-2 and 3): readily recognized by medium

size (FW alar 13.5 mm.), dorsum shiny pewter blue from wing base through postmedial area, thereafter abmptly

black; venter simply marked: over light beige ground color, FW with single, narrow and jagged, deep brown

postmedial band crossing entire wing; HW with highly contrasted, hoary and dark brown, basal disc followed distally

by lighter beige devoid of any outstanding markings in postmedial or limbal areas. Comparisons : among Palaearctic

Figs. 1-2 (d/v), 9 holotype Ahlbergia hsui Johnson. CHINA: GANSU Prov., [Longnan Dist.], Kang Xian, ca 1300 m, 8

July 1999, Y.F. Hsu, collector. Fig. 3, basic ventral (except b) wing pattern terminology from Johnson (1992): a.

elfins, the dorsal markings of A. hsui resemble A. chalybeia Riley (b and c areas of endemism, Fig. 5) a much larger

hsui resemble only A. bimaculata Johnson (b area of endemism, Fig. 5) a species with both sexes showing completely

r:rr.

lamella antevaginalis occurring only as a slight, secondary, lip-like structure,

unifurcate spine-like signa typical of numerous congeners.

lamella antevagionalis (Fig. ^ 3al) ^contrasing an insignificam lameha postvaginalis comprised of only a few,

DISCUSSION

Palaearctic Elfin Butterflies in the Diverse Scientific Literature of the World

In 1992 I published a thorough revision of the elfin butterflies of the Palaearctic Realm ( Ahlbergia ,

Cissatsuma, and Novosatsuma Johnson, 1992). Because of price, and European imprimatur, this monograph

was not widely distributed outside of institutional libraries among the world’s lepidopterists. In 1993

D’Abrera published a photofolio of Palaearctic Lycaenidae which has since been widely distributed. Even

though D’Abrera had been furnished a copy of Johnson 1992 while putting together his photofolio, he gave

only a cursory treatment of the region’s elfin butterflies, overlooking 21 species, and either misspelling or

wrongly attributing names to three others. Elfins, and other unglamorous lycaenids, are as scientifically

3

important as any other Lepidoptera. It is therefore imperative that these errors be corrected. Thus, the

corrective comments which follow are not meant as an attack on D’Abrera’s useful photofolio work, but are

only presented to correct his errors as required by the ICZN Code. Further, it is likely that had D’Abrera’s

publication not followed so soon after mine, he would have avoided many of these problems.

D’Abrera (1993: 436-437) did not recognize that, due to actual publication date, Ahlbergia Bryk

has priority over Ginzia Okano. He did state, however, that as of the time of his writing, the problem of

priority in generic names for elfin butterflies had not been “stabilised” [sic].

The following species were omitted by D’Abrera even tough their type material is at The Natural

History Museum (NHM) in London: Ahlbergia : bimaculata Johnson, korea Johnson, leei Johnson, arquata

Johnson, unicolora Johnson, caerulea Johnson, prodiga Johnson, caesius Johnson, lynda Johnson;

Novosatsuma : collosa Johnson, magnasuffusa Johnson, plumbagina Johnson, oppocoenosa Johnson,

magnapurpurea , Johnson, cibdela Johnson, monstrabila Johnson; Cissatsuma'. halos a Johnson, crenata

Johnson.

The following species were omitted by D’Abrera but do not have type material at the NHM:

Ahlbergia: pictila Johnson, haradai Igarashi; Novosatsuma : matusiki Johnson.

Given the fact that many of these species are spectacularly distinctive, D’Abrera may have

inadvertently overlooked the NHM drawers in which their type material is deposited. Or, he may not have

had the time to include them in his photography schedule. This appears likely because D’Abrera did include

the following three species described by Johnson with type material at the NHM: Cissatsuma tuba , C.

contexta, and Ahlbergia kansuensis.

D’Abrera misspelled the species name A. chalybeia as ‘ Ginzia chalybaea Two other names,

Ginzia “wells orum” and Ginzia “kimi were introduced by D’Abrera into the literature as nomen nudum

because they have never been described and, as such, have no validity under the International Code for

Zoological Nomenclature (ICZN Code). It should be noted that the specimen identified by D’Abrera as

“wellsorum ” is an A. bimaculata and the specimens noted by D’Abrera as “ kimi ” are A. korea.

Species Group Placement of A. hsui

The rudimentary invaginations comprising the lamella postvaginalis suggest relationship to the

diverse northern ferrea species group (Johnson 1992), in which such invaginations are strongly sclerotized

into a prominent lamellae postvaginalis. Since, due to their cryptic colorations, there is extreme homoplasy

in wing pattern and coloration in all elfin-like Eumaeini throughout the world, I tend to follow morphology

for placing species in groups. Accordingly, I would tentatively place A. hsui in the ferrea group noting that

its distribution represents a southern “outlier” for this group and its morphology may be primitive -

comprising characters of both the more southern chalybeia group and more northern ferrea group.

ACKNOWLEDGMENTS

Thanks are due Dr. Yu-Feng Hsu for providing the type material for study and description, and to

Ron Gatrelle (International Lepidoptera Survey) for photo processing of holotype for color reproduction.

LITERATURE CITED

D’ABERA, B. 1993. Ginzia Okano, pp. 436-437 in: Butterflies of the Holarctic Region. III. Nymphalidae

(concl.), Libytheidae, Riodinidae, and Lycaenidae. Hill House, Victoria (Australia).

JOHNSON. 1992. The Palaearctic “Elfin” Butterflies (Lycaenidae, Theclinae). Neue Entomologische

Nachricten 29. 142 pp.

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

TILS Purpose. TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific

discovery, determination, and documentation, without which there can be no preservation of Lepidoptera.

TILS Motto. As a world community, we can not protect that which we do not know. _

The Taxonomic Report is projected for publication at the rate of 10 issues a year. Subscription/

dues for Volume Two are $50 US for domestic and $65 US for overseas subscribers. The subscription year

follows the calendar year. All issues are mailed 1 st class. At the end of each year, subscribers receive that

year’s volume on a record-only compact disc (CDR) for permanent archiving and reproduction for personal

use (i.e. a museum or university may make as many copies as needed in whatever format desired). Non¬

members may receive individual issues in print any time for $10 per issue. Individual issues on CDR to

non-members are $25 per issue post paid. Subscriptions and individual issue orders should be made

payable to TILS; and mailed to: Scott D. Massey, Editor, 126 Wells Road, Goose Creek, SC USA 29445-

3413.

Articles for publication are sought. They may deal with any area of taxonomic research on

Lepidoptera. Before sending a manuscript, simply write TILS at the above address to set up discussion on

how to best handle your research for publication.

Everyday around the world, in jungles and urban areas alike, insect species and subspecies are

becoming extinct. Every year scores of taxa are lost that have not even been scientifically discovered and

documented. Thus, their extinction is unnoticed because their existence is unknown. They are unknown

simply because they have not been collected and systematically identified. Without systematic taxonomy

there is nothing. Without the collection, and exchange of specimens (information) there will be no

systematic taxonomy. Without amateur collectors the majority of the undiscovered species/subspecies will

vanish before they are discovered. Be it butterflies or moon rocks, collecting is the first step of access to all

other scientific information - and protection.

Donations are needed to support and further our efforts to discover and protect

butterflies worldwide. All donations are US tax deductible. Please help generously.

Donations should be mailed to: Scott D. Massey, Treasurer, 126 Wells Rd., Goose Creek, SC 29445.

Checks should be made payable to: TILS. Please indicate if you need an individual receipt.

TTR thanks Mr. Bryant Mather, 213 Mt. Salus Dr., Clinton, Mississippi 39056, for editing the text of this i

Volume 2

15 July 2000

Number 2

DESCRIPTION OF A NEW SUBSPECIES OF POANES AARONI

(HESPERIOIDAE: HESPERIINAE) FROM THE WEST CENTRAL

GULF COAST OF THE SOUTHERN UNITED STATES

RONALD R. GATRELLE 1

126 Wells Road, Goose Creek, South Carolina 29445

ABSTRACT. Poanes aaroni bordeloni is described from Jefferson County, Texas. This taxon is part of a

lepidopteran community endemic to the west central area of the US Gulf Coast between Mobile Bay, Alabama and Galveston

Bay, Texas. As such, bordeloni is defined taxonomically by its evolutionary development as much as by its facies. Bordeloni

is the largest aaroni subspecies. Bordeloni males are marked much like those of P. a. howardi both dorsally and ventrally.

Bordeloni females usually have the fulvous spotting on the dorsal forewings reduced as in P. a. minimus females while the

fulvous on their upper hindwings is more extensive than P. a. howardi females. Morphologically, the forewing outer margin

in both sexes of bordeloni is distinctly straighter and much less rounded than in the other aaroni subspecies. The bordeloni

holotype is deposited at Texas A & M University. Lectotypes are designated for Pamphila aaroni Skinner 1890 and

Pamphila howardi Skinner 1896 from specimens in the collection of the Carnegie Museum (NH), Pittsburgh.

Additional key words. Phenosyncronic subspeciation.

SOME MECHANICS OF LEPIDOPTERAN TAXONOMY

Species and Subspecies are Evaluated Differently

Lepidoptera populations are determined to be individual species when they meet certain objective

criteria - reproductive isolation, genetics, genitalia, mate location dynamics, host parameters, etc.

Evolutionary theory is not a factor in determining speciation because species (with or without subspecific

components) are the product of an evolutionary process past - a definitive plateau in the process. Thus,

when properly done, the taxonomic placement of populations at the species level is self evident and

unarguable because it is based solely on observable facts.

Conversely, the taxonomic placement of populations of Lepidoptera at the subspecific level must

deal with the past, present, and future of the organism’s evolution because subspecies are only subjectively

defined components of a species. Where phenetically (morphologically, biologically, ecologically, etc.) is a

subspecific population going relative to where it has been in its association with the other components of

the species. Evolutionary theory is a major factor in determining subspeciation because sub-species (as

sub-components) are the most observable manifestation of a species active evolutionary process - a semi¬

definitive step (toward stagnation/extinction or adaptation/speciation) in the plateau process.

Thus, at the subspecific level, the taxonomic classification of Lepidoptera is based largely on a

researcher’s assessment (hypothesis) of a population’s evolution. Therefore, the subjective analytical process

by which the researcher’s assessment was made is the primary element that must be weighed in determining

the validity of the proposed subspecific taxonomic conclusion - not some group of preset criteria. The

1 Research Associate Florida State Collection of Arthropods, Gainesville, Florida.

exception is non-clinal subspeciation where the set criteria of isolation is usually factored in. However, this

isolation can be established through either seasonality or geography. 2 Subspecific classification is

therefore often arguable because it is always subjective to some degree.

Mechanically, species level taxonomy can be accomplished without any evolutionary theory by

objectively weighing an organism against certain generally accepted criteria, while subspecific taxonomy

can not be properly accomplished without a subjective understanding (theory) of an organism’s evolutionary

past, present, and future. Species and subspecies are evaluated differently.

Thus, a taxonomist who utilizes the same system of analysis (either objective preset criteria or

subjective process and assessment) for both the specific and subspecific levels will inherently be in error at

one. This manifests itself in one of two ways: 1) A taxonomist will describe a population as a new species

but without sufficient proof because the research is too interpretive. This occurs when subspecific level

Figs. 1-4. Euphyes bayensis. 1/2 (d/v) d: Texas, Jefferson Co., Sabine Pass, 13 Sept. 1994, leg. Bordelon [Bordelon]. 3/4 (d/v)

9 paratype: Mississippi, Hancock Co., Bay St. Louis, 10 Oct. 1970, leg. Kergosien [MOTH]. Figs. 5-8 & 12. P. a. bordeloni.

5/6 (d/v) <j holotype: Texas, Jefferson Co., Sabine Pass, 12 May 1994, leg. Bordelon [TX A & M]. 7/8 (d/v) 9 allotype: 21 May

1994, then same data as 5.12 (d/v) 9 paratype: Texas, Jefferson Co., Sabine Pass, 25 May 1995, leg. Slotten [Slotten]. Figs. 9

& 10. P. a. minimus. 9 (d) d holotype: South Carolina, Orangeburg Co., Bull Swamp, 1 June 1992, leg. Gatrelle [CM]. 10 (d) 9

allotype: 6 June 1992, then same as 9. Figs. 11 & 19. P.a. howardi 11 (d) d: Florida, Volusia Co., 10 mi. w. Daytona, 24

April 1971, leg. Roman [FSCA]. 19 (d) 9: Florida, Volusia Co., Daytona, 25 July 1976, leg. Gatrelle [MOTH]. Fig. 20 (d/v) 9

E.pilatka : Mississippi, Hancock Co., Bay St. Louis, 27 Sept. 1970, leg. Kergosien [MOTH]. Figs. 13-18. P. aaroni subspecies

(leftFW margin). Fig. 13. d P. a. bordeloni. Fig. 14. d P. a. howardi. Fig. 15. d P. a. minimus. Fig. 16. 9 P. a. bordeloni.

Fig. 17. 9 P. a. howardi. Fig. 18. 9 P. a. minimus. (d - dorsal, v = ventral) [collection in which specimen is housed]

2 Subspecies can exist at the same time but not at the same location (i.e., both in March, but one in Texas and one in

California). Or, if absolutely univoltine, they can exist at the same location but not at the same season (i.e., both at the same

site, but one in the spring and one in the fall). When seasonal subspeciation also involves a shift in host specificity, the

organism is well on its way to full speciation (Pratt & Emmel, 1998).

process and assessment are misapplied to the species level. 2) A taxonomist will not describe (or not

recognize) a population as a subspecies because it does not meet certain objective criteria. This occurs

when specific level criteria are misapplied to the subspecies level. The result: species are weakly

described and valid subspecies go undescribed. The effect: subspecies are going unrecognized and therefore

unprotected from modem environmental pressures which in turn are causing their unnatural extinction.

An individual who frowns on subspecies (lumper) recently wrote me, “There is no objective way of

drawing a line between the degrees of variation in populational segregates.” Interestingly, that is exactly my

point! While species are objective and therefore largely unarguable, subspecies are subjective and often

arguable. While species are an end product (definitive plateau) of evolution, subspecies are the visible

components of a species continuing evolution (irregular steps between plateaus). In the sense that species

are easier to delineate, species level taxonomy is easier to do - and thus less controversial. The solution is

not to ignore subspecies or avoid subspecific taxonomy, but to find and define the variable taxonomic lines

that delimit various living organisms subspecifically.

Evolutionary Function of Subspeciation (Variation)

There are three factors by which the long term evolutionary vitality of a lepidopteran organism is

promoted: by its host acceptability - the more polyphagous a species larvae are the less apt it is to become

extinct; by its climatic adaptability - the more tolerant a species is of varying environments the less apt it is

to become extinct; and by its subspecific alterability - the more subspecific components within a species

the less apt it is to become extinct. The third factor is what I want to address here.

The world’s biota is composed of many more subspecies (animals), varieties (plants), and strains

(microbes) than species. Since the arrival of winged insects in the mid to late Paleozoic Era (270 to 350

million years ago) and modern insects in the Cretaceous period (70 to 135 million years ago), evolution has

insured the existence (by diversification) of Lepidoptera through time by means of subspeciation (that which

is adapting to new hosts, at different locations, in diverse climates, for a future time) rather than speciation

(that which has adapted to one host, at one location, in one climate, for a present time). Without variations

there will be no future evolution of tomorrow’s new organisms. Therefore, from the future evolutionary

perspective, a species with multiple subspecies (more variation) is more important (viable) than those

species with no subspeciation.

I therefore find two things very disconcerting about the current climate cf American lepidopteran

taxonomy. First, in reading the scientific literature, it is truly amazing how often subspecies have been

described without any accompanying evolutionary theory. Populations are deemed as subspecies just

because they “look different.” No wonder some taxonomists are paying so little attention to subspecies.

Second, I am disturbed by the increased number of taxonomists who have no interest in (see no value in)

recognizing or describing subspecies. How have some taxonomist’s apparently lost sight of the living,

evolutionary, theoretical, and future elements of taxonomic study?

Phenosyncronic Subspeciation

The word “color” is a term we often use in our day to day speech. However, color is composed of

several very different technical aspects such as shade, hue, value, and intensity. Thus, when one person is

talking to another about the “color” of a flower, the speaker may actually be focused on the hue while the

listener may be focused on the value. In which case, communication went forth but was not accomplished.

The word “morphology” is much like the word color. It is a broad term that includes all aspects of

an organism’s form and structure - wing and body contour, coloration, and variation. Thus, the words

morph and morphology often mean different things to different people. I always employ the term morphology

only to that aspect which deals with physical contour or shape (square, round, long, short, hairy, smooth,

etc.). When I deal with a population’s coloration pattern I use the word phenotype.

3

I do this because the root words morph and pheno are very different. Morph is primarily indicative

of that type of form which is physical in shape and structure. This is why butterfly larvae, pupae, and adults

are often spoken of as morphs. Conversely, pheno is from the Greek paino or phaninein which means to

shine reflectively - to appear via reflected light. Having conveyed my definition of terms, we can move on

in our current communication.

In subspecific development it is usual for sister subspecies to evolve distinctly different phenotypes

(which are produced by the interaction of the genotype with the environment). This is called phenotypic

subspeciation. In describing subspecies, lepidopteran taxonomists have historically focused primarily on a

taxon’s phenotypic evolution and secondarily on its biological, ecological, or morphological evolution. In

fact, many of our described subspecies, while looking very different, actually differ very little biologically or

ecologically (e.g., Limenitis arthemis arthemis (Dury, [1773]) and L. a. astyanax (Fabricius, 1775)).

In some cases however, while a sister subspecies is evolving in a divergent manner

morphologically, biologically, or ecologically, its phenotypic evolution is parallel - synchronous. I call this

phenosyncronic subspeciation. Thus, phenosyncronic subspeciation exists when the facies of one subspecies

differ very little from a sister subspecies (e.g., Glaucopsyche lygdamus xerces (Boisduval, 1852) and G. /.

pseudoxerces Emmel & Emmel, 1998; Anthocharis midea annickae (dos Passos & Klots, 1969) and A. m.

texana Gatrelle, 1998; Cercyonis sthenele sthenele (Boisduval, 1852) and C. s. hypoleuca Hawks &

Emmel, 1998).

The subspecific validity of a phenosyncronic subspecies is therefore based primarily on its unseen

evolutionary (present and future) genetic potential - as evidenced by its observable biological (inc. larval

host), environmental (inc. seasonal), or regional (inc. refugium) distinctives. Phenosyncronic subspecies are

a very important segment in the lepidopteran evolutionary chain. Yet, they seem largely overlooked. I

believe this is primarily because modem taxonomists have gravitated more and more into the error of

applying various species level criteria into the process of subspecific definition.

A phenosyncronic subspecies, by virtue of its divergent evolutionary course, is more evolutionarily

significant than a sister subspecies that differs only in its facies. For example, two subspecies which look

very different but occupy the same ecological niche, feed on the same host, and fly at the same time are not

nearly as evolutionarily important as a third sister subspecies which looks very similar to one of the first

two but occupies a different ecological niche, feeds on a unique host, or flies at a different time of year. For

example, the genus Euphilotes Mattoni, is rich in phenosyncronic subspecies 3 (Pratt & Emmel, 1998).

When a phenosyncronic subspecies evolves to the degree that it warrants recognition as a species,

the result is a sibling species pair (e.g., Euphyes dion (W. H. Edwards, 1879) and E. bayensis Shuey,

1989). Many northern specimens of E. dion are indistinguishable in coloration, male genitalia, and stigma

from E. bayensis. However, they usually differ in male stigma, greatly in habitat preference, and do not

interbreed when sympatric.

This paper employs the principles of subspecific taxonomic analysis (process, assessment, and

conclusion) as applied to the west central US Gulf Coast segregate of Poanes aaroni. The process, in this

case, is the examination of a skipper butterfly’s biogeographical history, integral biotic community, and

phenotype. The evolutionary assessment is that a phenosyncronic subspecies is determined. The conclusion

is the description of a new subspecies.

While these authors do not use the term phenosyncronic subspecies, the taxa they delineate primarily on the basis of

seasonal occurrence and host associations fall essentially into this type of subspeciation. In correctly emphasizing that these

taxa represent a higher degree of evolution than simple geographic subspecies, they referred to them as semi-species, which

is fine for the purpose of conceptual understanding but unacceptable within the Code of the ICZN (the Code only recognizes

organisms below the genus level as either species or subspecies). Therefore, Pratt and Emmel elevated several of these taxa

to species level - which may or may not be correct. Nonetheless, subspecies, whether biological, morphological, seasonal,

host specific, phenotypic, geographic, clinal, or convergent are still only SUBSPECIES. And all species, no matter how

marginal or complexly interrelated in groups, are still only SPECIES. The ICZN Code is wisely structured to accommodate

both scientific definition (by uniform restriction of technical terms) and evolutionary theory (by non-regulation of authors

textual presentation of organic interrelation).

POANESAARONIBORDELONI GATRELLE, NEW SUBSPECIES

Discovery

On 27 September 1992 Dr. Jeff Slotten of Gainesville, Florida discovered a population of Poanes

aaroni (Skinner, 1890) on the Gulf Coast at Bay Saint Louis, Hancock County, Mississippi. This record

was not only the first for Mississippi but extended the known range of aaroni about 600 km westward

(Opler & Krizek, 1984). This site is the type locality of Euphyes bayensis Shuey, 1989, and as such, had

been previously well sampled. We now know that this Gulf Coast segregate is an undescribed subspecies.

A year later on 30 September 1993 Mr. Charles Bordelon of Beaumont, Texas discovered a colony

of this segregate at Sabine Pass, Jefferson County, Texas. This location is near the Texas and Louisiana

state line, about 450 km to the west of Bay Saint Louis, and 1000 km from the nearest population of P.

aaroni howardi (Skinner, 1896). This skipper is now known to occur in Texas westward in Jefferson

County to McFaddin National Wildlife Refuge. Bordelon expects it to eventually be found along the coast in

neighboring Chambers County to Smith Point and, perhaps, into Harris County. The taxon is well

established in this region and is sometimes common.

E. bayensis (Figs. 1-4) is sympatric with this aaroni subspecies at all known locations. Euphyes

pilatka (W.H. Edwards, 1867) is also present with this aaroni subspecies at the Bay Saint Louis site.

In issue 1:10 of The Taxonomic Report (24 December 1999), I described a new subspecies, Poanes

aaroni minimus Gatrelle, from inland South Carolina. In the process of this description, I figured a pair of

this Texas aaroni and stated that I believed they represented an undescribed subspecies. I also suggested

that someone from Texas consider describing it. Shortly after the publication of TTR 1:10, I received a

phone call from Mr. Bordelon in which he offered to send me his specimens of this segregate and also

encouraged me to undertake the description of it. I have. This paper makes public the results of that study.

This subspecies is named for Mr. Bordelon in recognition of his many years of work with Lepidoptera and

his discovery of this taxon in Texas.

Overview of Poanes aaroni Subspecies

Poanes aaroni aaroni (Skinner, 1890) was described from seven males and one female collected

by Mr. Samuel Aaron at Cape May, New Jersey. The species was not figured and no type was designated.

Holland (1931) figures a male “paratype” on PL. XLVI, Fig. 37. It is typical of this subspecies and thus

represents it well. This specimen was assumed to be the type and located in the Carnegie Museum NH,

Pittsburgh. However, upon my correspondence with Dr. John Rawlins of the CMNH no such specimen

could be found. Dr. Rawlins relayed the following (emphasis his).

There is a single male specimen in the CMNH bearing a label stating it to be the specimen figured

in Plate LXVI, Fig. 37, but there is no question that it is NOT the specimen illustrated. A pin

label states that the specimen was obtained by Holland from Skinner in exchange for a pair of

Papilio brevicauda in 1890. The specimen bears a classic triangular “TYPE” label, but no

further data on locality or collector. It is probable but not certain that this is a syntype.

In my subsequent efforts to find a specific specimen specifically designated in the literature to be the

type of P. aaroni, Dr. Fred Rindge of the AMNH offered the following.

For P. aaroni : See both Skinner & Williams, 1924, Trans. Amer. Ent. Soc., 50:60, where they

state that the type is in the ANSP. And Gillham and Ehrlich, 1954, op. Cit., 80:112, where they

list the type as no. 7093 ANSP.

Dr. Rawlins has informed me that there is one other Skinner specimen at the CMNH. This specimen

is the one mentioned above as ANSP (Academy of Natural Science) type no. 7093.1 assume here that this is

also the specimen that Miller and Brown (1981 page 43) took as the “holotype” in their checklist.

Concerning this specimen and situation Dr. Rawlins has relayed the following.

Verbatim label data (labels separated by / with external comments in square brackets): New

Jersey [typeset in black ink] / Type 7093 Pamphila aaroni Henry Skinner [dark red label, typeset

except for “Pamphila aaroni” handwritten in black ink]... There is no reason to doubt that the

above male is a valid syntype. It is appropriate for lectotypification, if indeed this has not already

been done in the literature (Rawlins did not research this aspect thoroughly) as it was marked as a

Skinner type when the specimen was still at ANSP.

It seems to me that this specimen should have a lectotype label affixed to it if for no other reason

than to tie up any loose ends that may technically still exist. The word “type” on its label just means that it is

of the type series and not The Type - holotype. Thus, I here designate this specimen as lectotype of

Pamphila aaroni Skinner 1890.1 have sent a label that reads as follows to Dr. Rawlins to be placed on this

specimen. Handwritten on red paper in black ink: LECTOTYPE c? Pamphila aaroni SKINNER, 1890

Designated July, 2000, R.R. Gatrelle, TTR 2:2.

P. a. aaroni is the smallest and most brightly colored (especially females) of the aaroni subspecies.

I have examined 81 P. a. aaroni summarized as follows by state and county. DELAWARE: Kent; NEW

JERSEY: Burlington, Ocean, Cape May.

Poanes aaroni howardi (Skinner, 1896) was described from 12 specimens without mention of

their sex. All but two of these were only labeled as being from Florida. The two with site specific labels are

stated as being from Georgiana on the Indian River. Thus, a definitive type locality has never been

established. Accordingly, I here restrict the type locality to: vicinity of Georgiana, Brevard County, Florida.

This taxon was also not figured in the original description and no specimen was designated as holotype.

Holland (1931) figures a male syntype on PI. XLVI, fig. 38. It is typical of this subspecies and thus

represents it well. This specimen was also assumed to be housed in the Carnegie Museum NH, Pittsburgh.

Dr. John Rawlins has relayed the following concerning possible types of this taxon in the CMNH.

Verbatim label data (labels separated by / with external comments in square brackets): Florida

[handwritten in black ink] / Collection Dept. Agr. [typeset in black ink] / TYPE [large bold

typeset in black ink; a square label, but clearly the top third of the usual triangular Skinner label

for “TYPE”, meaning syntype] / Collect. Skinner [typeset in black ink] / type 7092 Pamphila

howardi Henry Skinner [dark red label, typeset except for “Pamphila howardi” handwritten in

black ink]...

Described from 12 syntypes of unspecified sex (but including both sexes), 8 specimens in Skinner

Collection and 4 from USNM via L.O. Howard. Two syntypes restricted to “Georgiana”, the

others without restriction (just “Florida”). The type above is one of the unrestricted ones, there is

no reason to doubt that this male is a valid syntype and it would be appropriate for

lectotypification if indeed this has not already been done in the literature (Rawlins did not

research this aspect thoroughly) as it was marked as a Skinner type when the specimen was still

at ANSP...

Rawlins is aware of only one other syntype, at CMNH in June 2000. It is a female labeled

verbatim as follows: Florida / Collection C.V. Riley/Collect. Skinner / ALLO-TPYE 9 Pamphila

6

aaroni Henry Skinner. There is no reason to doubt that this is a valid syntype, and with the

designation described above, it would become a paralectotype.

Holland (1898) in “The Butterfly Book” illustrates a male of this species on Plate 46, Figure 38,

as Phycanassa howardi. As of 15 June 2000 Rawlins could not locate a specimen (male or

female) in the CMNH bearing a label stating it to be the specimen figured in Plate LXVI, Fig. 38.

Because the type specimen situation is not absolutely clear in regard to this taxon, I here designate

the above mentioned male as lectotype of Pamphila howardi Skinner 1896.1 have sent a label that reads as

follows to Dr. Rawlins to be placed on this specimen. Handwritten on red paper in black ink: LECTOTYPE

c? Pamphila aaroni howardi SKINNER, 1896 Designated July, 2000, R.R. Gatrelle, TTR 2:2.

I have examined over 300 howardi from the following states (by county). VIRGINIA: Princess Ann;

NORTH CAROLINA: Currituck; SOUTH CAROLINA: Beaufort, Colleton, Charleston, Jasper; GEORGIA:

Bryan, Chatham, Glynn; FLORIDA: Baker, Dade, Duval, Levy, Putnam, Seminole, Volusia.

Poanes aaroni minimus Gatrelle, 1999 was described from 46 specimens from a fresh water

swamp in Orangeburg County, South Carolina. This is the only know colony of this unusual subspecies.

Minimus is the most phenetically distinct aaroni subspecies.

The Gulf Coast segregate of P. aaroni, described herein as P. a. hordeloni, is the largest aaroni

subspecies. Dorsally, it is more orange fulvous than yellow fulvous. Its most striking feature is the apically

elongate shape of its forewings. These have the outer margins fairly straight which makes the apex very

pointed and almost falcate in some specimens. (The outer fore wing margins of the three eastern subspecies

are distinctly rounded). The viator-spotting on the ventral hindwings of this segregate varies greatly from

prominent to absent. The ventral hindwing ground color is slightly darker than in howardi (both sexes).

I have examined 31 specimens of this subspecies from the following states (by county). TEXAS:

Jefferson. MISSISSIPPI: Hancock. These specimens make up the type series.

Process and Assessment

As I began to accumulate and study specimens, I first looked for reasons why this segregate should

not be considered subspecifically distinct from Floridian P. a. howardi. When I placed a few specimens of

Floridian howardi and the Texas segregate together and just took a glancing look at them they appeared

about the same (especially the males). This ephemeral examination of specimens, if coupled with a

conception that both populations are resident to a geologically ubiquitous southern United States, could

easily lead someone to the conclusion that only one subspecies is involved. However, when I moved

beyond a cursory examination of this skipper’s facies and also considered the differing biogeographical

development of the eastern and western area of the southern United States, a very different picture began to

emerge.

Looking closer. The Gulf Coast segregate (P. a. bordeloni) shares several coloration and spot

pattern characters with both minimus and howardi. Dorsally, hordeloni and howardi males are virtually

identical in their fulvous spot pattern with bordeloni tending to be more orange fulvous and howardi more

yellow fulvous. Ventrally, bordeloni males are more like those of minimus except in those males with

strong viator-spotting. Dorsally, female bordeloni usually have a reduced forewing spot pattern like

minimus females while the hindwing is more broadly fulvous than howardi females. They differ ventrally in

the same manner as the males. Thus, the Gulf Coast segregate differs phenotypically from howardi primarily

in the female and not the male. As stated above, the most noticeable visible difference between bordeloni

and the other aaroni subspecies are the shape of their fore wings.

If only these minimal phenotypic features and one strong morphological character were considered,

the Gulf segregate might warrant description as a subspecies. However, these are not the only factors that

must be considered.

The biogeographical evolution of the Gulf Coast segregate greatly distances it from Floridian howardi.

Like many other butterflies in the Gulf area, P. a. bordeloni is surely ascended from a Texas or Mexican

refugium while howardi is ascended from the Floridian refugium. As such, bordeloni belongs to the

southwestern US fauna while howardi is part of the southeastern US fauna. If this is so, then these two

southern taxa have been separated for thousands of years and are now moving toward each other. The

likelihood is good that they will meet one day in the Florida panhandle (if they have not done so already in

an undiscovered tension zone colony). Then again, they may never meet.

Evolutionally, we do not know the degree of variance contained in the future genetic potential of

either bordeloni and howardi. However, we do know that it is different - as evidenced by the observably

divergent shape of their forewings and minor phenotypic differences. This is regional genetic subspeciation.

According to Remington (1968) the lepidopteran subspecies of the southern United States are of

very recent origin. Remington postulates that most of these probably came into being in just the last 30,000

years during periods of glacial retreats and advances in the Pleistocene (1 million to 10 thousand years ago

which effected great climatic and geophysical changes in the Gulf Coast area). The last glacial maximum

(Wisconsin) is generally placed about 11,500 B.P.

During periods of glacial maxima ocean levels were lowered which resulted in shorelines being

extended. In the interglacial periods ocean levels rose which resulted in shoreline resorption. In the west

central Gulf Coast area this ebb and rise in sea level moved the shore line in or out many miles from its

present location. During the interglacial periods the plants and animals of the southern US became separated

into disjunct eastern and western species segregates (Remington, 1968). For numerous species, this break in

gene flow resulted in the evolution of eastern and western subspecies or sibling species.

As the land mass of the Southern United States increased toward its present day state, these

organisms were dislocated and adapted in concert with the changes in the region's ecology and geography.

This biogeographical evolutionary process has produced a unique community of biota that is presently

endemic to the west central Gulf Coast of the southern United States.

Dr. Richard Brown, Mississippi State University, states (pers. com.) that this region’s endemism is

limited to the off shore islands and in dune areas. The hurricanes and many river outlets which have shaped

the coastal area in modem times have fragmented these island, dune, and marsh habitats. This area of

endemism, though further southeast, correlates evolutionarily to Remington’s Louisiana-East Texas suture-

zone F. The following Lepidoptera are unique to this coastal region.

Automeris louisiana Ferguson & Brou, 1981, Asterocampa celtis alicia (W.H. Edwards, 1868),

Limenitis archippus watsoni (dos Passos, 1938), Euphyes bayensis, mdPoanes aaroni bordeloni.

Poanes aaroni bordeloni Gatrelle, new subspecies

Diagnosis. In both sexes of bordeloni the dorsal fulvous areas are usually a deeper color of orange than in

howardi. The dorsal facies of male bordeloni differ very little from howardi. However, the dorsal forewings of bordeloni

females usually have the fulvous areas much reduced, as in minimus females, while the fulvous on their hindwings is more

developed than in howardi females. (About 20% of females have extensive fulvous on their forewings.) Ventrally, the viator-

spots on the hindwings (in both sexes) of bordeloni are very variable. The ground color on this surface is also variable but

usually darker than in howardi. Ecologically, bordeloni is endemic to salt and brackish marshes as are subspecies a. aaroni

and a. howardi.

Description. Male (Figs. 5-6 ). Head: the coloration of the hair and palpi is the same as in howardi. Thorax (inc.

legs) and abdomen : colored as in howardi. Forewings', dorsally and ventrally, fulvous pattern the same as in howardi but

more orange, stigma less prominent than in howardi especially the upper segment. Hindwings: dorsally, the fulvous pattern

the same as in howardi but more orange and slightly more extensive; ventrally, the light streak in the cell and the viator-spots

may be very prominent or absent. Female (Figs. 7-8, 12). Head, thorax, and abdomen : as in male; Forewings: dorsally and

ventrally, without stigma, dark marginal border much wider than in male, usually having two light apical spots, in 80% of

8

females the fulvous is limited to a postmedian band of spots which gives these females a remarkable resemblance to Euphyes

pilatka females (Figs. 12 & 20). Hindwings : dorsally, broadly orange fulvous with the spot in cell CU2 usually well

developed; ventrally, as in male. General. The outer margins of the forewings of both sexes of bordeloni are only slightly

curved from the outer angle to the apex (apex of marginal curve at vein M2). In the other three subspecies the outer forewing

margins are rounded with the apical point of the curve being at about mid wing (vein M3). In some specimens the apical tip of

the forewings is slightly, but noticeably, falcate.

Types. Holotype d (Figs. 5-6 ): TEXAS: Jefferson County, Sabine Pass, 12 May 1994 (leg. Bordelon). Allotype 9

(Figs. 7-8 ): TEXAS: Sabine Pass, 21 May 1994 (leg. Bordelon). Paratypes : 16 d d and 13 9 9: TEXAS: Jefferson County,

Sabine Pass: 1 9, 30 September 1993 (leg. Bordelon); 5 d d, 8 May (2 leg. Bordelon, 3 leg. Rickard), 19, 12 May, 2 d d,

1 9,21 May, 1 9, 23 May (leg. Bordelon), 1 d\ 30 July (leg Knudson) 1994; 1 d, 5 9 9, 25 May, 2 c? d\ 2 9 9 27 May 1995

(leg. Slotten); 2 d d, 19 May (leg. Knudson), 1 d\ 24 May (leg. Bordelon) 1996. MISSISSIPPI: Hancock County, Bay Saint

Louis: 2 d d, 2 9 9,27 September 1992 (leg. Slotten). The holotype and allotype are deposited in the collection of Texas A

& M University, College Station, Texas. Paratypes are in the collections of Charles Bordelon, Beaumont, Texas ( 3 ); Ed

Knudson, Houston, Texas ( 4 ); Roy Kendall, San Antonio, Texas ( 7 ); Jeff Slotten, Gainesville, Florida ( 10 ); and the

Museum of the Hemispheres (MOTH), Goose Creek, SC ( 5 ).

Geoecological type locality. Coastal salt marshes of Jefferson County, Texas.

Etymology. Bordeloni is named for Mr. Charles Bordelon of Beaumont, Texas.

Remarks. Evolutionally, the reduced forewing and increased hindwing dorsal fulvous of female bordeloni connects

it with both minimus and howardi respectively. The dorsal color and pattern of the fulvous on male bordeloni strongly

connects it to howardi and aaroni and distances it from minimus. The highly variable ventral viator-spotting and dark ground

color connects it with both howardi and minimus and distances it from aaroni. The shape of the forewing margins distances

it from all other populations. I would interpret these character associations (or lack of association) to indicate that bordeloni

and minimus are the oldest subspecies and aaroni the youngest.

ACKNOWLEDGMENTS

I thank the following persons for their contributions without which the paper could not have been

produced. For the loan of specimens: Charles Bordelon, Jeff Slotten, Ed Knudson, and Roy Kendall. For

information of the syntypes of aaroni and howardi : Dr. John Rawlins, CMNH, Pittsburgh and Dr. Fred

Rindge, AMNH, New York. For critical textual review (pro and con): Dr. Richard L .Brown, Mississippi

State University. Manuscript review: Mr. Bryant Mather, Clinton, Mississippi. For photography of

specimens: Mr. Joseph Mueller, my son-in-law.

LITERATURE CITED

GATRELLE, R.R. 1999. Three New Hesperioidae (Hesperiinae) from South Carolina: New Subspecies of

Euphyes bimacula, Poanes aaroni, and Hesperia attalus. The Taxonomic Report, Vol. 1 (10), 13

pp. The Int. Lepid. Survey, Goose Creek, SC.

MILLER, L.D. & F.M. BROWN. 1981. A Catalogue/Checklist of the Butterflies of America North of

Mexico. Memoir No. 2, Lepid. Soc., Sarasota, FL. 280 pp.

OPLER, P.A. & G.O. KRIZEK. 1984. Butterflies East of the Great Plains, An Illustrated Natural History.

Johns Hopkins Univ. Press, Baltimore, MD. 294 pp.

PRATT, G.F. & J.F. EMMEL. 1998. Revision of the Euphilotes enoptes and E. battoides Complexes

(Lepidoptera: Lycaenidae). In: Systematics of Western North American Butterflies, Chap. 16, 207-

270. Mariposa Press, Gainesville, FI.

REMINGTON, C.L. 1968. Suture-zones of Hybrid Interaction Between Recently Joined Biotas. Evol.

Biology, Vol. 2(8), 325-413.

9

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

TILS Purpose. TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific

discovery, determination, and documentation, without which there can be no preservation of Lepidoptera.

TILS Motto. As a world community, we can not protect that which we do not know. _

The Taxonomic Report is projected for publication at the rate of about 10 issues a year. Subscription

for Volume Two is $50 US for domestic and $65 US for overseas subscribers. The subscription year

follows the calendar year. All issues are mailed 1 st class. At the end of each year, subscribers receive that

year’s volume on a CDR for permanent archiving and reproduction for personal use (i.e. a museum or

university may make as many copies as needed in whatever format desired). Non-members may receive

individual issues in print any time for $10 per issue. Individual issues on CDR to non-members are $25 per

issue post paid. Subscriptions and individual issue orders should be made payable to TILS and mailed to:

Scott D. Massey, Editor, 126 Wells Road, Goose Creek, SC USA 29445-3413.

Volume 2

15 July 2000

Number 3

The Taxonomic Report

yW OF THE INTERNATIONAL LEPIDOPTERA SURVEY f*

A NEW SPECIES OF ANGULO PIS (LYCAENIDAE, EUMAEINI) FROM

RELICT COASTAL FOREST IN EAST-CENTRAL ARGENTINA

GUSTAVO CANALS

Investigador Asociado, Departamento Cientifico de Entomologia

Museo de la Plata, Paseo del Bosque s/n., Argentina

AND

KURT JOHNSON

Environmental Department, The Ethical Culture Society,

53 Prospect Park West, Brooklyn, New York 11215

ABSTRACT. Angulopis puntalaraensis is described from a single female specimen collected between 1950 and

1955 in the National Reserve Punta Lara, Buenos Aires Province, Argentina. This site is a relict of coastal gallery forest. No

other specimens are known.

Additional key words: Unique tergite. Angusta species group

INTRODUCTION

The genus Angulopis Johnson 1991, though relatively new, has been the subject of numerous studies

- both locally and across the entire Neotropical Realm. The type species is the well known and widely

distributed “Red Banded Hairstreak” Thecla sangala Hewitson (senior synonym of the type species by

original description: Theda autoclea Hewitson which occurs from Mexico to Argentina [Austin and

Johnson, 1997]). In 1993, Johnson and Kroenlein elaborated the taxonomy and geographic ranges of twenty-

six taxa of Angulopis represented in various European and American museum collections. Subsequently,

Austin and Johnson (1997) studying voluminous collections from a lowland tropical rain forest locality in

Rondonia, Brazil, differentiated further taxa.

Worldwide, it is common for species representing isolated distributions or unique habitats within

the range of a genus to show “supralimital characters” (see Eliot, 1973; Shields 1996, and citations in this

latter work). Briefly, in colloquial terms, it is not unusual to find unusual specimens from unusual places,

especially in the Lycaenidae. Further, historical review of many lycaenid genera (especially from tropical

regions) reveals a significant number of taxa known only from early collections and/or small samples (e.g.,

see Balint 1993,D’Abrera 1995). This creates a predicament for contemporary systematists: such specimens

can either be ignored or described, depending on what one considers a “conservative” taxonomic approach

(Schwartz 1989).

The senior author has been studying Lepidoptera, and other biota, from a disturbed relict of coastal

gallery forest in Buenos Aires Province, Argentina, known since its inception in 1958 as the Natural

Reserve Punta Lara. Among old samples from this area was a specimen referable to Angulopis which is

unlike any known species in either wing pattern or genitalia. Considerable search of other material, from

both the vicinity of Punta Lara and across Latin American has turned up no similar specimen. Given the

breadth of previous work on Angulopis , and the constantly degrading status of the habitats of the Punta Lara

reserve, we have decided to describe the specimen here as a new species.

Terminology for wing pattern follows previously cited work on Angulopis , including for brevity the

abbreviations DFW, DHW (dorsal forewing, hindwing) and VFW, VHW (ventral forewing, hindwing) and

other thecline-oriented wing marking terms like “scent brand” (a DFW secondary sexual character) and

“Thecla-spot” (a large orb in the VHW limbal area typical of many hairstreaks). Morphological terms

follow previous work on Argentine hairstreaks by Johnson, Eisele and MacPherson (1988) including terms

for unique tergite and genitalic features not found in other Angulopis species. Additional abbreviated terms

include: OD (original description), TL (type locality), and regarding geography, N etc. (north, etc.), C (central), etc.

Fig. 1. Holotype: Angulopis puntalaraensis: ventral (left) and dorsal (right). (Specimen is three times natural size.) Fig. 2.

Genitalia (female) of holotype. Left, ventral view from terminus of lamella postvaginalis (‘Ipv”, top) to anterior end of ductus

bursae (“db”, anterior one-third rotated to lateral view). Immediate right of lpv—ventral view of sipc (“v”), then dorsal (“d”,

showing apophyses papillae anales ‘hp” within), and immediately below “d” the lateral view (“1”). Immediate right of ‘db”,

ventral view of corpus bursae signum. Fig. 3. Map of South America showing generalized, circum-Amazonian, distribution of

Angulopis species (light stippling for widespread, common, species; various larger symbols for regionals, endemics, or

poorly known taxa); compared to distributions of Argentine “White Wood” forests (black shading, marked with arrows at

“WW”) and type locality of A. puntalaraensis (star to left of Natural Reserve Punta Lara). Editors note: The hindwings of

this specimen are unnaturally mounted in that they are positioned on top of the forewings.

GENUS ANGULOPIS JOHNSON

Angulopis Johnson 1991: 41.

Species Group. The new species appears to belong to the angusta Species Group of Angulopis

(Johnson and Kroenlein 1993) although it is extreme in both wing and morphological features for that group.

Angulopis puntalaraensis Canals and Johnson, new species

DIAGNOSIS. Currently known only from the female; DFW, DHW uniform medium brown (FW without androconial

brand) and HW with two prominent tails. Distinguished from other generally similar Angulopis and/or Gigantrorubra

species (see Remarks) by VHW showing (1) costal element of a white-edged black medial band orbiculate and distally

disjunct from rest of band by a cell’s width and (2) entire limbal area basad the Thecla-spot marked from anal margin to cells

M3 and M2 with a clearly defined, brilliant bright red-orange patch. Female abdomen’s terminal tergite forming a well-

defined sipc (unique in genus), genitalia with (1) prominent, laterally lobate, lamellae postvaginalis membranously connected

to adjacent sipc, (2) ductus bursae relatively short for genus, angled 90 degrees dorsad in its terminal one-third and (3)

corpus bursae with two huge dendritic signa covering at least one-half of each bursal surface.

DESCRIPTION. (Fig. 1) DFW, DHW uniform medium brown (FW without androconial brand), HW with elongate

tail at terminus of vein CuA2, shorter tail at terminus of vein CuAl, margin of cell between and cells adjacent to each tail

lined light sky blue and with a small orange spot at anal angle and brown spot in cell CuAl. VFW, VHW ground beige, FW

with vague blackish discal slash, more prominent black postmedial line, whitish distally (costa through cell CuAl), submargin

with intercellular spaces showing black dashes; HW with vague blackish-brown discal slash, then prominent medial band

(colored blackish-brown basally, white distally); band showing (1) a prominent blackish brown costal orb disjunct from rest

of band by a cell’s width and (2) rest of its length jagged and forming prominent W-shaped configuration between cell CuAl

and the anal margin. Margin of cell between and cells adjacent to each tail lined white. Limbal area distinctly marked with (1)

Thecla-spot and surface adjacent to anal angle black and (2) rest of area from anal margin to cells M3 and M2 marked with a

clearly defined, brilliant bright red-orange, patch. FW Length: 11 mm (base of wing to apex).

Female Tergal Morphology and Genitalia (Fig. 2). Abdomen with terminal tergite sclerotized into a prominent

subchordate incised posterior cavity (“sipc”) showing at each lateral margin a prominent and ridged lobe, this sipc strongly

connected laterally to the lamellae postvaginalis of genitalia. Genitalia with robust and elongate lamellae postvaginalis (l.p.)

terminating at a prominent antrum in the posterior one-half of the genital length caudad of corpus bursae; l.p. with short

laterally protruding lobes in its terminal one-half, rest of robust terminal opening of ductus bursae strongly attached laterad

to the sipc with membranous material; ductus bursae anterior to the antrum narrow and inclined 90 degrees in its anterior

one-third, there connecting to the corpus bursae; corpus bursae with two extremely prominent dendritic signa, each covering

at least one-half of the respective bursal surface.

TYPE. Holotype female (Fig. 1), Breyer Collection, Museo de La Plata (MLP) labelled 'Punta Lara” (which refers,

in the Breyer Colletion at (MLP) to the Natural Reserve Punta Lara near La Plata, 60 km S Buenos Aires). Although the

collection date is not noted the specimen must have been collected between 1950-55 because the label is sky blue, the color

Breyer used for material collected during that time period. The collector per se is not noted; sometimes similar labels

include the additional data "Argentina, Prov. Bs. Aires, Breyer" but these are not attached to this particular specimen,

suggesting it was collected by an employee or colleague.

REMARKS

Habitat at Type Locality. Punta Lara is a small town located 12 kmN of La Plata (capital of Buenos

Aires Province). The climate is temperate, warm and wet (precipitation 1000 mm/year). Located nearby is

the Natural Reserve Punta Lara (3500 hectares) created in 1958. Both the Reserve and the town are located

in coastal lowlands along the Rio de la Plata. The Reserve was established to protect declining forest areas,

which here include gallery forest which occurs along the edges of adjacent rivers and streams. The gallery

forests are locally known as “white woods" because their flora is similar to forests in the Misiones

province of Argentina, of which they are considered the southernmost relict (Fig. 3). It also has a few little

spots of xerophilous wood with Tala (Celtis tala) and Coronillo (Scutia buxifolia ). Today the entire area

around Punta Lara is very disturbed by invading non-native plants, including large areas covered with Lirio

3

amarillo (Iris pseudacorus ), Cana india (Pleioblastus simonii ), Zarzamora (Rubus ulmifolius ), and

Ligustro (Ligustrum lucidum ); the latter two originally from China, Japan, and Korea. Today the Reserve

has been reduced to an island left behind by man’s environmental modifications (deforestation, introduction

of non-native plants, hunting, fishing, etc.) which continued uncontrolled for many decades. Historically, the

Reserve is known for significant biodiversity, including 770 species of vascular plants, 40 mammals, 300

birds, 25 reptiles and 23 amphibians. There is an ongoing effort to track the fate of these biota. About 75

species of butterflies have been recorded, representing 50 percent of the total butterfly fauna of Buenos

Aires Province. The Reserve is known for many relict populations, among them are the larger more well-

known butterflies, Morpho epistrophus argentina (the most austral Morphinae) and Adelpha syma ,

(Nymphalidae: Biblidinae: Limenitidini) which otherwise occur only northward in Misiones and Entre Rios

provinces. The butterflies of the grasslands in and surrounding the Reserve, however, include Hesperiidae,

Nymphalidae ( Vanessa, Anartia, Danaus) and Riodinidae ( Audre ) species which differ little from other

areas of the province. The only other area of distinctive habitat and biota in the Province is the Buenos

Aires Hills, known historically from collecting localities like Ventana, Tandil, Azul, and Olavarria.

Discovery of the Holotype. There is no other specimen of A. puntalaraensis among the Breyer

material or material from the Petrowsky and other local historical collections from the region. Further, the

senior author recently curated an extensive local collection and no specimens were located therein either.

Suspecting that the holotype specimen represented either extirpated species or a species of extremely

limited modem occurrence, outside specialist opinion was consulted, resulting in the present paper.

A, puntalaraensis Congeners and Phylogenetic Position. A .puntalaraensis only vaguely resembles

disparate congeners. There is no other known Angulopis with a large, well-defined, red-orange patch

covering nearly the entire limbal area. Some members of the angusta Species Group from the Amazon

region show suffusive red variously located in the limbal area but all these species have red VHW bands,

not black ones. Of species with black VHW bands, the detached costal orb of A. puntalaraensis resembles

only A. suarezensis Johnson and Kroenlein (TL Rio Suarez, Colombia) a species which is blue on the

DFW, DHW and on which the costal mark is a detached oblique slash. That the VHW red-orange patch in

A. puntalaraensis is most likely a unique supralimital character (sensu Eliot 1973) reflecting the isolation

of the species is suggested by the occurrence of a DHW metallic orange patch in an extremely peculiar and

isolated north Andean species A. constantinoi Johnson and Kroenlein (TL Remolinos, Meta, Colombia).

Although these wing patterns in other Angulopis species illustrate how supralimital characters occur in the

genus, it is most likely that the closest phylogenetic relative of A. puntalaraensis is the poorly known SE

coastal Brazil species A. obscurus Johnson and Kronlein (TL Pelotas, Brazil). The latter species is bigger

(FW alar 13.0 mm) with none of the spectacular VHW colors of A. puntalaraensis. However, one could

generally attain the overall look of A. puntalaraensis by simply adding its colorful markings to A. obscurus.

Unfortunately, genitalic comparison of the two species is not possible because A. obscurus is known only

from the male. However, it is certain that A. puntalaraensis is not the female of A. obscurus. The latter is

dorsally dull blue and in Angulopis the general pattern of sexual dimorphism is for blue to occur in females

only, or, when occurring in males, to have companion females even more expansively blue.

Taxonomic Relation of Angulopis . With over thirty taxa now recognized in Angulopis , it would

seem unlikely that other taxonomists would want to combine Angulopis with some other, previously

described, genus. However, taxonomists emphasizing external similarity over morphology, or just seeking

extremely simplified classification for complex and diverse groups like the Lycaenidae, may still refer this

entire assemblage and Gigantorubra (see Johnson and Kroenlein 1993) to either Electrostrymon Clench (a

“long shot” based on general external similarity) or the older genus Lamprospilus Geyer (based on some

generality of wing pattern and structure but overlooking the extremely unique, and apparently apomorphic,

features found in the taxa traditionally associated with this name [Draudt 1920]). While D’Abrera (1995)

placed some members in Electrostrymon and others in Thecla he also skipped over a number of drawers of

Angulopis in The Natural History Museum (London) including some type material, probably because the

group is so problematic. This explains why so few Angulopis-like taxa are figured in his book. We mention

all the above simply to further orient the reader to Angulopis in the context of the Eumaeini and current

taxonomic literature.

LITERATURE CITED

AUSTIN, G. and K. JOHNSON. 1997. Theclinae of Rondonia, Brazil: Gigantorubra and Angulopis , with

descriptions of new species (Lepidoptera: Lycaenidae). Insecta Mundi 11: 255-272.

BALINT, Zs. 1993. A Catalogue of Polyommatine Lycaenidae (Lepidoptera) of the Xeromontane Oreal

Biome in the Neotropics as Represented in European Collections. Rep. Mus. Nat. Hist. Univ. Wise.

(Stevens Point) 29, 42 pp.

D’ABERA, B. 1995. Butterflies of the Neotropical Region, Part VII, Lycaenidae. Victoria, Hill House,

Victoria (Australia), xi+1098-1270.

DRAUDT, M. 1920. Theda , pp. 794-811 in Seitz, A. (ed.), Vol. 5 [Plates, Vol. II], Die Gross-Schmetterlinge

der Erde, Band 1. - Stuttgart (Germany), Alfred Kemen Verlag.

ELIOT, J.N. 1973. The higher classification of the Lycaenidae (Lepidoptera): a tentative arrangement.

Bull. Brit. Mus. Nat. Hist. (Ent.), 28: 371-505.

JOHNSON, K. 1991. Neotropical Hairstreak Butterflies: genera of the “Calycopis/Calystryma grade” of

Eumaeini (Lepidoptera, Lycaenidae, Theclinae) and their diagnostics. Rep. Mus. Nat. Hist. Univ.

Wis. (Stevens Point) No. 21, 128 pp.

JOHNSON, K., R.C. EISELE and B. MACPHERSON. 1988. The “Hairstreak Butterflies” (Lycaenidae,

Theclinae) of northwestern Argentina. I. Introduction, Calycopis, Calystryma, Tergissima &

Femniterga. Bull. Allyn Mus. No. 123, 49 pp.

JOHNSON, K. and K.R. KROENLEIN. 1993. Revision of Angulopis (Lepidoptera, Lycaenidae, Theclinae).

Rep. Mus. Nat. Hist. Univ. Wis. (Stevens Point) 33: 38 pp.

SCHWARTZ, A. 1989. The Butterflies of Hispaniola. Gainesville, Univ. of Florida Press, 580 pp.

SHIELDS, O. 1996. Geographic isolation in southwestern North American butterflies (Lepidoptera,

Rhopalocera). Nachr. entomol. Ver. Apollo 17: 71-92.

TILS Note: This article was submitted to outside peer review.

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

TILS Purpose. TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific

discovery, determination, and documentation, without which there can be no preservation of Lepidoptera.

TILS Motto. As a world community, we can not protect that which we do not know.

The Taxonomic Report is projected for publication at the rate of about 10 issues a year. Subscription for Volume

Two is $50 US for domestic and $65 US for overseas subscribers. The subscription year follows the calendar year. All issues

are mailed 1 st class. At the end of each year, subscribers receive that year’s volume on a CDR for permanent archiving and

reproduction for personal use (i.e. a museum or university may make as many copies as needed in whatever format desired).

Non-members may receive individual issues in print any time for $10 per issue. Individual issues on CDR to non-members

are $25 per issue post paid. Subscriptions and individual issue orders should be made payable to TILS and mailed to:

Scott D. Massey, Editor, 126 Wells Road, Goose Creek, SC USA 29445-3413.

Volume 2

25 September 2000

Number 4

M The Taxonomic Report mm

OF THE INTERNA TIONAL LEPIDOPTERA SURVEY WM ^BB

A NEW NORTH AMERICAN SWALLOWTAIL BUTTERFLY:

DESCRIPTION OF A RELICT SUBSPECIES OF PTEROURUS TROILUS

(PAPILIONIDAE) FROM THE SOUTHERN TIP OF FLORIDA.

RONALD R. GATRELLE 1

126 Wells Road, Goose Creek, South Carolina 29445-3413

ABSTRACT. A neotype is designated for Papilio troilus Linnaeus, 1758 from Middleton Place Plantation,

Charleston County, South Carolina. A neotype is designated for Papilio ilioneus J.E. Smith, 1797 from Burke County,

Georgia. Abbot’s ilioneus figures in Smith are the first published representations of nominotypical Pterourus troilus

troilus. Papilio troilus variation texanus Ehrmann, 1900 was described from Houston, Texas. The texanus type is in the

Carnegie Museum, Pittsburgh. The name texanus was restricted (by original description) to a gray male form and is not

subspecifically available under ICZN article 45.5 or 45.6. This name has occasionally been misapplied subspecifically to

populations of troilus from Texas eastward along the Gulf coast to the Atlantic coast of north Florida and southeast Georgia.

All populations in these areas are Pterourus troilus troilus. Rothschild & Jordan (1906), Seitz (1924), and Tyler (1994) are

examined. Pterourus troilus fakahatcheensis Gatrelle is described from the vicinity of the Fakahatchee Strand, Collier

County, Florida. Its range is restricted to the Everglades ecosystem south of Florida Hwy. 80 at the southern tip of Florida.

Pterourus troilus fakahatcheensis is hypothesized to be a peri-Pleistocene relict of the central Florida island or Caribbean

faunas. The fakahatcheensis holotype and both neotypes are currently in the Museum of the Hemispheres (MOTH), Goose

Creek, South Carolina.

Additional key words: Papilio illioneus Cramer.

THE DELINEATION AND APPLICATION OF NAMES

Sometimes even the commonest of butterflies are found to be in need of taxonomic clarification.

Such is the case with Pterourus troilus (Linnaeus, 1758). About ten years ago I became interested in the

insect named Papilio ilioneus by Smith in Abbot, 1797. This eventually led to an examination of all of the

names associated with Pterourus troilus in the southern United States. This paper gives the results of that

investigation and taxonomic study. The names and taxa I researched are: Papilio troilus Linnaeus, 1758,

Papilio ilioneus J.E. Smith, 1797, and Papilio troilus var. texanus Ehrmann, 1900.

ILIONEUS

I began with ilioneus. The area of Screven and Burke counties, Georgia is the region from which

John Abbot depicted his examples of ilioneus (Figs. 1-3). Thus, in 1989 I began making expeditions into

that part of Georgia to observe and collect topotypical specimens for taxonomic examination. I soon found

that two primary forms of male troilus were prevalent in Screven and Burke counties. This was not

surprising as I had found this same variation in troilus males in south coastal South Carolina - which is

what triggered my interest in this taxon in the first place.

1 Research Associate, Florida State Collection of Arthropods, Gainesville, Florida.

The two forms in this area are: 1) individuals with bluish-white submarginal spots on the dorsal

hindwing margins (predominant February through April), and 2) individuals with greenish-blue spots along

the dorsal hindwings (predominant May through July). Both forms are found fairly equally from mid-August

to end of season. All green individuals occasionally occur (Fig 7.) The bluish-white spotted specimens are

Smith’s ilioneus. Beginning in August, the ilioneus form males are usually fresher and smaller than those of

the greenish-blue form which are becoming worn. I attribute the smaller size of ilioneus at that time of year

to heat stressed larvae/pupae in this hot arid region.

In Smith’s 1797 Volume One, the first species he presents are Papilio troilus and Papilio ilioneus .

However, the species Smith depicted as troilus is actually Papilio polyxenes asterius Stoll, 1782. It was

this misidentification and misapplication of Linnaeus’ name troilus to asterius that led Smith to describe

Abbot’s fine figures of typical troilus as an entirely different and new species - ilioneus. Smith’s textual

presentation of troilus (= asterius) contains information that helps us understand why he then introduced

ilioneus as a new species. Smith’s troilus text is as follows (my comments in brackets for clarity).

Figs. 1-3. Abbot’s Papilio ilioneus in Smith, 1797. Fig. 1. Male dorsal. Fig. 2. Female dorsal. Fig. 3. Male ventral, larvae and

pupae on Sassafras. Fig. 4. Neotype d Papilio ilioneus Smith, 1797: 3 August 2000, River Road, Burke County, GA. Fig. 5.

Neotype <5 Papilio troilus Linnaeus, 1758: 19 July 1970, vise. Middleton Place Plantation, Charleston County, SC.

dictory occurring in the two principal systematic authors above mentioned. We beg^leave only to remark that P.

P.E.T. alis caudatis nigris: posticis limbo caerule seentibus angulo ani fulvo; subms maculis bisariis subocellaribus.

[Another black, tailed, species: the background of the hindwings’ border is dark blue in the limbal area and the

Abbot had taken the same species in Virginia as well as Georgia and considered these as one population.

Abbot’s figures of this species, though artistically exaggerated, are accurate paintings of spring troilus with

creamy submarginal spots in the male. The fact that Abbot specifically mentioned rearing specimens from

October larva to spring adults is strong evidence that the light spring form is precisely what he painted. It is

of taxonomic import to note that Smith applied the name ilioneus with certainty only to Abbot’s male and

left open the possibility that Abbot’s female might be another species fide Jones’ assessment.

I have yet to find a valid type specimen (holotype or syntype) for any of Abbot’s Georgian taxa,

including ilioneus. In regard to Smith 1797, I refer the reader to Lucian Harris’ comments on Abbot’s

dealings with Smith in The Butterflies of Georgia (1972) page 7. We can be absolutely certain that a type

specimen for ilioneus never existed and that the name was based solely on Abbot’s figures and notes.

I have designated a male from Burke County with the following label data as a neotype of Papilio

ilioneus (Fig. 4). Small label with RONALD R. GATRELLE, COLLECTOR offset printed. Red label with hand

is died Sms

written inscription: NEOTYPE d\ Papilio ilioneus Smith, 1797, designated by R. Gatrelle 2000 in TTR

2:4. White label with hand written inscription: 3 August 2000, River Road, Burke County, GA. The

specimen is currently in the Museum of the Hemispheres (MOTH) collection, Goose Creek, South Carolina.

Although described as a species, ilioneus must be placed in the synonymy of Pterourus troilus

troilus because these two names are based on the same southeastern subspecific population. Because the

ilioneus form is as frequent as the greenish-blue spotted form in the region of troilus ’ type locality, either

could be considered as the “normal” male form.

TROILUS

Pterourus troilus was described by Linnaeus in a brief Latin statement without figures. The original

description is as follows - in Latin and then as translated into English. (My emphasis in bold.)

Troilus. 5. P.E. alis caudatis nigris: primoribus punctis marginalibus pa 11 id is; posticis subtus maculis fulvis.

M.L.U. Habitat in Indiis.

Troilus. 5. Another black, tailed, species: the margins of the upper wings with prominent pallid spots; the

maculation of the underside of the hindwings is yellowish-brown.

Habitat in [The] Indies [= the New World = southeastern America Colonies].

It is significant that the individual who translated this into English for me rendered Indiis as The

Indies - as in the West Indies. In other species’ descriptions Linnaeus uses India (as well as Indiis). It is

held here that Linnaeus’ India always means the country of India while his Indiis (in regard to troilus) is a

reference (albeit unclear today) to the New World. 3 But where in the 1758 New World? Obviously, the

specimens came from the eastern seaboard of Colonial America. But where in primitive Colonial America?

I am not an expert on Linnaeus’ work or by what channels he may have received specimens and

information. However, having lived for 30 years in Charleston, South Carolina, has given me an appreciation

of this cities’ Colonial history. Founded in 1670, it is one of the nations oldest cities. But more importantly,

Charleston soon became one of the most, if not the most, powerful economic and cultural center in Colonial

America. The College of Charleston (1770), is the oldest municipal college in the nation. The Charleston

Museum is the nation’s oldest Museum and the Charleston Library Society (1748) is the 3 rd oldest library in

the country. The gardens at Middleton Place plantation are America’s oldest landscaped gardens (1741).

Given Charleston’s prominence in the early 1700’s, it should be expected that not a few of the new

species of Lepidoptera being described by the educated and cultured European taxonomists would have

come from the vicinity of this preeminent Colonial city. In fact, it would be strange if this were not so. And

indeed, we find that Hubner, Godart, Fabricius, Cramer, and Linnaeus all described taxa from Charleston

area material. It was from Charleston that Linnaeus described Papilio (=Phoebis) sennae eubule in 1767.

We thus establish three key points. First, Linnaeus’ statement that troilus was resident to the Indies

(in the broad 1758 sense), necessitates that his description was based on coastal southeastern U.S.

Pterourus troilus. Second, in stressing pallid marginal spots as troilus ’ primary dorsal feature, he confirms

that his troilus was based on a phenotype found primarily in the Southeast. (The ibus suffix in primoribus

pluralizes the word to mean both the fore and hind wings.) Third, Linnaeus’ description of troilus precisely

fits Abbot’s (Georgian) ilioneus figure in Smith - which makes these two the exact same thing.

3 Linnaeus was a scientist who wrote technical papers in Latin. But like all of us, he thought in his common vernacular. At

times these two meet in verbal expression. At the time of Linnaeus, the New World, from the southeastern U.S. to upper

South America, was often referred to in the common language as “The Indies.” The proof. Under Proteus (#163 in this same

book), Linnaeus unequivocally lists its type locality as: ‘Habitat in Gramine Americes.” That is: “It lives in grassy America.”

Then, in the next line he states: “Varietes hujus numerosae sunt in Indiis..That is: “This variety is numerous [widespread] in

the Indies [New World - not India]...” Linnaeus was not stupid. He would not say in one line it was from “America,” and in

the next say it was found all over “India.” In some cases the old authors did incorrectly associate location and species. In

other instances [e.g. troilus ] they were only Latinizing their co mm on vernacular. There are Indians, then there are Indians.

Mr. M.J. Scoble of the Linnaean Society, London, has informed me that there is no type for Papilio

troilus there. He further states (in press; 2001) that the species was described from the collection of Queen

Ludovica Ulrika, which is now housed in the Zoological Museum at Uppsala, and that he and Martin Honey

have searched that collection and no type exists there or anywhere else. According to Smith’s 1797 remarks

under troilus, he could not locate a type specimen for troilus even then. Smith, in trying to accurately

determine what true troilus looked like, made a search of Linnaeus’ type specimens and found no troilus

stating: “Unfortunately the Linnaean cabinet possesses neither [ troilus or ilioneus ].” Remember that to

Smith asterius was troilus and troilus was ilioneus.

I do not think it inappropriate to designate a specimen from near Middleton Place, Charleston, South

Carolina as neotype of this species. In fact, it is well within the realm of possibility that the type of troilus

was actually collected at the already famous gardens at Middleton Place in the 1750’s. Accordingly, I have

designated a male from that location and bearing the following label data as neotype of Papilio troilus (Fig.

5). Small label with RONALD R. GATRELLE, COLLECTOR offset printed. Larger red label with hand written

inscription NEOTYPE <?, Papilio troilus Linnaeus 1758, designated by R. Gatrelle 2000 in TTR 2:4.

Second larger label, white with hand written inscription: July 19, 1970, Charleston Co., SC, vise.

Middleton Place Plantation. The specimen is currently deposited in the MOTH collection, Goose Creek, SC.

I chose an ilioneus form male as the troilus neotype because: 1) this phenotype best fits Linnaeus’

original description of a taxon with “prominent pallid [sallow] marginal spots,” and 2) because the ilioneus

phenotype is so frequent, throughout the entire flight season, in south coastal South Carolina that it can

accurately be referred to as the normal topotypical male troilus form.

Unfortunately, taxonomists in general have long misunderstood, and therefore disassociated, the

names troilus and ilioneus. To be blunt, the primary reason for this is that workers not based in the

Southeastern U.S. (which is almost everyone) have historically defined troilus by what was in their “back

yard” and not on the scientific record. Thus, for decades troilus troilus has been defined in their minds (and

published works) by atypical Northeastern or Midwestern populations dominated by blue-spotted (dark)

HW males - populations in which pale spotted males are nonexistent or rare. Therefore, when these

lepidopterists encountered light spotted troilus individuals in northern, eastern, or central Florida they

determined that they were subspecies ilioneus. However, they are actually nominotypical troilus in its

typical form - ilioneus . These workers should have noticed that Linnaeus’ chief dorsal feature was

prominent pallid (creamy) spots. They should have understood what Linnaeus meant by Indiis.

I believe that being familiar with a taxon as it occurs in the field is one of the taxonomists greatest

assets. Museum specimens can only reveal so much. In fact, sometimes they can even be misleading. This is

because many collectors often “target” only certain forms, or specimens of a certain size. This in turn results

in giving a false impression of the phenotypic percentages or average proportions in a wild population. But

mostly, there are many environmental and biological facts that can only be obtained by observing living

specimens in their natural environs. I believe I am as aquatinted with troilus in nature as anyone.

I first became familiar with Pterourus troilus troilus in 1966 while living in St. Louis, Missouri.

We often saw specimens of troilus in our back yard in urban St. Louis. I also observed scores of troilus in

south-central Missouri on our visits to my wife’s grandparents homestead on the banks of the Jacks Fork

River. I only encountered the greenish-blue spotted male phenotype in Missouri.

Since moving to Charleston in 1970 I have studied butterflies in all areas of the state. The only

subspecies in South Carolina is P. t. troilus, with form ilioneus being most frequent in the south coastal

area of the state. Since 1970 I have also often collected Lepidoptera in several counties in northern and

eastern Georgia. Only subspecies P. t. troilus is in Georgia with both male forms. (I have also collected P.

t. troilus in western and central North Carolina, central and eastern Kentucky, eastern Tennessee, northern

Mississippi, and eastern Arkansas.)

While serving in the US Navy, I was stationed in Pensacola, Florida from the early spring of 1968

to the spring of 1970. During that time I collected Lepidoptera extensively in the area between Fort Walton

Beach, Florida and the Mobile Bay in Alabama. P ter our us troilus was a common and widespread species

in that region. Over those two seasons I collected numerous troilus individuals and observed hundreds

more. The only subspecies I observed in that part of the Gulf coast was troilus troilus with greenish-blue

spotted males being dominant. I found only P. t. troilus in the following Gulf coast counties: Alabama:

Baldwin; Florida: Liberty, Bay, Walton, Okaloosa, Santa Rosa, and Escambia.

Over the last 30 years I have made many trips to sites in all of Florida. My experience with the

populations in Nassua, Duval, St. Johns, Clay, Putnam, Marion, Volusia, Levy, Citrus, and Hernando

counties reveals that these are also all referable to subspecies P. troilus troilus, with both male forms

being present in this area ( ilioneus dominant). Nominate troilus in this area are larger and have larger spots

but they are typical troilus in their coloration - especially the females. Male specimens begin to occur

regularly in the southern part of this area in transition toward the southern subspecies.

The tension zone 4 between the two subspecies in Florida becomes pronounced in Orange, Brevard,

Polk, Manatee, and Sarasota counties. I have collected/observed individuals from these areas which are

identical to topotypical troilus (Fig. 6) and others (at the same sites) which strongly lean toward the new

subspecies. In Manatee and Sarasota counties, most male specimens are closer to the new subspecies than

troilus, but are not the new subspecies. The females in these two counties remain close to normal troilus.

From at least 1951 (Klots) to 1994 (Tyler et al) all of the populations in Florida were referred to in

the literature as subspecies ilioneus. This is now know to have been incorrect for two reasons. First,

ilioneus is a synonym of troilus. Second, there are three distinct entities in Florida: P. t. troilus in northern

Florida, intermediates in middle and upper-southern Florida, and fakahatcheensis, Gatrelle nssp. (described

herein) in the southern tip of Florida. In 1994 Tyler (et al) erroneously applied the name texanus Ehrmann,

1900 to all populations from southeast Georgia throughout Florida (less the panhandle).

TEXANUS

In 1900 George A. Ehrmann of Pittsburgh, Pennsylvania published an article in The Canadian

Entomologist titled Variations in Some Common Species of Butterflies. The clear intent of this article was

to give names to variant individual forms of established taxa and not to name any new species or subspecies.

In addition to several other taxa, Ehrmann dealt with three swallowtails in his paper: Papilio polyxenes

asterius Stoll, 1782, Battus philenor (Linnaeus, 1771), and Pterourus troilus (Linnaeus, 1758). He named

a variant male form for each of these swallowtails as follows.

Papilio asterias, Fabr. Var. semi-alba, d , nov. var.

On July 31 st , 1899,1 captured a very interesting form which is out of the ordinary run of the variation which

prevails in this species. The size and markings are the same as the normal form, but all the maculations on the

primaries are pure white, while the markings on the secondaries are of a deep golden yellow. The underside is the

same, but not so conspicuous. Two males in my collection.

Hab.— S.W.Penn’a

Papilio philenor , Linn. Var. obsoleta, d , nov. var.

This form has no submarginal spots either on the fore or hind wings on the upper side; the underside of all the

wings is the same as the normal form. Two males in my collection.

Hab.— S.W.Penn’a

Papilio troilus , Linn. Var. Texanus, d , nov. var.

In this form the light suffusion on the hind wings between the submarginal lunules and the discoidal cell is

replaced by a well-decided band of ashen gray; the band is half an inch wide throughout; the submarginal spots,

both on the fore and hind wings, are much larger than the general form. Expands 4 V 2 inches. Male in my collection.

Hab. —Houston, Texas.

4 A “blend zone” is where two subspecies, one having arisen from the other, blend (a phenomenon of divergent evolution). A

“tension zone” is where two subspecies, of different refugia ancestors, meet and clash (a phenomenon of convergent

evolution).

6

All three swallowtail descriptions are worded similarly. The names given to these individual male

specimens are clearly based on aberrant, gender specific, forms within otherwise normal populations of

nominate subspecies. Under the rules of the ICZN, all of these names are infrasubspecific and unavailable.

To become subspecifically available, an infrasubspecific name would need to meet the condition of either

article 10.2 or article 45.6.4.1.1 do not see the name texanus as having ever met one of these conditions.

Rothschild and Jordan (1906), and Seitz (1924) are the only works that come near to satisfying

45.6.4.1, which had to be met before 1985, but they fall short. In examining these works we should

remember that there is both an extant type specimen (which defines/limits the phenotype) and a type locality

(which defines/limits the possible subspecific genotype). Thus, the name texanus Ehrmann should not be

arbitrarily moved about either geographically (to some other North America genotype - which is what

species and subspecies are all about) or morphologically (to some other distant phenotype).

Rothschild and Jordan (pg. 597) apply the name texanus so broadly as to generally fit all troilus

populations from coastal South Carolina southward. They place all Florida troilus under this name. They

thus include three phenotypes and two subspecies under texanus - troilus troilus in north and upper-central

Florida; intermediates in lower-central and upper-southern Florida; and undescribed subspecies in extreme

southern Florida. Their application of texanus is subspecifically indeterminate relative to Floridian troilus.

They state that Texas specimens are nominate troilus, but raise the preposterous idea that Ehrmann’s

Houston, Texas locality “is perhaps erroneous.” (Ehrmann surely knew where he caught the specimen!)

However, they fail to ask, or answer, the essential question. Is there a Houston, Florida? There is one

Houston in Suwannee County northern Florida. If this is the “Houston” type locality, it is nominate troilus.

To satisfy 45.6.4.1 they would have had to have definitively applied Ehrmann’s name to a single

subspecific population, which they did not do. In fact, they did just the opposite by associating the name on

too broad of a regional and phenotypic basis. Rothschild and Jordan associated the name texanus with three

taxonomically different Floridian entities - and (perhaps) a Texan one too. Article 45.6.4.1 requires

adoption (harmony) not ambiguity (discord).

Had 45.6.4.1 been met by Rothschild and Jordan, then Papilio troilus texanus Ehrmann, 1900

would stand as a subspecific name. But for what geographic subspecies - the one in east Texas (which is P.

t. troilus), the populations in north-central Florida which are also nominate troilus but containing

intermediate individuals between P. t. troilus and the subspecies in extreme southern Florida, or the

undescribed southern Florida subspecies? These populations are far from the same. The vast majority of the

populations Rothschild and Jordan associated this name with fall under the nominate subspecies or

intermediates. By original description, the type specimen limits the name to a gray male form.

If someone would propose that Rothschild and Jordan did meet (by adoption) article 45.6.4.1, then

we would be in the untenable ICZN position of having a type specimen for one taxon (a Florida “texanus ”)

whose aberrant type is representative of another taxon (Texas troilus)\ A taxon is defined first by its extant

biological representative(s), not a name. Also, if we eliminate the Ehrmann type from the equation, then we

would have a subspecies (Florida “texanus ”) with no delimiting description, deposited types, type locality,

or published figures.

The Papilio section in Seitz was written by Dr. Jordan. In this, Jordan simply repeats his 1906

theme. He establishes no type specimen or type locality. His key phrase is “[individual] specimens also

occur.” He thus again applies the name texanus in an uncertain manner. That is, if specimens “also occur”

in one form then they have to “also occur” in another form within his “Florida” population. His statement

is actually accurate as there are three forms, two troilus subspecies and an intermediate, in Florida. But

while it is accurate, it is not subspecifically correct. In actuality, he is only noting that within the

populations in Florida, and perhaps Texas, specimens (individuals) occur that have lighter spotting. His

textual application is nebulous and so out of sink with what occurs in nature that it renders his taxon without

phenotypic definition or geographic limitation (“Texas?”). This presents only more taxonomic confusion not

clarification. Article 45.6.4.1 is not met here.

From at least 1931 (Holland) to 1994 the taxonomy of troilus was fairly stable as virtually all of the

published works followed Ehrmann’s explicit intent and deposited texanus into nominate troilus synonymy.

Unfortunately, after 63 years of taxonomic stability relative to the non-subspecific use of texanus , Tyler et

al (1994) again incorrectly applied the name to all Florida and southern Georgia troilus. A few others have

now followed this (i.e. Mather, 1994 and Calhoun, 1997). None of these can meet article 45.6.4.1 as they

were published after 1985. I hold that the name texanus is subspecifically unavailable because it was

clearly introduced by Ehrmann as infrasubspecific and has never met article 45.6.4.1 or 10.2.

However, for the sake of taxonomic clarity, I here state that because the name Papilio troilus

variation texanus Ehrmann, 1900 (type locality Houston, Texas) was specifically applied, by all the above

authors (including Tyler et al), to Pterourus troilus troilus populations in either eastern Texas, southern

Mississippi, southern Georgia, and/or northern and central Florida, I here affix and confine this name, as a

subspecies (of authors), to the synonymy of nominotypical Papilio troilus Linnaeus, 1758 as this is the only

subspecies in the above mentioned regions. I further state that Papilio troilus variation texanus Ehrmann,

1900, as either a form or a subspecies (of authors) must, by this regional limitation and the ICZN rules of

priority (see below), also be placed in the synonymy of Papilio ilioneus Smith 1797. My position is not

new. This was the accepted (and correct) taxonomic position from 1931 to 1994.

FLORIDIAN TROILUS

The Florida populations of troilus have never been presented correctly in the literature. I believe

this is because many taxonomic workers (especially in the northeastern U.S.) have long misunderstood

nominate troilus as originally described by Linnaeus and Smith (as ilioneus). This is evidenced by the fact

that the nominotypical subspecies has rarely been attributed to the state except in the area of the panhandle

(e.g. Kimball (1965), Gerberg and Arnett (1989), Calhoun (1997)).

The populations in the northern half of Florida are the nominotypical subspecies, troilus. In the

lower half of Florida (except the southern tip) the populations are intermediate to the new subspecies

described herein. Those few workers who have seen specimens from the southern tip of the state have

usually viewed them as only aberrant forms, ecotypes, or extremes at the end of a cline. In the course of this

study I was surprised at how few people have actually collected this undescribed taxon.

The last major taxonomic presentation of Floridian troilus was by Tyler et al (1994). This work in

particular needs to be addressed here due to the serious errors in its presentation of Southeastern troilus.

First, they state (pp. 33, 208, & PI. 93) that they used the name texanus rather than P. ilioneus Smith

because they believed P. ilioneus Smith to be preoccupied by “ilioneus” Cramer (1776) making ilioneus

Smith a homonym. However, Cramer’s illioneus is spelled with two L’s not one as is ilioneus Smith. Under

the ICZN one-letter rule these names are not homonyms. Next, they are also not homonyms under 58.7

because article 58 applies only, “when the nominal taxa they denote are [currently] included in the same

genus or collective group.” Lastly, even if these names were identical, article 23.9.5 requires that ilioneus

Smith be retained because: 1) illioneus Cramer was used in conjunction with Caligo Hubner, 1816 long

before the required date of 1899, and 2) from 1899 to date all workers have employed these names as P.

ilioneus (Papilionidae) and C. illioneus (Nymphalidae). Thus, ilioneus Smith cannot be a disallowed

primary homonym under Caligo illioneus (Cramer) because 23.9.5 forbids it.

Second, the troilus on Plate 93 are replete with errors. Of the thirteen troilus specimens figured six

are in some type of error as follows. Fig. K: Actually a male - cited as a female (1 error). Fig. L: Actually a

dorsal female - cited as both a male and a female ventral (2 errors). Fig. N: Actually a female - cited as a

male on plate but female in appendix (2 errors). Fig. O: Actually a typical troilus form ilioneus with slightly

larger than normal spots, it is part of the Sarasota County tension zone population - presented as Florida

subspecies (1 error). Fig. P: Actually a male - cited as a female. The appendix lists this specimen as from

Texas. If this is so, it is a good example of a specimen of one subspecies ( troilus ) which looks like (but can

not be) another subspecies - which they say it is (2 errors) (it also demonstrates that Ehrmann’s type is

Texan). Fig. Q: Actually a very typical troilus troilus female from Jacksonville, Florida - cited as south

Florida subspecies (compare to J and R) (1 error).

Only figures 10, 12, and R are specimens of the new south Florida subspecies described herein. The

rest are either troilus troilus dark form, intermediates (K & O), or atypical specimens compared to the

norm in the populations from which they were taken (N & P). I also strongly suspect that female specimen N

is mislabeled. I have spent a lot of time in the area of New Smyrna Beach over the years and have never

seen a troilus in that area which even remotely resembled this specimen.

Tyler et al does not meet 10.2 in regard to the name texanus for several reasons, not the least of

which, is the fact that they apply the name, without type locality (except possibly Texas) to all Floridian

forms and subspecies (except in the panhandle).

A NEW SUBSPECIES OF PTEROURUS TROILUS

The preceding sections of this paper have been put forth to bring about the proper delineation and

application of those names which have already been published. However, even if the names troilus ,

ilioneus and texanus were to remain in place in their prior (incorrect) usage, it would have no effect on the

validity of the new subspecies about to be described in this section. This is because all of the above names

have been specifically applied to all of Florida. This paper is not the “renaming” of an already described

taxon because the Everglades ecosystem segregate has never before been recognized as subspecifically

distinct from the rest of Floridian troilus - nor has a name ever been proposed for this segregate before.

Yet, it is evident that for some time it has been known that the population of Pterourus troilus in

extreme southern Florida is at least phenotypically unique (e.g. Clark and Clark, 1951). Why someone has

not taken the time (and trouble) to fully research this is odd to me. Instead workers have either relied on the

opinions of others or sought to take the path of least resistance by haphazardly and broadly overlaying the

existent, but non-applicable, names ilioneus or texanus to specimens from this area of Florida.

This paper is the first evolutionary taxonomic investigation of the troilus populations in the extreme

southern tip of Florida. As such, the first focus is not the beauty of this taxon or its unusual phenotype. As

with all subspecies, the first inquire should be one of evolutionary origin. Where did this taxon come from?

The occurrence of intermediate forms in central Florida could indicate that the south Florida

phenotype is simply the extreme end of a long blend zone with no real definitive evolutionary origin. Or, the

presence of seasonally similar forms in other parts of the South (from Texas to Florida to South Carolina)

could indicate that this is simply an extreme expression of an ecotype. That is, a phenotype produced by

climate or environment and not evolution. Neither of these are the case.

Dr. Richard Boscoe of Lafayette Hill, Pennsylvania has reared this subspecies from Collier County

females. Boscoe’s females oviposited on Per sea borbonia (L.) (Red Bay), which is the primary host of this

subspecies. He then reared the larvae on Sassafras albidum (Nuttall) (Sassafras) in the northeastern U.S.

Eleven of these reared specimens are in the FSCA collection, Gainesville. They are all typical males of new

subspecies fakahatcheensis . This is firm evidence that fakahatcheensis is not an ecotype because its normal

phenotype was produced in specimens reared in the northern subspecies environment and on its host.

Dr. Jeff Slotten of Gainesville, Florida has a great deal of field experience, over several years, with

this segregate in the Fakahatchee Strand area of Collier County. (I have only three days experience with this

taxon in Collier County.) He has made a very important field observation relative to the timing of this

taxon’s adult flights. This subspecies acts much like Heraclides aristodemus ponceanus (Schaus, 1911) in

that its adult emergence is evidently tied to the advent of periods of rain. I have not found this phenomenon

to exist in any other part of troilus range. This is an evolutionarily unique event that gives evidence as to the

origin of this subspecies.

9

A precipitation related emergence pattern is common among tropical and subtropical Lepidoptera as

they have made the evolutionary adjustment to wet and dry seasons. The hypothesis here is that this survival

mechanism arose in a peri-Pleistocene Caribbean or Island Florida ancestor when this type of acclimation

was essential for species survival due to extended periods of dry weather. I lean toward a Caribbean

ancestor that arrived in Florida about the same time as ponceanus ’ ancestor. Museum specimens indicate

that ponceanus and fakahatcheensis were once sympatric on the mainland south of Miami.

This is reinforced by records of this subspecies from the Florida Keys and Cuba. Opler & Krizek

(1984) pg. 51 and Scott (1986) pg. 183 regarded the Cuban specimens as strays. However, they could also

be vestiges of a still extant population there, or the last specimens of a now extinct population. (The

nominate subspecies of the otherwise temperate Papilio polyxenes Fabricius, 1775 is a Cuban endemic.)

King in Kimball (1965) recorded this troilus as ranging into the Keys. In any case, fakahatcheensis was

once isolated from the northern (mainland) subspecies for at least a few thousand years.

Therefore, the intermediate forms in central Florida have come into being in only one of two ways.

If fakahatcheensis ’ origin was the central Florida island, then (beginning at the time of its contact with

mainland troilus) the northern troilus genes are diluting the fakahatcheensis gene pool and pushing it out

(further south). Or, if fakahatcheensis ’ origin was the Caribbean (which I believe is the case), then its

genes are incorporating into the troilus gene pool and moving the boundary of troilus further north. Either

way, fakahatcheensis is not an ecotype (as is ilioneus) or the end of a cline. Central Florida is a tension zone

where gene pools are struggling for dominance, stability, and subspecific identity.

There are several phenotypic characters in fakahatcheensis that manifest both its evolutionary

distance from nominate troilus and closeness to Pterourus palamedes (Drury, [1773]). All fakahatcheensis

males and females have the lower five VFW postmedian spots well developed with the great majority of

males having these five spots clearly defined on the DFW also. (There are two specimens in the type series

with eight DFW spots in this band.) In this they resemble palamedes more than nominate troilus whose

males only occasionally have two to four (rarely more) of these spots vaguely present. In troilus these spots

are bluish and rarely whitish, while in fakahatcheensis they are whitish and rarely bluish.

In the great majority of fakahatcheensis males the yellow line at the base of the VFW costal margin

is just as prominent as it is in palamedes. In troilus this feature is usually absent or lightly present. The

VHW yellow line in SM 2 of palamedes is occasionally represented in fakahatcheensis males by a diffuse

streak in the same area. If this occurs in troilus , I have not seen it.

The DHW submarginal row of spots in female fakahatcheensis is often yellow and thus exactly as

in the male. Thus, the fakahatcheensis sexes are much more alike than the troilus sexes are. In this they also

lean to palamedes whose sexes are similar. It is worth noting that the sexes of Cuban Papilio polyxenes are

much alike also. There is no mimicry of Battus philenor (Linnaeus, 1771) by fakahatcheensis females

which in flight look most like the dark females of Pterourus glaucus (Linnaeus, 1758).

These characters manifest fakahatcheensis as evolving between palamedes and nominate troilus.

Fakahatcheensis is not an ecotype or something at the “end of a cline.” It is a relict subspecies struggling

against the northern troilus gene pool for its subspecific integrity.

Pterourus troilus fakahatcheensis Gatrelle, new subspecies.

Diagnosis and Description. Female (Fig. 11). Subspecies fakahatcheensis and troilus are delineated most clearly

in the female - phenotypically and geographically. The regional shift in Florida from subspecies troilus to fakahatcheensis

is much more abrupt in females than in males. The males of these two integrate widely from Orlando to Lake Okeechobee. In

this same area, the great majority of females are typical troilus with yellow spotted females being virtually unknown. In

subspecies troilus, males are dimorphic while the females have only one form. In subspecies fakahatcheensis, the opposite

is found with fakahatcheensis males being relatively homogenous and the females variable. About 10% of fakahatcheensis

females have bluish-green DHW submarginal spots, another 25% have these spots intermediate, with the remaining 65%

having pale yellow DHW submarginal spots. These spots in troilus females are bluish. There are three primary differentiating

characters between the females of P. t. troilus and P. t. fakahatcheensis. First, in fakahatcheensis the submarginal spots are

10

11

ACKNOWLEDGMENTS

I thank John Heppner of the FSCA for reference information and access to that collection, Jeff

Slotten for field information and access to his specimens, J.D. Lafontaine for the OD of Cramer’s illioneus,

Malcolm Scoble of the Linnaean Society for information regarding the type of Papilio troilus , Ms Rebecca

McBee for Latin translation and application, the Charleston Library Society (Charleston, SC) for permission

to reproduce their copy of Smith’s figures of Papilio ilioneus. My son, Ben Gatrelle, and son-in-law, Joe

Mueller, for photography of specimens and many hours of help with the computer. I also thank the various

lepidopterists who, often without their even knowing it, greatly influenced this project through their advise,

criticism, questions, or anecdotal information.

LITERATURE CITED

CALHOUN, J.V. 1997. Updated List of the Butterflies and Skippers of Florida. Holo. Lepid. 4(2): pp. 39-

50. Gainesville, FI.

CLARK, A.H. &L.F. CLARK, 1951. The Butterflies of Virginia. Smithsonian Misc. Col. 116 (7) 239 pp.

CRAMER, P. 1776. Uitlandsche Kapellen (Papillons Exotique). 1: 52. Amsterdam.

EHRMANN, G.A. 1900. Variations in Some Common Species of Butterflies. Canadian Ent. 32:348.

GATRELLE, R.R. 2000. Description of a New Subspecies of Poanes aaroni (Hesperioidae: Hesperiinae) From

the West Central Gulf Coast of the Southern United States. The Taxonomic Report. Vol. 2 (2), 10 pp. The

Int. Lepid. Survey, Goose Creek, SC.

GERBERG, E.J. & H.R. ARNETT, Jr. 1989. Florida Butterflies. Nat. Sci. Pub., Inc., Baltimore, MD. 90 pp.

HARRIS, L., Jr. 1972. Butterflies of Georgia. Univ. of Okla. press, Norman OK. 326 pp.

HOLLAND, W.J. 1931. The Butterfly Book. Doubleday & Company, Inc., Garden City, NY. 424 pp.

HONEY, M.R. & M.J. SCOBLE. (2001, in press). Linnaeus’s butterflies (Lepidoptera: Papilionoidea and

Hesperioidea). Zool. J. of the Linnaean Soc., London.

INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE. 1 January, 2000. Fourth Ed. London.

KIMBALL, C.P. 1965. The Lepidoptera of Florida, an Annotated Checklist. In Arthropods of Florida and

Neighboring Land Areas. Vol.l. Gainesville: DPI. Fla. Dept. Agric. 363 pp, 26 pi.

KLOTS, A. B. 1951. A Field Guide to the Butterflies. Houghton Mifflin, Boston MA. 349 pp.

LINNAEUS, C. 1758. Systema Naturae per Regna Tria Naturae. Revised Tenth Edition.

MATHER, B. & E. DINGUS. 1994. Butterflies of Mississippi -A Field Checklist. Miss. Mus. of Nat. Sci.,

Miss. Dept, of WFP. Jackson, MS.

OPLER, P.A. & G.O. KRIZEK. 1984. Butterflies East of the Great Plains, An Illustrated Natural History.

Johns Hopkins Univ. Press, Baltimore, MD. 294 pp.

ROTHSCHILD, W. & K. JORDAN. 1906. A Revision of the American Papilios. Novitates Zoologica, Vol.

13. 341 pp. Tring, England.

SCOTT, J.A. 1986. The Butterflies of North America, A Natural History and Field Guide. Stanford Univ.

Press, Stanford, CA. 583 pp.

SEITZ, A. 1924. (Editor). The Macrolepidoptera of the World. Vol. V. The American Rhopalocera.

Various authors, Stuttgart.

SMITH, J.E. 1997. (in Abbot) Natural History of the Rarer Lepidopterous Insects of Georgia , Vol. 1:3

TYLER, H.A., K.S. BROWN, Jr. & K.H.WILSON. 1994. Swallowtail Butterflies of the Americas: a Study

in Biological Dynamics, Ecological Diversity, Biosystematics, and Conservation. Gainesville,

Florida Scientific Pub. 376 pp.

12

Fig. 6. Pterourus troilus d ( ilioneusj: 1 Mar. 1991, old Hwy. 37, S. of Mulberry, Polk Co., FL. Fig. 7. P. troilus d (green):

3 Aug. 2000, Brigham Landing., Burke Co., GA. Fig. 8. Holotype d P. t. fakahatcheensis: data in text. Fig. 9. P. troilus 9: 8

May 2000, Goose Creek, Berkeley Co., SC. Fig. 10. Paratype d (ventral) P. t. fakahatcheensis: 27 March 1999, Collier Co.,

FI. (leg. Slotten). Fig. 11. Allotype 9 P. t. fakahatcheensis: data in text. (All leg. R. Gatrelle, except 10.) All figures X-l.

13

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

TILS Purpose. TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific

discovery, determination, and documentation, without which there can be no preservation of Lepidoptera.

TILS Motto. As a world community, we can not protect that which we do not know.

The Taxonomic Report is projected for publication at the rate of 10 issues a year. Subscription/

dues for Volume Two are $50 US for USA and Canada, $65 US for overseas subscribers. The subscription

year follows the calendar year. All issues are mailed 1 st class. At the end of each year, subscribers receive

that year’s volume on a record-only compact disc (CDR) for permanent archiving and reproduction for

personal use (i.e. a museum or university may make as many copies as needed in whatever format desired).

Non-members may receive individual issues in print any time for $10 per issue. Individual issues on CDR

to non-members are $25 per issue post paid. Subscriptions and individual issue orders should be made

payable to TILS; and mailed to: Scott D. Massey, Editor, 126 Wells Road, Goose Creek, SC USA 29445-

3413.

Articles for publication are sought. They may deal with any area of taxonomic research on

Lepidoptera. Before sending a manuscript, simply write TILS at the above address to set up discussion on

how to best handle your research for publication.

Everyday around the world, in jungles and urban areas alike, insect species and subspecies are

becoming extinct. Every year scores of taxa are lost that have not even been scientifically discovered and

documented. Thus, their extinction is unnoticed because their existence is unknown. They are unknown

simply because they have not been collected and systematically identified. Without systematic taxonomy

there is nothing. Without the collection, and exchange of specimens (information) there will be no

systematic taxonomy. Without amateur collectors the majority of the undiscovered species/subspecies will

vanish before they are discovered. Be it butterflies or moon rocks, collecting is the first step of access to all

other scientific information - and protection.

Donations are needed to support and further our efforts to discover and protect

butterflies worldwide. All donations are US tax deductible. Please help generously.

Donations should be mailed to: Scott D. Massey, Treasurer, 126 Wells Rd., Goose Creek, SC 29445.

Checks should be made payable to: TILS. Please indicate if you need an individual receipt.

Catch us on the web at: www.tils-ttr.org

14

Volume 2

15 December 2000

Number 5

The Taxonomic Report

OF THE INTERNA TIONAL LEPIDOPTERA SURVEY

THE BIOLOGY, LIFE HISTORY, AND TAXONOMY OF CELASTRINA

NEGLECTAMAJOR (LYCAENIDAE: POLYOMMATINAE).

HARRY PAVULAAN 1

HISTORICAL SYSTEMATIC TREATMENT

“Azure” inhabiting the Appalachian Mountain region was first recognized as a valid taxon

Edwards, who intentionally redescribed it (Figs. 25-29) as Lycaena pseudargiolus

,e Conte) in his early work on West Virginia butterflies (Edwards, 1866, 1868). Edwards

i be distinct from his newly described species Lycaena neglecta (Edwards,

lieved to be of more northerly regions: “Massachusetts, New York, Wisconsin,

la violacea (Edwards, 1866) of the central Appalachian region. Edwards

rse second brood flew in July in West Virginia, which he believed was the

)lus. He did not adequately explain why he identified this brood with May

instead of his more northerly L. neglecta. In subsequent works he would

broods as neglecta. It is apparent to us that neglecta was not common in

i at the time, and Edwards’ faulty interpretation of his rearing results led him

oduced the small numbers of summer adults that he found in nature.

By 1875, Edwards changed his mind on the separate specificity of pseudargiolus [sensu Edwards], neglecta

and violacea in West Virginia. Edwards (1875) now identified violacea with pseudargiolus, but the status of neglecta

was left uncertain. At this time he still believed that the July and September flights in West Virginia were the progeny

of May pseudargiolus [sensu Edwards], and reported on a rearing experiment involving the offspring of September

adults: “...unexpected result shows violacea to be the spring form of pseudargiolus .” Of neglecta, he added: “I am

prepared to believe that neglecta may prove to be one of the summer broods of pseudargiolus in this latitude... There

are differences between the two forms sufficient to make me regard them as distinct...” At this point, Edwards still did

not identify the West Virginia summer broods with his neglecta of the north.

Another major writer at the time, Scudder (1876), basing his concept of Celastrina taxonomy on an entirely

different set of observations taken in New England, disputed Edwards’ outline of North American Celastrina.

Scudder believed that pseudargiolus represented large variant individuals of summer “form” neglecta in New

England. Writing about the three major “forms” of C. ladon found in the vicinity of Boston, Massachusetts, he states

of the summer brood: “a form...corresponding to the neglecta of W. Virginia; occasionally in midsummer large

specimens of this are taken, and these I have considered Pseudargiolus .” Thus, neglectamajor was immediately

misunderstood by at least one major writer of that time.

Edwards continued to rear West Virginia Celastrina and asserted (Edwards, 1878a): “The eggs laid by

violacea give larvae from which comes pseudargiolus last of May...” Further, “The female pseudargiolus lays

eggs...to produce perhaps violacea, but also perhaps the typical pseudargiolus again. But a small percentage...give

butterflies at irregular intervals during the year.” Edwards observed that these adults were always smaller than the

parents, an interesting phenomenon which we have also discovered in rearing neglectamajor. These “false” summer

brood adults are extremely similar in phenotype to neglecta, especially with their consistently smaller size. Thus

Edwards felt there were two regular annual broods of Celastrina in West Virginia, first violacea in April and then

pseudargiolus in May. Of the summer broods (neglecta), he felt their occurrence was too irregular to represent a

regular annual brood. Edwards apparently found populations of neglecta to be rather unpredictable in areas where he

studied them, no doubt a local ecological circumstance of heavily forested Appalachian habitats, “...in June, 1866, at

Coalburgh, neglecta appeared in large numbers...[i]n following years (1867 - 1868) neglecta has again been rare in

this district.” He did not report such fluctuations with ladon or neglectamajor.

Edwards was never single-minded on the issue of broods and repeatedly revised his own ideas. In 1881 he

reasserted that violacea produced pseudargiolus [sensu Edwards]. In the ensuing years Edwards (1883) rejected his

earlier conclusions: “...the fact that the hybemating chrysalids of the May generation (pseudargiolus) produce their

own form of butterfly... and also that the larger part of the chrysalids of violacea appear to hybemate, and that nearly

all the butterflies of the first generation must come in direct descent from their own form of the year before.” Edwards

had come full circle to believe that neglectamajor bred true. However, he was still confused by his observation that:

“Eggs laid by Pseudargiolus produce the same form the same year in very small numbers, but most of the chrysalids

hybernate to produce the same form the next spring. The few butterflies which emerge the same summer are

sometimes as large as the parent female, but are usually smaller.”

We have also discovered that in rearing larvae and maintaining chrysalides under artificial conditions, a false

summer brood can be “forced” out of some neglectamajor. Some of the resultant adults are phenotypically similar to

the natural form, but most are smaller and resemble neglecta to an extent. Edwards confused these false neglecta-Wke

summer individuals with true neglecta..

Edwards (1883) further stated: “The observations on the chrysalids and those on the appearance of the

butterflies in the field therefore agree, and together show that Pseudargiolus of May cannot proceed from violacea

butterflies of that year, but must come from hybemating chrysalids. The later butterflies, Neglecta... must come from

violacea butterflies of the same year. A small percentage of the chrysalids from violacea give butterflies at fifty or

more days from the eggs laid in April, which brings the emergence into June, and the result is Neglecta’, w hil e nearly

all the chrysalids hybemate...to give violacea the next spring.” He also added: 4 Pseudargiolus is an interpolated and

distinct generation, the first in the year of its series. It has no direct connection with the winter forms [violacea in

West Virginia], but an indirect one through the few individuals which spring from it late in the season.” Thus Edwards

felt that his pseudargiolus must produce some second brood neglecta-l\ke offspring in nature, which then bred with

neglecta to maintain a connection to species pseudargiolus. This erroneous conclusion was no doubt based on the

artificial rearing conditions that produced some summer adults from neglectamajor larvae raised on Cimicifuga

racemosa. In continuing, he stated: “The few late females spmng from Pseudargiolus, and which

3

emerge from chrysalis irregularly in August and September, lay eggs, and the chrysalids thereupon hybemate and

produce Violacea in the following spring.” These findings were based on wild larvae he found on a neglecta host,

Actinomeris alternifolia , in October. He never realized that his reared summer adults of neglectamajor were not the

same as the naturally occurring summer insect, neglecta, he found ovipositing on Actinomeris.

The next year, Edwards (1884) finalized his personal concept of all the North American races and forms. It

was plainly evident that he was still very confused over the status of pseudargiolus. Edwards said of neglectamajor.

“The chrysalids from the May generation, or Pseudargiolus, probably produce butterflies in small numbers in July

and later, after the June Neglecta have passed away, but most of them hybernate, and give Pseudargiolus the

following May, or earlier.” He ultimately realized that both larvae of neglecta and neglectamajor utilized the same

Cimicifuga host plants and are often found feeding together (though differing in age) on the same individual flower

clusters. Further, he states: “In the spring, there certainly is no connection between Violacea and Pseudargiolus... It

is only by a connection between Pseudargiolus and the other forms in the fall that any inter-relationship can be found;

that is, some chrysalids of Pseudargiolus give butterflies which unite with butterflies from chrysalids of the June

Neglecta to produce the fall larvae, from which come Violacea in April. Otherwise Pseudargiolus would be set in the

middle of the series, with no link in either direction. The true second generation of the year, in Virginia, is Neglecta,

appearing in June. Pseudargiolus is an interpolated spring generation, the first in the year of its series. Its second

comprises a part of the few butterflies which fly between July and October. If these late butterflies were suppressed,

Pseudargiolus would stand as a distinct species, with no trace of its relation to the other forms.” In Edwards (1885),

he repeated this statement and added: “No doubt that is one way in which species come to exist.”

Because of Edwards’ misinterpretation of the emergence of reared neglectamajor summer adults, he felt

there was some degree of reproductive/genetic connection to ladon and neglecta (through these partial summer

emergences). He was thus blind to the fact that the larvae of two separate taxa, neglecta and his pseudargiolus (=

neglectamajor ), were actually feeding on the same plant. Rather, he maintained a belief that they were merely two

separate forms. He therefore falsely concluded (Edwards, 1884) of the several distinct species of North American

Celastrina that: “...their history has come to be thoroughly known, and it is found that they...constitute one

polymorphic species, which has possession of the broad continent, from the boreal regions to Mexico.” With these

words, the mistaken concept of a single species of North American Celastrina species was cast. A concept that

prevailed for over a century! Neglectamajor roamed within this conceptual boundary in unquestioned synonymy,

misunderstood by one author after another. The account of its rediscovery has taken on the flavor of fo lkl ore.

Scudder (1889) continued to consider pseudargiolus [sensu Edwards] as merely “large examples of the

summer form.” The summer insect found in New England is neglecta. It is only slightly variable in size, but not to

the extent that Scudder alluded. Others proceeded to follow Scudder’s concept without explanation or further

investigation, reinforcing this incorrect alignment. In Comstock and Comstock (1904), neglectamajor was given a

brief description as the typical, late spring form (ladon) of Cyaniris ladon, which the authors felt to be the older

name. The Comstocks cited Scudder, but misunderstood his concept: “Scudder does not regard neglecta as distinct

from C. ladon ladon, or pseudargiolus... according to this view neglecta is one of the spring forms as well as the

summer form.”

Tutt (1908) applied the name neglecta-major to Edwards’ insect, which he considered to be a distinct size

form of Celastrina argiolus pseudargiolus (Boisduval & Le Conte). His description was extremely brief and

misleading. In making a comparison of Edwards’ (1884) illustrations of form pseudargiolus to form neglecta, Tutt

described form neglecta-major as: “Exactly parallel pair only larger. Underside white, with contrasting black spots,

many, however, obsolete ^pseudargiolus, Edw., pi. ii., figs. 8-9 ^neglecta, large form, Scudder. (No doubt a

summer form, which may be called neglecta-major, though Edwards insists that it comes from overwintering pupa).”

This incorrect placement was repeatedly reinforced by subsequent writers. Skinner (1915), no doubt

following Scudder, wrote of Lycaena argiolus : ‘Pseudargiolus and neglecta are the same thing.” Field (1938) stated

of Kansas Azures: “The form neglecta-maor [sic] (Tutt) represents the common summer form.” Comstock (1940)

wrote of Lycaenopsis argiolus pseudargiolus in New Jersey: “Form vernalis neglecta-major Tutt. This form follows

the early spring forms and is intermediate in occurrence between them and the first summer brood.” Klots (1951),

under Lycaenopsis argiolus pseudargiolus, mentioned: “In some regions occurs a partial second brood (neglecta-

major)... This is ‘spotty’ in its occurrence, but I know some places where it occurs very constantly.” Clark and Clark

(1951), referring to the late spring emergence of Cyaniris argiolus pseudargiolus in the Piedmont and Coastal Plain

regions of Virginia, stated: “in the mountains...it is largely replaced by a larger form (neglecta-major )...”

n of it is found west of Denver.” The inclusion of B

en C. ladon sidara and C. neglecta, analogous to Appalachian neglectamajor. It ft

w (Hop Vine) and Lupinus argenteus (Lupine). Shull’s Indiana work (1987) entire

c’s treatment of C. neglectamajor, placing it under Celastrina ladon ladon : “Paul O

Iftner et.al. (1992) provided an extensive P description of the biology of neglectamajor in Ohio. Opler & Malikul (1992)

adopted Appalachian Azure as the common name. The host plant was liste

York City area. Wright (1995)^inttoduce/a new^concept ^of

Id & Burger (1997)

^npre^CoSfgW

(1884) and the writings of Tutt (1

REGIONAL OBSERVATIONS

hybrid features. By the ti

it is on the wane and tf

Will neglectamajor, which flies earlier in shaded woodhuKfFlight dates in southe

d-May (extremes April 5 - May 24); n,

d in the vicinity of stands of th

Is and a variety of oj

;r period of spring (May l:

»■ and neglecta fly n

it it had not yet emerged or perhaps was not pi

jalized climatic conditions^cool air and late sp:

peered wom y a nd aged, like those in the Great Sr

. On May 25, a fresh pair of atypical, large

e (May 25), in the sar

mpression is that the ft

L Only a single very a

on July 7,10,11, & 22. L

1983) and

timing of th/adult flight and started^he divergence

BIOLOGY AND LIFE HISTORY

(2) northern

TABLE 1.

Population Group

Number of

Minimum

Maximum

Mean

Median size

Mode

specimens

size (mm.)

size (mm.)

(mm.) (S.E.)

(mm.)

(mm.)

Southeastern Pennsylvania

60

11.5

17.0

14.7 (±0.2)

14.7

15.0

Northern Virginia

272

11.0

17.5

14.9 (±0.2)

15.0

16.0

Western North Carolina

192

13.0

18.5

16.2 (±0.1)

16.1

17.0

The smallest adults (11-14 mm) from northern Virginia (Figs. 8 & 9) frequently display deeper dorsal violet-blue in

collected on May 13, 2000 near Berrys, VA

(Fig. 12) measures 14 mm. and the female (Fig. 13)

variation found in eastern populations. However, the ft

have a considerable degree of white sc

es have extensive areas of white on the hi

length (range 10.0 mm - 15.01

12.6-17.0 mm).

11.0

> submarginal band. Many neglecta females have a faded scalloped

ss of the ventral hindwing. This character is generally lacking in m

while*that^of ar^oto'tendst^sharply^gidtde

uniformly smooth in neglecta, but in neglectam <

ajor (Fig. G2) the inferior surface of the spine is thick and irregular.

10

(Fig. DUn ted *h® g round^°lor wi^es from bright^green, olive green^dull pea green.

cease their feeding activity and prepfreto pupate, the coIot flways fades to a cream pink. The head is disproportionately

small, located at the end of a long extendable neck. The neck is attached to the anterior of the prothorax, which forms a

the interior of the host buds, ^completely hidden from view. Young larvae rest wholly on the floral bud, curving about it

completely. Older larvae seem to rest wherever they can maintain a hold, curving the forward part of their body around the

Tutt by Pavulaan and Wright. The corresponding drawings in Edwards (1884) are shown (Figs. 25-27, 29). Since Tutt

d his description of form neglecta-major on Edwards’ figures (1884: pi. “Lycaena II” figs. 8, 9), the types had to be

ards’ originals from which the figures were created (Art. 74.4, 2000ICZN Code)). The labels read as follows:

/therefore lectotype (male) / of ne

|; (3) “probably the model / of Edw. pi. “Lycaena II” / fig. 8 and un fig. 9

major / Tutt 1908: 407 / H. Clench 1975.” [handwriting of H.C. Clench];

of / Edw ( . plate '^Lycaena II” / fig. 9 (up), and therefore / lectotype (female) of / neglecta-mtjor Tuft / 1908: 407 /

(1884: pi. “Lycaena II”, fig. 22) (Fig. 29) and subsequently given the form name “obsoleLfunulata ” by Tutt (1908: 426).

?d by Opler & Krizek (1984), is

er&Krizek (1984) treated theta

(Map 2). Essentially Appalfchian, with separated coloniis in the Ozlrk region. Northernmost records near Albany, N.Y., and

Murray, Rabun, Union, White; INDIANA: Ripley; KENTUCKY: Floyd, Graves, Jackson, Jefferson, Harlan, McCreary,

bers of chrysalides may break diapause e.

le for breaking diapause is not fully unde.

e Code’s rules of a'

s 11-18) to give Tutt ai

did not do. Opler, in personal communication to Wright, stated that he and Krizek did not intend to author a species name in

ip is Tutt’s from 1908. If they di

E&F). Edwards was th

14

Shapiro (1966) listed the host plant as being 1

Pratt and Ballmer (1991) reported that neglectamajor larvalcan be successfully reared on Lotus scoparius in the

lab. This legume plant is a nearly universal alternative host for a wide variety of lycaenid butterflies. Our rearing

the abdomen (A7). The gland secretes a’sweef fluid droplet whichthe ants consume. This is known as “milking”. A pair of

cs Co., PA, 6/23/87: F

*. P. M. Marsh, R. W. Carlson, D. L.

Agryponcushmani{ Dasch), a

within the mature larvae and

s the presence of A

of Arthropods (John Heppner), II

Natural History (Julian Donahue),

Furth and Stephan Cover), New .

State University Museum of Bi(

te of Science (John

it of Forestry (John

y of Delaware (Dale Bray and Tom Wood), University of Guelph (;

Michigan Museum of Zoology (Mark

i, Tom Carr, Charles V. Covell, Harry N. Darrow, Link I

Richard Heitzman, John Hyatt, David Iftner, John Hyatt, Phil Kean, Harry King, Ron King, Marc M

g in retrieval of the published works of W. H. Edwards and Tutt. We are grateful to Jim

;y Butterfly Club) and Steve Walter (New York City Butterfly Club) who kindly shared

their respective databases. Our appreciation is extended to the expertise and efforts of R. W. Carlson, T. J. Henry, P.

M. Marsh, S. R. Shaw, D. R. Smith and D. L. Vincent of the U.S. Dept, of Agriculture, Systematic Entomology

N.C. during the period of May 30 - June g 1, 1

i, R. 1993. A D

and Long Island. New York City B

k, A.H., and L.F. Clark. 1951. The B

ich, H.K. 1972. Celastrina ebenina, ,

I, est. 1974-78). Celastrina pi

f, nrDNA ITS, and cpDNA trnL-F S<

Taxon 47:593-634.

14:54-58.

218 pp., 50 pi. CLycaenal”

r, D.C., J.A. Shuey, a

Vol. 9, No. 1, xii +

s, A.B. 1951. A Field

Mifflin Company, xvi + 349 pp.

Miller, L.D., and F.M. Brown. 1981. A c<

2, 280 pp.

North American Butterfly Association. 1993. English names for North American Butterflies. Amer. Butt. 1:21-29.

Opler, P.A., and G.O. Krizek. 1984. Butterflies East of the Great Plains. Baltimore: J. Hopkins Univ. Press, 294 pp.

Opler, P.A., and V. Malikul. 1992. Eastern Butterflies. New York: Houghton Mifflin Company, xvii + 396 pp.

a. Ent. News 26 (7):329.

Figures A, B, G, & H. Mature larvae of C. neglectamajor on host. Fig. A. Mottled morph, 22 June 1987, St. Game Lands #157

Bucks Co., PA. Fig. B. Red morph (most common in cent. & n. Appalachians) mid June, 1992, Elkins, Randolph Co., WV. Fig.

G. Green morph, June 1987, same site as A. Fig. H. Light green morph attended by Camponotus pennsylvanicus ants, 23 June

1994, same site as A Fig. C. 1st instar C. neglectamajor (yellow with short blunt-end dorsal setae) on racemosa bud, 23 June

1987, same site as A. Fig. D. 1st instar C. neglecta (green with long pointed dorsal setae) on racemosa bud, 23 June 1987, same

site as A. Fig. E. Ova of C. neglectamajor on bud spike of C. racemosa, 1 June 1990, same site as A. Fig. F. Flowering spike of

Cimicifuga racemosa. Spring Valley Pk., Montgomery Co., PA, 9 July 1994. Photos: B by Tom Allen, all others by Wright.

18

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

TILS Purpose. TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific

discovery, determination, and documentation, without which there can be no preservation of Lepidoptera.

TILS Motto. As a world community, we can not protect that which we do not know.

The Taxonomic Report is projected for publication at the rate of 10 issues a year. Subscription/

dues for Volume Two are $50 US for domestic and $60 US for overseas subscribers. The subscription year

follows the calendar year. All issues are mailed 1 st class. At the end of each year, subscribers receive that

year’s volume on a record-only compact disc (CDR) for permanent archiving and reproduction for personal

use (i.e. a museum or university may make as many copies as needed in whatever format desired). Non¬

members may receive individual issues in print any time for $10 per issue. Individual issues on CDR to

non-members are $25 per issue post paid. Subscriptions and individual issue orders should be made

payable to TILS; and mailed to: Scott D. Massey, Editor, 126 Wells Road, Goose Creek, SC USA 29445-

3413.

Articles for publication are sought. They may deal with any area of taxonomic research on

Lepidoptera. Before sending a manuscript, simply write TILS at the above address to set up discussion on

how to best handle your research for publication.

Everyday around the world, in jungles and urban areas alike, insect species and subspecies are

becoming extinct. Every year scores of taxa are lost that have not even been scientifically discovered and

documented. Thus, their extinction is unnoticed because their existence is unknown. They are unknown

simply because they have not been collected and systematically identified. Without systematic taxonomy

there is nothing. Without the collection, and exchange of specimens (information) there will be no

systematic taxonomy. Without amateur collectors the majority of the undiscovered species/subspecies will

vanish before they are discovered. Be it butterflies or moon rocks, collecting is the first step of access to all

other scientific information - and protection.

Donations are needed to support and further our efforts to discover and protect

butterflies worldwide. All donations are US tax deductible. Please help generously.

Donations should be mailed to: Treasurer, 126 Wells Rd., Goose Creek, SC 29445.

Checks should be made payable to: TILS. Please indicate if you need an individual receipt.

19

Volume 2

15 December 2000

Number 6

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

A TAXONOMIC STUDY OF, AND KEY TO, THE LECITHOCERIDAE

(LEPIDOPTERA) FROM GUIZHOU, CHINA

CHUNSHENGWU

Institute of Zoology, the Chinese Academy of Sciences

Beijing 100080, China

ABSTRACT. This paper provides a key to twelve species (in ten genera and three subfamilies) of Lecithoceridae

from Guizhou Province, China. Among them, three species are unnamed and eight are new Guizhou Province records. The

female of Opacoptera ecblasta Wu is known for the first time and its genitalia is illustrated for the first time.

Additional key words. Taxonomy, Lepidoptera, Lecithoceridae, fauna, Guizhou

INTRODUCTION

The family Lecithoceridae is widely distributed throughout the world, with approximately 860

known species in over 100 genera. About 90% of the described species are known from the Oriental and

the southern border of the Palaearctic regions. This area extends from southern China to the southern

Himalayas and beyond to the entire Oriental region, with some being distributed in the Mediterranean

subregion, including Asia Minor and southeastern Europe. Another 84 species are known from Australia,

and 73 species from South Africa (Gaede 1937, Clarke 1965, Gozmany 1978, Wu 1997, Park 1999, Wu and

Park 1998-1999).

In China, 46 genera with 219 species in 3 subfamilies have been reported by Wu (1997), and Park

and Wu (1997). Among them, only one species, Quassitagma glabrata Wu and Liu, has been recorded for

Guizhou Province. Guizhou is on the eastern section of the Yunnan-Guizhou Plateau in southwestern China.

This paper gives a key to the 10 genera and 12 species in 3 subfamilies from Guizhou Province. Eight of

these species are new records for Guizhou Province. The female of Opacoptera ecblasta Wu is known for

the first time and its genitalia is illustrated. In addition, the Atrichaozancla sp., Tegenocharis sp., and

Odites sp. are certainly new species. However, I have not yet named them as each is known from only one

male specimen. These descriptions will follow when more specimens become available.

KEY TO GUIZHOU PROVINCE SPECIES

1. Antenna short, less than 2/3 length of forewing.2

Antenna long, more than 3/4 length of forewing .5

2. Fore wing with Cu 2 arising far from lower corner of cell; male genitalia with well developed gnathos .. 3

Forewing with Cu 2 arising from near the lower corner of cell; gnathos lacking or reduced in male

genitalia. Odites sp.

3. Forewing only with one spot on end of cell. Sycthropiodes sp.

Forewing with 2 spots on cell and its end.4

4. Forewing with a spot at anal fold. S. issikii

Forewing without spot at anal fold. S.jiulianae

5. Hindwing without vein M 2 . Atrichozancla sp.

Hindwing with vein M 2 .6

6. Abdominal tergites no spinose.7

Abdominal tergites spinose. 10

7. Forewing with Cui and Cu 2 separate at base.8

Fore wing with Cui and Cu 2 stalked.9

8. Fore wing with spots, M 3 and Cui coincident. Lecitholaxa thiodora

Forewing without spots, M 3 and Cui separate at base. Homaloxestis mucroraphis

9. Forewing with M 2 and M 3 stalked. Quassitagma glabrata

Forewing with M 2 and M 3 separate. Lecithocera palmata

10. Forewing with Cui and Cu 2 stalked. Tor odor a manoconta

Forewing with Cui and Cu 2 separate .11

11. Forewing with M 2 and M 3 coincident. Halolaguna sublaxata

Forewing with M 2 and M 3 separate.12

12. Hindwing with M 3 and Cui coincident. Opacoptera ecblasta

Hindwing with M 3 and Cui stalked . Tegenocharis sp.

Subfamily Lecithocerinae

The subfamily Lecithocerinae is characterized by the male genitalia with a bridge-like structure

connecting the tegumen and the valva, and the uncus almost always vestigal with two lobes at the dorsal

base, only exceptionally united into a broad plate, but never as a thorn or spine.

L Lecithocera palmata Wu and Liu, 1993.

Lecithocerapanmata Wu and Liu, 1993: 332; Wu, 1997: 134.

Material examined: Guizhou, Chishui Co., Jinshagou, Id*, June 2, 2000, Wu Chunsheng.

Distribution: Guizhou, Hainan.

2. Homaloxestis mucroraphis Gozmany, 1978.

Homaloxestis mucroraphis Gozmany, 1978: 71; Wu, 1997: 147.

Material examined: Guizhou, Xishui Co., Sanchahe, 1 9, May 28, 2000, Wu Chunsheng.

3. Quassitagma glabrata Wu and Liu, 1992 (Fig. 1).

Quassitagma glabrata Wu and Liu, 1992: 445; Wu, 1997: 209.

Material examined: Guizhou, Xishui Co., Sanchahe, Id, May 28, 2000, Wu Chunsheng.

Distribution: Guizhou, Jiangxi, Fujian, Hunan, Yunnan.

4. Opacoptera ecblasta Wu, 1996.

Opacoptera ecblasta Wu, 1996: 12; 1997: 157.

Female genitalia (Fig. 3): Eight stemite long and wide, caudal margin almost straight; apophyses

anteriores longer than 1/2 length of apophyses posteriors; antrum funnel-shaped; ductus bursae longer than

corpus bursae, narrow at base, then widening toward corpus bursae; ductus seminalis arising beyond

middle of ductus bursae; corpus bursae with 2 signa, one with 5 big dents, the other densely with minute

spines on upper 1/3.

Material examined: Guizhou, Xishui Co., Sanchahe, 3d, 19, May 28, 2000, Wu Chunsheng.

Distribution: Guizhou, Sichuan.

Remarks: This species is endemic in Southwestern China. The female is known for the first time and

the genitalia is newly illustrated.

5. Lecitholaxa thiodora (Meyrick, 1914) (Fig. 2).

Lecithocera thiodora Meyrick, 1914, Supplta. Ent. 3:51.

Lecithocera leucoceros Meyrick, 1932, Exot. Microlepidopt. 4: 204.

Lecitholaxa thiodora (Meyrick, 1914), Gozmany, 1978: 124; Wu, 1997: 185; Park and Lee, 1999: 123.

Material examined: Guizhou, Xishui Co., Sanchahe, Id, May 28, 2000, Wu Chunsheng.

Distribution: Guizhou, Beijing, Henan, Jiangsu, Zhejiang, jiangxi, Fujian, Hunan, Guangdong, Sichuan,

Hainan, Taiwan; Japan, Korea.

6. Atrichozancla sp.

Wing expanse 10mm. Antenna grayish yellow, with brown annulations. Second segment of labial palpus

grayish white, with brown scales on outer surface of base; 3 rd segment slender, pale grayish yellow. Head,

thorax and tegula grayish white. Forewing long and narrow, apex pointed, termen oblique; ground color

pale grayish yellow, with a large brown spot at end of cell. Hindwing pale grayish yellow.

Male Genitalia (Fig. 5): Gnathos small, basal half wide; lobes of uncus short and wide, bearing hairs;

valva long, broad in basal 2/5, apical 3/5 even to a finely rounded apex, bearing hairs and bristles; sacculus

wide and long; juxta large, bottle shaped, base with a long thorn; aedeagus as long as valva; vesica with a

group of small spines in apical half.

Material examined: Guizhou, Xishui Co., Sanchahe, Id, May 28, 2000, gen. slide no. WZ20004, Wu

Chunsheng.

Distribution: Guizhou.

Remarks: The Genus Atrichozancla Janse includes 2 named species from Africa. The venation of this

species agrees with that of Atrichozancla , but the labial palpus with smooth scales on second segment is

inconsistent with the description of the genus, which bears tufts of rough scales on second segment. The

characters of labial palpus are important in establishing genera in family Lecithoceridae. Thus, this species

may present a new genus and a new species, but it is unnamed and included in Atrichozancla because we

have only one male specimen.

7. Tegenocharis sp.

Wing expanse 13 mm. Antenna pale orange, with annulation on segments. Second segment of labial

palpus relatively narrow, pale yellowish brown; 3 rd segment slender, slightly longer than 2 nd , pale

yellowish brown. Head, thorax, and tegula pale yellowish brown. Forewing yellowish brown, with a silky

luster, no pattern. Hindwing grayish yellow.

3

Male genitalia (Fig. 4): Gnathos small; lobes of uncus short and wide, bearing hairs; valva long, broad

at base, tapering to middle, then even width to a rounded apex, bearing hairs and bristles; sacculus long,

base wide, middle with a row of bristles; juxta shield-shaped, a pair of caudal processes short and pointed;

aedeagus as long as valva; vesica with a spine apically.

Material examined: Guizhou, Chishui Co., Jinshagou, Id, June 2, 2000, gen. slide no. WZ20016, Wu

Chunsheng.

Distribution: Guizhou.

Remarks: Genus Tegenocharis Gozmany contains 2 named species respectively from Nepal and China.

This species is related to T. tenbrans Gozmany from Nepal, but differs from the latter by the valva with a

broad basal half and juxta laterally pointed on caudal margin.

Subfamily Torodorinae

Members of the subfamily Torodorinae have no bridge-like structure in the valva of the male genitalia,

while the uncus is directed caudally and is thorn-like.

1 . Torodora manoconta Wu and Liu, 1994.

Torodora manoconta Wu and Liu, 1994: 164; Wu, 1997: 67.

Material examined: Guizhou, Chishui Co., Jinshagou, 4d, June 2, 2000, Wu Chunsheng; Xishui Co.,

Sanchahe, 1 d, May 28, 2000, Wu Chunsheng.

Distribution: Guizhou, Jiangxi, Yunnan, Taiwan.

2. Halolaguna sublaxata Gozmany, 1978.

Halolaguna sublaxata Gozmany, 1978: 238; Wu, 1997: 90; Park and Lee, 1999: 127.

Material examined: Guizhou, Chishui Co., Jinshagou, Id, June 2, 2000, Wu Chunsheng.

Distribution: Guizhou, Jiangsu Zhejiang; Korea.

Subfamily Oditinae

The genus Odites-g roup, including Scythropiodes , comprises about 150 species in the world. It has

been commonly placed in the family Lecithoceridae, but sometimes placed previously in the Xyloryctidae.

Lvovsky (1996) supported the opinion that the genus Odites and its allied genera belong to Lecithoceridae,

and proposed a new subfamily Oditinae.

The wing venation and the gnathos in the male genitalia of the genus Scythropiodes agree with those of

the Lecithoceridae, but the shorter antennae and the shape of wings more resemble to these of the family

Xyloryctidae, as stated by Gozmany (1978). Park and Wu (1997) placed the Scythropiodes in the family

Lecithoceridae rather than Xyloryctidae.

1. Scythropiodes issikii (Takahashi, 1930).

Depressaria issikii Takahashi, 1930, Kaju gaityu kakuron, 1:285.

Oditesplocamopa Meyrick, 1935: 84; Clarke, 1955: 478.

Odotesperissopis Meyrick, 1936: 27; Clarke, 1955: All.

Odotess issikii (Takahashi); Saito, 1961: 51.

Scythropiodes issikii (Takahashi); Lvovsky, 1996: 650; Park and Wu, 1997: 42.

Material examined: Guizhou, Chishui Co., Jinshagou, 3 d, June 2, 2000, Wu Chunsheng.

Distribution: Guizhou, Liaoning, Beijing, hebei, Shandong, Shaanxi, Zhejiang, Anhui, Hunan, Jiangxi,

Fujian, Guangxi, Yunnan, Sichuan; Japan; Korea.

Hosts: Gardenis jasminioides Ellis, Maluspumila Miller, Pyrus spp., Populus nigra L. Salix spp.,

Smilax china L., Ulmuspavifolia Jacquin, Viburnum awabuki K., and Weigela coreaensis Thunberg.

2. Scythropiodes jiulianae Park and Wu, 1997.

Scythropiodes jiulianae Park and Wu, 1997: 37.

Material examined: Guizhou, Chishui Co., Jinshagou, Id*, June 2, 2000, Wu Chunsheng.

Distribution: Guizhou, Sichuan, Jiangxi.

3. Odites sp.

Wing expanse 20 mm. Antenna yellowish brown, with brown annulations. Second segment of labial

palpus brown on basal 3/4 outer surface, and yellowish white on apical 1/4 and inner surface; 3 rd segment

slender, yellowish white. Face yellowish white; vertex, thorax and tegula yellowish gray. Forewing grayish

yellow, with a dark brown spot at end of cell, another small one at apex; clila long, grayish white;

underside with a row brown spots along termen. Hindwing pale yellow.

Male genitalia (Fig. 6): Valva elliptical, bearing long hairs, with acute apex; basal process with leaf¬

like basal part, and horn-shaped apical part. Transtilla with a pair of long bar bearing a dent apically. Juxta

long, elliptical, with a digitate lobe on caudal margin, and a pair of lateral lobes in middle. Aedeagus

longer than valva, vesica with 2 strong comuti.

Material examined: Guizhou, Chishui Co., Jinshagou, Id, June 2, 2000, gen. slide no. WZ20014, Wu

Chunsheng.

Distribution: Guizhou.

Remarks: Genus Scythropiodes Matsumura was erected by monotypy, based on S. seriatopunctata

Matsumura, 1931, originally placed in the family Yponomeutidae. However, Inoue (1954) included it in the

Gelechiidae, treated it as a junior synonym of Odites Walsingham. Lvovsky (1996) separated the genus

Scythropiodes Matsumura from the genus Odites , and combined 16 previously known species of Odites

into Scythropiodes. The genus Scythropiodes differs from the genus Odites by the forewing with Cu 2

arising far from the lower corner of cell, and male genitalia with well developed downturned gnathos,

which is lacking or reduced in Odites.

This species is similar to Odites notocapna Meyrick in superficial characters, but differs from the

latter by the shapes of valva and juxta.

ACKNOWLEDGMENTS

This project was supported by a grant for systematic and evolutionary biology, Chinese Academy of

Sciences, Beijing.

LITERATURE CITED

Clarke, J.F.G. 1965. Catalogue of the Type Specimens of Microlepidoptera in the British Museum (Natural

History) described by Edward Meyrick, Vol. 5. London: British Museum, pp. 1-255.

Gaede, M. 1937. Gelechiidae. In F Bryk, ed. Lepidopterorum catalogues. Vol. 79. Berlin: Gravenhage, pp.

1-630.

Gozmany, L. 1978. Lecithoceridae. In H.G. Amsel, F. Gregor and H. Reisser, eds. Microlepidoptera

Palaearctica. Vol. 5. Wien: Geore and Co., pp. 1-306.

Park, K.T. 1999. Lecithoceridae (Lepidoptera) of Taiwan (I): Subfamily Lecithocerinae: Genera

Homaloxestis Meyrick and Lecithocera Herrich-Schaffer. Zoological Studies 38 (2): 238-256.

Park, K.T., and C.S. Wu. 1997. Genus Scythropiodes Matsumura in China and Korea (Lepidoptera,

Lecithoceridae), with Description of Seven new species. Ins. Koreana 14: 29-50.

Park, K.T., and S.M. Lee. 1999. A Review of the Lecithocerinae and Torodorinae (Lepidoptera,

Lecithoceridae) in Korea. Ins. Koreana 16 (2): 119-129.

Wu, C.S., and Y.Q. Liu. 1992. Lepidoptera: Lecithoceridae. In Huang Fusheng, ed. Insects of Wuling

Mountains Area, Southwestern China, pp. 445-447.

_. 1993. A Study of the Chinese Lecithocera Herrich-Schaffer, 1853 and Descriptions of

New Species (Lepidoptera: Lecithoceridae). Sinozoologia, 10: 319-346.

Wu, C.S. 1996. Revision of the genus Opacoptera Gozmany (Lepidoptera: Lecithoceridae). Entomologia

Sinica, 3 (1): 9-13.

_. 1997. Lepidoptera Lecithoceridae. In Editorial Committee of Fauna Sinica, Academia

Sinica, ed. Fauna Sinica. Insecta Vol. 7. Beijing: Science Press, 306 pp.

Wu, C.S., and K.T. Park. 1998-1999. A Taxonomic Review of the Family Lecithoceridae (Lepidoptera)

from Sri Lanka. I., Tinea 16 (1): 61-72, 1999; H, Ins. Koreana 15: 1-22, 1998; III., Korean Joum.

Syst. Zool. 15 (1): 1-9, 1999; IV., Insecta Koreana 16 (1): 1-14, 1999; V., Korean Joum. Syst. Zool. 15

6

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

TILS Purpose. TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific

discovery, determination, and documentation, without which there can be no preservation of Lepidoptera.

TILS Motto. As a world community, we can not protect that which we do not know.

The Taxonomic Report is projected for publication at the rate of 10 issues a year. Subscription/

dues for Volume Two are $50 US for domestic and $60 US for overseas subscribers. The subscription year

follows the calendar year. All issues are mailed 1 st class. At the end of each year, subscribers receive that

year’s volume on a record-only compact disc (CDR) for permanent archiving and reproduction for personal

use (i.e. a museum or university may make as many copies as needed in whatever format desired). Non¬

members may receive individual issues in print any time for $10 per issue. Individual issues on CDR to

non-members are $25 per issue post paid. Subscriptions and individual issue orders should be made

payable to TILS; and mailed to: Scott D. Massey, Editor, 126 Wells Road, Goose Creek, SC USA 29445-

3413.

Articles for publication are sought. They may deal with any area of taxonomic research on

Lepidoptera. Before sending a manuscript, simply write TILS at the above address to set up discussion on

how to best handle your research for publication.

Everyday around the world, in jungles and urban areas alike, insect species and subspecies are

becoming extinct. Every year scores of taxa are lost that have not even been scientifically discovered and

documented. Thus, their extinction is unnoticed because their existence is unknown. They are unknown

simply because they have not been collected and systematically identified. Without systematic taxonomy

there is nothing. Without the collection, and exchange of specimens (information) there will be no

systematic taxonomy. Without amateur collectors the majority of the undiscovered species/subspecies will

vanish before they are discovered. Be it butterflies or moon rocks, collecting is the first step of access to all

other scientific information - and protection.

Donations are needed to support and further our efforts to discover and protect

butterflies worldwide. All donations are US tax deductible. Please help generously.

Donations should be mailed to: Treasurer, 126 Wells Rd., Goose Creek, SC 29445.

Checks should be made payable to: TILS. Please indicate if you need an individual receipt.

Volume 2

31 December 2000

Number 7

The Taxonomic Report 1

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

RECORDS FOR THE UTILIZATION OF PRUNUS AS A LARVAL FOODPLANT

BY 71 SPECIES OF LEPIDOPTERA IN NORTHEAST CALIFORNIA

LAURENCE L. CRABTREE

P.O. Box 181, Austin, Nevada 89310, USA

AND

RON LEUSCHNER

1900 John Street, Manhattan Beach, California, 90266, USA

ABSTRACT. Twenty-six sites in five northeastern California counties (Shasta, Modoc, Plumas, Lassen, Tehama)

were surveyed from 1991 - 1999 for the presence of lepidopteran larvae on naturally occurring shrubs of the genus Prunus.

To date, a total of seventy-one species of Lepidoptera from seventeen families have been documented to utilize one or more

of the area’s three Prunus species (P. emarginata, P. subcordata, and P. virginiana var. demissa).

Additional key words. Host plant, rearing, beating method.

INTRODUCTION

There are numerous articles in the lepidopterous literature that discuss the utilization of various

larval foodplants by a specific family, genus, or species of butterfly or moth. However, little has been

published where the focus is on a specific genus or species of plant to determine the extent of its generalized

lepidopterous utilization. We have undertaken such a study and here publish its results 1 .

We selected the genus Prunus L. (Rosaceae) because 1) it is widely distributed in our study area of

northeastern California and 2) was suspected to be used by a significant number of species. It is also often

found growing in thickets, which increases the likelihood that the larvae found on it actually use it as a

foodplant in nature. We selected a five county region in northeastern California for our study area. These

counties are: Shasta, Modoc, Lassen, Plumas, and Tehama.

In this northeastern California region Prunus is represented by three native species of deciduous

shrubs or small trees - P. emarginata (Hook) Walp, P. virginiana L. var. demissa (Nutt) Torrey, P.

subcordata Beuth. These plants grow under a wide range of conditions in this diverse area where the

Cascades merge with the Sierras, the Modoc Plateau, and Great Basin. A forth species, P. andersonii Gray,

also occurs in northeastern California but is restricted to a small, unique area of Lassen County and was not

included in this study.

1 Editor’s note: Host association is an important taxonomic indicator, both specifically and subspecifically. This is because a

shift in host preference (which often offers a habitat shift opportunity) is a precursor to more significant evolutionary shifts.

At TTR we feel that documenting the larval hosts of today will greatly help in constructing the taxonomy of tomorrow.

PROFILE OF STUDY AREA PRUNUS SPECIES

The Prunus in this study were determined to species by the senior author by examining their branch

characteristics, leaves, flowers, and fruits as they progressed through the stages of their seasonal cycle.

Prunus emarginata (Bitter Cherry) is a 4-12 foot shrub with high slim, flexible branches. Its

flowers are white and form rounded clusters of 3-12. The leaves are 0.75 to 2.00 inches long, oblong, and

have finely toothed margins. Its fruit is oval, 0.50 inches in diameter, and turns from red to black when ripe.

It inhabits mountain ridges, moist slopes, and stream banks from 500 to 9,000 feet in the southern California

mountains, Coast Ranges, and Sierra Nevada. It often forms large thickets on damp slopes and in canyons.

Prunus virginiana var. demissa (Western Chokecherry) is usually found in the form of a shrub 3-12

feet high with flexible branches, but occasionally becomes a small tree reaching up to 20 feet high. The

white flowers form racemes, making Chokecherry easy to distinguish from the other area Prunus species.

The leaf-blades are oval, 1.5 to 3.5 inches long, and finely serrate on the margins. The fruits are round, 0.25

to 0.5 inches in diameter, and turn from red to dark purple when mature. It occupies moist sites in the

mountains throughout California, northward to British Columbia, and eastward to Nevada and Arizona. It is

often found with P. emarginata.

Prunus subcordata (Sierra or Klamath Plum) is a shrub or small tree 6-20 feet tall with stiff, short,

thorn-like, crooked branches. Its leaves are ovate, 0.75 to 2.0 inches long, and usually heart shaped at their

base. Its flowers are white or pink in clusters of 2-4. The fruits are oblong, 0.75 to 1.0 inches in length, and

become bright red or yellow when mature. This Prunus inhabits moist or dry foothill rocky slopes in

middle elevations of the Sierra Nevadas north into southern Oregon. It is often found with P. virginiana and

occasionally P. emarginata.

COLLECTION OF IMMATURES AND RESULTS

Methods

The best way to determine the number of lepidopterous species utilizing Prunus is to collect eggs,

larvae, and pupae directly from wild plants and rear them to adults. All other methods are greatly inferior

(e.g. trying to observe oviposition by wild females).

The best technique for gathering immatures is the long employed, but simple, method called

“beating.” Beating is simply the act of suddenly jarring a branch so that resting larvae are loosed from their

grip on the plant and then fall harmlessly onto a light colored cloth beneath the branch. The immatures are

then gathered into rearing containers where they complete their metamorphosis. Beating does little

immediate damage to these shrubs/trees and inflicts no long term harm to them.

From 1991 through 1999 the senior author made many field trips into the northeastern California

study area to gather immature Lepidoptera from indigenous Prunus. Twenty-six separate locations were

established and surveyed for immatures in the five county study area (Table 1). These sites were widely

dispersed to provide the maximum amount of species. Although there was no sampling design, efforts were

made to collect all sites at various times of the year. The vast majority of gathered specimens were larvae

in all stages of development. Occasionally, pupae were also obtained.

Specimens were usually collected mid-morning and mid-afternoon. After a brief visual search for

larvae, the branches of the foodplant were struck with a stick and the larvae were collected from a canvas

sheet being held under the branch. The larvae were then reared on the same species of Prunus from which

they were gathered in small plastic containers or in glass lamp chimneys until pupation. Most of the larvae

were photographed.

Results

This nine year study has shown that the Ihree Prunus species growing in northeastern California are

utilized as food plants by a large and diverse assemblage of Lepidoptera. Seventy-one species representing

seventeen lepidopteran families (Table 2) were collected from and reared to adults on these native Prunus.

One hundred and nineteen foodplant records were documented by this project. Most of these were new food

plant records. Sixty-six of the specimens represent new county records for northeastern California.

Approximately half of the Lepidoptera recorded in this study utilized more than one species of

Prunus. Thirty-six of the 71 species (51%) were recorded on two or more species of Prunus. Ten of these

were recorded on all three of the Prunus hosts studied.

Thirty-three of the 71 species (46%) were recorded on only one Prunus hostplant. These host

specific species were divided almost equally between the three species of Prunus , indicating that none of

the hostplants were more (or less) important as a larval substrate. Eight species (23%) were found only on P.

emarginata , thirteen species (38%) were found only on P. virginiana , and fourteen species (40%) occurred

only on P. subcordata.

As a way to evaluate the completeness of this study, published hostplant information for three of the

best studied families of Lepidoptera (Papilionidae, Lycaenidae, and Nymphalidae) was compared to our

results. Six species from these families are known to utilize Prunus as a larval host. We found five of these

during random collecting for this study and reared them to adults. The larvae of Satyrium titus occidentalis

(Austin & Emmel, 1998), an uncommon butterfly also know to feed on Prunus , was not recorded. However,

Deciduphagus augustinus iroides (Boisduval, 1852) was discovered feeding on both P. emarginata and P.

subcordata which were previously unrecorded hostplants for this taxon.

This evaluation method suggests that this study recorded most of the common Lepidoptera utilizing

Prunus as a larval hostplant. Collecting larvae at night, at additional locations, and with improved rearing

techniques would undoubtedly result in several additions to these records from the study area. The 230

reared adults are deposited in the Essig Museum of Entomology, University of California, Berkeley.

3

| TABLE 2. Prunus host(s) utilized by Lepidoptera species. Prunus abbreviations and location numbers from Table 1.

ACKNOWLEDGMENTS

We thank the following for their taxonomic identifications: Dr. Ronald Hodges, Chionodes thoraceochrella ; Dr.

James Tuttle, Sphingidae larvae; Ron Robertson, Oncocnemis species; Dr. Paul Tuskes, Hemileuca larvae; Dr. Jean-

Francois Landry, Coleophora irroratella; Debora Matthews Lott, Pterophorids; Dr. Jerry Powell, Acleris aenigmana.

We thank Ron Gatrelle for his encouragement and patience in editing assistance. We also thank Denice and Timothy

Crabtree for their many hours of diligently searching for larvae, feeding them and keeping records.

Bibliography

Austin, G. &J.F. Emmel. 1998. A Review of Papilio multicaudatus Kirby (Lepidoptera: Papilionidae). Systematics

of Western North American Butterflies. Gainesville, FI: Mariposa Press. Pp. 691-700.

Covell, C.V. 1984. A Field Guide to the Moths of Eastern North America. Boston, MA: Houghton Mif flin Co.

Crabtree, L.L. 1998. Discovering the Butterflies of Lassen Volcanic National Park, Mineral, CA: Lassen Loomis

Museum Assoc.

Hickman, J.C. Ed. 1993. The Jepson Manual: Higher Plants of California. Berk. & L.A., CA: Univ. of CA Press.

Hodges, R.W., et al. 1983. Checklist of the Lepidoptera of America North of Mexico. London: Curwen Press.

Tietz, H.M. 1972. An Index to Described Life Histories, Early Stages and Hosts of the Macrolepidoptera of the

United States and Canada. Vol. III. Allyn Museum of Entomology, Sarasota, Florida.

Figures 1-17. Sample of study area species. Site numbers from table one. Fig. 1. Orgyia cana, site 1. Fig. 2. Nematocampa

brehmeata, site 25. Fig. 3. Acronicta mansueta, site 5. Fig. 4. Schizura unicornis conspecta, site 25. Fig. 5. Sparganothis

senecionana, site 5. Fig. 6. Dasyfidonia avuncularia, site 10. Fig. 7. Hesperumia fumosaria impensa, site 25. Fig. 8.

Hesperumia sulphuraria, site 26. Fig. 9. Drasteria adumbrata, site 14. Fig. 10. Drasteria strechii, site 4. Fig. 11.

Leptarctia californiae, site 1. Fig. 12. Itame umbriferata, site 15. Fig. 13. Hesperumia latipennis, site 14. Figs. 14.

Erannis tiliaria vancouverensis, site 7. Fig. 15. Synaxis hirsutaria, site 5. Fig. 16. Synaxis cervinaria, site 4. Fig. 17.

Synaxis barnesii, site 21. All photos by L. Crabtree. Specimens enlarged to 1.20 natural size.

6

Editor’s note. A reviewer offered these comments: The beating method of collecting misses about half of

the Lepidoptera fauna. The few micro’s listed in this paper are mainly external feeders (and leaf-rollers),

which do sometimes come out on beating, but all the leaf-miners, of which there will be lots, and stem

borers, are completely missed by the beating method. Some of the Leps listed have never had their host

plants recorded before, some are known already but are restricted to Prunus (and often Salix also) and

others feed on almost anything. Some type of code as to which are new, restricted, and general feeders

would be useful in this type of paper. The paper’s OK as is.. .it does have good info in it!

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

TILS Purpose. TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific

discovery, determination, and documentation, without which there can be no preservation of Lepidoptera.

TILS Motto. As a world community, we can not protect that which we do not know.

The Taxonomic Report is projected for publication at the rate of 8-10 issues

Volume Three are $45 US for domestic and $55 US for overseas subscribers. The

calendar year. All issues are mailed 1 st class. At the end of each year, subscribers rec

year. Subscription/ dues for

Volume 2

31 December 2000*

Number 8

The Taxonomic Report

NINE NEW SPECIES OF LACINIPOLIA (NOCTUIDAE)

FROM ARIZONA, CALIFORNIA AND VICINITY

CHARLES L. SELMAN

626 Grub Road, Patriot, Ohio,45658 USA

AND

RON LEUSCHNER

1900 John Street, Manhattan Beach, California, 90266, USA

ABSTRACT. This article formalizes the taxonomic conclusions of the senior author’s 1975 successful doctorate

dissertation. The nine new species defined in that 1975 dissertation are here formally described in compliance with the rules

of the International Code of Zoological Nomenclature (ICZN). The authors have essentially abstracted the original 447 page

dissertation and designated types. The new Lacinipolia are: delongi, aileenae, triplehorni, bucketti, baueri, sharonae,

martini, fordi, and franclemonti. The purpose of this paper is simply to make these names available to the scientific

community and professional researchers by making them ICZN compliant.

Additional key words. Original dissertation, revision of names, Colorado, Oregon, New Mexico, Texas.

BACKGROUND INFORMATION

In the early 1960’s. Lloyd Martin, then Curator of Lepidoptera at the Los Angeles County Museum of

Natural History (LACM), began a study of the Noctuid genus Lacinipolia. In this study, he identified a

number of new species that had been collected in the Southwest and Pacific Coast. At that time he proposed

changes to the nomenclature of existing species in this genus. Unfortunately, this work was not completed

when he retired and subsequently moved to Prescott, Arizona in 1969.

In the 1970’s the senior author was a doctoral candidate at Ohio State University. Selman obtained

the loan of Martin’s specimens and genitalic slides for the purpose of completing this revision. Selman’s

doctoral dissertation on the genus Lacinipolia was completed in 1975, leading to the granting of a Ph.D.

This 447 page document, with numerous drawings and photographs, was too long for regular Journals, and

no other method of publication was found. Thus, the information contained has only been available on

microfilm reproduction, which does not satisfy ICZN requirements for the publication and validation of

taxonomic names.

The lack of properly described names for these easily recognized new species has long been a

source of frustration to those who prepare regional lists and Lepidopterists’ Society Season Summary

inputs. To resolve this problem at long last, the junior author has separated the new species descriptions

from dissertation and prepared them for formal publication in this paper. A certain amount of revision was

required to make each description stand by itself without reference to other parts of the original study not

contained here. The designation of paratypes has extended the range of some species beyond that given in

the original paper. All type photographs have been re-done as the originals were unsuitable. Since all types

were returned to the LACM and are available to the junior author, this was not a problem.

The authors both desire to dedicate this paper to the memory of Lloyd Martin, who provided

encouragement and collecting companionship to so many people, and who provided the original study

which is the basis of this paper. The manuscript names chosen by Selman have been retained to avoid

confusion by those who have long been familiar with these taxa based on the original manuscript. The

synonymies and name status changes contained in the revision are not included here. These would require a

much more extensive paper, and would thus further delay publication. It is the hope of both authors that a

revision of the entire genus may some day be published.

Deposition of type specimens referred to in this paper are: LACM: Los Angeles County Museum of

Natural History; USNM: National Museum of Natural History, Smithsonian Institution, Washington, DC.

THE GENUS LACINIPOLIA

In 1937, McDunnough erected the genus Lacinipolia for a group of related Hadenine species within

the genus Polia , based on male genitalic characters. In his diagnosis, McDunnough states that these species

possess "...a large, very hairy membraneous flap, lightly attached to the median section of the valves near

the ventral margin. This flap apparently takes the place of the peniculus, which is well-developed in the true

Polia's but practically wanting in this group."

The genotype of Lacinipolia is Mamestra illaudabilis Grote, 1875. In Poole's 1989 Noctuid

catalog, 76 species are listed, confined to North, Central and South America. Of these, 58 are found in

North America (MONA, 1983 plus Leuschner, 1992). One additional species was described by Mustelin in

2000; this paper adds nine more, so the above totals are increased by ten to 86 and 68 respectively.

Figures 1-10. New Lacinipolia species. Fig. 1. Paratype c?: L. delongi. Fig. 2. Paratype d : L. aileenae. Fig. 3.

Holotype d : L. triplehorni. Fig. 4. Allotype ?: L. triplehorni. Fig. 5. Paratype d : L. bucketti. Fig. 6. Paratype d : L.

baueri. Fig. 7. Paratype d : L. sharonae. Fig. 8. Paratype d : L. fordi. Fig. 9. Paratype d : L. franclemonti. Fig. 10.

Paratype d : L. martini. All photographs by R. Leuschner.

NEW SPECIES DESCRIPTIONS

Lacinipolia delongi Selman and Leuschner, new species.

Description. Male (Fig. 1). Head : front, gray; palpi, smoky with inner edge and third segment paler; antennae,

serrate. Thorax and abdomen : light gray, concolorous; collar with faint traverse band subapically; many dark scales of

the thorax are silvery tipped. Forewing : concolorous with thorax and markings somewhat distinct; basal line obsolete,

antemedial (AM) line faint, postmedial (PM) line bent outwards around reniform, then straight to inner margin,

denticulate on veins; subterminal line vague, marked by pale scales; reniform upright, constricted in the middle, and

black outlined; orbicular oblique; claviform long, extending from AM to PM lines, concolorous. Hindwing’, dull smoky,

paler at base with concolorous fringes. Female. Similar to male but darker and more contrasting.

Male genitalia. As shown in Figure 11.

Types. All: ARIZONA. Holotype d: Santa Cruz County, Madera Canyon, Santa Rita Mts., 1770 m., 22 June

1955, leg. Lloyd M. Martin. Genitalia on slide 61-4, L.M.M. Allotype 9: Cochise County, Upper Camp, Pinery

Canyon, Chiricauhua Mts., 26 June 1955, leg. Lloyd M. Martin. Genitalia on slide 503 L.M.M. Paratypes: 4d d, 19.

2d d, same data as holotype, leg. L.M. Martin & W. Rees; Id, Yavapai County, Prescott, 14 June 1970, leg. R.

Leuschner; Id, Coconino County, Oak Creek Canyon, 11 July 1988, leg. R. Leuschner. 19Cochise Co., Pinery

Canyon, 2075 m., 13 August 1964, leg. R. Leuschner. The holotype, allotype, and 1 paratype are in the LACM. One

paratype in R. Leuschner collection.

Diagnosis. Superficially similar to L. prognata, but males have distinctly serrate antennae, while prognata has

simple or weakly serrate antennae. The orbicular is oblique in delongi and rounded in prognata; delongi has a basal

dash while prognata has none.

Distribution. L. delongi is known only from Arizona.

Etymology. This taxon is named for Dr. Dwight M. Delong, in recognition of his help during the four years

leading to the senior author’s doctorate.

Lacinipolia aileenae Selman & Leuschner, new species.

Description. Male (Fig. 2). Head: palpi, fuscous, much lighter on inner lateral sides; front, luteous with dark bar

extending between the eyes; antennae, simple. Thorax : admixture of gray, silvery and yellowish scales. Collar : same

color as thorax, with trace of a transverse median band. Forewing : ground color same as thorax, basal line barely

traceable, denticulate, antemedial line geminate, pale filled, nearly straight from costa to orbicular, then curved under

obicular to inner margin, with an outward tooth at vein 2A; postmedial line geminate, pale filled, sinuate, the outer line

obsolescent on upper half, curved inward below reniform; reniform black outlined, incomplete at top, with contrasting

pale filling, not constricted at middle; orbicular oblique, with pale filling; claviform long, black outlined, extending from

AM to PM line; basal dash long and distinct; subterminal area light below apex to vein Cu2, a dark patch fills the

tornal area from vein Cu2 to vein 2A, from PM line to outer margin. Hindwing : sordid, gradually paler toward base,

veins contrastingly dark; fringe is cream with median wavy band. Female. Similar to male, but darker in coloration,

causing markings to be more contrasting.

Male genitalia. As shown in Figure 12.

Types. All: ARIZONA, Cochise Co. Holotype d: South Fork Cave Creek, Chiricauhua Mts., 23 May 1962, leg.

L.M. Martin, genitalia on slide 570 L.M.M. Allotype 9: same location as holotype, 21 May 1962, genitalia on slide

572 L.M.M. Paratypes : 4dd. 3dd, same location and data as holotype; Id, Cochise Co., 28 April 1992, leg. N.

McFarland. The holotype and allotype are deposited in the LACM. Paratypes in LACM and R. Leuschner.

Diagnosis: The horizontal black mark near the tomus of the fore wing is quite distinctive, and separates aileenae

from all other Lacinipolia known to occur in western North America.

Distribution. L. aileenae is known only from southern Arizona.

Etymology. L. aileenae is named for the senior author’s mother, whose financial and moral support through

eight years of college were much appreciated.

3

Lacinipolia triplehorni Selman & Leuschner, new species.

Description. Male (Fig. 3). Head : palpi, yellowish brown; antennae, simple, pubescent. Front, thorax and

abdomen : yellowish brown with only a slight sprinkling of a few dark hairs. Collar, similar, with trace of a transverse

median band. Forewings : concolorous with body, marking hardly traceable. Hindwing s: nearly immaculate with only a

trace of yellowish brown scales along outer margin and veins. Female (Fig. 4). Much darker than male with most lines

traceable. Forewings : subterminal line is quite wavy; postmedial line is sinuate, but does not touch the reniform.

Hindwings: white with yellowish brown along outer margin and veins.

Male genitalia. As shown in Figure 13.

Female genitalia. As shown in Figure 21.

Types. All: ARIZONA. Holotype d: Pima Co., Babonquivari Mts., 15 May 1924 (no collector), genitalia on

slide 639 L.M.M. Allotype ?: Santa Cruz Co., Madera Canyon, 13 September 1950, leg. H.R. Reid, C.W. Kirkwood,

genitalia on slide 642 L.M.M. Paratypes : 34 specimens. 7, same data as holotype; 15, same data as holotype except

(2) 1-15 June 1924, (3) 1-15 September 1923, (10) 15-30 September 1923, leg. O.C. Poling; 2, same as allotype; 2,

Santa Cruz Co., Pena Blanca, Oro Blanca Mts., 21-27 September 1963. leg. L. Martin; 5, Cochise Co., Ash Canyon,

(3) 23-30 September 1990, leg. N. McFarland, (2) 28 April 1992, leg R. Leuschner; 1, Portal, Chiricahua Mts., 3

June 1964, leg. M. Cazier; 2, Pueblo del Sol. Huachuca Mts., 12 May 1982, leg. R. Wielgus. Holotype and Allotype

in LACM; Paratypes in LACM, USNM, and personal collection of R. Leuschner.

Diagnosis. L. triplehorni is similar to L. erecta, but the male is much lighter - so light that the maculation is

hardly traceable. The female of L. erecta has a less sinuate subterminal line, and the postmedial line is quite close or

touches the lower edge of the reniform, where in triplehorni it is distinctly separated. In the male genitalia, the clasper

of triplehorni is large and robust, where in erecta the clasper is thin and narrow.

Distribution. L. triplehorni is known only from Arizona.

Etymology. This species is named for Dr. Charles A. Triplehom, the senior author’s graduate advisor.

Lacinipolia bucketti Selman & Leuschner, new species.

Description. Male (Fig. 5). Head : palpi, ash gray with third segment lightest; front: ash gray with dark band

extending between the eyes; antennae, simple; Thorax and abdomen : light gray with many scales silvery tipped.

Collar: ash gray with narrow transverse median band. Forewings : light gray with most lines geminate, pale filled,

traceable but not contrasting; basal line present; antemedial line oblique to vein Cu-M, then outwardly convex to inner

margin with a slight tooth on vein 2A; postmedial line sinuate, denticulate; reniform with dark outline and light silvery

filling; orbicular oblique, roundish, with dark outline but filling not contrasting; claviform concolorous with dark

outline; subterminal line indicated by a difference in shade only. The median area of the wing is darkest, while basal

and subterminal area are silvery, basal dash short but distinct; fringe concolorous. Hindwings: uniformly dull luteous,

with a lunule at end of discal cell; veins slightly darker; fringe light with a trace of transverse median band. Female.

Slightly darker overall than male, but median area not contrastingly dark; reniform filled with ground color, not silvery.

Male genitalia. As shown in Figure 14.

Types. Holotype d: CALIFORNIA: Mendocino Co., 4 miles E. of Point Arena, 5 July 1958, leg. W.R. Bauer,

J.S. Buckett, genitalia on slide 61-410 L.M.M. Allotype ?: CALIFORNIA: same location and collector as holotype,

18 July 1958, genitalia on slide 772 L.M.M. Paratypes : 40 specimens. CALIFORNIA: 1, same data as holotype; 2,

Mendocino Co., Ft. Bragg, 24 August 1973, Leg. R. Leuschner; 6, Del Norte CO., Crescent City, 11 August 1971and

23 August 1962, leg. Leuschner; 21, Humboldt Co., Areata, Beach Pine Woods, 8-19 July 1972, leg. J.W. Johnson;

OREGON: 10, Curry Co., Humbug Mtn. State Park,22 July 1962, leg. R. Leuschner. The holotype and allotype are in

the LACM; Paratypes in LACM and personal collection of R. Leuschner.

Diagnosis. L. bucketti is quite similar to L. olivacea, but has silvery terminal area where olivacea does not.

From L. sharonae, which also has a silvery subterminal area, bucketti is separated by being smaller with a median

area that is not as dark. In the male genitalia, the clasper of bucketti is tapered to a point, while in sharonae it is

uniformly thick throughout its length.

Distribution. L. bucketti has been found in coastal areas

i of northern California and Oregon.

Etymology. L. bucketti is named for Steve Buckett, who collected the types and made significant contributions

to the study of western Noctuidae.

Lacinipolia baueri Selman & Leuschner, new species.

Description. Male (Fig. 6). Head : front & palpi, light reddish brown; antennae, simple. Thorax and abdomen :

light reddish brown; collar, light brown with trace of median band. Forewings : light reddish brown with ordinary lines

indicated by shades only - no dark scales, basal line not traceable, antemedial line nearly straight to vein 2A, then

inwardly oblique, slightly toothed on some veins; postmedial line sinuate for its entire length, slightly denticulate;

subterminal line wavy; reniform upright and indicated largely by the paler filling; orbicular and claviform obsolescent;

fringe concolorous. Hindwings : same shade as forewing but slightly paler toward the base. Female. Slightly darker

than the male causing maculation to be more contrasting; reniform more prominent, with white outline; orbicular

slightly traceable; basal line stronger and wavy.

Male genitalia. As shown in figure 15.

Types. All: CALIFORNIA. Holotype d : Marin Co., Point Reyes peninsula, McClure Beach, 15 June 1960, leg.

J.S. Buckett, genitalia on slide 61-411 L.M.M. Allotype 9: same location as holotype, 17 June 1958, leg. W.R. Bauer

& J.S. Buckett, genitalia on slide 770 L.M.M. Paratypes : 6 specimens. 1, same location as holotype 15 June 1955; 4,

same location as holotype, 12 July 1956, leg. R. Leuschner; 1, San Luis Obispo Co., Cambria Pines, 20 July 1962, leg.

R. Leuschner. Holotype and allotype are in the LACM; paratypes in LACM and collection of R. Leuschner.

Diagnosis. Although the forewing maculation of L. baueri is quite similar to several other species in the genus

(e.g. L. olivacea), the light reddish brown coloration of the forewings, with the lack of any black scales on the

ordinary lines, will easily distinguish it.

Distribution. L. baueri is found in coastal California from Marin Co. south to San Luis Obispo Co. in isolated

conclaves of coastal pines and cypress.

Etymology. L. baueri is named for William R. Bauer of Petaluma, CA who, with Steve Buckett, made important

collections of moths from coastal and northern California, and was a collecting companion of the junior author.

Lacinipolia sharonae Selman & Leuschner, new species.

Description. Male (Fig. 7). Head : front, ash gray with some silvery tipped hairs; palpi, dark ash gray, only

slightly lighter inside laterally, but tip of third segment distinctly lightest; antennae, simple, pubescent. Thorax and

abdomen: ash gray with some hairs silvery tipped; collar, basal half with yellowish tinge, above with silvery transverse

median band. Forewings : concolorous with thorax with overall silvery tinge, lines geminate and pale filled; basal line

barely traceable, wavy; antemedial line starts one-third along costa from base, is quite outwardly oblique to vein Cu2,

then makes a convex curve to mid-point of inner margin; the median area is darker than the rest of the wing; reniform

distinct, contrastingly light, a rounded vertical oblong with thin lines in the center; obicular oval with dark outline but

not paler than rest of median area; postmedial line sinuate, first outwardly convex, then concave to inner margin;

beyond the median area there is silvery white with dark-lined veins in most specimens, with this silvery area extending

to the outer margin in some examples, while in others, it is confined between the PM and subterminal lines; claviform

concolorous, dark outlined; basal dash short, broad; fringe concolorous. Hindwings : sordid, only slightly lighter

toward base, with veins slightly darker; fringe concolorous with traceable median line. Female. Forewings : distinctly

darker than the male, differing in that the median area is lighter, not darker, than the rest of the wing; the reniform has

a dark filling followed by pale, then dark, outlines. Hindwings : slightly darker with the basal half of the fringe sordid.

Male genitalia. As shown in figure 16.

Types. All: ARIZONA. Holotype d: Gila Co., Christopher Creek, Mogollon Rim, 1770 m., 22 June 1957, leg.

L.M. Martin, R.J. Ford & W.A. Rees, genitalia on slide 61-72 L.M.M. Allotype 9: same location as holotype, 27 June

1957, genitalia on slide 682 L.M.M. Paratypes : 7dd, 29 9. 2 dd, same data as holotype; 3 dd, 19, Apache Co.,

Greer, White Mts., 2530 m., 26 June 1968, leg. R. Leuschner; Id, Cochise Co., Barefoot Park, 2440 m., 1 July

1992, leg. K.M. Leuschner; Id, Cochise Co., Rustler Park, 2590 m., 16 July 1998, leg. R. Leuschner & K. Richers;

1 9, Pima Co., Summerhaven, Catalina Mts., 2500 m., 1 June 1997, leg R. Leuschner. The Holotype and allotype are

in the LACM; paratypes are in the LACM and R. Leuschner collection.

Diagnosis. L. sharonae is similar to L. olivacea and davena. The highly oblique AM line with a sharp curve to

the inner margin separates sharonae from both of these, where the AM line is greatly rounded. L sharonae is larger

(27-29 mm wingspan) than olivacea (23-26 mm). L davena has a large light red-brown patch at the tornus which is

replaced by a more extensive silvery white area in sharonae.

Distribution. Thus far, L. sharonae is known only from northern Arizona at elevations above 1700 meters.

Etymology. L sharonae is dedicated to the senior author’s wife, Sharon A. Selman, whose support was vital to

this study.

Lacinipolia fordi Selman & Leuschner, new species.

Description. Male (Fig. 8). Head’, front, with a dark spot near each eye; palpi, an admixture of cream, brownish

and black scales, slightly lighter inside; antennae, simple and pubescent. Thorax and abdomen : blackish brown with

some light gray-tipped scales; collar with a black band across the top. Forewings: ground color blackish brown like

thorax with silvery tinge; basal line obsolescent, other lines geminate; antemedial line concave on upper half, convex

on lower; postmedian line straight, not denticulate; subterminal line broadly waved, indicated by a few pale scales but

more so by the difference in the shades of the terminal and subterminal areas, and marked with white scales above

anal angle; subterminal area lightest, the silvery tinge especially noticeable; reniform upright, black outlined, silvery

filled; orbicular roundish with incomplete black outline, somewhat filled; claviform concolorous, with irregular outline;

basal dash indistinct; fringe concolorous. Hindwings: white with scattered sordid scales along outer margin and some

veins; fringe white, with trace of a dark line. Female. Forewings: darker than male. Hindwings: as in male. The white

of the subterminal line at the anal angle and filling on lower portion of postmedial line contrast markedly against the

darker background.

Male Genitalia. As shown in figure 17.

Female Genitalia. As shown in figure 22.

Types. Holotype d: ARIZONA: Gila Co., Christopher Creek, Mogollon Rim, 1770 m., 17 June 1957, leg. L.M.

Martin, R.J. Ford & W.A. Rees, genitalia on slide 61-74 L.M.M. Allotype 9: ARIZONA: Coconino Co., Parks, 2165

m., 27 June 1957, leg. L.M. Martin, R.J. Ford, W.A. Rees, genitalia on slide 763 L.M.M. Paratypes: 56 specimens.

ARIZONA: 1, same data as holotype; 41, same data as allotype; 6, Coconino Co., Fort Meadows, 9 July 1959, leg

W.A. Hammer; COLORADO: 7, Teller Co., Florissant, Big Springs Ranch, 19 June - 8 July 1960, 1962, leg. T.C.

Emmel. The holotype, allotype and paratypes are in the LACM.

Diagnosis. L. fordi and L. franclemonti are dark brown species unlike any other species in the genus in North

America. L. fordi lacks the reddish brown suffusion found on the forewings of L. franclemonti.

Distribution. L. fordi is found at higher elevations in Arizona and Colorado.

Etymology. L. fordi is named for Robert Ford, a frequent collecting companion of Lloyd Martin on trips to

Arizona, including the 1957 trip when the types were collected.

Lacinipolia franclemonti Selman and Leuschner, new species.

Description. Male (fig. 9). Head: palpi, uniformly dark ash; front, concolorous with palpi, no trace of typical

dark spot near the eyes; antennae, simple, somewhat bristled. Thorax: dark ash but with overall reddish-brown tinge;

collar, with narrow transverse band. Forewing: color like thorax but reddish-brown tinge more prominent, especially in

subterminal area; most lines blurred but traceable, geminate; basal line obsolescent; antemedial line straight, deeply

denticulate, brownish filled; postmedial line dark, slightly convex around reniform, then oblique to inner margin,

followed by brown outline, denticulate; subterminal line traceable as a difference in shades, broadly waved, dentate on

some veins, ending at anal angle; some veins in subterminal area marked with black scales; reniform upright, kidney

shaped, pale followed by dark thin outline, reddish-brown filled; orbicular oval, oblique, otherwise similar to reniform;

claviform obsolescent; fringe concolorous with terminal area. Hindwing: white with scattered sordid scales along outer

margin and veins; fringe white, with fragments of a medial band. Female. Forewings: slightly darker than male.

Hindwings: white, with sordid scales at least on distal third, pale at base; fringe with distinct median band.

Male Genitalia. As shown in Figure 18.

6

Types. All: ARIZONA: Holotype d : Gila Co., Christopher Creek, Mogollon Rim, 1770 m., 22 June 1957, leg.

L.M. Martin, R.J. Ford, & W.A. Rees, genitalia on slide 61-77 L.M.M. Allotype 9: same as holotype, 18 June 1957,

genitalia on slide 766 L.M.M. Paratypes: 94 specimens. 93, same data as holotype except 17-22 June 1957; 1,

Coconino Co., Oak Creek Canyon, 1525 m., 13 June 1970, leg. R. Leuschner. Holotype and Allotype in LACM.

Paratypes in LACM and collection of R. Leuschner.

Diagnosis. L. franclemonti is similar only to L. fordi in the genus, but franclemonti has a reddish brown

suffusion which is lacking in fordi.

Distribution. L. franclemonti has only been found at higher elevations above 1500 m. in Arizona thus far.

Etymology. This new species is named for Dr. J.G. Franclemont, a leading authority on the Noctuidae who

provided help to the senior author on numerous occasions.

Lacinipolia martini Selman and Leuschner, new species.

Description. Male (Fig. 10). Head: front, luteous with dark spot near each eye; palpi, cream suffused with

blackish scales on outer lateral sides; antennae, simple, pubescent. Thorax: admixture of cream, blackish and brown

scales; collar, with prominent black transverse median band. Abdomen: darker brown-black than the thorax.

Forewing: heavily suffused with dark scales with a reddish brown tinge; antemedial line not noticeably dentate except

on vein 2A, outwardly oblique from costa to Cu2, then inwardly oblique to inner margin; postmedial line nearly

straight for its length, beginning near apex and running obliquely to the inner margin, distinctly dentate; basal line

obsolete; reniform upright, not constricted, pale filled; orbicular oval, oblique, pale filled; claviform concolorous, with

dark outline; fringe concolorous. Hindwing: immaculate. Female. Forewings : more contrasting than male.

Male Genitalia. The apex of the aedaeagus (base of vesica) is shown in figure 19.

Female Genitalia. As shown in Figure 23.

Types. Holotype d: ARIZONA: Pima Co., Baboquivari Mts., 23 April 1938, leg. J.A. Comstock, genitalia on

slide 61-398 L.M.M. Allotype 9: ARIZONA: Gila Co., Jones Water Camp nr. Seneca, 18 May 1961, leg. R. Reid &

W. Rees, genitalia on slide 749 L.M.M. Paratypes : 293 specimens. ARIZONA: Pima Co.: 2, same site as holotype,

April and November; 9, Sells P.O., 15 April - 10 May 1923; 78, same site as holotype, 1 Sept. - 30 November 1923;

7, Catalina Mts., Peppersauce Cyn., 4 June 1935, leg. J.A. Comstock. Gila Co.: 11, same data as Allotype; 2, Payson,

E. Verde R., 26 Oct. 1959, leg. L. Martin, & F. Truxal; 2, Mogollon Rim, 26 June 1957, leg. J.Comstock & W. Rees;

1, Sierra Anche Exp. Sta., 3 June 1956, leg. R. Leuschner & C. Hill. Santa Cruz Co.: 97, Madera Cyn., April -

September, leg. L. Martin, C. Kirkwood, T. Davies & R. Leuschner; 9, Oro Blanco Mts., Pena Blanca, 27 May

1963, leg. L. Martin; Maricopa Co.: 3, Wickenburg, 1 June 1959, leg. K. Stange. Cochise Co.: 36, Chiricahua Mts,

Cave Creek, May - Sept., leg. Martin, Kirkwood & R. Leuschner; 2, Willcox, 10 May 1993, leg. K.M. Leuschner, 7

Oct. 1955, leg. Martin; 3, Dragoon Mts., Cochise Stronghold, 11 Sept. 1958, leg. Menke & Stange; 2, Portal, 3 June

1964, leg. R. Leuschner; 2, Huachuca Mts., Pueblo del Sol, 12 May 1986, leg. R. Wielgus; 1, Ash Canyon, 5 May

1984, leg. Leuschner & McFarland. NEW MEXICO: Eddy Co.: 1, Whites City, 1160 m., 6 Sept.1982, leg. R.

Leuschner; 1, Dona Ana Co.: Las Cruces, 3 May 1992, leg. R. Leuschner. TEXAS: Randall Co.: 5, Palo Duro Cyn.,

11 May 1961, leg. L. Martin. Jefferson Davis Co.: 18, Davis Mts. St. Pk., 14 May 1962, leg. L. Martin & 8 May

1993, Leg. R. Leuschner; 1, Alpine, 7-15 May 1926. The holotype and allotype are in the LACM; paratypes are in

the LACM, NMNH and personal collection of R. Leuschner.

Diagnosis. The base of the vesica of L. martini differs from that of L .quadrilineata Grote (Fig. 20) in that

martini has one diverticulum, and the spine at the tip of the aedaeagus is only slightly longer than the aedaeagus is

wide. In quadrilineata, there are two diverticula and the spine is twice as long. The forewing of male quadrilineata

has the basal area (out to the diagonally sloping AM line) noticeably paler than the median area, where in martini the

basal area is only slightly lighter or the same shade as the median area.

Distribution. L. martini occurs throughout Arizona, and ranges through New Mexico into West Texas,

replacing L. quadrilineata which is confined to California and the Pacific Coast.

Etymology. This species is named for the late Lloyd M. Martin, who made the original study of the genus

Lacinipolia and first recognized all the new species described in this paper.

LITERATURE CITED

GROTE, A.R. 1873. Descriptions of Noctuidae Principally From California. Bull. Buffalo Soc. Nat. Hist.

Vol. I, pp. 129-155 (P.140, desc. of L. quadrilineata ).

_. 1875. Preliminary List of the Noctuidae of California. Canad. Ent., Vol.7, Pp.25-28 (P.27,

desc. of L. illaudabilis ).

HODGES, R.W. et al. 1983. Checklist of the Lepidoptera of America North of Mexico. E.W. Classey, Ltd.,

London. 284 pp.

LEUSCHNER, R. 1992. An Overlooked Record of Lacinipolia rodora (Noctuidae) from the United States.

J. Lepid. Soc., Vol. 46, pp. 79-80.

McDUNNOUGH, J. 1937. Notes on North American Noctuid Genera. Canad. Ent., Vol. 69, pp. 40-47.

MORRISON, H.K. 1874. Descriptions of New Noctuidae. Proc. Boston Soc. Nat. Hist., Vol. 17, pp.131-

166 (P. 143, desc. of L. olivacea).

MUSTELIN, T., R.H. LEUSCHNER, K. MIKKOLA, & J.D. LAFONTAINE. 2000. Two New Genera and

Thirteen New Species of Owlet Moths (Noctuidae) Mainly From Southern California. Proc. San Diego

Soc. Nat. Hist., No. 36. 18 pp.

POOLE, R.W. 1989. Lepidopterum Catalogus (new series), Fascicle 118: Noctuidae. E.J. Brill, Leiden,

The Netherlands. 1314 pp. (in three volumes).

SELMAN, C.L. 1975. Revision of the Genus Lacinipolia McDunnough of America North of Mexico

(Lepidoptera: Noctuidae). The Ohio State University, Ph.D. in Entomology, 1975.

SMITH, J.B. 1891. Contributions Toward a Monograph of the Noctuidae of Temporate North America:

Revision of the Species of Mamestra. Proc. USNM, Vol. 14, pp. 197-276 (many now in Lacinipolia).

__. 1901. Notes on Mamestra olivacea Morr. and Its Allies. Trans. Am. Ent. Soc. Vol. 27, pp.

230-240 (P. 237, desc. of L. davena ).

* The actual publication date of this issue is 12 January 2001.

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

TILS Purpose. TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific

discovery, determination, and documentation, without which there can be no preservation of Lepidoptera.

TILS Motto. As a world community, we can not protect that which we do not know.

The Taxonomic Report is projected for publication at the rate of 8-10 issues a year. Subscription/ dues for

Volume Three are $45 US for domestic and $55 US for overseas subscribers. The subscription year follows the

calendar year. All issues are mailed 1 st class. At the end of each year, subscribers receive that year’s volume on a

record-only compact disc (CDR) for permanent archiving and reproduction for personal use (i.e. a museum or

university may make as many copies as needed in whatever format desired). Non-members may receive individual

issues in print any time for $10 per issue. Individual issues on CDR to non-members are $25 per issue post paid.

Subscriptions and individual issue orders should be made payable to TILS; and mailed to: Treasurer, 126 Wells

Road, Goose Creek, SC USA 29445-3413.

Articles for publication are sought. They may deal with any area of taxonomic research on Lepidoptera.

Before sending a manuscript, simply write TILS at the above address (C/O Editor) to set up discussion on how to

best handle your research for publication.

9