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

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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

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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.

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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

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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.

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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!

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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.

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