Volume 1

1 October 1998

Number 1

f V ■? The Taxonomic Report "■' + '

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

AN ADDENDUM TO ANTHOCHARIS MIDEA DOS PASSOS AND KLOTS 1969.

(DESCRIPTION OF A NEW SUBSPECIES FROM TEXAS.)

RONALD R. GATRELLE 1

126 Wells Road, Goose Creek, South Carolina 29445

ABSTRACT. This paper verifies and expands upon the information presented by dos Passos and Klots in their 1969

systematic revision of Anthocharis midea. Their restriction of the range of A. midea midea to the coastal islands and

immediate coast of Georgia and South Carolina is confirmed and expanded north into southern coastal North Carolina. Their

range of A. m. annickae is expanded southward from Virginia to central Georgia. A new subspecies, A. midea texana, is

described from Texas. Texana is annickae’s ancestor. It differs from annickae only in having greatly reduced black scaling

at the base of the dorsal wings. Klots’ theory that midea midea is descended from an extinct Florida ancestor is confirmed.

Midea (coastal) and annickae (Sandhills) occur within 40 km of each other in Georgia and South Carolina. Midea is

confirmed to not only be characterized by expanded orange of the dorsal FW of males, but also by frequent orange on the

dorsal HW apex of males, and by females with frequent orange scaling on the dorsal FW apex.

Additional key words: Biogeographical evolution, convergent evolution.

C. F. dos Passos and A. B. Klots (1969) surveyed the phenetic variation of Anthocharis midea

(Hubner) throughout its range. This present article is an addendum to their research, and merely confirms

and expands their original findings. The sum of their research revealed the existence of four regionally

distinct phenotypic populations of A. midea as follows:

1) along the coast of Georgia and South Carolina, characterized by extensive dorsal orange apical

patches in males. They restricted the nominate subspecies, A. m. midea, to this area.

2) from Massachusetts to Virginia, characterized by males with a small orange apical patch. This

they named as a new subspecies A. midea annickae.

3) Texas, also characterized by restricted male orange apical patches. They believed these to be

indistinguishable from annickae in phenotype, but not synonymous with that subspecies.

4) remainder of species range, polytypic and thus characterized by no defining character(s).

Nearly 30 years have passed since their research was published. Having lived for the last 29 years

in the area of the type locality of A. midea midea, I have been provided with the unique opportunity to

compare hundreds of typical A. midea midea from various sights in Charleston, Colleton, Beaufort, and

Jasper counties, South Carolina, and Chatham County, Georgia, against dos Passos and Klots’ observations.

Further, I have collected/observed hundreds of midea from inland Georgia and South Carolina, which has

revealed significant information regarding the subspecies annickae and its relation to midea midea. I have

also examined typical annickae from New Jersey and series of midea from Missouri, Texas, Kansas,

Mississippi and other states. These specimens provide a clear picture of not only midea’s subspeciation,

but of its biogeographical evolution.

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

FIGS. 1-8. Anthocharis midea subspecies. 1, holotype d Anthocharis midea texana, 16 March 1989, 6 mi. w. of Buffalo,

Freestone Co., Texas. 2, d A. m. annickae 2 May 1971 Lakehurst, NJ. 3, d A. m. annickae 11 March 1997 Burke Co., GA.

4, e A. m. midea 29 March 1975 Edisto Island, Colleton Co., SC. 5, allotype 9 A. m. texana 16 March 1989, 6 mi. w. of

Buffalo, Freestone Co., Texas. 6, 9 A. m. annickae 1 May 1971 Lakehurst, NJ. 7, 9 A. m. annickae 10 April 1992 Oconee

Co., SC. 8, 9 A. m. midea 29 March 1975 Edisto Island, Colleton Co., SC.

REEXAMINATION OF MIDEA MIDEA AND MIDEA ANNICKAE

In their research, dos Passos and Klots were only able to examine a very small number of male

specimens of A. m. midea from coastal Georgia (23) and South Carolina (13). They did not mention how

many coastal females were examined, but it was surely less than the number of males. Nonetheless, they

were able to document the three characters that separate midea midea from the other populations.

The foremost character was the larger size of the orange patch in the apical area of male fore wings.

The remaining two characters were noted, but their significance underrated due to the limited series they

examined. On their page 10 they stated, “It may be noted that in this region the males often show tinges,

sometimes strong, of yellow about the apex of the hindwing and that females sometimes show a tinge

of yellow about the apex of the forewing. This is the population to which the names A. genutia

(Fabricius), midea (Hiibner), Iherminieri (Godart) and flavida Skinner must be applied” (Emphasis mine).

The several hundred midea midea specimens I have collected/observed over the years confirm that

approximately 25 to 50% of males (Fig. 4) possess the yellow HW apical character and about 25 to 30% of

females have yellow scales in their FW apical area (Fig. 8). (% varies from colony to colony.) These two

characters should also be considered as primary diagnostic traits along with the broad orange apical patch

in males. Further, these three diagnostic characters of midea midea are of great significance because it is

now known that they are genetic and not environmentally induced ecoforms.

We know this because the A. midea populations only a few kilometers inland in Screven, Burke, and

Richmond counties, Georgia, and Aiken, Barnwell, Allendale, Bamberg, and Orangeburg counties, South

Carolina are identical to northeastern annickae from New Jersey and are referable to that subspecies.

These Georgia and South Carolina A. m. annickae populations never possess the yellow apex in females

and very rarely the yellow dusting on HW apexes of males. It is also rare in these Sandhill counties to find

any males with midea midea- like expanded apical orange areas.

Several years ago, Mr. Bob Cavanaugh informed me that this same situation existed in southern

North Carolina. South coastal North Carolina specimens were A. m. midea and specimens only 30-40

kilometers inland were clearly A. m. annickae.

Dos Passos and Klots examined no specimens from North Carolina, inland South Carolina, or from

the Sandhills of Georgia. This enormous geographical “gap” in material led them to the false assumption

that a dine existed from south to north - from midea to annickae. This is not the case. No clinal variation

exists along the eastern US seaboard. (Isolated pockets of genetic mixture may exist in north coastal NC

and southern VA through intersubspecific contact.)

Dos Passos and Klots limited the range of A. midea midea to the coastal islands and immediate

coast of Georgia and South Carolina. I extend this to include the south coast of North Carolina in the range

of A. m. midea

They described annickae from a mere 8 specimens from West Rock, New Haven, Connecticut and

limited the range of A m. annickae to Massachusetts south through Virginia. Having examined large numbers

of A. m. annickae from the above mentioned Georgia and South Carolina counties, and also Lancaster

(boarders North Carolina) and Oconee (mountains) counties, South Carolina; I herein extend the range of A.

midea annickae from New England south to at least the Sandhill counties of eastern Georgia.

REEXAMINATION OF TEXAS MIDEA

Concerning the midea populations in Texas, dos Passos and Klots stated on page 10, “The material

from Texas shows a very different picture. The 49 specimens examined come from a wide range (Dallas,

Harris, Harrison, Brazos, Bexar, Kerr, Comal, San Patricio, and Smith counties). Among them there are no

specimens with very extensive [orange FW] patch (groups A & B), and by far the largest group (38/49 = 77

%) has the patch greatly reduced. This is all the more surprising because of the dominance of large-patched

populations in most of the southern and western range of the species. It would be difficult, in fact, to find

any consistent points of difference between the series from Texas and those from the most distant

northeastern part of the species’ range in New Jersey, New York and Connecticut.”

And on page 17, “The great phenetic similarity of the populations in Texas and in the northeast

(annickae) is by no means evidence that they are genetically so similar that they should be considered

subspecifically congruent. To do so would, in fact, contravene everything that is now known about the

evolutionary differentiation of populations on the specific and subspecific level during periods of spatial

isolation from each other.”

I have examined 50 Texas midea (40 ## & 10 9 9) from Freestone, Comal, and Smith counties.

These range over a distance of 500 km. from northeast to south central Texas. These specimens confirm the

observations of dos Passos and Klots in relation to the small size of the male apical orange patch.

However, all 50 examples differ definitively from A. m. annickae in having much less black scaling at the

base of the dorsal wings. I have also examined a small series of 10 specimens (7 d d\ 3 9 9) from Johnson

and Douglas counties in eastern Kansas which match this Texas phenotype.

The amount of black scales at the base of the wings is a diagnostic subspecific character between A.

sara sara Lucas and/l. sara flora W. G. Wright (Layberry, Hall and Lafontaine 1998). It is also diagnostic

for species in the related genus Euchloe. Accordingly, I therefore describe these Texas/prairie populations

as a new subspecies.

Anthocharis midea texana Gatrelle, new subspecies

Description and diagnosis. Male (Fig. 1): In all phenotypic aspects as in subspecies annickae except that the

black scaling at the base of the dorsal wings adjoining the abdomen is generally 50 % or less that of annickae. Female (Fig.

5) as in male but black basal dusting even more restricted somet im es almost immaculate white at base of wings.

Types. Holotype S (Fig. 1): 16 March 1989, 6 mi. w. of Buffalo, Freestone Co., Texas. Allotype 9 (Fig. 5): 16

March 1989, 6 mi. w. of Buffalo, Freestone Co., Texas. Paratypes: 39 <J <J, 9 9 9: all TEXAS: FREESTONE COUNTY, 6

mi. w. of Buffalo, 1 <J, 13 March 1986; 4 <j <j, 3 9 9,16 March 1989. SMITH COUNTY, Tyler St. Pk., 3 d d, 14-15 March,

4 6 d\ 18-19 March 1987. COMAL COUNTY, Landa Pk., New Braunfels, 2 9 9,8 March, 4 d cJ, 1 9, 20 March 1982; 20

dd*, 1 9, 14 March 1983; 3 rfd*, 23 April 1988; 2 9 9,3 April 1995. The Holotype, Allotype, and 6 d* paratypes are

deposited in the Florida State Collection of Arthropods, Gainesville, Florida. 6 rf paratypes are deposited in the American

Museum of Natural History, New York, with the remaining paratypes deposited in the author’s collection.

Etymology. Texana is named for the great state of Texas.

Remarks. The range of texana needs to be worked out by local collectors/researchers in the region. It apparently

does not occur in Missouri. However, Missouri specimens tend to have the restricted basal black scaling at the juncture of

the wings with the abdomen, and thus resemble texana in this trait more then annickae or midea. Texana does occur in the

Lawrence area of Kansas, then south and westward. Colonies in Louisiana should be closely documented and observed since

they occupy the tension zone between the convergent phenotype in Mississippi and Texan texana.

BIOGEOGRAPHICAL EVOLUTION

The evolutionary cause for the odd geographical phenotypic dispersal of midea was a mystery dos

Passos and Klots left unanswered. However, they were not far from answering this question. They stated on

page 10, “The concentration of large-patched individuals in coastal South Carolina, Georgia, and

Mississippi and also in the northwestern part of the range of the species, presents a special problem. It may

very well be, as one of us has postulated (Klots, 1965, p 462-463) that the southeastern coastal plain large-

patch characteristic arose in peninsular Florida during the Pleistocene when, due to changes in ocean level,

this area was an island separated from the mainland; and that the character later spread both northeastward

and northwestward, chiefly along the coast, but not into Texas, while the refugium population died out in

Florida. The small-patch Texas population would then be the descendants of a Pleistocene population in a

different refugium, perhaps in Texas itself and Mexico. We are unable to surmise how or where the

northwestern (i.e., Missouri) large-patch character arose.

“Northward from Georgia and South Carolina in the Coastal Plain and Piedmont the populations

show a sharp diminution of the amount of orange. Unfortunately, adequate material from northern South

Carolina and North Carolina is lacking.”

Now that adequate material has been examined from Georgia and South Carolina, the solution seems

fairly clear. Midea midea and annickae are from different ancestors. While annickae and texana have the

same ancestor.

First, we now know that midea and annickae exist subspecifically in nature only a few kilometers

apart in the coastal areas of Georgia, South Carolina and south coastal North Carolina. In fact, they

probably occur in adjacent counties in this region (i.e. Beaufort and Hampton counties SC). They parallel

each other along the coast for several hundred kilometers.

Second, climate and environment play no part in the coloration of these subspecies because

specimens from Burke County, Georgia look exactly like specimens from Connecticut. While specimens

from Chatham County, Georgia are vividly distinct from both. These phenotypes are genetic not

environmental.

I believe Klots was exactly correct in postulating that midea midea ascended from an isolated

Florida population. He was also accurate in stating that at the same geological time (the Pleistocene)

texana was an isolate in Mexico. What I believe occurred was that the western population colonized itself

eastward along the then existent coast (today’s Black Belt Prairies of Mississippi and Alabama, and the

Sandhills of Georgia, South Carolina and North Carolina), while midea was still isolated in island Florida.

This eastward movement is substantiated by the research of Dr. Richard L. Brown of Mississippi State

University. He has documented the eastward movement, and subsequent influence, of western species of

Lepidoptera into and on the Lepidoptera of the Black Belt region (personal communication).

After Florida was joined to the mainland, those large-patch midea which moved northwestward

(unhindered by habitat barriers) met the texana/annickae phenotype which was already present in that area.

This gave rise to the mixed phenotypes north and northwest of the Florida panhandle. Those midea which

moved northeastward became isolated on the coastal islands while annickae was isolated in the Sandhills.

Before colonial times, these two southeastern subspecies were kept apart in the region by thick maritime

forest swamps which were the dominant feature of the mainland coast.

Today, midea midea and midea annickae are becoming accessible to each other as the result of 300

years of deforestation and human environmental alteration. The convergent evolution we see taking place in

the mixed phenotype populations (which now occupies most of the species range in the south and lower

Midwest) will eventually cause the extinction of midea midea. Over the last 30 years I have observed

midea further and further inland and annickae closer and closer to the coast. There may be undiscovered

colonies in Jasper and Hampton Counties, SC, which are now only separated by 20 km or less.

Biogeographical evolution is a slow process. We seem to usually think in terms of divergence, but

convergence is a product of evolution also. Without the intervention of unforeseen isolating factors, the day

will surely come when midea will be one polytypic species.

LITERATURE CITED

DOS PASSOS, C.F. & A.B. KLOTS. 1969. The Systematics of Anthocharis midea Hubner (Lepidoptera:

Pieridae). Entomologica America. Vol. 45: 34 pp.

LAYBERRY, ROSS A., PETER W. HALL & J. DONALD LAFONTAINE. 1998. The Butterflies of

Canada. Univ. of Toronto Press, Toronto, Canada 279 pp.

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

126 Wells Road, Goose Creek, SC 29445-3413

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Volume 1

1 November 1998

Number 2

%i|| The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY ^ **

THE REDISCOVERY, TAXONOMY, AND BIOLOGY OF CHLOSYNE

GORGONE GORGONE AND CHLOSYNE ISMERIA (NYMPHALIDAE)

IN BURKE COUNTY, GEORGIA.

RONALD R. GATRELLE 1

ABSTRACT. On 24

;ofC.

TYPE LOCALITIES AND HISTORICAL INFORMATION

Chlosyne gorgone gorgone (Hubner) was described in 1810 from a John Abbot painting (Fig. 1)

without any text. In the absence of a text, and because the figures of the type were produced by Abbot, it has

traditionally been presumed that the Burke County area of Georgia is the type locality of C. gorgone.

Chlosyne ismeria (Boisduval and LeConte) was described in 1833 from a poor copy of an Abbot painting

(Fig. 4) and was accompanied by a brief textual annotation. The annotation established the range of ismeria

as “Carolina and Georgia.” The traditional assumption is that “coastal” Georgia and South Carolina is

meant. The range of ismeria may actually be wider once its status in nature is better understood. Further,

Burke County, Georgia has been established throughout the historical literature as the type locality of

FIGS. 1-7. 1, HUbner’s d Dryas reticulata Gorgone, dorsal and ventral surfaces from his plate 41. 2, Neotype d Dryas

reticulata Gorgone, 27 April 1993 Burke Co., GA, River Rd., dorsal and ventral surfaces. 3, Topotype 9 Chlosyne gorgone

gorgone , 21 April 1994 Burke Co., GA, River Road, dorsal and ventral surfaces. 4, Boisduval and LeConte’s 9 Melitaea

ismeria, dorsal and ventral surfaces from their plate 46. 5, d C. g. carlota, 22 April 1989 Oconee Co., SC, Brasstown Creek

Rd., dorsal and ventral surfaces (leg. Watson). 6, 9 C. g. carlota, 25 June 1988 Oconee Co., SC, Rt. 107 S. of Oconee St. Park,

dorsal and ventral surfaces (leg. Watson). 7, Neotype d Melitaea ismeria, 20 August 1989, Burke County, GA, at Savannah

River, dorsal and ventral surfaces. All specimens collected by R. Gatrelle unless otherwise noted.

I have not found any valid historical records for either of these species from or near their type

locality since their discovery - a period of about 180 years. Harris (1972) lists several records of C.

gorgone from various inland and upland Georgia counties. Burke County is 180 km from Harris’ nearest

record.

Forbes (1960) under gorgone , sites personal knowledge of “a few males in the British Museum

[NH] from Abbot”. His text structure is unclear, but the statement seems to refer to ismeria. Dr. Ian James

Kitching of the British Museum (NH) kindly searched their holdings and found no Georgia specimens of C.

gorgone, C. ismeria or Chlosyne nycteis (Doubleday and Hewitson). Nor did they find any Abbot

specimens in the BMNH. Thus, Forbes statement is in error.

Opler (1984) has a dot on his range map for C. gorgone along the coast of South Carolina. After

communicating with Dr. Opler on this matter, I contacted the individual from whom he received this record.

This individual informed me that the record was in error, and he had not taken nor observed C. gorgone at

any South Carolina location. Dr. Opler’s record is also in error.

F. Martin Brown (1974) in number 16 of the Bulletin of the Allyn Museum , provides the singular

most extensive historical and thorough taxonomic treatment of C. gorgone, C. gorgone carlota (Reakirt),

and C. ismeria. He too found no existent coastal Georgia specimens of either C. gorgone or C. ismeria.

REDISCOVERY OF BURKE COUNTY CHLOSYNE

On 24 April 1993 I stopped to fix a flat tire on an Orangeburg County, South Carolina road.

(Localities will remain vague due to the rarity of species concerned.) While fixing the tire, a non-

lepidopterist friend, Scott Massey, who occasionally accompanies me and helps with collecting, caught a

butterfly by the car and asked what it was. It was a Chlosyne gorgonel By chance we had stopped in the

middle of a flourishing colony of C. gorgone. I had been actively collecting in South Carolina for 23 years,

and this was my first encounter with this long sought after species.

Both sexes were numerous and fresh. A small series was collected. Adults were nectaring at the

diminutive Coreopsis auriculata L. The habitat at the Orangeburg locality reminded me of an area I had

collected a few years earlier in Burke County, Georgia.

On 27 April 1993, I made a trip to this location in Burke County, and upon finding a patch of C

auriculata, parked the car. Within a minute of exiting the vehicle, I had rediscovered this rare butterfly at

its type locality. 200 years had passed since John Abbot had found this butterfly in this same general area

and it was still there! While C. gorgone was not as numerous here as in Orangeburg County, the suitable

habitat was more extensive and continuous. Consequently, C. gorgone was more dispersed at the Burke

County site.

The rediscovery of the insect named Melitaea ismeria by Boisduval and LeConte was much less

dramatic but more significant.

On 20 August 1989 at about 1800 h, while observing several species of butterflies gathered at mud

along the banks of the Savannah River, I noted that among them were some fresh males of Chlosyne nycteis.

Having found nycteis in many states over the years, I initially gave it no thought. In fact, earlier that day I

had been collecting in the mountains of Georgia where nycteis had been a virtual pest. Remembering I was

in Burke County, it occurred to me that this species may be unusual for that part of the state. So I netted three

as vouchers. These nycteis were not only a new record for Burke County, but for southeastern Georgia, and

over 140 km from the nearest record for this species in Harris (1972).

These sat unexamined (I only mounted one) in my collection from 1989 until 1997. By then I had

rediscovered gorgone and was putting together a paper on it. One day while working on gorgone, I was

struck by the obvious - Abbot had found two species of what we now call Chlosyne in Burke County. I had

also found two. Could these two be Abbot’s gorgone and ismerial I herein propose that they are.

TAXONOMY OF THESE SPECIES

Brown’s aforementioned paper is a detailed and definitive work on the literary history and

taxonomy of the species which we are examining. While Brown’s work is foundational to this present

paper, I will only mention herein a few key quotations and points, and leave it to the reader to search out the

full record.

Melitaea ismeria Boisduval and LeConte

Brown demonstrated that the modem misuse of ismeria being synonymous with gorgone was the

result of an identification error by Scudder and A. G. Butler subsequently communicated to W. H. Edwards

and others. On page 2 Brown states, “Scudder’s concept [of ismeria = gorgone ] is untenable.” On page 3

he says, “Careful study of the use of the name ismeria indicates that its acceptance as the equivalent of

gorgone male hinges on Scudder’s statement about the [faulty] manuscript plate of Abbot in the British

Museum (N. H.).” Then in Brown’s concluding statements on page 10 he unequivocally states, “In

conclusion, I believe that the name ismeria Blv. & LeC. cannot be used in association with gorgone

Huebner.”

Brown’s determination was that while in much of the literature ismeria and gorgone have been

associated as the same species, in nature they are not.

Brown’s quandary was not in determining what ismeria wasn’t, that is, not gorgone , but in what it

was. To Brown, ismeria was most likely either Chlosyne harrisii (Scudder) or C. nycteis. His personal

leaning was toward harrisii, chiefly because of his preoccupation with one feature, the orange margin of the

ventral wings. Consider the following quotes from Brown in reference to the original painting of ismeria.

On page 2, “On the underside the most striking feature is broad orange brown margins on both wings, much

as on harrisii Scudder.” On page 3, “It cannot be considered even a crude representation of either of the

insects called gorgone by Huebner. It can be considered a cmde representation of the butterfly called

harrisii by Scudder, but this insect is not now found in Georgia. The larvae figured... does

3

not conform to the mature larva of car lota (gorgone male Huebner). It does suggest the mature larvae of

harrisii... [or] early-stage larvae of... nycteis.” On page 10, “Again the evidence, poor as it is, points

toward harrisii (or nycteis) rather than carlota [ = gorgone] being the closest known taxon to ismeria.”

Brown’s conclusion on ismeria was that he considered it a nomen incognitum.

However, now that it is known that two Chlosyne species are living today in Burke County,

Georgia, two of Brown’s statements stand out. On page 8, “Taxonomically there are two, possibly three,

species involved.” On page 3, “There is no way today to judge the accuracy of Abbot’s representations in

Boisduval and LeConte’s book, except by comparison with known butterflies from [Georgian] America.”

(Emphasis mine.) The comparison can now be made.

The consideration of the following facts and comparison of species characters clearly reveals just

what ismeria was and is.

1) Brown established ismeria as a valid (but unidentified) species separate from gorgone and

postulated that it could well be C. nycteis.

2) Abbot found two species of Chlosyne in Burke County, Georgia. 200 years later there are still

two (and only two) species of Chlosyne in Burke County.

3) One of these species has always been readily identifiable as C. gorgone. The logical

probability is that the remaining species is the poorly depicted, but valid, ismeria.

4) The range of C. harrisii is hundreds of miles to the north of Burke County.

5) C. nycteis, as the only other Chlosyne in the eastern United States, is the only other species

available for consideration as ismeria.

6) Abbot stated that his ismeria fed on sunflower, and depicted it on Helianthus strumosus L. ( =

trachelifolius Miller). This plant occurs in Burke county, and is a known host for nycteis.

7) The only known host (Opler 1984) of harrisii , Aster umhellatus Miller, is not known in eastern

Georgia. Aster umbellatus is know from South Carolina only as variation brevisquamus Fernald and in

only a few extreme northeast counties adjacent to North Carolina (Radford 1968). Thus, harrisii should be

eliminated from consideration.

8) Abbot stated that his ismeria pupated on May 16 and eclosed on May 26. This coincides with

the first brood records of nycteis in Georgia.

9) Burke County gorgone emerge from mid to late April, a full month before ismeria.

10) Lastly, Burke County specimens of nycteis possess all the major phenotypic characters of the

original painting of ismeria and twice as many of these characters than does harrisii. They differ from

northern specimens of nycteis and C. harrisii as detailed in Table I.

While characters F and G on the original ismeria painting match in size harrisii better than Burke

County nycteis (=ismeria), it must be remembered that the painting is a copy and a very poor rendition.

Which means we should first be looking for the presence of a character and secondarily at the accuracy of

the size and shape of the character. What is important is that in nature, Burke County nycteis (= ismeria) do

possess all characters A - G.

There is one other character on the Boisduval and LeConte plate of ismeria which is of interest.

That is the complete row of silver spots along the margin of the hind wing. This picture is of a female. As

yet we have no topotypical females for comparison. However, I do have a female nycteis from Spartanburg

County, South Carolina with silver spots all along its hind wing margin. It is very probable that as we go

further south and to the coast this character becomes more pronounced and more frequent, especially for

females.

TABLE I

Character

C. i. ismeria

C. i nycteis

C, h. harrisii

(A) Dorsal hindwing black

postmedian line

Narrow across the entire

area

Often wide to very wide

Usually only partial, but very

variable

(B) Dorsal forewing sub¬

marginal spots

Well developed to ¥i size of

postmedian spots

Often just dots, usually less

then Va size of postmedian

Variable, dots to well devel-

(C) Eyespots on apex of

ventral forewing

3-4 well defined pupiled

eyespots on light to medium

brown background

3 - 4 eyespots less defined

on dark background.

Usually no eyespots, if so,

only one or two and weak

(D) Inner three fourths of

ventral forewing

Nearly solid light orange,

light striations

Medium orange striated

with brown lines.

Orange-red, well striated

with black-brown lines

(E) Inner two thirds of

ventral hindwing

Light buff dominant, some¬

what silvered, usually with

medium to dark brown stria-

Same as ismeria , but tend¬

ing to less silvered and

brown striations lighter.

Dark, contrasting, orange-

brown and white checkered

(F) Ventral hindwing sub¬

marginal black spots

Surrounded by some narrow

orange in dark field.

Nearly always in dark field

In broad orange band in dark

field

(G) Ventral margins

Orange, narrow, clearly de-

Mainly cream or buff, may

have some orange.

Orange, wide, well defined

It should also be remembered that my Burke County specimens are from the fall brood. The spring

brood (which we know Abbot reared and perhaps his figures were made from) should be expected to differ

in appearance and may more fully fit the form painted.

Based on the above, C. nycteis is synonymous with C. ismeria in Burke County, Georgia.

Accordingly, I have designated a male (Fig. 7) taken 20 August 1989 along the Savannah River, Burke

County, Georgia as neotype of Melitaea ismeria Boisduval and LeConte (1833). This specimen is

deposited in the Allyn Museum of Entomology, Sarasota Florida and is labeled: NEOTYPE, Melitaea

ismeria Boisduval and LeConte, Det. R. R. Gatrelle. One male topotype is also deposited in the Allyn

Museum. The remaining male is in the authors collection.

The new nomenclature is:

Chlosyne ismeria (Boisduval and LeConte) MELITAEA . (1833)

a. i. ismeria (Boisduval and LeConte) MELITAEA. (1833)

b. i. nycteis (Doubleday and Hewitson) MELITAEA . (1847) [NEW COMBINATION]

c. i. drusius (W. H. Edwards) PHYCIODES. (1884) [NEW COMBINATION]

d. i. reversa (F. and R. Chermock) MELITAEA. (1940) [NEW COMBINATION]

Having made only one attempt to collect this insect, I am sure this subspecies, while virtually

uncollected, is not that rare in nature. It should be looked for along watercourses in southern Georgia and

South Carolina. Harris’ record of nycteis from Laurens County, Georgia, is likely C. ismeria also. The

nycteis I have from Spartanburg County tend to have well defined narrow orange boarders also. So the final

decision as to the range of Chlosyne ismeria ismeria and C. ismeria nycteis remains to be worked out. For

now, I think it best to limit the range of C. ismeria ismeria to the coastal plain of Georgia and southeastern

South Carolina.

Chlosyne gorgone (Hubner)

John Abbot’s painting of C. gorgone published by Hubner is excellent. In it we see a dark, boldly

marked gorgone phenotype. Its two most distinguishing characters are the lack of a white pupil in the dark

submarginal spot in cell M 3 , both dorsally and ventrally , and the restricted white chevrons on the margin of

the ventral hind wing. We now know that this is in fact exactly what typical gorgone gorgone looks like in

nature (Figs. 2, 3). I have designated a male (Fig. 2) taken 27 April 1993 Burke County, Georgia as

neotype. It is labeled: NEOTYPE, Dryas reticulata gorgone Hlibner, Det. R. R. Gatrelle. It has been

deposited in the Allyn Museum of Entomology, Sarasota, Florida.

C. gorgone carlota is a valid subspecies. Ventrally, car lota is generally lighter, nearly always has

a white pupil in the submarginal black spot in cell M 3 , both dorsally and ventrally (especially in females),

and bolder marginal chevrons. Its range is primarily Midwestern, northern and western. A number of

gorgone have been collected in upland Georgia and South Carolina and most workers who have seen them

have rightly concluded that in appearance they are so close to Midwestern carlota as to be the same

subspecies (Figs. 5, 6). Thus the range of subspecies carlota extends into at least the upper piedmont of

Georgia and South Carolina.

The existence of carlota in north Georgia has caused most modem workers to wrongly sink carlota

into synonymy. Why is this the case?

In spite of the accuracy of the original depiction of gorgone , subsequent writers of popular butterfly

literature have consistently transposed the ismeria phenotype (large pale washed out below more well

marked above) onto the name gorgone. In other words, their term was “gorgone” but their concept was

“ismeria.” This false concept of gorgone has also affected the concept and taxonomic validity of carlota.

Two conceptual errors have become orthodoxy. 1) The name gorgone is used but the concept is

ismeria and 2) the name carlota is no longer used because the concept is gorgone.

As pointed out by Brown, this conceptual confusion all began with the misidentification and

misassociation of C. ismeria with the totally dissimilar species C. gorgone by Scudder and A. G. Butler

who in turn relayed this error to W. H. Edwards, and he to others. Since Edwards, many have seemed to

just “take the word” of the previous worker and perpetuated the error. The treatment of C. gorgone by

Opler (1984) and Scott (1986) are examples of this continuing “hearsay” taxonomy. (It appears that neither

Scott nor Opler were aware of Brown’s 1974 paper.) Howe (1975) is one modern writer to have the

taxonomy correct.

Opler defined ismeria as a large pale (dominant in coastal Georgia?) form of gorgone. Then, by

not mentioning carlota, demonstrated that he considered carlota and gorgone to be phenotypically the

same. He held both conceptual errors.

Scott took Scudder’s error to its logical conclusion. He only mentioned gorgone, demonstrating that

his concept of carlota was gorgone and his concept of ismeria was as a form of gorgone. He held both

conceptual errors.

Howe correctly treated C. gorgone as a valid subspecies limited to the coast of Georgia and C. g.

carlota as the inland and westward subspecies. He did this correctly, but probably for the wrong reason.

Howe seemed under the impression that ismeria and gorgone were synonymous and did not mention

ismeria for that reason. If this is correct, he also associated the painting of ismeria as being what coastal

Georgia gorgone looked like. In other words, conceptually, his gorgone was still phenotype ismeria. He

held only one of the conceptual errors. So he had the right taxonomy, but part of the old concept.

There may or may not be some avoidable errors here. But by far, the primary, most unavoidable and

problematic factor has been that up until now there were no actual topotypes in existence of either C.

gorgone or C. ismeria.

As mentioned above, gorgone gorgone varies slightly, but consistently, from gorgone carlota in

phenotype. These differences alone are enough to validate subspecific status of both entities. However, my

attempts to rear nominate gorgone revealed that it is strongly univoltine. I have attempted to rear

approximately 800 larvae and have only kept two out of diapause. (Larvae were subjected to the usually

very effective long photoperiod (continuous light) method of breaking diapause. Roughly 600 were from the

Orangeburg sight. The two kept out of diapause were also from the Orangeburg site.) I also sent a few

larvae to Dr. Tom Allen of the West Virginia DNR (in connection with his book on W.Va. butterflies) and

6

returned to their

he too was unable to keep larvae out of diapause or break it. (All diapausing larvae were

place of origin and securely released back into the population.)

Adults were found at both locations in April of 1993 and 1994. No adults were ever seen in

repeated visits to these areas in those years at the appropriate time for subsequent flights of adults. And

perhaps more importantly (since larvae are easier to find than adults) no other immatures were ever found

later in those years.

C. g. carlota is multiple brooded throughout its range, including Canada (Catling and Layberry

1998), upstate Georgia (Harris) and Pickens County, South Carolina (Dr. Charles Watson, personal

communication). Thus, one would expect the populations of gorgone at its southern extreme to be

multibrooded - especially if we were dealing with clinal subspeciation. But it is only single brooded. I

consider this difference in genetic biology to be the strongest reason to consider gorgone as a distinct

subspecies.

It is also significant that in rearing Canadian gorgone carlota, Catling and Layberry found that H.

divaricatus L. was rejected as a host. At both sites where gorgone gorgone was found, intensive searching

located larvae and ova on H. divaricatus only.

Gorgone biogeographical evolution

The habitat of C. gorgone in Burke County is xeric, turkey oak, rolling sandhill in the vicinity of the

Georgia Power Nuclear Facility. These areas in Burke County change very little from year to year, and

slowly decade to decade. Thus, species in this type of habitat have less of a need to continually be

colonizing new areas. Multiple broods are obviously advantageous in species relocation dynamics. (Fire is

a medium factor here in species survival strategy.)

The habitat at the Orangeburg County site is open mixed hardwood and some pine in hilly sandhill

dispersed amid wetter areas and agricultural fields. This habitat may vary greatly decade to decade.

Lepidoptera in these areas need to be more flexible for relocation purposes. (Fire is a major survival

factor here.) However, larvae of gorgone from both sites were virtually impossible to keep out of

diapause.

Thus, both colonies are genetically linked by a common ancestor. An ancestor which evolved into a

univoltine subspecies to insure its survival due to hostile environmental conditions (lack of surface water?).

Both of these locations are usually very hot and dry in June and July. While the larval host, H. divaricatus ,

is leafed out at that time, conditions are nearly always such that acceptable nectar sources for adults are

virtually nonexistent.

I believe C. gorgone gorgone is a remnant of a biogeological relic whose univoltine reproductive

cycle developed under the climatic pressure of an annual dry season in Florida when it was an island. It is

pertinent to note that Falcapica midea midea (Hiibner) which is endemic to the Georgia and South Carolina

coastal islands, has gone as long as three years as pupae when kept inside in dry conditions and then

emerged when exposed to high spring humidity (Gatrelle 1985). This genetic trait, though currently

unnecessary in midea’s present ecology, is still retained by midea from its Florida ancestor.

This theory would indicate that the multivoltine carlota populations in Georgia and South Carolina

are descendent from ancestors from the west. Dr. Richard L. Brown has documented the eastward

movement, and subsequent influence, of western species on the Lepidoptera of the Black Belt region

(remnant prairies) of Mississippi and Alabama (personal communication). If the above is so, convergent

evolution is taking place as these two formerly isolated taxa move toward each other. Something which is

surely enhanced by human environmental alteration through deforestation and agriculture.

We see this same type of convergent evolution occurring between F. m. midea and F. m. annickae

(dos Passos and Klots) in this same region (Gatrelle, 1998). Midea midea is from a Florida ancestor.

Annickae is from a Mexican/Texas ancestor which moved eastward along the then coast into Georgia and

then northeastward, while midea midea was still isolated in Island Florida. After Florida was rejoined to

the mainland, those midea which moved northwestward (unhindered by habitat barriers) met the annickae

phenotype and gave rise to the mixed phenotypes north and northwest of the Florida panhandle. Those

midea which moved northeastward became isolated on the coastal islands. Before colonial times, these two

subspecies were kept apart in that region by thick maritime forest swamps.

Today, a nnickae ranges from the upper coastal plain and Sandhills of east central Georgia into New

England. Midea was once limited to the Islands of Georgia, South Carolina, and south coastal North

Carolina. However, they may now be found within 60 km of each other in Chatham (midea) and Burke

(i annickae ) counties, Georgia, and within 50 km in adjacent Jasper and Hampton counties of South Carolina.

Three centuries of environmental alteration and destruction is allowing midea and annickae to move

toward each other, and eventually genetically absorb each other in the region. Perhaps, this is also

occurring with gorgone and carlota.

ACKNOWLEDGMENTS

Thanks go to: Mr. Scott Massey for capturing gorgone and computerization of photos, my son Ben

Gatrelle for photographs and computer input, the late C. F. dos Passos for encouraging me in taxonomy and

supplying hard to find publications, and SC DNR staff botanist Bert Pittman for assistance on plant

identification.

LITERATURE CITED

BROWN, F.M. 1974. The Butterfly Called ismeria by Boisduval and LeConte. Bull, of the Allyn Mus.

Bull. No. 16. 12 pp.

CATLING, P.M. & R.A. LAYBERRY. 1998. Distribution and Biology of Chlosyne gorgone carlota

(Nymphalidae) at its Northeastern Limit. J. Lepid. Soc. 52:98-104.

DOS PASSOS, C.F. & A.B. KLOTS. 1969. The Systematics of Anthocharis midea Hubner (Lepidoptera:

Pieridae). Entomologica Americana. Vol. 45: 34 pp.

FORBES, W.T.M. 1960. Lepidoptera of New York and Neighboring States. Memoir 371 part IV (188

pp.). Cornell Univ. Agric. Expt. Sta. New York St. Col. of Agric., Ithaca, NY

HARRIS, L., Jr. 1972. Butterflies of Georgia. Univ. of Okla. press, Norman OK. 326 pp.

HOWE, W.H. 1974. The Butterflies of North America. Doubleday & Co., Inc. NewYork, NY. 633 pp.

GATRELLE, R.R. 1985. The Papilionoidea of the South Coastal Area of South Carolina. Bull. Southern

Lepid. Soc. Bull. No. 2. 15 pp.

-1998. An addendum to Anthocharis midea dosPassos and Klots 1969 (description of a new

subspecies from Texas). The Taxonomic Report Vol. 1, No 1, TILS, Goose Creek, SC. 5 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.

RADFORD, A.E., H.E AHLES & C.R. BELL. 1968. Manual of the Vascular Flora of the Carolinas. Univ.

of North Carolina Press, Chapel Hill, NC. 1183 pp.

SCOTT, J.A. 1986. The Butterflies of North America, A Natural History and Field Guide. Stanford Univ.

Press, Stanford, CA. 583 pp.

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 a publication of The International Lepidoptera Survey (TILS),

126 Wells Road, Goose Creek, SC 29445-3413

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

Every year scores of these taxa 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

(i.e. information) there will be no systematic taxonomy. Without amateur collectors the majority of the undiscovered

species/subspecies will die out before they are discovered.

Please support the environment, support collecting. Be it moon rocks, urine samples, or butterflies, collecting is the

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The Taxonomic Report is projected for publication at the rate of at least 10 issues a year. Subscription is $65 US

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sending a manuscript, simply write TILS at the above address to set up discussion with the head of our research dept, on how

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preserved intact and be available to researchers in this form. For information on this write to: Ronald R. Gatrelle, MOTH

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9

Volume 1

1 December 1998

Number 3

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

TWO NEW NYMPHALIDAE FROM WESTERN NORTH CAROLINA:

NEW SUBSPECIES OF SPEYERIA APHRODITE

AND PHYCIODES BATESII.

RONALD R. GATRELLE 1

126 Wells Road, Goose Creek, South Carolina 29445

ABSTRACT. Two new subspecies are described from Macon County, North Carolina. Speyeria aphrodite

cullasaja is characterized by dorsal melanism, especially in females, by usually lacking the dorsal FW basal spots in cell Cui

and Cu2, and by a narrowed buff marginal band on the ventral HW (sometimes absent in females). Phyciodes batesii

maconensis males are characterized by wholly yellow ventral hind wings devoid of any dark marginal dusting or whitish

crescents; and females occasionally possessing, lightly dusted, brown margins and rarely having a silvered marginal ventral

HW spot. P. batesii maconensis larvae are gregarious Aster feeders. Females are known to oviposit only on Aster undulatus.

Both subspecies are presently known only from the drainage of the Little Tennessee River in western North Carolina and

northern Georgia at altitudes above 4,000 feet. Both are common in this area.

Additional key words: batesii syntypes, Speyeria atlantis, high altitude, Great Smoky Mountains National Park

A new subspecies of Speyeria aphrodite Fabricius

L. Paul Grey was North America’s expert on Speyeria. In our correspondence over the years he had

expressed his belief that the southernmost populations of both Speyeria aphrodite in north Georgia and

western North Carolina (Figs. 1, 2) and Speyeria atlantis Edwards in West Virginia (Figs. 9, 10) were

undescribed subspecies. Unfortunately, I never found the aphrodite he wanted from that area until after his

health failed.

On 8 July 1988,1 located a population of S. aphrodite in the area of Jones Knob, Macon County,

North Carolina. This population was visited in 1988 through 1994 and in 1998 with aphrodite present all

years. In 1990, another population was located in the vicinity of nearby Scaly Mountain, Macon County;

and in 1994 aphrodite was discovered in the area of Rabun Bald in adjacent Rabun County, Georgia. All

known colonies are resident at elevations above 4,000 feet.

A total of about 100 specimens have been collected from these sites. They confirm Dr. Grey’s

speculation and represent a new subspecies.

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

Speyeria aphrodite cullasaja Gatrelle,

species

Speyeria diana (Cramer) .^Secondly, 75 - 80% of specimens (both sexes) haw the basal spots in cell Cu, and/or Cu* either

is dark brown dorsally, concolorous light brown ventrally. Forewings: black

Si

to postmedian black spots in Cm and Cu 2 , frequently into base of cell M 3 , and to second black

Types. Holotype <? (Fig. 1): trail to Scaly Mountain, 4200’, Macon Co., North Carolina, 5 July 1990, Coll. Ronald

R. Gatrelle. Allotype 9 (Fig. 2): meadow near Jones Knob, 4150', Macon Co., North Carolina, 15 July 1994, Coll. Ronald R.

Gatrelle. Paratypes: 58c? <?, 25 9 9: all MACON COUNTY, North Carolina: JONES KNOB: Id, 2 9?, 8 July, 2c? c?,

49 9, 9 July 1988; 4c?c?,7July, 2c? c?, 19, 8Julyl989; 1 c?, 19, 4 July 1990; 1 c?, 5 July 1991; lc?, 2 9 9, 6 July 1993;

2c?, 10 June, 10c? c?, 17 June, 2c? c?, 26 June, 4c? <?, 29 9, 2 July, lc?, 99 9, 15 July 1994; 39 9, 12 July, 19, 13 July

Dr “laughing water.” 1

the area and is threaten

have the typical basaf spots in Cu, &

and none of the females approach cullasaja in degree of basal darkless. They are

acts as an island isolate in a cluster of peaks and connecting ridges above 4000 feet in the southern end of the Blue Ridge. I

expect it to inhabit other Blue Ridge mountains between Highlands and Asheville, North Carolina. Since it has not been

found in the western Appalachian Mtns., it may be a southern Blue Ridge endemic.

A new subspecies of Phyciodes batesii (Reakirt)

Scott (1994) presented a thorough study of the biology and taxonomy of the genus Phyciodes. In that

work, he described three new subspecies of Phyciodes batesii (Reakirt), of which P. b. lakota Scott, TL

Pine Ridge, Sioux County, Nebraska is one. Scott lumped lakota into his clinal concept of “eastern” batesii

- which he sees extending from Alberta, Canada, across southern Canada and the northern United States to

the Atlantic Ocean, and then down the Appalachian Mountains terminating in northern Georgia. A vast area

overlapping many life zones, habitats, elevations, etcetera.

In this clinal concept, he correctly sees two phenotypic “extremes” (his term) and a variable

intermediate. He correctly extends the range of his western extreme, lakota (Fig. 16), eastward into

Michigan and central Ontario. He accurately limits his other extreme phenotype to the high mountains of

the southern Appalachians in northeast Georgia and western North Carolina. He then incorrectly associated

this second extreme with the intermediate variable phenotype, northeastern US nominate batesii (Figs. 17,

18, 19), which he extended into eastern Ontario.

His taxonomic verdict should have been based foremost on any existent type specimens and an

accurate understanding of the type locality (location, habitat, altitude etc.). Then without prejudice, allow

the facts to dictate the conclusions. He did not. His treatment of P. batesii in the eastern and southern US is

prejudicially incorrect. His statements reveal his lakota taxonomy is based heavily on in his own

predisposed personal agenda. “Because I wish to use only two names in this cline, I want to have the TLs

[type localities] as near as possible to the ends of the cline.” “I have used taxonomic license to force the

[his] concept.”

Type specimens

In his batesii research, he refers, directly or indirectly, to information supplied to him by myself no

less than 7 times. His taking “taxonomic license” included not mentioning photos of the syntypes (one from

Winchester, VA and one from Colorado) in the FMNH that I informed him of in a 17 Nov. 1993 letter,

(photos provided by the Allyn Museum of Entomology.) The Winchester syntype clearly shows a P. batesii

batesii type specimen (Fig. 14) with dark shading along the outer margin of the ventral HW. In his study,

he did mention that I had informed him (in the same letter) that no Gloucester, New Jersey (type locality)

specimens could be found. However, he neglected to mention that I had located and obtained the loan of

Shipiro’s Philadelphia, County, PA, batesii taken VI-23-65, Wissahickom. A specimen taken only 15 miles

from Gloucester (and thus a virtual topotype) with quite a bit of shading along the ventral margin of the

HW. (Specimen now housed in the Bohart Museum, University of California Davis.)

Since both the existent syntype and topotype possess brown scaling (and of equal importance,

crescent spots) in their margins, Scott’s attempt to present nominate batesii as a taxon that doesn’t posses

dark scaling along the ventral margin of the HW is false (Figs. 17, 19). Scott states, “My separation of ssp.

batesii and lakota is based on this [brown] patch...” But, this patch in lakota isn’t consistently prominent

(Fig. 16). Scott, on page 51, mentions “families” (colonies? adults from larval groups?) of lakota that “in

wing pattern resembles N Appalachians [= batesii batesii ] adults.”

3

In actuality, Scott’s lakota is either an east-west clinal subspecies of batesii or should be

considered a synonym of nominate batesii. The three major characters on the ventral HW mentioned by

Scott (silver crescent, brown dusting, heavier marking) are shared by lakota and eastern batesii in classical

clinal gradation. Further, there is no barrier or break (isolation) in his east-west cline. He has both

subspecies in Ontario Qakota central and batesii eastern). The syntypes themselves connect the western

0 lakota ) and eastern ( batesii ) population, as one is from Colorado (Fig. 15) and the other Virginia!

Conversely, none of those three characters, by Scott’s own research, are found in northern Georgia

and western North Carolina batesii. Furthermore, batesii in north Georgia and western North Carolina are

isolated on elevational Islands above 4,000 feet in the southern section of the Appalachians. They are

isolated broadly from northern populations by valleys at about 2,000 feet (followed by the routes of 1-81

and 1-75) in southwestern Virginia. They may be more narrowly isolated by elevation, as is Speyeria

aphrodite cullasaja (described above), which occupies the same ecological niche.

Type locality

Scott continually refers to his restricted P. b. batesii type locality of Winchester, VA as being in the

northern Appalachian “mountains.” The elevation of Winchester is 714 feet - basically the same as

Charlotte, North Carolina, and 100 feet lower than Spartanburg, South Carolina. Winchester is on the edge

of the Virginia piedmont (see Clark and Clark, 1951). Winchester is only 38 miles south of Pennsylvania

and 145 miles southwest from Gloucester, NJ.

Scott (1986) shows Winchester, VA, Gloucester, NJ, and Sioux County, Nebraska, to be in the

Upper Austral faunal zone. Both batesii and lakota are inhabitants of primarily once glaciated and/or Upper

Austral faunal areas. His same 1986 map shows his unique batesii extreme in North Carolina to be in the

Canadian zone. In nature, P. batesii batesii, and lakota, are non-isolated, non-montane. The heretofore

undescribed western North Carolina batesii is an isolated, high altitude, montane subspecies.

I see no reason to “restrict” the type locality to the immediate vicinity of either city mentioned in the

original description (as did Klots (1951, Gloucester), or Scott (1994, Winchester)). Though to do so, in my

opinion, would have no effect on the systematic taxonomy of P. batesii batesii. Thus, I consider the type

locality to be the relatively small 145 mile Upper Austral geographical area from sea level (coastal plain)

to 714 feet (piedmont) as stated in the original description. (State lines are artificial boundaries that have

nothing to do with taxonomy.)

Phyciodes batesii maconensis Gatrelle, new subspecies

Diagnosis. As described in Scott (1994), “In southern Appalachians the unh of males always lack a marginal brown

patch and pale crescent, and the unh of females generally lacks a marginal brown patch (or it is very weak) and usually lacks

the crescent.” (Emphasis mine.) When a crescent is found on females, it is rarely silvered (I have seen only 2 silvered 9 9

out of 90 examined). Nominate batesii from northeastern Virginia (TL) and the northeastern US differs from maconensis as

per Scott (1994), “The brown unh marginal patch and pale unh marginal crescent vary esp. in males (females often have both

patch and crescent).” (Emphasis mine.) Further, the crescent is often silvered in both males and females of batesii (Fig. 18).

The theme throughout the literature is that P. batesii batesii may be distinguished by its diminished or absent VHW brown

patch. Distinguished from what? From P. tharos (Drury). Regarding the brown patch, maconensis is to batesii as batesii is

to tharos. In comparison to maconensis, many batesii may be said to have a prominent brown patch. These same batesii in

comparison to tharos may be said to have a light patch. The average FW radius of male maconensis is 18 mm (expanse 35

mm), in females it is 21 mm (expanse 38.5 mm). My largest female has an expanse of 41 mm. In four pair of NY batesii I

examined, the male FW radius averaged 16.5 mm (expanse 33.5), the female FW radius averaged 18 mm

s, the butterfly is out, but the collector 8 is in. My own discovery of it was by “accident.” The year 1992 was a very late

Conclusion

Speyeria aphrodite cullasaja and Phyciodes batesii maconensis occupy what seems to have been a

relatively uncollected niche in the southern Blue Ridge and Appalachian Mountains. Both of these species

have, until now, been considered rare in north Georgia and western North Carolina. They are actually

rather common. It is not surprising therefore, that other species which are generally considered uncommon

or rare in that region (Harris 1972) also inhabit this same niche - with annual regularity. The following are

resident at Scaly Mountain (S), Jones Knob (J), Rabun Bald (R): Autochton cellus (Boisduval and

LeConte) (J); Polites coras (Cramer) (S,J,R); Poanes hobomok (Harris) (S,J,R); Artogeia virginiensis

(below J); Lycaena p. americana Harris (J,R); Parrhasius m-album (Boisduval and Leconte) (S,J,R);

Erora laeta (W. H. Edwards) (S,J); Polygonia faunus smythi A. H. Clark (R,J); Clossiana bellona

(Fabricius) (R,S); Ceryconis pegala Carolina (F. & R. Chermock) (status per Gatrelle 1985) (S,J). Of

these, coras, hobomok and americana are common; laeta, smythi, bellona and Carolina generally present

in small numbers; and virginiensis, m-album and cellus , infrequent.

Acknowledgments

Thanks go to my son, Ben Gatrelle, for photography of specimens and Scott Massey for

computerization of said photos. To L. Paul Grey for pointing me in the right direction and putting me on the

trail. Dr. James Scott for specimens and exchange of thought. To Dr. Lee Miller and the Allyn Museum for

continued support of my research by the supply of information and photos of type specimens. And to Dr. Art

Shapiro for the loan of his Philadelphia batesii.

Literature cited

CLARK, A. H. & L. F. CLARK, 1951. The Butterflies of Virginia. Smithsonian Misc. Collect. 116 (7) 239

pp.

GATRELLE, R.R. 1985. The Butterflies of the south coastal area of South Carolina. Bull. Southern Lepid.

Soc. Bull. No. 2. 15 pp.

HARRIS, L., Jr. 1972. Butterflies of Georgia. Univ. of Okla. Press, Norman, OK. 326 pp.

KLOTS, A. B. 1951. A Field Guide to the Butterflies. Houghton Mifflin, Boston MA. 349 pp.

SCOTT, J. A. 1986. The Butterflies of North America, a natural history and field guide. Stanford Univ.

Press, Stanford, CA. 583 pp.

- 1994. Biology and Systematics of Phyciodes. Papilio (new series), by author, Lakewood, CO. 120 pp.

FIGS. l-2. Type specimens of Speyeria aphrodite cullasaja. 1, d holotype, dorsal and ventral surfaces. 2, 9 Allotype,

dorsal and ventral surfaces. Data in text.

6

FIGS. 3-19. 3, Topotype d Speyeria aphrodite, 3 July 1983 Long Island, NY, dorsal and ventral surfaces. 4, 9 S. aphrodite, 3 July 1981

Ulster Co., NY, dorsal and ventral surfaces. 5, d S. cybele novascotiae, 28 July 1990, Halifax Co., N.S. Canada, dorsal surface. 6,

Topotype 9 S. atlantis capitanensis, 1 July 1989, 8000’, Lincoln Co., NM, dorsal surface. 7, d S. atlantis, 16 July 1983 Vilas Co., WI,

dorsal surface. 8, 9 S. atlantis, 15 July 1983 Florence Co., WI, dorsal surface. 9, d S. atlantis ssp., 3 July 1987, 4200’, Randolph Co., WV,

dorsal surface. 10, 9 S. atlantis ssp., 28 June 1987, 4200’, Randolph Co., WV, dorsal surface. 11, d holotype Phyciodes batesii

maconensis, dorsal and ventral surfaces, data in text. 12, 9 allotype Phycoides batesii maconensis, dorsal and ventral surfaces, data in text.

13, 9 paratype 077 P. b. maconensis, 10 June 1994 Macon Co., NC, ventral surface. 14, VA d syntype of P. batesii. 15, CO d syntype of

P. batesii. 16, d paratype P. b. lakota, ex pupa 10 August 1994, Sowbelly Cyn., Sioux Co., NE. 17-18, d d P. batesii. 19, 9 P. batesii.

17-19, 10 June 1974 Jamesville, Onondaga Co., NY.

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

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.

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

Every year scores of these taxa have not even been scientifically discovered and documented. Thus, their extinction is

Volume 1 1 January 1999 Number 4

SUBSPECIFIC STATUS OF SOUTHEASTERN U.S. MEGATHYMUS COFAQUI AND M

YUCCAE : RENAMING OF THE FLORIDA SUBSPECIES OF M COFAQUI .

RONALD R. GATRELLE 1

126 Wells Road, Goose Creek, South Carolina 29445

ABSTRACT. Megathymus cofaqui and M yuccae are both represented in the southeastern U.S. by two subspecies.

The type locality of both M. y. yuccae and M c. cofaqui is the area of Burke/Screven counties Georgia. Each of their

subspecies are primarily Floridian, with M. y. buchholzi extending along the immediate coast of Georgia into southern South

Carolina. Topotypes of M cofaqui from Burke County, Georgia, and Aiken County, South Carolina are phenotypically

indistinguishable from both the holotype of M cofaqui and topotypes of M. c. harrisi. Thus, M. c. harrisi is synonymous

with M. c. cofaqui. This leaves the Florida subspecies of M cofaqui without a valid name. Megathymus cofaqui slotteni is

proposed as a new name for the Florida subspecies. The holotype of M c. slotteni is deposited in the Florida State Collection

of Arthropods, Gainesville, Florida.

Additional key words: type localities.

OVERVIEW AND DELINEATION OF TAXONOMIC PROBLEMS

At least 33 species of butterflies were originally described from east coastal Georgia or south

coastal South Carolina. A broad taxonomic problem exists because most of these 33 taxa are not

represented by any type or topotypical specimens in any of the world’s institutional or private collections.

A few of these are known in their nominate form from only a unique type specimen. Megathymus cofaqui

(Strecker) is an example of the latter. Further, many of these 33 species/subspecies have been wrongly

assumed to occur in their nymotypical form in their Floridian components. Unfortunately, this common

erroneous assumption was applied to M. cofaqui.

Having personally collected nearly all of these 33 species in Burke or Screven counties over the

last ten years, this researcher can attest to the fact that the only reason topotypes of these taxa are not

represented in collections, and are thus unavailable to researchers, is simply because few lepidopterists

have gone there to collect them. Most of them are not uncommon in that area. Over the decades, northern

lepidopterists literally drove right past scientifically important specimens along highways 17 and 301 in

Georgia on their way to collect the subtropical species of south Florida.

Lucian Harris, Jr. (1972) presents a concise historical overview of Megathymus cofaqui and

Megathymus yuccae (Boisduval and LeConte). Both species were first discovered in and described from

the area of Burke and Screven counties, Georgia. M. yuccae was described from a John Abbot painting. M.

cofaqui was described from a female collected by Morrison. Harris was not enthused with H. A.

Freeman’s designation of Aiken County, South Carolina as the type locality for Megathymus yuccae; or

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

Freeman’s affirmation of Grossbeck’s 1917 assignment of Boca Grande, Lee County, Florida as the type

locality of M. cofaqui. Harris’ consternation is especially understandable in relation to M. cofaqui, for the

9 holotype (fig. 5) is existent in the Field Museum in Chicago, and Strecker had clearly designated Georgia

as the type locality in the original description.

COFAQUI

At the time of Harris’ writing, it was not yet known that Megathymus karrisi H. A. Freeman was in

fact only a cofaqui progeny and not a valid species. Up to the 1950’s, the only known Georgian cofaqui

was the type. About 85 years passed between the capture of the cofaqui type specimen and the next capture

of a Georgian specimen (in the Atlanta area). During this time some supposed cofaqui were being collected

in various locations in central/southern Florida. The assumption was that what was being collected in

Florida was the same thing as what had originally been discovered in eastern Georgia. It was further

assumed by some (Freeman) that since no other cofaqui had been found in Georgia in 85 years that perhaps

the type really did not come from there. These two errors - viewing harrisi as a full species and the

accepting of southwestern Florida as the type locality of cofaqui - have created an unallowable taxonomic

situation.

Miller and Brown (1981) recognized the type locality error and reestablished Burke County,

Georgia as the type locality of M. cofaqui , but left cofaqui and harrisi as distinct species in the Lepid. Soc.

checklist. Ferris (1989) retained harrisi and cofaqui as separate species in the checklist revision. Hodges

(1983) also retained these two as separate species in his checklist. This is unfortunate since Howe (1974),

Pyle (1981), Opler and Krizek (1984), Scott (1986), and Gerberg and Arnett (1989), all correctly

recognized that the northern and southern cofaqui phenotypes were but north/south clinal subspecies and not

species.

Figs. 1-6. Megathymus cofaqui subspecies. Fig. 1, Paratype d M. cofaqui slotteni, ex pupa 13 Aug. 1990 Interlocked

Putnam County, Florida (leg. Dr. Jeff Slotten). Fig. 2, Holotype 9 M cofaqui slotteni, 20 Sept. 1988 vise, of Willliston,

Levy County, Florida (leg. Dr. Jeff Slotten). Fig. 3, Topotype 9 M cofaqui harrisi, ex pupa 24 July 1977 Atlanta, Georgia.

Fig. 4, Topotype d M. cofaqui, ex pupa 20 Aug. 1998 Burke County, Georgia. Fig. 5, Holotype 9 Megathymus cofaqui,

Georgia (dorsal & ventral). Fig. 6, 9 M cofaqui, ex pupa 16 Aug. 1990 New Ellenton, Aiken County, South Carolina.

It has taken me 25 years to locate a few Megathymus cofaqui colonies in Columbia, Burke, and

Screven counties, Georgia, and Aiken, Orangeburg, Sumter, and Lancaster counties, South Carolina. With

several of these records based only on empty larval tents. All adult specimens from these localities are the

same subspecies. Topotypes of M. cofaqui from Burke County are nearly identical to topotypical harrisi.

Thus, there is only one subspecies of M. cofaqui that occurs in Georgia and South Carolina.

The holotype of M. cofaqui (fig. 5) has far too dark of a ground color to be from southern or central

Florida. All of its markings are within the typical range of Burke County cofaqui , especially the marking of

the dorsal fore wings. The presence of yellow scales along its dorsal hindwing margin is a character more

prevalent in females of the Florida subspecies (Freeman 1969). However, some Georgia females also

exhibit this character. The Fulton County, Georgia harrisi female figured by Harris (plate 10 fig. 19) and

the holotype of cofaqui could almost pass as the same specimen. It is documented in Harris (pg. 28) that

Morrison was collecting in Burke/Screven counties at the proper time of year to have found his cofaqui

there. Morrison stated that he found it there. We know, from specimens collected by myself, that cofaqui is

still found there. There is absolutely no historical or phenotypic evidence to suggest that the type of cofaqui

came from any other population than that in the area of Burke/Screven counties in Georgia.

Topotypes of cofaqui from Burke County, Georgia (fig. 4) and adjacent Aiken County, South

Carolina (fig. 6), and topotypical specimens of harrisi from the Atlanta, Georgia area (fig. 3) reveal that

cofaqui and harrisi are synonymous. This dictates that the taxon named after Harris (M. c. harrisi) must be

dropped into the synonymy of cofaqui. It also necessitates that the name cofaqui applies only to the

northern (Georgian) subspecies and can not be used for the long recognized southern (Floridian)

subspecies. This in turn leaves the Florida cofaqui subspecies (figs. 1 & 2) without a name. Accordingly, I

herein describe the southern race as a new subspecies, Megathymus cofaqui slotteni Gatrelle.

YUCCAE

The problem concerning Megathymus yuccae is slight but should at least be mentioned. H. A.

Freeman designated a neotype for M. yuccae from Aiken County, South Carolina simply because no Burke

or Screven County, Georgia specimens were known at that time. There is no conflict in Freeman’s

designation of a biological neotype from adjoining Aiken County, South Carolina. The ecology of Burke

County, Georgia has more in common with adjoining Aiken County, South Carolina than with most of

adjoining Screven County, Georgia. This is because most of both Burke and Aiken counties are upper

coastal sandhill habitats, while most of Screven County is lower coastal plain maritime forest.

I do not think that national, state, county, or any other subjective political boundary lines, have any

bearing on systematic taxonomy. If the environmental factors are the same, a specimen caught one inch or

one mile on one side or the other of such a line should not matter in biologically defining a type locality, or

what constitutes a biological topotype. For example, I have a specimen of Asterocampa celtis (Boisduval

and LeConte) which I caught as it landed on the Burke County, Georgia bank of the Savannah River after it

appeared to have just flown 40 yards across the river from South Carolina! Is this a South Carolina or a

Georgia specimen?

Humans divide up areas and affix names to places in order to establish their ownership and legal

jurisdiction. (Where is Ceylon or the USSR today?) All type localities are geopolitical and geoecological.

The latter is all that should matter scientifically, because the former is totally artificial. In practice,

taxonomists use geopolitical names to easily reference a type locality. But scientifically, the type-locality

can only be the geoecological area (which may be very small or quite large depending on the parameters set

by the author — a colony, population, or phenotype) occupied by the single colony, population, or

phenotype from which the representative type specimen(s) was/were taken. This is analogous to the use of

3

common and scientific names. Common names have absolutely no scientific standing (any person or

organization can make up their own list or book of butterfly common names and it is just as valid as anyone

else’s.) Geopolitical type localities are like common names. They only serve as an easy and brief way to

reference (in an inherently inaccurate manner) areas occupied by taxa which have a much more involved

and scientific geoecological type locality. I accept the region of Burke County, Georgia as the type locality

of the nominate population of Eudamus yuccae Boisduval and LeConte and, conditionally, accept

Freeman’s designated neotype from that region and population (Aiken County, South Carolina) as a valid

topotype.

The condition of my acceptance stems from the artificial environment from which the neotype came.

I have a problem with the fact that Freeman’s neotype came from an urbanized, artificially established,

decorative stand, of Yucca aloifolia L. If Harris’ determination is accurate, these plants were far from their

natural range. Y. aloifolia is indigenous to South Carolina only in a few coastal counties (Radford et. al.

1968). Thus, we can not be sure that the Megathymus yuccae specimens found there were not from larvae

or eggs imported into the area (from Florida?) with the plants.

I did not feel it necessary to figure specimens of typical M. yuccae yuccae or M. y. buchholzi. They

do not differ a great deal and not always consistently. However, I feel that most specimens can be

distinguished from each other without looking at locality labels. Further, M. yuccae becomes more distinct

as one goes north and west from the Georgia coast. Unfortunately, the type locality is barely outside of what

may be the blend zone of these subspecies. The Burke, Screven, and Aiken material I have from wild stands

of Y. filamentosa L., have smaller, lighter, more yellowish median spots on the dorsal FW. Specimens

from south coastal South Carolina have larger, richer, more orange spots. My coastal South Carolina

specimens generally match individuals I have collected or examined in various personal and institutional

series of Floridian M. y. buchholzi and are thus either referable to buchholzi, or represent an intermediate

population at the northern end of the range of buchholzi.

A 9 specimen of M. y. buchholzi I collected on Edisto Island, Charleston County, South Carolina is

figured by Scott on plate 57, Figure 417 d. This specimen clearly shows the orange spotting that is

characteristic of buchholzi. This specimen can be contrasted against the typical yellow spotted M. y.

yuccae figured by Harris on plate 10, figures 14 & 15.

Megathymus cofaqui slotteni Gatrelle, new subspecies

Diagnosis, slotteni, having passed for decades as typical cofaqui, is well depicted under that name throughout the

popular and scientific literature. Figures of slotteni may be found under the name cofaqui in Holland (plate LIV, fig. 34 &

35), Klots (plate 40, figs. 3 & 4), Harris (plate 10, figs. 20 & 21), Howe (plate 82, figs. 7 & 8), and Scott (plate 57, fig.

420). Harris’ figures perfectly depict and contrast these two subspecies as described below. The dorsal brown ground color

is lighter in specimens further south (slotteni), and darker in specimens further to the north (cofaqui). The key distinguishing

characters are the shape and relative size of the three spots in the postmedian spot band on the dorsal forewings of both males

and females, and the size of the postmedian spot band on the dorsal hindwings of females. In slotteni the spot in cell Cu2 is

nearly always at least half the width of the spot in cell Cui or larger. In cofaqui this spot is usually only one third the width

(at the vein) and rarely over half (both sexes). In cofaqui the spots in M 3 and Cui usually tend to be elongated and fused with

the spot in the distal end of the fore wing cell (both sexes). In slotteni this is not often the case, and when it is, the spot in Cu 2

is also expanded (esp. in females). The extent of the light yellow spotting on the ventral hindwings of females is so variable

throughout the range that it should not be considered as a diagnostic character. Only at the extremes of the cline is this

spotting consistently stronger in the south and nearly absent in the north. The tendency of females to have yellow dorsal

hindwing margins is typical in slotteni and atypical, but occasional, in cofaqui. The dorsal hindwing spots on cofaqui females

are usually smaller while on slotteni they are larger (heavily spotted cofaqui females look like lightly spotted slotteni

females). In general, one could say that females of cofaqui have a greater size difference between their dorsal forewing and

hindwing spots, while slotteni females have less of a size difference between their dorsal forewing and hindwing spots.

These subspecies are clinal, and individual specimens within each subspecies are variable. They are not greatly distinct.

Description. Male (Fig. 1): Head, thorax, abdomen, and appendages with gray and brown scaling slightly lighter

than in nominate subspecies. Forewings: dorsally, ground color dark to medium brown with golden brown scaling at base

lighter than in nominate subspecies, postmedian spots light yellow, with spot in Cu 2 not strikingly smaller then those in Cui

and M3, spots in Cui and M 3 not often fused with spot in discal cell; ventrally, as in dorsal except no light basal scaling.

Hindwings: dorsally, dark to medium brown with golden brown scaling a base lighter than in nominate subspecies; ventrally,

as in nominate subspecies, except that black postmedian spots often more pronounced. Female (Fig. 2): Head, thorax,

abdomen, and appendages with gray and brown scaling slightly lighter than in nominate subspecies. Forewings: dorsally,

ground color dark to medium light brown, lighter than in nominate subspecies (especially in southwestern Florida), golden

brown scaling at base lighter than in nominate subspecies (especially in southwestern Florida), postmedian spots light orange

yellow, with spot in Cu 2 not strikingly smaller then those in Cui and M3, spots in Cui and M 3 not usually fused with spot in

discal cell; ventrally, as in dorsal except no light basal scaling. Hindwings: dorsally, dark to medium brown with golden

brown scaling at base lighter than in nominate subspecies (especially in southwestern Florida), row of orange yellow

postmedian spots often large, outer margins usually with extensive yellow; ventrally, with light yellow/whitish basal spots

usually present and sometimes prominent, yellow/whitish spots and black spots of postmedian spot band often prominent.

Types. Holotype 9 (Fig. 2): Vicinity of Williston, Levy County, Florida, 20 September 1988, leg Dr. Jeff Slotten.

Paratypes: 3 d* d\ 2 9 9: all FLORIDA: 1 d* near Interlocken, Putnam County, 13 Aug. 1990; 1 d* Hernando County, 7 Sept.

1989; 1 d* near Williston, Levy County, 6 Sept. 1989; 1 9 Clay County, 1 March 1991; 1 9 near Williston, Levy County, 5

Sept 1989. All were collected by Dr. Jeff Slotten. The holotype is deposited in the FSCA, Gainesville Florida. The paratypes

are in the authors collection in Goose Creek, South Carolina.

Etymology. Slotteni is named after Dr. Jeff Slotten, a prominent amateur Florida lepidopterist.

Remarks. Because these two subspecies have been recognized for over 40 years, I see no need in designating a long

type series. In fact, 5 paratypes might be considered excessive by some. I chose a female as the holotype of slotteni for two

reasons. First, the type of cofaqui is female, and second, females of the two subspecies differ subspecifically more than their

males. There is probably a broad blend zone between these subspecies roughly parallel to the Georgia/Florida state line. I

feel the type locality of slotteni is sufficiently to the south of this blend zone. It is assumed here that no M. c. cofaqui occur

in Florida. If it does, it would be expected only in the northwestern part of the state. M. c. slotteni does not occur north of

Florida. I have examined all the cofaqui in the FSCA collection in Gainesville, and three private Floridian collections. I

have not seen any of the cofaqui from the apparently large population that exists in the mountains of North Carolina. The one

(and to my knowledge only) collector who has had a series of these, for years, has published nothing. With the known

tendency of Megathymus to evolve into almost micro geographical subspecies, these North Carolina cofaqui (as well as the

Tennessee population) need to be collected, in a large enough sample, so they can be examined by a competent taxonomist -

and their status, whatever it may be, published.

ACKNOWLEDGMENTS

Thanks to my son Ben Gatrelle for photographs and computer input, Dr. Lee Miller and the Allyn

Museum for photo of type specimen, Jeff Slotten for specimens, and to the FSCA, Gainesville, for library

support.

LITERATURE CITED

FERRIS, C.D. ed. 1989. Supplement to: A Catalogue/Checklist of the Butterflies of America North of

Mexico. Memoir No. 3, Lepid. Soc. 103 pp.

FREEMAN, H.A. 1969. Systematic Review of the Megathymidae. Supplement 1, J. Lepid. Soc. 58 pp.

GERBERG, E.J. & H.R. ARNETT, JR. 1989. Florida Butterflies. Natural Science Publications, Inc.,

Baltimore, MD. 90 pp.

HARRIS, L., Jr. 1972. Butterflies of Georgia. Univ. of Okla. Press, Norman OK. 326 pp.

HODGES, R.W., et al. 1983. Check List of the Lepidoptera of America North of Mexico. MONA series.

E.W. Classey Ltd. & Wedge Ento. Research Foundation, Oxfordshire, England. 284 pp.

HOLLAND, W.J. 1930. The Butterfly Book. Doubleday & Company, Inc., Garden City, NY. 424 pp.

HOWE, W.H. 1974. The Butterflies of North America. Doubleday & Comapany, Inc. NewYork. 633 pp.

KLOTS, A.B. 1951. A Field Guide to the Butterflies. Houghton Mifflin, Boston MA. 349 pp.

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. 294 pp.

PYLE, R.M. 1981. The Audubon Soc. Field Guide t> North American Butterflies. Alfred A. Knopf, Inc.,

New York. 294 pp.

RADFORD, A.E., H.E. AHLES & C.R. BELL. 1968. Manual of the Vascular Flora of the Carolinas. Univ.

of North Carolina Press, Chapel Hill. 1183 pp.

SCOTT, J.A. 1986. The Butterflies of North America, A Natural History and Field Guide. Stanford Univ.

Press, Stanford, California. 583 pp.

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

126 Wells Road, Goose Creek, SC 29445-3413

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.

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

Every year scores of these taxa 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

(i.e. information) there will be no systematic taxonomy. Without amateur collectors the majority of the undiscovered

species/subspecies will die out before they are discovered.

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6

Volume 1 1 January 1999 Number 5

A COMMENT ON FRIEDLANDER’S ASTEROCAMPA (NYMPHALIDAE,

APATURINAE): DESIGNATION OF NEOTYPES FOR A. CELTIS AND A. CLYTON.

RONALD R. GATRELLE 1

126 Wells Road, Goose Creek, South Carolina 29445

ABSTRACT. Friedlander’s 1986 [1987] treatment of the southeastern U.S. taxa of Asterocampa is examined.

Neotypes of A. celtis and A. clyton are designated from Burke County, Georgia to help stabilize the status of these taxa and

aid in any future research. A. celtis is currently known to exist within at least 40 km of A. c. reinthali in Georgia. Their

proximity, without integration, suggests evolutionary distance, perhaps even speciation. It is proposed that A. celtis is

descended from A. c. alicia, mdalicia from a Mexican refugium. A. clyton and A. c. flora intergrade in eastern Georgia and

are subspecific.

Additional key words: cryptic subspecies.

UNSTABLE TAXONOMY

Up to 1975, Asterocampa was usually thought of in North America as two or three species with five

to seven subspecies. From 1975 (Howe fide Reinthal) until 1986 (Scott fide Friedlander) Asterocampa was

presented in the literature as consisting of twelve North American species with only one subspecies. In

1987, Friedlander’s revision of the genus left us with four species having six subspecies.

When I received number 25:4 of The Journal of Research on Lepidoptera in early 1988 (the date of

publication is Dec. 31, 1987) devoted entirely to Friedlander’s revision of Asterocampa, I immediately sat

down to read it. I then spent the next several weeks comparing it against the literature and specimens in my

collection. I, like many others, found his treatment much more acceptable than the hyper-splitting of

everything into distinct species that had existed for a decade. However, it seemed to me, then and now, that

his criteria for sinking taxa into subspecies or synonymy was not consistent and somewhat extreme in the

opposite direction. Further, while he properly affirmed the type locality of Asterocampa celtis (Boisduval

& Le Conte) to be Screven County in the lower coastal plain of Georgia, it seemed very risky to me that he

did so without examining any topotypical specimens. Why?

All of the Georgia A. celtis celtis he examined were from locations relatively far from the type

locality. Four locations were in the Georgia piedmont, one was montane, and two (Decatur County in

southwestern GA and Chattahoochee County which borders Alabama) were in the upper coastal plain. The

closest specimens he had to the type locality were from Bibb and Clarke counties, which are both over 170

km away. All of the specimens he had seen from the counties just to the east of the type locality (in both

Georgia and South Carolina) were his new subspecies, A. celtis reinthali Friedlander, and were only 40

km from the celtis type locality. Without specimens, how could Friedlander know that what occurred in

nature in Screven County was not reinthali too? The answer is he couldn’t.

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

Figs. 1-10 eastern U.S . Asterocampa. 1, tfc ? A. celtis reinthali. Top: 3 July 1980,

Gainesville, Alachua Co., FL; bottom: 6 Sept. 1992, Goose Creek, Berkeley Co., SC

(ventrals) 2, Top: Neotype d Apatura celtis celtis, 6 Aug. 1998, at Savannah River,

Burke Co., GA; bottom: same as 1 bottom (ventrals). 3, Dorsal surfaces of speci¬

mens in figure 2. 4, Top: same as 2 top; bottom: d A. celtis celtis, July 1960, Zering,

Story Co., IA (dorsals). 5, Topotype 9 A. clyton clyton, 6 Aug. 1998, at Savannah

River, Burke Co., GA. 6, Top: d A. clyton flora, 21 July 1992, Gainesville, Alachua

Co., FL; bottom: Neotype d Apatura clyton clyton, 6 Aug. 1998, at Savannah River,

Burke Co., GA (dorsals). 7, 9 A. celtis reinthali, 4 Sept. 1992, Goose Creek, Berkeley

Co., SC. (drosal & ventral). 8, 9 A. clyton flora (dark form), 9 Sept. 1991, Newnans

Lake, Alachua Co., FL (dorsal). 9, 9 A. celtis reinthali, 1 July 1980, Gainesville,

Alachua Co., FI (dorsal). 10, 9 A. clyton flora (light form) 24 July 1992, Gainesville,

Alachua Co., FI (dorsal).

Friedlander accepted Abbot’s paintings as being adequate to represent the type. However, Abbot’s

artistic renditions can not be trusted as being so accurate, either to phenotype or place of origin, as to be

equivalent to actual specimens - especially type specimens. (To me, the elevation of the artistic products of

the early workers on Lepidoptera as being definitive scientific types is equivalent to accepting prehistoric

cave paintings as being on a par with the fossil record.)

If it could be demonstrated, through a series of specimens from Screven or Burke counties, that

what occurred at the celtis type locality was the reinthali phenotype, then reinthali would have to be

dropped into synonymy and celtis would become the name applied to his new south coastal South Carolina

to southern Florida subspecies. This would also mean that A. alicia (W. H. Edwards), as the next oldest

name, would have to be applied to the inland and northern subspecies long know as celtis. If Friedlander

was wrong about what phenotype (subspecies) existed in Burke and Screven counties, he had created a

taxonomic mess. As it turns out, he was correct.

LOCAL OBSERVATIONS

I had been collecting and observing Asterocampa celtis and A. clyton (Boisduval & LeConte) in

south coastal South Carolina for 18 years when Friedlander’s paper came out. All I had ever encountered in

this area was his reinthali phenotype. Dr. Richard T. Arbogast had found only reinthali and an occasional

A. clyton with a flora (W. H. Edwards) facies in his many years as a resident of Savannah, Georgia.

Charleston was further north than Screven County. In Friedlander’s study, the reinthali he examined from

coastal Georgia were four times closer to celtis ’ type locality than any celtis he had seen from Georgia.

Thus, I felt there was a strong possibility that what occurred in Burke and Screven counties was a

nondescript blend zone population, or perhaps the coastal reinthali - but not celtis celtis. So I set out to

collect and rear both A. celtis and A. clyton from the south coastal area of South Carolina, and to collect

topotypical specimens of A. celtis and A. clyton to compare against Friedlander’s position.

From 1988 to 1995,1 netted hundreds of adults and reared about 3,000 specimens of both of these

species from Berkeley and Charleston counties SC. (Nearly all netted and reared specimens were

released.) All local celtis were reinthali (Figs. 1-3 bottoms, & 7) and all clyton were clyton clyton.

Random reinthali larvae were checked for antler scoli AB5. All keyed out with this and the other reinthali

larval characters set forth by Friedlander. Celtis laevigata Wild, was the local host of both A. c. reinthali

and A. clyton.

Of anecdotal interest, A. c. reinthali eclosed after midnight and before dawn. If a light was on in the room, newly

emerged adults (especially females) would often come to the light. Perhaps they were fooled into thinking it was dawn and

moved into the light to make themselves more available to males (this may be an explanation for Friedlander’s question at the

bottom of page 240). For those who find wild Asterocampa at light, they should note if the females are usually fresh and the

males old. If so, this could indicate early morning mate locating behavior.

In August of 1998 I located sizable populations of topotypical A. celtis (Figs. 2-4, tops) and A.

clyton (Fig. 5 & Fig. 6 bottom) near the Savannah River in Burke County, Georgia and Allendale County,

South Carolina. All the clyton were clyton clyton and, to my surprise, all the celtis were identical to

specimens of celtis celtis I had collected as a teenager in my home state of Iowa (Fig. 4 bottom)! I had

expected to find a reinthali/celtis intermediate at the type locality. And indeed, if we were dealing with a

clinal subspecies, that is what should have been found in this area.

These A sterocampa were in association with Celtis occidentalis var. georgiana (Small) Ahles.

Both males and females were found of A. clyton but only males of A. celtis. Both species were very

common and about two dozen of each were netted and examined. (C. occidentalis var. georgiana does not

occur along the coast where A. c. reinthali is found. I did not observe C. laevigata in the area of the A.

celtis colonies, but it probably occurs in the vicinity. To this point in time, these plant associations only

indicate occurrence and do not necessarily indicate, or eliminate, regional host specificity.)

3

POSTULATING A CONCLUSION

Specimens, adults and immatures, from southeastern Florida and south coastal South Carolina (700

km apart) are phenotypically (genetically?) the same - reinthali. Specimens, at least adults, from the warm

climate of east coastal Georgia and the cold climate of Iowa (1,500 km apart) are phenotypically

(genetically?) so similar as to be virtually the same - celtis. Yet, with only one county between them (40

km), these celtis and reinthali exist in nature in eastern Georgia as very distinct phenotypes, with no known

intermediate populations or specimens from this area. Friedlander only “presumed hybridization” in this

area (page 245.)

It is strikingly odd that specimens of a single species should be so subspecifically related on both a

distant and proximate geographical basis. Add to this: the aspect that these reinthali populations along the

coast may be host specific to C. laevigata, and the Burke/Allendale county celtis populations may be host

specific with C. occidentalis var. georgiana; the morphological differences in larval characters; and

enough circumstantial evidence is accumulated to reasonably indicate speciation rather than subspeciation.

This would be considered weak evidence by some, perhaps many, but we are only postulating here. We are

simply giving probable cause to keep this research open until the subspeciation question can be absolutely

concluded one way or the other. It should be noted that Anthocharis midea (Hubner) and A. m. annickae

dos Passos & Klots function as distinct subspecies in the exact same area.

I propose that the taxon closest to Georgian A. celtis is the Gulf Coast endemic A. celtis alicia and

not A. c. reinthali. In noting the differences between alicia and the Floridian population, Friedlander not

only named the Florida population ( reinthali ) but sank alicia into celtis synonymy. In my opinion, this is

unfortunate. However, this is consistent with the rest of his revisional rationale (pg. 232). I consider alicia

a valid subspecies with the same basic Gulf Coast range as Basilarchia archippus watsoni dos Passos. At

first glance, alicia merely looks like a very large celtis. And indeed their markings are about the same, with

one important exception. I have only seen a few topotypical alicia, but even in this small series they show a

strong tendency to have an additional partial dorsal FW eyespot in cell M 3 . This character was not

mentioned by Friedlander.

The evolutionary significance of this tendency should not be minimized. This single feature in alicia

would seem to tie southeastern celtis together with the western A. c. antonia (W. H. Edwards) and its many

allies, which are double or partially double eyespotted entities. Alicia is positioned, both geographically

and phenotypically, as the “link” connecting western and eastern celtis. I agree with Friedlander, that A. c.

reinthali is a taxon ascended from the Florida refugium. However, I see celtis, by way of alicia, as

descended from the Mexican/Texas refugium. (Friedlander, in his prepublication review of this paper,

agrees that this hypothesis of the origin of celtis is plausible and warrants further investigation.)

The celtis - reinthali taxonomic question seems three fold. First, while the celtis and reinthali

ancestors were separated in different refugium, did they evolve into close but separate species which may

now be sympatric in some part of Georgia or South Carolina? Second, are celtis and reinthali just

subspecies in unusually close proximity simply because celtis, at the terminus of it evolutionary tract, has

now abutted a long disjunct relative? If so why are they not interbreeding? Third, are they subspecies that

due to evolutionary distance, say by host specificity, now cryptically act as species in this area? Until one

of these, or some other option, is proved correct by collecting and/or breeding experiments, reinthali and

celtis may correctly be thought of as separate species - the status they had before Friedlander’s revision.

The status Dr. Reinthal held to be correct. However, I am by no means embracing specific status. I am

saying that the exact relationship of these two is still in question.

The situation with southeastern A. clyton is not complex. Occasional specimens from Savannah,

Georgia I’ve seen in the collection of R. T. Arbogast tend to have features characteristic of both flora and

clyton. Burke County clyton seem to be solidly clyton. All south coastal South Carolina clyton I’ve seen

from Charleston, Colleton, Dorchester and Berkeley counties are true clyton. Thus, a blend zone between

clyton and flora can be observed in north coastal Georgia. Further, there is a dark female form of flora

(Fig. 8) that is very similar to the light form of female clyton (Fig. 5).

Neotypes are definitely called for here. Accordingly I have designated a male of A. celtis and A.

clyton each from Burke County, Georgia as neotypes. They have been deposited in the Carnegie Museum of

Natural History, Pittsburgh, PA. They are labeled as follows: NEOTYPE Apatura celtis, 6 August 1998, at

Savannah River, Burke County, Georgia (Figs. 2-4 top); NEOTYPE Apatura clyton, 6 August 1998, at

Savannah River, Burke County, Georgia (Fig 6 bottom)

WESTERN SUBSPECIATION

A. leilia codes (Lintner), A. celtis montis (W. H. Edwards), and A. clyton subpallida (Barnes &

McDonnough) were considered species by most writers in the 1970’s and 80’s. Today, via Friedlander,

they are not even considered valid subspecies. I have all the western taxa in my collection and have reared

subpallida. They seem to be perfectly good subspecies to me. I think this bears a second look. I would

hope some western researchers would revisit this situation. Bailowitz and Brock (1991) had the best

opportunity to reexamine montis and subpallida but chose to strictly follow Friedlander.

I am not sure if the criteria Friedlander used in defining his western subspecies (pg. 232) was

applied in the same way to his eastern subspecies. However, based on his review comments of this paper, I

think many may have read a finality into his Asterocampa revision that he absolutely did not intend.

He and I are certainly in agreement that, “very little is written in stone about Asterocampa His

lumping of western taxa was not an indication of a taxonomic simplicity or finality on his part. It was just

the opposite. It was (is) the complex relationships of the western populations, coupled with his

conservative approach, in the face of the unknown, that resulted in what he produced on paper. Let me relay

a little of what he has written to me:

The rest of the iceberg is those western populations I sunk under the name of A. celtis antonia.

There are dozens of distinctive, geographically isolated antonia- like A. celtis in the West, and I

don’t have a clue about their phylogeny. I can guess that A. celtis celtis arose from their ancestors,

but how are the antonia- like A. celtis related one to another? Some may be on the line to A. celtis

celtis, some may be persistent ancestors, some may be on entirely different lines of evolution. It will

become useful to have separate names for them only when we have sufficient data to do the

phylogeny work. For now we have no reason to believe that A. celtis antonia in New Mexico is a

single entity, nor do we know how A. c. antonia from Arizona, Colorado, Texas, Oklahoma,

Mexico, etc. relate to them - better to wait so the critical research can be done, and I think it will

have to be molecular. So, if you need a name, the A. celtis antonia population from “X” will do, but

be sure to voucher your material! In this sense, vouchering is as important as naming, maybe more

so, for it’s clear we need genetic material vouchered, whereas defining types is still largely

morphological.

I sunk subpallida because it is the Arizona version(s) of texana, both of which probably are not

single entities and require further investigation. I almost named subgenera for the two groupings, but

of what use? They are distinctive enough to have come over from Asia separately, then separated

into tropical/temperate pairs.

To sum up, I guess I’m emphatic that subspecific names not be raised or given until the critical

research is done to explain the phylogeny of the populations. I’m not debating the distinctiveness or

isolation of the western populations - they will all eventually need formal names, if they survive

extinction. But there are many, many more such populations than subpallida, montis, codes, and

the one A celtis I left unnamed in Mexico (and A. clyton louisa). But if it helps to have a name [to

provide protection to taxon], resurrect it now! I’d prefer to wait until the phylogeny is done right.

Two things are clear from Friedlander’s remarks. First, much more research needs to be done.

Second, the last word on Asterocampa subspeciation has not been spoken. My reason for writing this paper

is to stimulate more research on the Asterocampa. I feel some of the named taxa were sunk prematurely.

Why sink a subspecies we know will one day be validly resurrected just because it needs to be defined in a

more accurate way? If some of the demoted taxa are not truly synonymous with (the same thing as) the

subspecies they were placed under, then the only thing accomplished in sinking them was the replacement of

an inaccurate subspeciation with an inaccurate synonymy.

If Philotes sonorensis extinctis Mattoni is a valid subspecies in half a canyon, and Icaricia

icariodes missionensis (Hovanitz) on half a hill (with I. i. pheres (Boisduval) at the bottom of the hill),

then A. c. subpallada and A. c. montis can be valid in half a state (this sentence contains hyperbole but it

makes the point). As I stated, I hope someone from the area of these to-be or not-to-be Western

Asterocampa subspecies will further research the question. It seems Tim Friedlander feels the same, though

for different reasons.

ACKNOWLEDGMENTS

Thanks to Tim Friedlander for his review of this paper and input. To my

excellent photos.

LITERATURE CITED

son, Ben, for his usual

BAILOWITZ, R.A. & J.P. BROCK. 1991. Butterflies of Southeastern Arizona. Sonoran Arthropod Studies,

Inc., Tucson, AZ. 342 pp.

FRIEDLANDER, T.P. 1987. Taxonomy, Phylogeny, and Biogeography of Asterocampa Rober 1916

(Lepidoptera, Nymphalidae, Apaturinae). The Journal of Research on the Lepidoptera, Santa

Barbara, CA. 25:4, 119 pp.

HOWE, W.H. 1974. The Butterflies of North America. Doubleday & Co., Inc. New York, NY. 633 pp.

SCOTT, J.A. 1986. The Butterflies of North America, A Natural History and Field Guide. Stanford Univ.

Press, Stanford, CA. 583 pp.

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6

Volume 1

1 March 1999

Number 6

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

AN EVOLUTIONARY SUBSPECIFIC ASSESSMENT OF DECIDUPHAGUS HENRICI

(LYCAENIDAE) BASED ON ITS UTILIZATION OF ILEX AND NON-ILEX HOSTS:

DESCRIPTION OF A THIRD ILEX ASSOCIATED SUBSPECIES. DESIGNATION OF A

NEOTYPE AND TYPE LOCALITY FOR DECIDUPHAGUS IRUS.

RONALD R. GATRELLE 1

126 Wells Road, Goose Creek, South Carolina 29445

ABSTRACT. Deciduphagus henrici subsists as six subspecies which have evolved into two distinct larval host

associated groups of three taxa each. Group one is composed of those taxa which utilize primarily Ilex (holly - various

species) as their host. They inhabit the non-montane area of the southeastern United States and up the Atlantic coast to at

least southern New Jersey. They are: D. h. margaretae (TL Deland, Volusia Co., FL) occupying east-coastal to south central

Florida; D. h. viridissima (TL Nag’s Head, Dare Co., NC) occupying the upper Outer Banks area of NC north through the

Chesapeake and Delaware Bays to perhaps Rhode Island; and a new subspecies herein described as D. h. yahwehus (TL

Orangeburg Co., SC) occupying the area from south central North Carolina south through Georgia then westward to perhaps

Mississippi and the upper west coast of Florida. This Ilex host association is believed to be peri-Pleistocene and Floridian in

origin. Group two consists of two subspecies which use primarily non -Ilex plants, and one taxon which is oligophagous

(including Ilex). The primarily non -Ilex subspecies are: D. h. solatus (TL Blanco Co., TX) occupying south central Texas

westward; D. h. turneri (TL Cowley Co., KS) occupying the area from north Texas northeast to at least Indiana; and (the

oligophagous) D. h. henrici (TL Philadelphia, PA) occupying southeastern Canada south through the Appalachians. Nominate

D. h. henrici is the most recently evolved of the six henrici subspecies. In parts of the northeastern U.S. D. h. henrici is in

tension with D. h. viridissima, both phenotypically and biologically. D. henrici and D. irus were often confused. To stabilize

the taxonomy, the false holotype of D. irus is designated as neotype. Its type locality is restricted to southern NJ.

Additional key words: sibling species, original descriptions.

EARLY HISTORICAL AND BIOLOGICAL INFORMATION

John Abbot found two species of Deciduphagus Johnson (1992) in coastal Georgia. Abbot sent his

paintings and life history information regarding these taxa to Boisduval and Leconte who in 1833 published

Abbot’s renditions and information. Today, we know these species as Deciduphagus henrici (Grote &

Robinson, 1867) and Deciduphagus irus (Godart, 1824) (see appendix). However, at the time of Boisduval

and Leconte’s publication, henrici had not yet been described, and Godart had only recently described

Polyommatus irus from a single male specimen, of uncertain origin, which he had deposited in the Museum

Nationale, Paris (see appendix). Thus, upon the receipt of Abbot’s paintings and information, Boisduval

and Leconte had to determine which of Abbot’s depictions was Godart’s P. irus and which was an

undescribed species. After examining Godart’s type (which they said was so “defected by dilapidation, it

would not have been possible to recognize this type, if we had not seen it”), they determined one Abbot

species to be Godart’s irus (Figs. 1-2) and described the other as a new species, Thecla arsace Boisduval

and Leconte (Figs. 3-4). However, they associated the wrong Abbot species with Godart’s irus.

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

** M

MMMMMM

VfWWWMW

WXHMMM

19 20 21 22 23 24

MMWMMM

25 26 27 28 29 30

NNNMMM

Figs. 1-36. 1-2 (d/v), c? Thecla irus (=henrid) reproduced from Boisduval & Leconte 1833. 3-4 (d/v), d Theda arsace from Boisduval & Leconte

1833. 5 (d/v), false d holotype Polyommatus irus herein designated as neotype. 6 (v), d topotype D. irus, 15 May 1971, Lakehurst, Ocean Co., NJ, leg.

Stanford. 7-8 (d/v), d holotype D. h. yahwehus, 10 March 1994, Nr. Bull Swamp, Orangeburg Co., SC. 9-10 (d/v), 9 allotype D. h. yahwehus, same data

as 7. 11 (v), d D. h. yahwehus, 29 March 1986, Edisto Is., Colleton Co., SC. 12 (d/v), red HW d D. h. yahwehus, 19 March 1977, Ft. Stewart, Bryan

Co., GA. 13-14 (d/v), d topotype D. h. margaretae, 7 March 1976, Deland, Volusia Co., FL. 15-16 (d/v), 9 topotype D. h. margaretae, same data as 13.

17 (v), topotype d D. h. margaretae, 9 March 1975, Deland, Volusia Co., FI. 18 (v), dark topotype 9 D. h. margaretae, same data as 17. 19-20 (d/v), d

holotype D. h. viridissima, 3 April 1991, Bodie Is. lighthouse, Dare Co., NC, leg. Pavulaan. 21-22 (d/v), $ allotype D. h. viridissima, same data as 19. 23

(v), d int. morph D. h. viridissima, 15 April 1992, Sussex Co., DE, leg. Pavulaan. 24 (v), d brown morph D. h. viridissima, same data as 23. 25-26 (d/v),

d D. h. turned, 21 April 1979, Ernie Miller Pk., Johnson Co., KS, leg. unknown. 27 (d/v), d D. h. henrici, 10 April 1977, Lee Co., VA, leg. Hyatt. 28 (d/v),

d D. h. henrid, 29 April 1988, Great Swamp, Washington Co., RI, leg. Pavulaan. 29-30 (d/v), d topotype D. h. solatus, 24 February 1976, 8 Mi. E. of

Blanco, Blanco Co., TX, leg unknown. 31-32 (d/v), 9 D. h. turned, 18 March 1973, Tyler, Smith Co., TX, leg. unknown. 33 (d/v), 9 D. h. turned, 8 March

1987, Nr. Dow, Pittsburg Co., OK, leg. unknown. 34 (d/v), 9 D. h. turned, 6 May 1975, Zaleski St. For., Vinton Co., OH, leg. Parshall. 35-36 (d/v), 9

topotype D. h. solatus, same data as 29. Specimens leg. R. Gatrelle unless otherwise noted; (d) = dorsal, (v) = ventral.

Abbot provided paintings and basic information to Boisduval and Leconte. Boisduval and Leconte

made the determinations and wrote the descriptions. These authors saw no actual Abbot specimens. We

know now that the Abbot painting to which Boisduval and Leconte assigned the name irus was in fact not

Godart’s irus, but the species Grote and Robinson would describe as Thecla henrid 34 years later. We

know this because: 1) Abbot’s adult figures depict henrid (no male scent patch), not irus; 2) his

description of the larva fits henrici, not irus; 3) his larval hosts (Ilex L. and Vaccinium L.) are that of

henrid, not irus; and 4) the verbal description (see appendix) is in all aspects, except one, henrici and not

irus. Conversely, Boisduval and Leconte’s arsace is an irus in these same four points (see appendix).

Abbot’s male figure of what Boisduval and Leconte called arsace has scent patches on the forewings.

Boisduval and Leconte’s error of associating Godart’s name irus with the as yet undescribed

henrici, resulted in their describing the true coastal Georgian irus under the name T. arsace. Thus, Thecla

irus Boisduval and Leconte 1833, should be listed under the synonymy of Deciduphagus henrici and noted

as preoccupied by Polyommatus irus Godart, 1824.

To preclude the possibility that anyone might ever suggest that Godart’s irus was a henrici , I wrote

the Museum Nationale in 1975 and asked if Godart’s P. irus type was extant. Dr. P. Viette not only

confirmed that the holotype was there, but also sent me the specimen for taxonomic verification (Fig. 5). It

is a D. irus. (See the appendix under Polyommatus irus for additional information.)

Boisduval and Leconte are not the only ones to have confused D. henrici with D. irus. Scudder

(1889) lists both T. henrici and T. arsace in his synonymy of D. irus. He devotes seven pages of narrative

to what is actually an irus/henrici composite. There is a wealth of information, but it is untrustworthy

because he unknowingly goes back and forth between data relating to henrici and irus. However, for the

reader who knows that Scudder was actually dealing with two species, this information is enlightening.

Scudder’s comments under the heading “variations” manifests this confusion. He muses as to why

some male irus nearly lack DFW scent spots (being henrici, they actually had none) and how these same

males had noticeable HW tails “twice as long as usual [for irus].” He also notes that some of these same

tailed (irus = henrici) specimens from Albany, NY were “decidedly olivaceous” in hue. This reference to

decidedly olivaceous (greenish) specimens in upstate New York in 1889 is significant now that the often

greenish D. h. viridissima Pavulaan has been described (1998).

Scudder quotes Abbot as stating that irus (actually henrici not irus) is often found in swampy areas.

I too have found southeastern henrici in very wet areas. I have also found it curious that henrici in this area

have the trait of firmly attaching their pupa to holly leaves or bark. I have wondered if this wetland habitat

association might have caused these henrici to develop this trait - because the ground is often inundated

with water. Or, is this characteristic of other populations in dryer environs also?

Holly (Ilex) and Lupine (Lupinus perennis L.) are the only plants Scudder confirmed as definite

hosts for his “irus.” We know now that the former only applies to D. henrici and the latter to D. irus.

Scudder mentioned other plants, but stated he could not confirm any of them as being oviposited on by wild

females. Scudder noted that larvae (of which species?) accepted wild plum fruit in captivity. He also

documented that females (of which species?) refused to lay eggs when confined with Vaccinium

(blueberry), Quercus L. (oak), or Cyrilla racemifolia L.

His comments on Cercis canadensis L. (redbud) are noteworthy. He records that Abbot only noted

C. canadensis as an adult nectar source and not a larval aliment. I have often found local henrici adults

around redbud (never irus). But I have never succeeded in getting henrici larvae from this area to eat, or

females to oviposit on, C. canadensis. This plant is documented (in lit.) as a primary host of D. h. henrici

and D. h. turneri Clench, in the Appalachian, northeastern, and western areas of the United States.

This confusing of D. henrici and D. irus adults, coupled with the unclear references to plants as

either larval hosts or adult nectar sources by the early workers, should make the long list of henrici larval

hosts as recorded in much of the modem literature suspect (i.e. Scott 1986). A lot of field work needs to be

carried out throughout the range of D. henrici to accurately reassess and document its actual hosts.

3

BIOGEOGRAPHICAL EVOLUTION

AND SUBSPECIATION

The six subspecies of Deciduphagus henrici can be divided into two groups of three subspecies

each according to their known larval host associations. Three subspecies appear to be nearly monophagous

toward Ilex. I call this group the Ilex group. The other group of subspecies is composed of two taxa which

are primarily non-ilex feeders and one which is oligophagous (including Ilex). I call this the non-ilex group.

The Ilex group is found in the non-montane areas of the southern and eastern United States. The three

taxa which comprise the Ilex group are: D. h. margaretae dos Passos, type locality Deland, Volusia

County, Florida occupying east-coastal to south central Florida; D. h. viridissima, type locality Nag’s

Head, Dare County, North Carolina occupying the upper Outer Banks area of NC north through the

Chesapeake and Delaware Bays and in scattered coastal colonies to perhaps Rhode Island; and a new

subspecies herein described as D. h. yahwehus Gatrelle (TL Orangeburg County, South Carolina)

occupying the area from south central North Carolina south to north Florida then westward through Georgia

to perhaps Mississippi and the upper west coast of Florida.

The non -Ilex subspecies are: D. h. solatus Cook & Watson, type locality Blanco County, Texas

occupying south central Texas westward; D. h. turneri Clench, type locality Cowley County, Kansas

occupying the area from north Texas northeast to at least Indiana; and the oligophagous (including holly) D.

h. henrici, type locality Philadelphia, Pennsylvania and occupying the southeastern area of Canada south

through the Appalachians.

Ilex Group

Ilex is the primary, perhaps exclusive, larval substrate of this group. After nearly 30 years of

experience with D. henrici in Florida, coastal Georgia, and South Carolina I have only found it in

association with Ilex. Ilex opaca Aiton, I. cassine L., and I vomitoria Aiton are known larval aliments of

margaretae, yahwehus and viridissima. Where more than one of these hollies grow in the same location,

they are all utilized by the local henrici population. I. opaca and I. vomitoria are found from Virginia south

to Florida and west to Mississippi, with vomitoria primarily along the coast. I. cassine is found from south

coastal North Carolina south to Florida and west to Mississippi (includes var. myrigloia Aiton). This

mutual host association and shared morphological characters of prominent tails and limited red scaling

dorsally is consistent with a common ancestry of these three subspecies. This Ilex host association is

therefore evidently peri-Pleistocene and Floridian in origin. Thus, they have arisen from a Floridian

Pleistocene relict.

I. cassine prefers mesic, even paludal, habitats more than the other known hosts of this group. Thus,

I. cassine’s range is restricted to the wet areas of the lower coastal plain. I cassine is the primary host of

margaretae in east coastal Florida. I. opaca is tolerant of both mesic and xeric habitats. Thus, I opaca is

by far the most upland of the known larval hosts. I opaca is the primary host of the inland populations of

yahwehus. I. vomitoria is the primary host of yahwehus on the coastal islands of South Carolina. D. h.

viridissima utilizes I. opaca and I vomitoria in the same fashion as yahwehus does.

In my evolutionary model, this group of subspecies arose from an ancestor adapted to lowland Ilex

during the Pleistocene in island Florida with margaretae being the oldest most direct descendent. By this

model, the species then followed the host(s) and invaded low wetlands northeastward and westward from

Florida adapting to other hollies in the process. Those which followed I opaca’s adaptation to xeric and

upland environs moved inland with it and gave rise to yahwehus.

D. h. yahwehus is both phenotypically and geographically closest to D. h. margaretae. D. h.

viridissima is equally distinct in phenotype from both of these. It is not possible to determine, without an

involved study, if viridissima arose from margaretae or yahwehus. However, it is likely that yahwehus,

based on its upland host adaptation, is the most recently evolved; and viridissima, by its continued lowland

coastal acclimation, is older than yahwehus and younger than margaretae having simply moved northward

from Florida along the coast.

However, regardless of viridissima’s evolutionary ancestry, I disagree with Pavulaan’s idea that

viridissima’s green morph may have stemmed from environmental factors. If this were true, it would only

be an ecotype (form) and not a subspecies. Further, if this were the case, why are not all coastal North and

South Carolina populations predominantly greenish? All North and South Carolina maritime populations

feed mostly on I. vomitoria and are subject to the same basic ecological conditions. It is far more likely that

the great frequency of the green morph arose over thousands of years as the direct result of green gene

selection and its eventual genetic dominance within the isolated population on the upper Outer Banks of

North Carolina. I see \iridissima as a genetically distinct Ilex group subspecies, with green, brown, and

intermediate morphs, which now extends along the northeast coast from the upper Outer Banks of North

Carolina to perhaps the Great Swamp of Rhode Island.

Regardless of what evolutionary models are eventually demonstrated to be the most likely, these

three biologically similar Ilex associated henrici subspecies 1) form a group most probably evolved from a

Floridian ancestor, and as such are 2) separated by thousands of years from the non -Ilex associated taxa.

Non-7/ex Group

I am by no means as familiar with this group as I am with the Ilex group. However, it seems fairly

probable that the non-IIex group is 1) from a common ancestor, and 2) from a different refugium than the

Ilex group. The range of the three subspecies in this group, their host associations, and phenotypic

similarities indicate that their ancestor is from a Texan or Mexican population.

Cercis canadensis seems to be the primary larval host of D. h. turneri and D. h. henrici (Pavulaan,

1998) from Texas northeastward to southeastern Canada and eastward to the southern Appalachians. This

includes the northern glaciated and southern non-glaciated areas within this range. This dictates that C.

canadensis became the primary host of this group before it came to occupy the formerly glaciated regions.

Hypothetically then, there should be two biological situations in place in the Midwest, eastern US, and in

southeast Canada which need to be investigated through careful field observations.

Bio-situation A. The older (more biologically conformed) populations in the non-glaciated areas

would be expected to be monoalimentary in larval host selection. One plant genus, or species, would be the

dominant larval aliment (i.e. Cercis). However, occasional colonies within this large geographic primary

host area might utilize plants of one or two other genera (including Ilex) as secondary hosts; or even as a

primary host in certain micro populations within the non- Ilex group’s total range. D. h. turneri is the

expected taxon of bio-situation A (Figs. 25-26, 31-34).

Bio-situation B. The newer (more biologically unstable) populations in formerly glaciated areas

would be expected to be largely polyalimentary in larval host selection. They would equally utilize plants

of multiple genera as larval hosts. D. henrici’s recent adaptation to Rhamnus frangula (Layberry et al.,

1998), is indicative of this type of ongoing host adaptation by populations still in ecological and geographic

advance. Situation B populations should be endemic to Remington’s (1968) suture zone one. D. h. henrici is

the expected taxon of bio-situation B (Figs. 27-28). D. h. henrici was the first taxon of henrici to be

named; however, by my model, it is the most recently evolved and is the eastern extension of turneri.

Nevertheless, this does not mean that D. h. turneri is not also present in some situation B areas. I

agree with Pavulaan (1998) that D. h. turneri extends from Texas well into the eastern United States and is

probably the best name for henrici in much of Ontario, Canada. D. h. turneri was determined as the

subspecies in Missouri by Heitzman (1987), and in Indiana by Shull (1987). Specimens I have from Vinton,

Co., Ohio (Fig. 34) also fit best with turneri. It is the frequently extensive amount of red scaling on the

dorsal surface of turneri that differentiates it from henrici henrici. Ventrally, they are very similar. The

name henrici should be limited to the oligophagous, dorsally dark (compared to turneri) populations of the

northeast and Appalachians. Nominate henrici has more red dorsally than any of the Ilex group subspecies.

Diospyros texana (persimmon) is apparently the exclusive foodplant of D. h. solatus in Texas. This

host association and solatus ’ distinct phenotype (Figs. 29-30, 35-36), indicates that it is either from a third

ancestor (further to the west), or is an early subspecific split from the Texan ancestor of it and the other two

taxa in the non-Ilex group. A breeding experiment between solatus and viridissima would be of interest to

test if these two, by evolutionary distance, act as sibling species.

The Ilex and non-ilex groups already act as sibling species in some aspects stemming from their

host association. For example, Pavulaan was not able to get females of viridissima to oviposit on Cercis.

And his viridissima larvae also exhibited difficulty in accepting Cercis. The viridissima larvae he was

able to rear through to pupae on Cercis had extremely high pupal mortality. I have found this same condition

to exist in nature here in South Carolina. The possibility exists that the taxa in these groups do not just act in

some ways as sibling species, but are in fact such.

Subspecies In Tension

Mr. Harry Pavulaan and Dr. Dale Schweitzer have provided me with a great deal of information and

offered their personal opinions and interpretations regarding henrici/viridissima in the northeastern United

States. Dr. Schweitzer’s input was specifically offered relative to his review of this paper. These two

workers agree on the relevant biological and ecological data. However, they differ, sometimes

dramatically, in their interpretation of these data. The facts and advice provided by Schweitzer and

Pavulaan constitutes virtually all of the specific information in this section. Accordingly, they deserve

special acknowledgment here. The subspecific theory is mine.

Cryptic species look exactly alike, but the factors of reproductive isolation and biological diversity

distinguish them. Subspecifically, because there is no reproductive isolation, many butterfly taxonomists

have come to rely almost entirely on morphological wing differences to determine subspeciation. I believe

this is incorrect. As with species, major biological differences are of greater subspecific importance than

wing coloration.

Pavulaan, in his description of viridissima, documented many populations which contained various

percentages of green morph viridissima and/or intermediate viridissima like individuals meshed with

predominately brown populations (which he called henrici) ranging through the entire Chesapeake Bay area

and reaching as far west as West Virginia and northeast to New Jersey. According to both Schweitzer and

Pavulaan, all Chesapeake Bay populations are Ilex feeders be they green, brown, or intermediate. They also

agree that virtually all populations along the east coast from the Outer Banks through southern New Jersey

and then in scattered colonies north to the Great Swamp of Rhode Island, are holly feeders.

However, Pavulaan limited the range of his viridissima to only that area of the upper Outer Banks of

North Carolina containing the highest frequency of green individuals. I understand why he did this;

however, I feel his approach is too narrow and simplistic. In actuality, his delineation of viridissima nearly

renders viridissima as little more than a form. The situation in the northeast is about much more than green

verses non-green specimens or even populations.

According to Pavulaan’s original description, viridissima specimens vary in the degree to which the

subspecies’ three ventral green scaling characters are manifest - including brown topotypical individuals

with no green scaling. While the green characters are the most visible viridissima attribute, subspeciation is

primarily about evolutionary factors and not just how something appears to the human eye. Thus, Pavulaan

errs throughout his article by using the subspecific name viridissima in relation to reported “greenish”

individual specimens (forms) in the Midwestern populations of turneri and other areas. The term

viridissima is a subspecific name, not a tag for a variation or form.

He misuses the subspecific name henrici in the same way for the “brownish” form within the

topotypical viridissima population. The all-brown individuals at the type locality of viridissima are just as

much viridissima as the green ones. I embrace viridissima as being subspecifically distinct from henrici

6

because of their different evolutional ancestry, larval host associations, and morphological characters.

Viridissima also differs phenotypically from its nearest relatives - margaretae and yahwehus.

Mr. Pavulaan did not figure or describe the brown viridissima morph. The following is a brief

The relationship of the populations in the northeastern area of the United States is very complex

because two very distinct and evolutionarily distant subspecies are now in direct contact. One is D. h.

viridissima of the Floridian parented Ilex group, and the other is D. h. henrici of the western parented non-

Ilex only group. These two subspecies while in contact geographically, are separated evolutionally by

thousands of years. The morphological and biological characteristics of each of these subspecies, which

have taken tens of thousands of years to evolve, have doubtlessly been clashing in this tension zone 2 for a

long time. A great deal of detailed field work needs to be done in this tension zone area.

The correct understanding of viridissima , based on the information supplied to me by Schweitzer

and Pavulaan, seems to be that all of the monophagous Ilex populations along the east coast from about New

York City south to the upper coastal area of North Carolina should be considered viridissima. These

populations vary greatly in the percentage of green and brown morphs. However, this is the only henrici

subspecies to have any significant amount of green specimens. Thus, the green morph remains as the most

distinctive phenotypic character of this subspecies.

The limit of viridissima’s inland range, and where it meets and is in tension with henrici, is beyond

my knowledge and remains for others to work out. Dos Passos’ lectotype of henrici is from Philadelphia,

PA. According to Schweitzer, the shift to holly is east of the type locality in New Jersey. The fact that D. h

this situation. In the tension zone, non-ilex associated green henrici specimens may occur on occasion due

to the interbreeding of these two subspecies.

DESCRIPTION OF A THIRD ILEX ASSOCIATED SUBSPECIES.

I have been collecting butterflies extensively throughout the southeastern United States for 31 years.

Occasionally, I have come in contact with colonies of D. henrici in this region. It became apparent to me

about 1973 that two henrici subspecies were present in this area. In 1975 I submitted an article to the

Bulletin of the Allyn Museum describing the South Carolina population as a new subspecies. This paper

was accepted and an initial time frame was set for publication. However, several factors unrelated to the

research itself, resulted in that and several other projects, being put on hold - until now.

D. h. margaretae

Dos Passos described Incisalia henrici margaretae in 1943 from only four Florida specimens: two

males taken at Deland, Volusia County, in the central coastal region, and two females collected near

Aubumdale, Polk County, in the south central part of the state. The holotype was a male from Deland, so that

has become the type locality, though dos Passos did not specify it as such. His subspecies was based on four

characters as contrasted against nominate D. henrici. His description is as follows. I have highlighted these

characters in bold. In his comments section, dos Passos stated that the AMNH had no specimens from

Georgia or South Carolina. The AMNH did have two specimens from Southern Pines, North Carolina,

which he considered intermediate in dorsal color (more HW red) and length of tails.

“In Florida a very distinct race occurs, which, while having certain characters of henrici , is

easily differentiated by the length of its tails. These are more than twice as long as in typical henrici.

It is also somewhat larger, especially the females, and is uniformly dark grayish brown on the

upperside. The underside is a more uniform color because the basal area is lighter and the limbal

area darker than in henrici .”

From 1973 to 1988 I made several excursions to the type locality of D. h. margaretae on Hwy. 44

just east of Deland. The habitat is usually very wet, almost marshy, in the spring. Depending on the arrival

of warmer weather, topotypical margaretae could be on the wing as early as the end of January or be found

as late as the first of April. The topotypical population is very homogeneous in appearance.

During this time, I accumulated a large number of specimens of topotypical margaretae which was

usually abundant about its larval host, I. cassine. I. cassine grows profusely in this forested area of

Florida. D. h. margaretae fly high in the trees and only occasionally drop to within 10 feet of the ground,

which is usually to nectar at low I. cassine or Salix L. (willow) flowers. I employed a series of pole

extensions that enabled me to net specimens up to 30 feet high. Even with this method, the majority of

observed individuals were too high and could not be caught. Specimens could often be seen about the tops

of the highest pines. In this respect, collecting margaretae is very similar to collecting Mitoura hesseli. A

good idea of its abundance is illustrated by the fact that even with this general unobtainability, 50 specimens

could be captured in just 3 - 4 hours. This abundance was normal and did not represent a “population

explosion” as hairstreaks are occasionally known to have.

The only other Florida henrici I have personal experience with is an I. cassine associated

population I came across in March of 1988 near the Jet. of roads 337 and 326 in Levy County. I collected

17 specimens which all have the ventral basal area markedly darker than topotypical margaretae.

In addition to my own specimens, I have seen Florida henrici in the collections of Rick Gillmore,

Jeff Slotten, Dave Baggett, and the Florida State Collection of Arthropods, Gainesville.

There is one striking observation to be made about these Florida henrici - only specimens from the

central east coast to the south central area of Florida seem to possess all four of dos Passos characters and

are thus true D. h. margaretae. I consider the range of margaretae to be limited to this basic area. Exactly

how far inland this subspecies may be found is not known by me. I do know that specimens from Nassau and

Duval counties (Jacksonville) are atypical of margaretae and are probably best considered a blend zone

population near the new subspecies, or the new subspecies described herein.

A New Subspecies

To lepidopterists who are only familiar with henrici from outside the deep south, the first thing

noticed about specimens from central North Carolina south are their very long tails. (These tails may often

be six mm long on some Floridian males.) This striking character is undoubtedly why some popular

butterfly book authors have extended the range of margaretae well north of central Florida into southern

Georgia (Harris, 1972 & Scott, 1986), and South Carolina (Howe, 1975). However, there are three other

characters which typify margaretae, and these seem to have been largely ignored by those observing

southern specimens. Even many Florida lepidopterists are apparently only assessing margaretae by the

length of their tails.

A comparison of Southern and Floridian specimens against the four characters dos Passos listed as

being definitive of margaretae, reveals that two phenotypes are present in this region. One is margaretae,

from within the previously restricted range. The other extends from central North Carolina south through

Georgia to Jacksonville and apparently down the west coast of Florida to Levy County and (evidently) west

to Mississippi. Specimens from throughout this broad area are quite similar, are very different from

margaretae in two characters, and moderately different in one. The moderately different character is that

specimens from this area are noticeably larger (character noted by Pavulaan, 1998). They differ markedly in

that they have more red at the margin of the dorsal HW (as noted by dos Passos), and most noticeably, have

very contrasting ventral basal and limbal areas which are often partially delineated by a white line (as

pictured by Abbot). The tails are about the same, reducing as one goes northward.

I first encountered this undescribed subspecies in Givhans Ferry State Park, Dorchester County,

South Carolina in 1971. Since then I have collected it from the following locations. GEORGIA: Bryan

County, about 30 specimens nr. Fort Steward, 1976 (mid March); SOUTH CAROLINA: Aiken, Barnwell,

Berkeley, Charleston, Colleton, Dorchester, and Orangeburg counties, about 90 specimens accumulated

from 1971 to 1998 (mid February - late April); NORTH CAROLINA: Hoke County, 1 worn specimen near

Raeford, 1974 (early April).

I have also seen additional specimens from Bryan County, Georgia in the collection of R. T.

Arbogast, and a long series collected by the late R. B. Dominick at the Wedge Plantation, Charleston

County, South Carolina (now at the University of South Carolina).

Biologically, this new subspecies differs from margaretae in that it is the result of larval host

adaptation to Ilex opaca which, in turn, allowed the species to expand its range, both environmentally and

geographically, into dryer and upland environs. It is distinct morphologically, as noted above and in its

description as follows. It should be remembered that this new subspecies, as a part of the 7/ev-group, has

never been directly related to the nominotypical D. henrici, and as such, can have no clinal or blend zone

relation with henrici. At any place where these two may be found to be in contact (and thus producing

phenotypically intermediate offspring), these populations should technically be referred to as tension zone

populations, not blend zone populations.

Deciduphagus henriciyahwehus Gatrelle, new subspecies

Diagnosis. There is very little variation in margaretae (Figs. 13-18). The overall general appearance of both sexes

of margaretae is about the same. They are a warm gray brown dorsally. Females occasionally have a few red scales on the

DHW margin at the tails, males rarely do. The ventral HW basal and limbal browns of margaretae are much less

contrasting than in the other subspecies (especially in males). Specimens of margaretae are often encountered with a good

bit of rusty red-brown suffusion over the ventral forewings. The anterior portion of the submarginal line of basally pointed

chevrons that boarders the gray marginal area on margaretae’s ventral HW may also contain patches of rusty red. The tails

are quite long, as can bee seen from the figures. Yahwehus differs in that the ventral HW basal and limbal areas are much

darker and more contrasting. The basal area in yahwehus is blackish and the limbal area brown. As pointed out by

9

Pavulaan, fresh specimens of yahwehus sometimes have a purplish wine tint to the ventral surface. Dorsally, yahwehus is a

darker brown than margaretae and have browner, less checkered margins. Specimens of yahwehus often have a small patch

of red along the outer margin of the hindwings at the tails (both sexes). The tails are not as long as in margaretae, varying

from 2 to 4 mm throughout its range. Generally, these tails are always twice as long as in henrici. The average forewing

radius (from base of wing to apex, right FW) of the 49 specimens comprising the type series is 14 mm (same for both sexes)

- nearly all specimens are 13 to 15 mm.

Description. Male (Figs. 7-8): Head: face, eyes, palpi, and antennae slightly darker than margaretae. Thorax and

abdomen: dark blackish brown dorsally, dark gray ventrally, with legs and ventral thoracic hairs dark charcoal gray (in

margaretae, the ventral thoracic hairs are light gray and the legs are a markedly lighter gray.) Forewings: dorsally, uniform,

dark brown, slightly grayish, with restricted white checkering along the fringe of wing and usually limited to the apical

margin; ventrally, rusty brown with some specimens very lightly dusted with green scales, postmedian line prominent often

black inwardly and highlighted with white outwardly, with the fringe more checkered looking on this surface. Hindwings:

dorsally, same color as forewing, with about 30% of specimens having a small amount of red scaling along the outer margin

in cells Cui and Cu 2 in the area of the tail, average tail length 2.8 mm; ventrally, basal area dark blackish brown to black, often

edged with white except at the middle of wing, fringe not checkered, marginal bluish gray area outlined by a row of

occasionally prominent basally pointing black chevrons. Female (Figs. 9-10): Head: as male. Thorax and abdomen : ventral

slightly lighter gray than male. Forewings', dorsally, ground color as in male, may have a very small amount of red scaling in

postmedian area; ventrally, as in male. Hindwings: dorsally, color as in male except that about 75% of specimens have red

scaling along the outer margin in cells Cui and Cu 2 in the area of the tail, average tail length 2.5 mm; ventrally, as in male.

Types. Holotype d (Figs. 7-8): vicinity of Bull Swamp, Orangeburg County, South Carolina, 10 March 1994, coll.

Ronald R. Gatrelle. Allotype 9 (Figs. 9-10): vicinity of Bull Swamp, Orangeburg County, South Carolina, 10 March 1994,

coll. Ronald R. Gatrelle. Paratypes : 33d d, 14 9 9, all coll. R. R. Gatrelle unless otherwise noted: SOUTH CAROLINA:

AIKEN COUNTY: White Cedar bog north of Aiken State Park, 1 9 (worn), 21 April 1984; CHARLESTON COUNTY: Wedge

Plantation, Id, 1 April 1971; 1 9, 30 March 1968 (both leg. R. B. Dominick); COLLETON COUNTY: Edisto Island on /.

vomitoria, 1 d, 1 9 (worn), 4 April 1980; 4 d d, 29 March 1986; DORCHESTER COUNTY: Givhans Ferry State Park, 1 d,

ex pupa 10 February, 1 9, ex pupa 1 March, 3 9 9, 21 March 1976; on Hwy. 61 3 mi. east of Givhans, Id, 27 March 1986;

ORANGEBURG COUNTY: Hwy. 172 1 mi. east of Hwy. 672,1 d, 5 March, 1 d, 27 March 1992; Bull Swamp 2 mi. north of

North off Hwy. 178, 3dd, 29 9, 10 March 1994, Id, 4 March 1997; GEORGIA: BRYAN COUNTY: Hwy. 204 nr

Ogeechee River, Id, 19, 20 March 1976, 8d d, 19 March 1977; Hwy. 204 3 mi. north of Morgans Bridge, Id, 17 March

1980 (leg. R. T. Arbogast); 3.3 miles south of Ellabelle, 3dd, 17 March 1982 (leg. R. T. Arbogast); Pine Barrens Rd. Id,

2 9 9,29 February 1992 (leg. R. T. Arbogast); 5 d d, 29 9, 2 March 1992 (leg. R.T. Arbogast). The holotype, allotype, and 2

paratypes are deposited in the American Museum of Natural History (AMNH), New York, where the type series of D. h.

margaretae is located. 2 d and 2 9 paratypes are deposited in the Carnegie Museum (CMNH), Pittsburgh, where the bulk of

the paratypes of D. h. viridissima are located. The remaining type specimens are in the Museum Of The Hemispheres

(MOTH), Goose Creek, South Carolina.

Geoecological type locality. I. opaca groves in 50 mile radius of Bull Swamp, Orangeburg County, South

Carolina.

Etymology. YHWH (translated as Jehovah in English) is an ancient Hebrew name for God as Creator. This is the

mono-Deity of the world’s Jews, Moslems, and Christians. From this perspective, yahwehus is named for The Architect of

the natural world. Its common name can be “The Architect,” or “Architect’s Elfin.” Johnson established Deciduphagus as

masculine. The us ending is correspondingly masculine m yahwehus.

Remarks. The parameters of D. h. yahwehus ’ range are not known. According to Pavulaan, it evidentially extends

westward to Mississippi. The populations in the panhandle of Florida are yahwehus. In 1969, while living in Pensacola,

Florida, I observed (but did not net) what was probably yahwehus flying about some Ilex opaca trees near Cantonment in

Escambia County. When I moved a year later, the area was being considered for development as a park and zoo. That same

year, I also observed (but was unable to net) two tailed elfins flying about a large planted I. opaca in a cemetery just east of

Foley, Baldwin County, Alabama. How far down the west coast of Florida yahwehus extends is unknown. My specimens from

Levy County (except for tail length) are certainly closer to topotypes of yahwehus than topotypes of margaretae. Likewise,

specimens I have seen from Jacksonville, Florida seem to be closer to yahwehus than margaretae. Florida lepidopterists

need to correctly define the range of margaretae subject to dos Passos characters, specifically the lightly contrasting ventral

HW. Too much emphasis has been placed on tail length. The northern limit of yahwehus ’ range is probably north central

North Carolina. However, too much attention can be placed upon the length of tails of yahwehus also. All biological and

morphological characters must be considered together in accessing populations at the outskirts of its range. Yahwehus will

undoubtedly prove to be the primary subspecies throughout the non-montane southern and southeastern US.

10

Volume 1

APPENDIX

Number 6

Polyomr

The Original Descriptions

and Key Notations

' irus Godart, 1824. Here translated from the original French into English (Fig. 5).

dorsal of the wings of the male is ;

is more brown, with the extremity a fawn-red, forming on the primaries a large spot a little mellow by ii

i a general tint, and on the secondaries a spot (more small) situated fairly near the anal angle. Beyond that, th

by

is rare. It lives in Virginia;

aval & Leconte, 1833 (= henrici). Here translated

the original Fr

English

(Figs. 1-2).

“EEE

The cate

2-A

ACKNOWLEDGMENTS

Thanks to Harry Pavulaan for information and loan of specimens, inc. the viridissima types, Dr.

Dale Schweitzer for critical textual review and life history information, Dr. Kurt Johnson for technical

review, Ms Anne Latimore for translation of the original descriptions, and Ben Gatrelle for photographs.

LITERATURE CITED

BOISDUVAL, J.A., & J.E. LECONTE. 1833. Historic generate et iconographie des Lepidopteres et des

chenilles de l’Amerique Septentrionalis. Vol. 1. Paris 228 pp.

CLENCH, H.K. 1943. Two New Subspecies of Incisalia (Lepidoptera: Lycaenidae). Canadian Ent.

75(10): 182-185.

COOK, J.H., & F.E. WATSON. 1909. Incisalia (Lepidoptera) from Texas. Canadian Ent. 41(6): 181-182.

DOS PASSOS, C.F. 1943. Some New Subspecies of Incisalia from North America. (Lepidoptera,

Lycaenidae) American Museum Novit. No. 1230: 5 pp.

3-A

GOD ART, J.B. 1824. Encylopedie Methodique Paris. 9: 674 pp.

GROTE, A.R., & C.T. ROBINSON. 1867. Descriptions of American Lepidoptera, No. 2. Trans, of the

Amer. Ent. Soc. 1 (2): 171-192.

HARRIS, L., Jr. 1972. Butterflies of Georgia. Univ. of Okla. press, Norman OK. 326 pp.

HEITZMAN, J.R., & J.E. HEITZMAN. 1987. Butterflies and Moths of Missouri. Missouri Dept, of

Conserv., Jefferson City, MO. 385 pp.

HOWE, W.H. 1974. The Butterflies of North America. Doubleday & Co., Inc. New York, NY. 633 pp.

JOHNSON, K. 1992. The Palaearctic ‘elfin’ butterflies (Lycaenidae, Theclinae). Neue Ent. Naschr. 29:1-

141, ill.

LAYBERRY, ROSS A., PETER W. HALL & J. DONALD LAFONTAINE. 1998. The Butterflies of

Canada. Univ. of Toronto Press, Toronto, Canada 279 pp.

PAVULAAN, H. 1998. A New Subspecies of Incisalia henrici (Grote & Robinson) (Lepidoptera:

Lycaenidae) from the Outer Banks of North Carolina. Maryland Ent. Vol. 4 (2) 1-16.

RADFORD, A.E., H.E. AHLES & C.R. BELL. 1968. Manual of the Vascular Flora of the Carolinas. Univ.

of North Carolina Press, Chapel Hill. 1183 pp.

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

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

SCOTT, J.A. 1986. The Butterflies of North America, A Natural History and Field Guide. Stanford Univ.

Press, Stanford, CA. 583 pp.

SCUDDER, S.H. 1889. The Butterflies of the Eastern United States and Canada with special reference to

New England. Publ. by author. Cambridge, MA. Pages 834-841.

SHULL, E.M. 1987. The Butterflies of Indiana. Indiana Academy of Science, U. of Indiana Press,

Indianapolis, IN. 262 pp.

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that year’s volume on CD for permanent archiving and reproduction for personal use (i.e. a museum or university may make

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at $9 per issue post paid. Checks should be made payable to TILS, and mailed to: Scott D. Massey, Editor, 126 Wells Road,

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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.

TILS has established the Museum Of The Hemispheres (MOTH). The MOTH collection will be a collection of

collections. Each individual sponsor, upon their death or retirement, will have their personal collection housed in a

personalized cubical. Thus, their personal collection (specimens, storage setup, library, desk, etc.) will forever be preserved

intact and be available to researchers in this form. For information write to: Ronald R. Gatrelle, MOTH Curator, 126 Wells

Road, Goose Creek SC USA 29445.

4-A

Volume 1

15 April 1999

Number 7

The Taxonomic Report

OF THE INTERNATIONAL T.EPTDOPTERA SURVEY

A NEW SUBSPECIES OF BREPHIDIUM ISOPHTHALMA (LYCAENIDAE:

POLYOMMATINAE) FROM COASTAL SOUTH CAROLINA.

HARRY PAVULAAN

494 Fillmore Street, Herndon, Virginia 22070

AND

RONALD R. GATRELLE 1

126 Wells Road, Goose Creek, South Carolina 29445

ABSTRACT. Brephidium isophthalma pseudofea was described in 1873 from the Florida Keys. The ventral

ground color of the wings in topotypical pseudofea is a yellowish brown. The antennal clubs of pseudofea have prominent

orange red tips. The ventral surface of the abdomen of pseudofea is broadly white to very light gray and usually extends

halfway around the sides of the light to medium brown abdomen. Specimens of Brephidium isophthalma from coastal South

Carolina differ greatly from topotypical pseudofea. Accordingly, Brephidium isophthalma insularus is described as a new

subspecies from coastal South Carolina. The antennal clubs of insularus are only slightly tipped in red, with many specimens

having the antennae completely black. The light area on the ventral abdomen of insularus is medium to dark gray and does not

extend up the dark brown sides of the abdomen, giving it a very dark appearance. The ventral surface of the wings of insularus

are medium chocolate brown with prominent white markings. The spring brood of insularus is especially unique and was

given the name Carolina (Pavulaan, 1993). The northern and southern limits of insularus ’ range is undetermined. However,

we expect insularus to be the resident subspecies in at least South Carolina, Georgia, and northern Florida. Its common

name is Island Pigmy Blue. It is possible that insularus is a distinct species. It is likely that pseudofea is conspecific with B.

Additional key words: subtropical species, resident species.

SUBSPECIFIC STATUS OF THE EASTERN PIGMY BLUE

Lycaena pseudofea was described by Morrison in 1873 (without figures) from three specimens

collected at Key West, Florida. The Pigmy Blue is a local, but common, butterfly. However, its proper

taxonomic status is unsettled. It is in need of a definitive study to determine its true specific/subspecific

relationships. Scott (1986) treats pseudofea as a subspecies of Brephidium exilis (Boisduval). Calhoun

(1997) treats pseudofea as a subspecies of isophthalma Herrich-Schaffer, but states that it may be a

subspecies of exilis. Opler (1984) treats B. i. pseudofea and B. exilis as separate species. Our opinion is

that all these taxa are probably part of the same species - exilis.

One of us, Gatrelle, was a resident of San Diego, California from 1967 - 1969 and is very familiar

with exilis (Figs. 3-4) in southern California. Occasional specimens of California exilis (Fig. 5, collected

by David Hawks in Riverside, California) are similar to topotypical pseudofea. Morrison referred to the

ventral ground of pseudofea as being concolorous. However, some specimens of typical pseudofea from

the Florida Keys tend to have light suffusion at the base of the ventral wings. Specimens like these lend

credence to Scott’s taxonomic alignment. However, the environmental, biological, and morphological

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

differences of the eastern and western US taxa are so great that genetic analysis and/or breeding studies are

needed before a nomenclatural shift should be formally adopted.

We have not examined the genitalia of our new subspecies or pseudofea , but significant differences

in their antennal clubs (shape and color), adult size, and morphological characters indicates that they very

well may be different species. Thus, not only may the subspecific alignment of our new taxon eventually

need to be changed, its specific status may need reconsideration also. We have followed the most

conservative course and have placed our new taxon under isophthalma.

NORTHERN RESIDENCY AND RANGE OF THE PIGMY BLUE

Populations of Brephidium are common along Florida’s coasts. In southern Florida B. i. pseudofea

flies year round (Gerberg, 1989). Up to 1986, specimens of B. isophthalma ssp. from the northern area of

its range in coastal South Carolina had been collected only sporadically and only in the summer and fall

(Harris, 1972 & Gatrelle, 1985). These flight dates had led Gatrelle to assume that isophthalma, regardless

of its tiny size and weak flight, was only a nonresident summer migrant in south coastal South Carolina.

In March of 1986 Pavulaan found the Pigmy Blue fresh and common on Hunting Island, Beaufort

County, South Carolina. The presence of a large number of fresh specimens that early in the year is strong

evidence that isophthalma is a resident species in coastal South Carolina. This is further supported by log

notations (as follows) made by Gatrelle regarding the winter of 1985. These notations indicate that the

winter of 1985 was earlier and colder than normal. We also see that the spring of 1986 was only slightly

early and was otherwise normal.

Entry of December 20, 1985: “Dec. has been very cold - freezing most nights.” This is unusually

cold for Charleston at that time of year. Spring appears to have arrived about two weeks early as the entry

on February 7, 1986 reads: “has been warm for about a week 60 - 70.” The entry on February 23 reads:

“weather has been very warm for about a week 70’s and 80’s.” It was also noted that temperatures dipped

to freezing the nights of March 20 and 21, then warmed up again. It is typical in Charleston to have extended

periods of warm weather from mid-February to mid-March, then one last freeze in March before spring

completely sets in. (The freezes of March 20 and 21 are especially significant as Pavulaan found specimens

of the Pigmy Blue common and fresh just six days later on March 27 th .) Prolonged freezing temperatures in

South Carolina are usually threatening to most subtropical wildlife like the Pigmy Blue.

w nf w

1 2 3 4 5 6

Figures 1-12, Brephidium subspecies. Fig. 1 (d/v), d B. i. pseudofea, 15 May 1978, Big Pine Key, Monroe Co., FI. Fig. 2 (d/v),

9 B. i. pseudofea, same data as 1. Fig. 3 (d/v), dB. exilis, 11 May 1968, Balboa Park, San Diego Co., CA (leg. Gatrelle). Fig. 4

(d/v), 9 B. exilis, same data as 3. Fig. 5 (v), d B. exilis, 25 June 1975, Santa Ana River, Riverside Co., CA (leg. Hawks). Fig. 6

(v), d B. i. insularus, 9 June 1988, Cedar Key, Levy Co., FI (leg. Gatrelle). Fig. 7 (v/d), d holotype B. i. insularus, 3 Oct. 1993,

Rt. 21 Nr. Beaufort, Beaufort Co., SC. Fig. 8 (d/v), 9 allotype, B. i. insularus, 27 March 1986, Rt. 21 Nr. Beaufort, Beaufort Co.,

SC. Fig. 9 (v), d paratype B. i. insularus, same data as 8. Fig. 10 (v), d paratype B. i. insularus, same data as 7. Fig. 11 (v), light

9 paratype, B. i. insularus, 24 Sept. 1972, Hunting Island, Beaufort Co., SC (leg. Gatrelle). Fig. 12 (v), 9 B. i. insularus, same data

as 6. All specimens leg. Pavulaan except where otherwise noted, (d) = dorsal, (v) = ventral.

Snow, which would help insulate delicate wildlife on the ground, is very unusual in coastal South

Carolina. (The most recent snow was ten years ago in 1989.) The climate, flora, and fauna of south coastal

South Carolina and Pensacola, Florida is about the same. The authors thus state with certainty that B.

isophthalma is confirmed to be resident along the immediate coast of South Carolina at least as far north as

Fort Johnson in the Charleston harbor and probably up to at least south Myrtle Beach.

B. isophthalma has only been found around salt and brackish marshes on the Islands and immediate

coastal mainland in coastal South Carolina. Several other butterfly species inhabit this narrow coastal area

which are also primarily subtropical in their range and habitat associations. Papilio Palamedes (Drury),

Phoebis sennae eubule (Linnaeus) (TL Charleston), Eurema daira (Godart), Heliconius charitonius

tuckeri Comstock & Brown, Agraulis vanillae nigrior (Riley), Phyciodes phaon (W.H. Edwards), Danaus

gillipus (Cramer), Heriargus ceranus antibubastus (Hubner), Leptotes cassius theonus (Lucas), Anartia j.

guantanamo Munroe, Calpodes ethilus (Stoll), and Urbanus dorantes (Stoll) are common residents or

usual summer transients on these coastal islands.

RECOGNITION AND DESCRIPTION OF A NEW SUBSPECIES

The Brephidium species/subspecies seem to have been largely ignored taxonomically. Perhaps this

is because of their small size. We have placed them under magnification and determined that they deserve

much more study. Pavulaan (1993) was the first to note the phenotypic differences between the island

population in South Carolina and typical pseudofea from the Florida keys. The spring brood from South

Carolina is especially distinct and he gave these the form name Carolina. Further comparison and

consideration has now led the authors to conclude that two subspecies, if not two species, exist in the

southeastern US.

B. i. pseudofea was described from the Florida Keys (Figs. 1-2). We have examined 6c? and 39

topotypes collected by Richard Anderson from Key West, and 6 specimens from Big Pine Key collected by

Pavulaan. The ventral ground color of the wings in pseudofea is decidedly yellow brown. Pseudofea also

exhibits lighter basal shading ventrally and a diminishing of the ventral forewing spots toward the outer

margin. The short spoon shaped antennal clubs of pseudofea have prominent orange red tips (Morrison

mentioned this as a major diagnostic character in his original description). The ventral surface of the

abdomen of pseudofea is broadly white to very light gray and usually extends halfway around the sides of

the light to medium brown abdomen. We have found the Brephidium from coastal South Carolina (Figs. 7-

11) to differ significantly from topotypical pseudofea. Accordingly, Brephidium isophthalma insularus

Pavulaan and Gatrelle is herein described as a new subspecies from coastal South Carolina.

We do not know how far south the new taxon ranges, but specimens we have from Levy County in

northwestern Florida (Figs. 6 & 12) seem to be good insularus. They differ from insularus only in tending

to have the ventral white FW spotting more faded toward the outer margins. It is very possible that the name

pseudofea properly applies to only the populations in the Florida keys or extreme southern Florida.

Populations in the remainder of Florida need to be studied to determine their proper taxonomic status.

Lepidopterists need to check their series of Florida Pigmy Blues for sympatric populations.

Brephidium isophthalma insularus Pavulaan & Gatrelle, new subspecies

Diagnosis and Description. The male (Fig. 7) and female (Fig. 8) of insularus are marked alike. B. i. pseudofea

and B. i. insularus are similar dorsally except that insularus averages a darker shade of reddish brown. Ventrally, they are

quite distinct. The ventral ground color of pseudofea is distinctly yellowish brown and occasionally has the basal third of the

wings lightly washed with white. In insularus, the ground color is a rich concolorous medium or dark brown in all broods

with the basal area always uniformly brown. The white lines which give a spotted appearance to this species are usually faint,

and sometimes absent, toward the outer margin of the forewings of pseudofea; while in insularus they are usually present all

the way to the forewing margin. This white marginal spotting is especially strong in the insularus spring form Carolina. In

the early brood(s) of insularus there is also a considerable amount of whitish clouding in the submarginal band of the ventral

HW. The light band along the outer margin of the HW of insularus is more orangeish and is more extensive than in

pseudofea. The antennal clubs of insularus are elongate and completely black (most specimens) or only slightly tipped in

3

orange-red. In pseudofea the antennae are shorter and spoon shaped with prominent orange-yellow tips (Fig. 1). The light

area running the length of the ventral abdomen of insularus is usually medium gray, narrow, and does not extend up the side

of the dark brown abdomen, giving it a very dark appearance. We describe the South Carolina population as a new subspecies

based on these characters. The geoecological type locality is: coastal tidal flats in Beaufort County, South Carolina.

Types. Holotype d* (Fig. 7): Rt. 21 near Beaufort, Beaufort County, South Carolina, 3 October 1993, leg. H.

Pavulaan. Allotype 9 (Fig. 8): Rt. 21 near Beaufort, Beaufort County, South Carolina, 27 March 1986, leg. H. Pavulaan.

Paratypes : 12 <j c?, 5 9 9: all SOUTH CAROLINA: BEAUFORT COUNTY: Rt. 21 nr. Beaufort, 8 3 <J, 2 9 9, 27 March 1986,

leg Pavulaan; 2d'd', 1?, 3 October 1993, leg. Pavulaan; Hunting Island, 1 d, 1?, 24 September 1972, leg. Gatrelle;

COLLETON COUNTY: Edisto Island, Id", 19, 9 April 1988, leg. Gatrelle. The holotype and allotype are deposited in the

Carnegie MNH, Pittsburgh, PA. Paratypes are in the MOTH, Goose Creek, SC, and Harry Pavulaan, Herndon, Virginia.

Etymology. The name insularus means, of the island. Its common name is Island Pigmy Blue.

Remarks. The striking differences in the antennae of insularus and pseudofea necessitates that their specific status

eventually be reevaluated. In pseudofea the l A to V 3 orange tipped clubs are short stemmed and spoon shaped. In insularus

the dark clubs are elongate, narrow, and tapering toward the shaft. These differences can be seen on the CD figures when

“zoomed.” Glassberg’s (1999) figure 1 on plate 22 from Black River, Savannah, GA is of insularus not pseudofea. Pavulaan

has examined 12 specimens of insularus in the Robert Gardner collection taken 24 March 1994 on Tybee Island, Chatham

Co., GA. Topotypes of pseudofea collected in the winter months (December - February) tend to be less yellow brown than

those from the summer broods but are still noticeably lighter than insularus. Specimens figured are about twice their natural

size but are proportionate to each other. All TILS photos are taken outside in natural light which often brings out different

hues than one sees in the same specimens viewed indoors in artificial lighting.

ACKNOWLEDGMENTS

Thanks to: Dr. John Rawlins and the Carnegie Museum for the copy of the OD of L. pseudofea ;

Richard Anderson, the loan of topotypical Key West specimens; Ben Gatrelle, photographs.

LITERATURE CITED

CALHOUN, J.V. 1997. Updated List of the Butterflies and Skippers of Florida (Lepidoptera: Papilionoidea

and Hesperioidea), Holo. Lepid. 4(2): 39-50.

GATRELLE, R.R. 1985. The Papilionoidea of the South Coastal Area of South Carolina. Bull. Southern

Lepid. Soc. Bull. No. 2. 15 pp.

GERBERG, EJ. & H.R. ARNETT, JR. 1989. Florida Butterflies. Natural Science Publications, Inc.,

Baltimore, MD. 90 pp.

GLASSBERG, J. 1999. Butterflies Through Binoculars - The East: A Field Guide to the Butterflies of

Eastern North America. Oxford: Oxford Univ. Press. 242 pp. + 71 color plates.

HARRIS, L., Jr. 1972. Butterflies of Georgia. Univ. of Okla. press, Norman OK. 326 pp.

MORRISON, H.K. 1873. Notes on North American Lepidoptera. Bull. Buffalo Soc. Nat. Sci., 1: 186-187.

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.

PAVULAAN, H. 1993. Some Observations on South Carolina Butterflies and Description of a New Form

of Brephidium isophthalmapseudofea. S. Lep. News Vol. 15 (4): 34-39.

SCOTT, J.A. 1986. The Butterflies of North America, A Natural History and Field Guide. Stanford Univ.

Press, Stanford, CA. 583 pp.

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

The Taxonomic Report is published at the rate of about 10 - 12 issues a year. Subscription for vol. 1 is $65 US. The subscription year

begins in August. All issues are mailed 1 st class. At the end of each year, subscribers receive that year’s volume on CD for permanent archiving

and reproduction for personal use. Checks should be made payable to TILS, and mailed to: Scott D. Massey, Editor, 126 Wells Road, Goose

Creek SC USA 29445. Articles for publication are sought.

Volume 1

15 July 1999

Number 8

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

HUBNER’S HELICTA: THE FORGOTTEN NEONYMPHA.

THE RECOGNITION AND ELEVATION OF NEONYMPHA HELICTA

(NYMPHALIDAE: SATYRINAE) TO SPECIFIC STATUS.

THE DESIGNATION OF NEOTYPES FOR N. HELICTA AND N. AREOLATUS.

THE SUBSPECIFIC TRANSFER OF SEPTENTRIONALIS TO HELICTA AND THE

DESCRIPTION OF A THIRD HELICTA SUBSPECIES FROM SOUTH FLORIDA.

RONALD R. GATRELLE 12

126 Wells Road, Goose Creek, South Carolina 29445

the AME Septentrionalis Davis, 1924 is recognized as the northern US subspecies of helicta not areolatus. An apparently

HUBNER’S HELICTA

In 1806 Hubner established the nami

a southeastern US butterfly in Sammlung e.

description. However, by 1816 Hubner ha

Neonympha areolatus (J.E. Smith, 1797), stating under number <

Schmettlinge : “Neonympha Helicta. Areolatus Abbot. Lepid. 13 Hut

In dos Passos’ 1964 checklist, Hubner’s helicta is listed i

areolata and N. areolata septentrionalis Davis, 1924. In their 1981

checklist, Miller and Brown list helicta in the synonymy of areolai

might be more correctly associated with N. areolatus septi

inting of

written

in his 1816 Verzeichness bekannter

ti. Oreas fimb. Helicta

l the synonymy of both N. areolata

evision of the Lepidopterists’ Society

us. However, they noted that helicta

is. Miller and Brown also suggested

y Museum, Vienna, Austria - or lost. r

own printed labels, it took years to find out that some of the material in Mazzola’s collection is in fact original HUbner

material. However, although the Mazzola collection is in Vienna, there is only European material involved. There are

no possible Hubner types of American butterflies existing. I got this inf ormation from Dr. Sabina Gaal,

Naturhistorissches Museum Wien (NHMW).” (Emphasis mine.)

This situation calls for the designation and deposition of an appropriate neotype. Accordingly, I

have designated a male specimen (Fig. 4) taken 1 June 1990 in the vicinity of Aiken State Park, Aiken

County, South Carolina as neotype of Oreas fimbriata helicta Hubner, 1806. It has been appropriately

labeled and deposited in the Allyn Museum of Entomology (AME), Sarasota, Florida. This specimen

becomes the type for Neonympha helicta. I restrict the geoecological type locality of Oreas fimbriata

helicta to the upland sandhill habitats of Aiken County, South Carolina and Burke County, Georgia.

DAVIS’ SEPTENTRIONALIS , AN HELICTA

In Davis’ 1924 description of septentrionalis he briefly mentions helicta on page 106 and figures a

specimen from Southern Pines, Moore County, NC which he correctly determined as helicta. However, he

regarded this specimen as “a rather uncommon variation” since it was the only specimen of this phenotype

he had seen. The specimen has its VHW eyespots rounded and the two VHW central brown lines rather

close together, which indeed correlates well to the original figures of helicta (Fig. 2).

However, it should have been obvious to Davis that his New Jersey septentrionalis specimens

were also close to Hubner’s figures of helicta. They were certainly much closer to Hubner’s plate than to

any areolatus Davis had at his disposal. Davis expected septentrionalis to be variable and made the

following statement at the end of his paper: “The writer does not mean to imply that specimens of areolatus

from New Jersey and vicinity may not occasionally show spots resembling those from Florida and Georgia.

He simply wishes to point out that there is a rather constant difference between those from the north and

south.” Helicta certainly falls well within Davis’ expected degree of variation.

Figs. 1-12. Neonympha subspecies (ventral surfaces). 1, Neotype d Papilio areolata, 2 Sept. 1989, Pine Barrens Road, Chatham

Co., GA. 2, d O.f. helicta from Hubner, 1806. 3, 9 (9. / helicta from Hubner, 1806. 4, Neotype d Oreas fimbriata helicta, 1 June

1990, nr. Aiken State Park, Aiken Co., SC. 5, topotype 9 N. h. helicta, 8 June 1980, nr. Aiken State Park, Aiken Co., SC. 6, Holotype

d Neonympha helicta dadeensis, 22 Nov. 1989, Carde Sound Road, Dade Co., FL (leg. Koehn). 7, 9 N. areolatus, 13 May 1973, Nr.

Jet. 1-26/17-A, Berkeley Co., SC. 8, <j N. areolatus, 27 March 1989, Collier Co., FI (leg. Koehn). 9, d N. h. helicta, 23 June 1968,

Foley, Baldwin Co., AL. 10, topotype d N. h. septentrionalis, 29 June 1970, Lakehurst, Ocean Co., NJ (leg. ?). 11, 9 N. h.

septentrionalis, 2 July 1989, Lebanon St. For., Burlington Co., NJ (leg. ?). 12, Allotype 9 Neonympha helicta dadeensis, 24 Nov.

1989, Carde Sound Road, Dade Co., FI (leg. Koehn). All specimens leg. R.R. Gatrelle unless otherwise noted. All figs, natural size.

Given the very limited understanding of the inland fauna of the Carolinas and Georgia in Davis’

time, it is perhaps understandable why Davis gave his northern specimens a new name. However, given the

closeness of his New Jersey specimens to the original Hubner figures, one is left to wonder why he made

absolutely no connection between helicta and his new taxon, septentrionalis.

Because helicta and areolatus were both described from specimens collected by John Abbot in the

southeastern US (and thus assumed to be consubspecific), and because septentrionalis was described from

the northeastern US, it is understandable, but unfortunate, that taxonomists have mostly associated the name

helicta with areolatus and given little consideration to the idea that helicta and septentrionalis were the

close relatives. Two things should have led taxonomists to conclude that septentrionalis and helicta were

the “same thing.” First, the Abbot depiction of the species named helicta by Hubner is a more accurate

depiction of septentrionalis than areolatus (especially in the male). Second, workers should have placed

more confidence in Abbot’s skills as a naturalist.

Abbot was a keen observer who knew what he was doing. While we today have had difficulty, in

some cases, figuring out what Abbot had, he surely knew that his helicta and areolatus were two species -

just as he knew that his Chlosyne gorgone (Hubner, 1810) and C. ismeria (Boisduval & Leconte, 1833)

were two species (Gatrelle, 1998); and that his "irus” in Boisduval & Leconte, 1833 (= Deciduphagus

henrici (Grote & Robinson, 1867)) andD. arsace (Boisduval & Leconte, 1833) (= D. irus (Godart, 1824))

were two species (Gatrelle, 1999); and his two Azure Blues {TTR 1:9, in press).

Davis’ lone specimen of helicta came from the south central North Carolina sandhills. Today, we

know that the Sandhills region from North Carolina south through Georgia and west through Mississippi is

the stronghold of Hubner’s helicta. Further, by their shared wing patterns and genitalia, we now know that

septentrionalis and helicta are two subspecies of the same species - helicta. We also know now that

differences in genitalia, wing pattern, habitat association, and flight characteristics demonstrate that

areolatus and helicta are distinct species.

Assimilating helicta and septentrionalis

While living in Pensacola, Florida I collected four septentrionalis- like specimens on 23 June 1968

east of Foley, Alabama along the Gulf coast. I loaned three of these to the Allyn Museum for comment in the

late 1970’s, but they subsequently lost the specimens. Fortunately, I still have one worn male (Fig. 9).

In 1976 I found a Neonympha phenotype that was widely distributed in the upland sandhills around

Aiken State Park in Aiken County, South Carolina. I have visited this area sporadically over the last twenty

three years to collect and observe this species. In wing pattern and genitalia these Neonympha are very

close to topotypes of septentrionalis I have from New Jersey. The only meaningful taxonomic difference

between the New Jersey and South Carolina populations is that those from New Jersey are much darker in

their ventral ground color and the orange brown lines on the ventral HW are usually more brown than

orange. (In both populations the two ventral forewing bands are nearly always brown.) When I first

encountered this population, I thought they were just a very disjunct, lightly colored colony of what I then

understood to be N. areolatus septentrionalis.

Later in the 1970’s while visiting Irvin Finkelstein at his home in Atlanta, Georgia, I learned that he

had taken a few septentrionalis -like specimens in central Georgia. I remember how struck I was with one

specimen in particular because it so closely matched New Jersey septentrionalis. In the 1980’s I became

aware of a population of “ areolatus ” south of Miami, Florida with large rounded eyespots (Fig 13). Then

in the early 1990’s I became aware of septentrionalis- like specimens from Mississippi. I have now seen

specimens in series from both of these areas.

By the early 1980’s I had concluded that the southeastern populations were at least subspecifically

distinct from both New Jersey septentrionalis and typical areolatus, and deserved recognition as such. At

this point the Miller and Brown notation regarding helicta became a central issue to this research.

3

Unfortunately at that same time, certain personal matters necessitated that all of my taxonomic

research be placed on hold. Well over a decade passed with little collecting, and less research, being done.

However, in June of 1990 I did have the time to collect and dissect several fresh males of these taxa. The

differences in the genitalia of Berkeley County, SC areolatus and topotypical Aiken County, SC helicta

were found to be significant and consistent (see below). The differences in genitalia confirm that the two

distinct phenotypes which occur throughout the southern and eastern US are two sibling species.

Early in 1998 I again picked up my research on these taxa. In late December of 1998, Dr. Gerhard

Tarmann of Innsbruck, Austria supplied me with the long sought after copy of the color plate of Hubner’s

helicta and the above quoted statement regarding the absence of a type for that taxon. This original helicta

plate clearly portrays what we have come to know, in the broad sense, as the septentrionalis phenotype and

not that of areolatus.

The occurrence of colonies of septentrionalis- like specimens from New Jersey south through

Georgia to Mississippi, and disjunctly in extreme south Florida, certainly demonstrates that this taxon is

what John Abbot had based his helicta paintings upon. (Note the exactly similar shape of the V median

bands in Figs. 2 & 4 and 3 & 12.) As the senior name, helicta has priority over septentrionalis and

becomes the proper specific name for all allied populations that comprise Neonympha helicta.

All of the populations of helicta south and west from North Carolina are of the same basic

phenotype (except in the Miami, Florida area). They differ significantly from N. h. septentrionalis in the

New Jersey vicinity only in having the ventral ground color a much lighter brown; and secondarily in a

tendency to have the VHW eyespots larger (more elongate) and the lines on the VHW more orange than the

New Jersey subspecies. The yellow rings around the eyespots are also bolder in helicta helicta.

NEONYMPHA AREOLATUS AND NEONYMPHA HELICTA

Separating helicta and areolatus

Helicta (all subspecies) and areolatus remain consistently distinct in size, wing pattern, and

overall habitat preference throughout their respective ranges. These differences, coupled with their

differences in genitalia, lend strongly toward defining these taxa as distinct species. The following will

serve as a basic guide to help lepidopterists properly distinguish and separate these two species. However,

it should be noted that some phenotypically extreme N. helicta helicta individuals can only be positively

separated by their genitalia. Also, the septentrionalis figured by Howe (1975) is areolatus- like in the

shape of its spots.

Flight pattern. Dr. Richard Arbogast was the first to make the observation that the flight patterns of

areolatus and helicta differed. Dr. Arbogast, as a long time resident of Savannah, Georgia, has collected/observed

hundreds of areolatus about the marshes and swamps in the Savannah area over the years. After accompanying me

on a collecting trip to Aiken County, he pointed out that the Aiken County helicta flew higher, faster, and straighter

than areolatus. Being familiar with areolatus in coastal South Carolina, I acknowledged that this was indeed true.

Areolatus flies either just above the sedges and grasses or down in them. It also has a rather slow, but darting

flight pattern. Female areolatus are reclusive and often have to be stirred up to be found. Conversely, helicta often fly

up to three feet above the grass. Their flight is swift for a satyr, and tends to be much less darting. Female helicta are

encountered about as often as males.

Habitat. I have found areolatus only around wet marshy (at least soggy) areas in South Carolina, North

Carolina, Georgia, Florida, and Alabama. Ricky Patterson has informed me that this is also his observation for

areolatus in Mississippi. Leroy Koehn has written that this is the situation in south Florida also. I have found helicta

only in open, dry, upland sandhill in South Carolina. My Alabama helicta were taken in fairly dry, open pine flat

woods. Ricky Patterson has also usually found helicta in dryer upland habitats in Mississippi. Whereas areolatus

seems to be limited to wet areas, helicta does not seem to be limited to dry areas. In New Jersey, in particular, helicta

“''SKEW

Mather (1965) <

)r of the two bands on ft

t. All of the VHW lines are m

As there is no type for Papilio areolata, I have designated a male I collected 2 September 1989 at

Pine Barrens Road, Chatham County, Georgia as neotype of Papilio areolata J.E. Smith, 1797 (Fig. 1). It

has been appropriately labeled as neotype and deposited in the Allyn Museum of Entomology (AME),

Sarasota, Florida. This specimen becomes the type for Neonympha areolatus. I restrict the geoecological

type locality of Papilio areolata to the marshy sedge forests of coastal Georgia.

A NEW HELICTA SUBSPECIES FROM DADE COUNTY, FLORIDA.

There are two populations of Neonympha in southern Florida of uncertain taxonomic standing. Both

of these populations have the anal angle of the HW angulate to the degree of tending to be lobed. They also

tend to have the anal margin of the VHW moderately to heavily covered with whitish scales (especially the

southwest FL areolatus segregate). The lobed HWs of the these Neonympha are reminiscent of tropical

Satyrinae in the genera Cissia Doubleday, 1848 and Cyllopsis Felder, 1869.

The helicta isolate in the southeast comer of Florida south of Miami is mentioned by Scott (1986).

He refers to this “strangely” disjunct population in his discussion of the range of septentrionalis on page

238. This population is phenotypically distinct from neighboring populations of typical N. areolatus just to

the north, but less so from the atypical areolatus west of it. In its morphological features, it appears to be a

subspecies of helicta with large “blind” eyespots (Fig. 13) reminiscent of Enodia portlandia floralae (J.R.

Heitzman & dos Passos, 1974). Its ventral ground color is much darker than in helicta helicta. Occasional

specimens vary toward areolatus - YHW orange bands meeting at (Fig. 13).

The variation in this population leads me to wonder if it is either a relict link between, or actual

ancestor of, both areolatus and helicta. Further, in its distinction from, or connection with, both helicta and

areolatus it may not have followed the same evolutionary path as the populations that moved north. This

helicta population solicits a great deal of evolutionary study. It is entirely possible that while helicta and

areolatus have clearly evolved into distinct species in the non-Floridian part of their range, they may not

have evolved far beyond a subspecific relationship in extreme south Florida. If this is the case, it will be

problematic to those taxonomists who approach their craft with fossilized rigidity rather than living fluidity.

The N. areolatus populations of southwest Florida may also represent a weak, but evolutionarily

definable, subspecies (Fig. 8). They differ from topotypical areolatus primarily by having the anal margin

of the VHW strongly and broadly overlaid with whitish scaling; and are helicta- like in that the bands of the

VHW tend to meet at vein Rs and notMi. The light anal margins are a striking trait. Because these areolatus

are not isolated from the other areolatus in the rest of the Florida peninsula, their possible subspecific

status will need to be evaluated against the examination of a large sample of specimens from throughout the

state. I do not see a problem in the identifiability of the southwest Florida areolatus. The problem is in

determining the boundaries of this likely clinal entity. I hope someone will undertake this in the near future.

Like Papilio aristodemus ponceanus Schaus, the southeast Florida helicta isolate’s range is very

restricted. It is definitely known only from the vicinity of Florida City, specifically the area of Carde Sound

Road. Leroy Koehn has seen one specimen from north Key Largo, but considers this a stray. However, and

hopefully, it may also occur sympatricly with areolatus across extreme south Florida to the Fakahatchee

Strand, Collier County. I have seen only about 20 specimens from this area, but a couple of them look like

good helicta to me. I have not examined the genitalia of any of these.

Koehn and Jeff Slotten have both informed me that its numbers appear to be in decline in the Carde

Sound area. Its greatest threat is from urban development and agricultural encroachment - not butterfly

collectors. It is amateur butterfly collectors that discovered it and are trying to protect it!

It may be premature to describe this isolate as a subspecies now. However, I feel this population

needs official scientific recognition (to help qualify it for environmental protection) more than it needs

additional study (which could take years). One of the primary goals of The International Lepidoptera

Survey is to discover, determine, and document taxa before they become extinct. We would hate to see this

become another Philotes sonorensis extinctis Mattoni, 1991 (a well known, but very geographically

restricted, Sonora Blue subspecies which was described only after becoming extinct as a result of

governmental land mismanagement). If in time it is determined that this population is not subspecificly

distinct from helicta helicta someone can always sink it later. Thus, for the above stated reasons, I now

describe this population as Neonympha helicta dadeensis .

6

Neonympha helicta dadeensis Gatrelle, new subspecies.

Diagnosis and description. Male (Fig. 6): All appendages and markings as in helicta helicta except as follows.

Ventral forewing: brown ground color much darker than in helicta, but not quite as dark as in septentrionalis, darker than in

areolatus; transverse median bands similar to areolatus - tending to orange and not prominent. Ventral hindwings :

marginal and median bands tending to areolatus - more orange, seldom open at the costal margin, and usually meeting at vein

Rs; eyespots ranging from as in helicta to very broad and often touching each other at the veins, never with yellow pupils; the

anal margin always with more white scaling than in nymotypical helicta, but rarely as heavy as in southwestern Florida

areolatus ; anal angle quite angular, pronounced, and often slightly lobed. Female (Fig. 12): As in the dadeensis male except

as follows. Ventral forewing : brown ground color lighter with the transverse bands a little more prominent; may have one or

two tiny eyespots in the submarginal area. Ventral hindwings : marginal and median bands more orange, occasionally open at

the costal margin, usually meeting at vein Rs, and rarely at Mi; whitish scaling along the anal margin not as pronounced; the

anal angle angulate but not as lobed.

Types. Holotype d* (Fig. 6): Carde Sound Road, Dade County, Florida, 22 November 1989. Allotype 9 (Fig. 12):

Carde Sound Road, Dade County, Florida, 24 Nov. 1989. Paratypes: 27c? c?, 11 9 9: all FLORIDA, Dade County, Carde Sound

Road: 4c? <j, 29 9, 10 May 1991; 11 <J <J, 49 9, 18 Nov., 5c? <?, 1 $, 22 Nov., 1 c?, 3 9 9, 24 Nov. 1989; 3<j c?, 19, 10 May

1990 (leg. Slotten); 3c?c?, 9-11 March 1987 (leg. ?). The Holotype and Allotype are deposited in the Allyn Museum of

Entomology, Sarasota, Florida. Paratypes are distributed as follows: Jeff Slotten, Gainesville, FL (4); MOTH, Goose Creek,

SC (3); Leroy Koehn, Lake Worth, FL (31). All type specimens were collected by Leroy Koehn unless otherwise noted.

Etymology. Dadeensis is named for Dade County, Florida - the only area from which it is currently known. I

suggest Miami Helicta as its co mm on name. Helicta Satyr is the co mm on name of Neonympha helicta. Northern

Helicta is the proper common name of Neonympha helicta septentrionalis (, septentrionalis means northern).

Remarks. The holotype is somewhat atypical in that its VHW eyespots are slightly smaller

than average. I utilized it because of its excellent condition. Figure 13 is a male paratype with larger

fused eyespots. The type locality of N. h. dadeensis is Dade County, FL. At present, the ranges of

the three helicta subspecies are not known to come into contact. There are no known blend zone

populations. Leroy Koehn first found dadeensis on 9 May 1972. The two males he collected then,

plus four pair he took in 1982, are now in the Carnegie Museum NH, Pittsburgh. I emphasize again

that the taxonomic relationship of the south Florida dadeensis and areolatus populations is unsure

and needs more study. All photos were taken outside in full sunlight which brings out the natural

colors and highlights of butterflies.

Fig. 13

ACKNOWLEDGMENTS

Special thanks to: Dr. Gerhard Tarmann for supplying a color copy of the original helicta plate;

Leroy Koehn, Jeff Slotten, and Ricky Patterson for their field observations and loan of specimens; Ben

Gatrelle for photography and computerization of photos.

LITERATURE CITED

DAVIS, W.T. 1924. Northern Form of the Butterfly Neonympha areolatus. J. New York Ent. Soc., 105.

DOS PASSOS, C.F. 1964. A Synonymic List of the Nearctic Rhopalocera. Memoir No. 1, Lepid. Soc.,

Yale University, New Haven, CT. 146 pp.

GATRELLE, R.R. 1998. The Rediscovery, Taxonomy, and Biology of Chlosyne gorgone and Chlosyne

ismeria (Nymphalidae) in Burke County, Georgia. TTR, Vol. 1:2, 9 pp. TILS, Goose Creek, SC.

_ 1999. An Evolutionary Subspecific Assessment of Deciduphagus henrici (Lycaenidae)

Based On Its Utilization of Ilex and Non-//ev Hosts: Description of a Third Ilex Associated

Subspecies. Designation of a Neotype and Type Locality for Deciduphagus irus. TTR, Vol. 1:6, 14

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

GOCHFELD, M. & J. BURGER. 1998. Butterflies of New Jersey. Rutgers Univ. Press, New Brunswick,

NJ. 329 pp.

HOWE, W.H. 1975. The Butterflies of North America. Doubleday & Co., Inc. New York, NY. 633 pp.

HUBNER, J. 1806. Sammlung exotischer Schmetterlinge. Augsburg.

MATHER, B. 1965. Euptychia Areolata : Distribution and Variation, with Special Reference to

Mississippi (Satyridae). J. Lepid. Soc., Vol. 19:3, pp. 139-160.

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.

SCOTT, J.A. 1986. The Butterflies of North America, A Natural History and Field Guide. Stanford Univ.

Press, Stanford, CA. 583 pp.

SMITH, J.E. 1797. The Natural History of the Rarer Lepidopterous Insects of Georgia. London. 2 Vols.

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Volume 1

15 August 1999

Number 9

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

CELASTRINA IDELLA (LYCAENIDAE: POLYOMMATINAE):

A NEW BUTTERFLY SPECIES FROM THE ATLANTIC COASTAL PLAIN.

DAVID M. WRIGHT

100 Medical Campus Drive, Lansdale, PA 19446

AND

HARRY PAVULAAN 1

494 Fillmore Street, Herndon, VA 22070

ABSTRACT. A new species of Polyommatinae, Celastrina idella, is described from the sandy Atlantic coastal plain

of the eastern US. It is presently known to occur from southern New Jersey through Georgia. C. idella larvae have been

recorded from four species of Ilex (holly). C. idella is distinguished from sympatric C. ladon and C. neglecta by differences

in larval host, flight period, pupal diapause, and adult size and wing characters. In the New Jersey pine barrens the butterfly is

univoltine and flies in the spring between the flights of its sympatric congeners.

CONCEPTS OF CELASTRINA SYSTEMATICS

Two hundred years ago John Abbot, the famous Georgian naturalist, found larvae of a species of

Celastrina Tutt, 1906 on Ilex L. (holly). He reared these to adults and in 1792 wrote: “The butterfly is not

common, but is more frequent in Hammocks and near swamps... tied itself up 30th April, changed into

chrysalis 2 May, bred March 12 following [year].” Abbot’s historical description is remarkable on several

accounts. We believe it to be the first reference to a heretofore undescribed species of Celastrina which we

now describe in this paper. It is also one of the earliest insect life histories from the New World.

The above quote was not included with Abbot’s plate in Smith (1797). It was uncovered later by

Scudder (1872, 1876) who examined the original plates and unpublished manuscript in London. In Smith

(1797), Abbot described a different Celastrina caterpillar found in June. This taxon fed on a wild legume

and gave rise to a second brood without diapausing.

Unfortunately, Abbot’s differing life history accounts have been largely overlooked taxonomically.

Abbot’s two very different life history descriptions indicated that at least two species of Celastrina were

present in the eastern US. Specifically, an earlier flying univoltine species whose larvae feed on holly, and

a later flying multivoltine species whose larvae feed on a different host(s).

Concepts of Celastrina systematics have emerged gradually. Prominent 19th century worker, W.H.

Edwards of Coalburgh, WV, provided the first systematic study of American taxa. He reared many of them

and correctly recognized them as distinct from Old World C. argiolus (Linnaeus, 1758). Edwards (1883)

discovered that pupae derived from eggs of the spring generation hibernated, and accurately concluded the

first generation was single-brooded. He then arbitrarily, and incorrectly, reasoned that all generations were

interrelated and lumped them together as “one polymorphic species” - pseudargiolus. Again, accurate life

history information was taxonomically overlooked. This view prevailed for nearly a century.

1 Staff member, The International Lepidoptera Survey, Goose Creek, SC.

Figs. 1-2 (d/v), d holotype Celastrina idella, 11 May 1987, nr. Chatsworth, Burlington Co., NJ. Figs. 3-4 (d/v), 9 Allotype

Celastrina idella, 19 May 1990, nr. Chatsworth, Burlington Co., NJ. Fig. 5 (d/v), d paratype C. idella, 16 April 1995, Bevan

WMA, Cumberland Co., NJ. Fig. 6 (d/v), d* C. idella f. lucia, 6 May 1987, Chatsworth, Burlington Co., NJ. Fig. 7 (d/v), d C.

idella, 24 March 1989, Green Swamp, Brunswick Co., NC. Fig. 8 (d/v), 9 C. idella, same data as Fig. 7. Fig. 9 (d/v), d C.

ladon lucia, f. marginata, 10 April 1992, Chatsworth, Burlington Co., NJ. Fig. 10 (d/v), 9 C. /. lucia, f. marginata, same data

as Fig. 9. Fig. 11 (d/v), d C. 1. lucia, f. lucia, same data as Fig. 9. Fig. 12 (d/v), d Celastrina undescribed sp., 24 May 1997,

Pocono Pines, Monroe Co., PA. Fig. 13. (d/v), d C. ladon ladon, 22 April 1988, near Rancocas State Park, Burlington Co.,

NJ. Fig. 14 (d/v), 9 C. /. ladon, 16 April 1995, Alloway, Salem Co., NJ. Fig. 15 (d/v), d C. neglecta, 23 July 1988, Red

Lion, Burlington Co., NJ. Fig. 16 (d/v), 9 C. neglecta, 20 June 1992, nr. Chatsworth, Burlington Co., NJ. Fig. 17 (d/v), d C.

neglectamajor, 15 May 1990, Fork Creek PHA, Boone Co., WV. Fig. 18 (d/v), d C. neglectamajor, 4 June 1997, Mt. Joy,

Hunterdon Co., NJ. Fig. 19 (d/v), d C. neglecta, spring form, 21 April 1999, Sumneytown, Montgomery Co., PA.

All figures are enlarged to 1. 5 natural size. _ (d) = dorsal, (v) = ventral.

Twentieth century advancements in evolutionary theory have propelled the idea of sympatric

speciation and have offered a new context in which to view co-occurring entities. Re-examination of many

of Edwards’ forms and “generations” have proven them to be isolated breeding populations. Several have

been resurrected to species level or redescribed as new species.

Host plant adaptation plays a key role in the evolution of phytophagous insects, especially

Celastrina (Pratt et.al. 1994). Their larvae are adapted to feed almost exclusively on flowering parts of

their hosts, a short-lived ephemeral resource. By necessity, adult flights are phenologically coupled to their

host’s flowering period in order that eggs may be laid on the seasonally limited provision. Because host

plants often bloom at different periods, host specialization can result in asynchrony of adult flights and

isolated mating periods.

Further, genetic isolation through seasonal isolation is believed to be an important mechanism in the

evolution of species in sympatry (Smith 1988; Wood and Keese 1990; Pratt 1994). In eastern North

America, Celastrina have diversified through host specialization. Local populations in nature often occur

asynchronously to one another and gene flow between them appears limited to non-existent. Morphologic

markers characterize several of these populations and formal description is needed to delineate them for

future studies in molecular systematics, phylogenetics, and possible conservation.

In the late 1970’s it became apparent to us that two separate entities flew in the spring in the New

Jersey pine barrens. Given the changing status of Celastrina systematics and the recognition of new

sympatric species in the Appalachians, we suspected a sibling species pair occurred in the springtime in

southern New Jersey. The first flight, consisting of larger dark individuals (f. “lucia” and ‘fnarginata”),

flew in April when its blueberry host was in bud; the second flight, consisting of smaller brighter

individuals (f. “violacea”), followed in May.

The host of the latter was unknown until one of us (DW) witnessed “violacea” females ovipositing

on unopened buds of inkberry holly Ilex glabra (L.) near cedar bogs in mid-late May, 1989. Viable larvae

were eventually found on inkberry and other Ilex species in the pine barrens. After two decades of field

studies we were convinced of the distinctness of the two spring taxa. The major distinguishing features

between them are contrasting adult phenotypes, asynchronous flights, different larval hosts with staggered

flowering periods, and experimental evidence of segregated pupal eclosion times. (A third species, the

summer flying C. neglecta (W.H. Edwards, 1862) is extremely rare and absent from much of the pine

barrens. When present, it occurs well after the spring siblings.) We felt these differences merited sibling

species status and formal description of the Ilex feeding taxon.

A NEW CELASTRINA SPECIES

The Holarctic genus Celastrina consists of small polyommatine (blue) butterflies whose larvae feed

on the flowering parts of a diverse variety of plants. Adult flights vary from early spring to late summer and

fall. Their biology and systematics have been the subject of recent investigation (Pratt et.al. 1994; Pavulaan

and Wright 1994; Wright 1995; Scott and Wright 1998). To this point in time, the Celastrina complex in

eastern North America consists of four recognized sympatric species; C. ladon (Cramer, 1780), C.

neglecta, C. nigra (Forbes, 1960), and C. neglectamajor Opler & Krizek, 1984. Additional distinct

biological races have been described, from other areas, for which species level designations have been

suggested (Wright 1995; Gochfeld and Burger 1997; Allen 1997; Layberry et.al. 1998; Glassberg 1999).

Common species ladon and neglecta are polyphagous and occur in widespread overlapping ranges.

Once thought to be different seasonal forms of the same insect (Edwards 1883), ladon is now known to be a

univoltine spring species and neglecta a multivoltine summer species. Recently discovered spring species

nigra and neglectamajor are uncommon monophagous species that fly in restrictive ranges in the

Appalachians and Ozarks. Their ranges match those of their unique hosts.

3

We divide the common eastern spring species, C. ladon, into two broad subspecies. Northern ssp.

C. ladon lucia (Kirby, 1837) occurs from the Canadian subarctic southward through New England to the

pine barrens of coastal southern New Jersey (Figs. 9-11), where its larvae feed on highbush blueberry

Vaccinium corymbosum L. A high percentage of heavily melanized adults (forms “lucia” and “marginata”)

appear in lucia populations. Males have androconia and lack long transparent scales. Southern ssp. C.

ladon ladon occurs from eastern Texas to northern Florida, northward through the central Appalachian Mts.

to the Piedmont of Pennsylvania and north central New Jersey (Figs. 13-14). Throughout its range,

flowering dogwood Cornus florida L. is the principal host. Subspecies ladon adults normally are lightly

marked on the venter and lack excessive melanization (form “violacea”). Males lack androconia and have

long transparent scales overlaying the blue scales of the forewing (Pratt et.al. 1994; Wright 1995; Wright

1998). Interestingly, ssp. ladon is often completely absent or rare on the outer coastal plain, presumably

due to competitive exclusion by a different spring Celastrina species.

We describe here Celastrina idella n. sp., a distinctive spring univoltine Ilex- feeding species from

the middle Atlantic seaboard (Figs. 1-8). It is distinguished from other Celastrina by its smaller size, wing

color, flight period, pupal diapause, and larval host. At the northern extent of its range, in the pine barrens

of southern New Jersey, it is sympatric with C. ladon lucia. There, it comprises the second of two easily

recognizable allochronic spring flights. From Delaware southward to Savannah, Georgia, it is virtually

parapatric to inland C. ladon ladon , and serves as the sole spring flight near the coast. C. idella flights are

completed before the flights of the ubiquitous sympatric summer species, C. neglecta (Figs. 15-16).

Celastrina idella Wright and Pavulaan, new species

Description. Male (Figs. 1-2, 5-7). Forewing length 10-15 mm (n=167). Southern NJ males average smaller (12.2

mm, n=68) than those from coastal North Carolina (13.2 mm, n=54). Dorsal color uniform light blue; some individuals with

distinct purplish-blue tint (especially NJ pine barrens). White insuffusion between veins on DHW common. Androconia

present. Wing fringes white; black checkering minimal to absent. Ventral color uniform light gray to white. Black

maculations greatly reduced. All are lightly marked, similar to form “violacea”, except in southern NJ where a few (3-6%)

have partially fused maculations on the VHW disc (near form “lucia”). Female (Figs. 3-4, 8). Forewing length 11-14.5 mm

(n=36). NJ females average smaller (12.2 mm, n=19) than those from North Carolina (13.5 mm, n=ll). Dorsal color

lustrous metallic light blue; many NJ individuals (pine barrens) with purplish tint. Black on DFW costa and outer margin.

DHW with series of submarginal black dots. White insuffusion on DHW common; occasionally also on DFW. Wing fringes

white; black checkering minimal to absent. Ventral color and pattern as in male.

Types. Holotype d* (Figs. 1-2): 2.2 km. s. of Chatsworth, Burlington Co., NJ, 11 May 1987, cedar bog in Risley

Branch of Wading River, leg. D.M. Wright. Allotype 9 (Figs. 3-4): 8 km. e. of Chatsworth, Burlington Co., NJ, 19 May 1990,

cedar bog in Pope Branch of Wading River, ovipositing on I. laevigata (Pursh) Gray, leg. D.M. Wright. Holotype and

allotype deposited in The Academy of Natural Sciences of Philadelphia (ANSP), Philadelphia, PA. Paratypes : 101 d* d* and

35 9 9: GROUP I (in pine barrens from 26 April - 28 May, 1987-1995): 50dc? and 20 9 9 from Chatsworth, NJ, and

vicinity; GROUP II (from outside the pine barrens 15 March - 20 May, 1979 - 1997): 45 d d and 13 9 9 from Bevan WMA

and Bear Swamp East, Cumberland Co., NJ; 1 d, 2 9 9 from N. Dennis and Eldora, Cape May Co., NJ; 5 d d nr. Pleasant Mills

and Egg Harbor Twp., Atlantic Co., NJ. Paratypes deposited in ANSP, United States National Museum (Washington, D.C.),

American Museum of Natural History (New York, NY), and collections of authors and contributors (DS, DI, JP).

Etymology. Idella is a feminine name derived from several possible Latin roots (id, ida, idea). We interpret it as a

corrupted form of Latin idealis, meaning “model embodying perfection.” Its common name should be Holly Azure.

Synonymy. Papilio argiolus L. in Abbot (1792), “Caterpillar feeds on...Holly &c. [F]requent in Hammocks and

near swamps.” [GEORGIA]; Cyaniris pseudargiolus (Bsd. & Le C.) in Scudder (1876, 1889), “...Ilex, on which Abbot

found it.” [GEORGIA]; Lycaena ladon (Cramer), form “violacea” Edw. in Smith (1910), “occurs in April and May...

foodplants...are Ilex.” [NEW JERSEY]; Lycaenopsis argiolus pseudargiolus, form vem. pseudargiolus in Comstock

(1940), “...spring forms facia and marginata ] occur with pseudargiolus... Lakewood [pine barrens]...May 18” [NEW

JERSEY]; L. a. pseudargiolus, form vem. neglecta-major (Tutt) in Comstock (1940), “This form follows the early spring

forms and is intermediate in occurrence between them and the first summer brood...Jamesburg [pine barrens] May 30.”

[NEW JERSEY]; Lycaenopsis argiolus, form violacea, in Shapiro (1966), “...majority of spring specimens are form

marginata...[V]iolacea increases in frequency as the spring emergence progresses.” [NEW JERSEY, in part]; Celastrina

ladon (Cramer), form “violacea” in Gatrelle (1986), “from all south coastal counties...form violacea emerges at end of

February and flies throughout March.” [SOUTH CAROLINA]; Celastrina ladon, “violacea” in Glassberg (1993) “In our area

there may be at least three species -[including] a ‘ violacea ’ type Azure appearing in the pine barrens in early May.” [NEW

JERSEY]; Celastrina ladon, “Pine Azure” in Walter (1993), “Pine Azure...is associated with pine barren habitats.” [NEW

JERSEY]; Celastrina ladon, “Late Spring Azure” in Pavulaan and Wright (1994), “especially co mm on in New Jersey white

cedar swamps...also very common in New Jersey deciduous swamps, Cape May County.” [NEW JERSEY]; Celastrina ladon

ladon (Cramer), race ‘ \iolacea ” III in Pratt et.al. (1994), “type III feeds on male flowers of Ilex species...on coastal plain”

[NEW JERSEY to GEORGIA]; Celastrina sp., “Cherry Gall Azure” and “Eastern Spring Azure” in Wright (1995), “...sizable

populations of Cherry Gall Azures can be found in cedar swamps of the New Jersey pine barrens, where...[they] feed on male

flowers of Inkberry (Ilex glabra),” and “...along outer coastal plain from Cape May, New Jersey to Georgia, there occurs a

brightly-colored, holly-feeding, ‘ violacea ’ variety of the Eastern Spring Azure.” [NEW JERSEY to GEORGIA]; Celastrina

n. sp., “Late Spring Azure” in Iftner and Wright (1996), as “Holly Spring Azure” [Fig. 1] [NEW JERSEY]; Celastrina sp.,

“Pine Barren Azure” in Gochfeld and Burger (1997), “...this distinct entity may be either a small race...or a distinct species.

Forested Coastal Plain including the Pine Barrens...[f]eeds on Inkberry...and American Holly.” [NEW JERSEY]; “Coastal

Holly Azure” in Williams (1998), “...coastal holly azures have been discovered so recently that they have yet to be assigned

scientific name.” [NEW JERSEY]; Celastrinajadon, “Pine Barrens Spring Azure” in Glassberg (1999), “...may eventually

merit full species status” [EAST]; “ Celastrina sp.” Holly [Spring] Azure in LeGrand and Howard (1999)...“This species found

in a wide variety of wooded or semi-wooded sites, in the vicinity of hollies...Coastal Plain only” [NORTH CAROLINA].

Biology. Flight period: In the north (type locality), a four-week flight from end of April to end of May and

following the flight of C. ladon lucia (extreme dates: 25 April - 4 June). To the south (inc. Delaware Bay counties of

southern New Jersey), flight is correspondingly earlier (extreme dates: 26 February - 20 May). Adult Nectar Sources: Open

flowers of highbush blueberry Vaccinium corymbosum L., sand myrtle Leiophyllum buxifolium (Berg.) Ell., chokeberries

Aronia arbutifolia (L.) L.F. and A. melanocarpa (Michx.) Willd., black cherry Prunus serotina Ehrh., red maple Acer

rubrum L., and rock cress (Arabis sp.) Hosts: Male flowers of dioeious Ilex species (Aquifoliaceae). Eggs and larvae

recorded on American holly I. opaca Ait., inkberry I glabra (L.) Gray, and smooth winterberry I. laevigata (Pursh) Gray in

southern New Jersey. Recorded from yaupon holly I. vomitoria Ait. in the outer banks of North Carolina and I. opaca Ait. in

coastal southern Virginia. Possibly tall gallberry holly I. coriacea (Pursh) Chapm. in North Carolina. [Note: Wild larvae

reared from flowers of Virginia Willow Itea virginica L. at Virginia Beach, VA, in May, diapaused and produced idella

adults. This may be an alternative host, especially near the end of the adult flight when unopened Ilex buds are dwindling.]

Immature Stages. Egg white, similar to other Celastrina : placed on floral buds of Ilex hosts. First instar light

green: bores into floral buds. Mature larva (4th instar) variable, light green to creamy white; some lightly contrasted with

dorsal white chevrons, maroon prothoracic shield, or rarely with faint rosy mid-dorsal stripe. (Lighter and less boldly marked

than larvae of C. ladon lucia.} Myrmecophilous larvae on Ilex glabra (L.) Gray attended by worker ants of Camponotus

ferrugineus (F.), C. nearcticus Emery, and Lasius alienus (Foerster). Hymenopterous parasitoids recovered during rearing

include Trichogramma sp. from eggs and a small wasp Cotesia cyaniridis (Riley) from mature larvae. Pupa light brown

(7.2-8.2 mm, n=10), obtect. Pupation dates, June 14-July 3 in New Jersey. Pupae diapause.

Habitat. Wet areas (cedar swamps, cypress swamps, pocosins, bottomlands, wet thickets) and upland dry maritime

forests of sandy coastal plain and barrier islands, with holly (Ilex).

Frequent Associates. Mitoura hesseli Rawson & Ziegler, 1950, in Atlantic white cedar bogs and the Ilex

associated subspecies (Gatrelle, 1999) of Deciduphagus henrici (Grote & Robinson, 1867) in coastal holly woodlands.

Range (Fig. 22). Known localities include: NEW JERSEY: ATLANTIC CO.: Atlantic City International Airport,

Brigantine, Folsom, Forge Pond, Hammonton, Mays Landing, Mizpah, Pleasant Mills, Pomona, Weekstown; BURLINGTON

CO.: Batsto, 2.2 km. s. Chatsworth, 8 km. e. Chatsworth, Duke’s Bridge, High Bridge, Jenkins, 6 km. e. Medford, Ong,

Oswego Lake, Penn SF, Quaker Bridge, Red Lion, Speedwell, West Plains; CAMDEN CO.: Atco, Gibbsboro, Kirkwood Lake;

CAPE MAY CO.: Belleplain SF, Cape May Point, Dennisville, Eldora, Five Mile Beach, Goshen, Hidden Valley, Higbee

Beach, Jake’s Landing, North Dennis, Petersburg, Seaville, Woodbine (Great Cedar Swamp); CUMBERLAND CO.: Bevan

WMA, Bear Swamp, Center Grove, Dividing Creek, Hansey Creek, Millville, Manumuskin, N. Port Norris; GLOUCESTER

CO.: Iona; MONMOUTH CO.: Allaire SP; OCEAN CO.: Lakehurst, Lakewood, Manahawkin, Ridgeway, Waretown, Warren

Grove; SALEM CO.: Alloway, Quinton. DELAWARE:. KENT CO.: w. Milford (Mispillion River); SUSSEX CO.: nr. Bethel,

nr. Lincoln, Nanticoke WMA. MARYLAND: ANNE ARUNDEL CO.: Annapolis, Riva; CALVERT CO.: Lusby; CAROLINE

CO.: nr. Templeville; CHARLES CO.: Cedarville SF; PRINCE CHARLES CO.: Cedarville; WICOMICO CO.: n. Salisbury

(white cedar swamp); WORCESTER CO.: Wango (John’s Tract). VIRGINIA: CHESAPEAKE: Northwest River Park.;

FAIRFAX CO.: Mason Neck SP; JAMES CITY CO.: Lakewood (nr. Jamestown); NEW KENT CO.: New Kent; PRINCE

WILLIAM CO.: Triangle (Prince William Forest); SUFFOLK: Dismal Swamp, nr. Corapeake, NC; VIRGINIA BEACH:

Seashore SP. NORTH CAROLINA: BEAUFORT CO.: Locality not given; BLADEN CO.: Bladen Lakes SF (nr.

Elizabethtown), Jones Lake SP (nr. Elizabethtown); BRUNSWICK CO.: Green Swamp; CAMDEN CO.: Great Dismal Swamp;

CATERET CO.: Moorhead City; COLUMBUS CO.: Green Swamp; CRAVEN CO.: Croatan NF (nr. Croatan); DARE CO.: Kill

Devil Hills (Outer Banks), Nag’s Head (Outer Banks), Whalebone (Outer Banks); DUPLIN CO.: Kenansville; GATES CO.:

nr. Sunbury (Great Dismal Swamp); JONES CO.: Croatan NF (nr. Maysville); NEW HANOVER CO.: Locality not given;

ONSLOW CO.: Locality not given; PASQUOTANK CO.: Elizabeth City; PENDER CO.: Angola Swamp; PITT CO.: Locality

not given; ROBESON CO.: Locality not given; SAMPSON CO.: Turkey; WAYNE CO.: Locality not given. SOUTH

CAROLINA. AIKEN CO.: A

CO.: Charleston; COLLETON CO.: L

CO.: Locality not given; CHARLE!

Givhans Eerry SP; GEORGETOWN

overlap of ffightem certam years. Mean peak flights of lucia (April 16) and ideUa { Ma/l| are neariya month apart in the

fully two weeks earlier. C. IdeUa (Figs. 1-2) is distinctly smaller and lighter in color tha 1 lucia. The wings of both sexes of

lucia (Figs. 9-10) are larger (FW 13.1 mm, n=42), solid blue in color with no white insuffusion, and strongly outlined in

le (7.5-8.6 m

unique to this subspecies 2 . C. /. ladon males often have a satiny matte appearance (not seen in \della) because of these

scales. A dissecting scopeor hand lens is often needed to diagnose this scale pattern. The ultrastructure (SEM) of C. /. ladon

host plants. In the surrounding Delaware Bay counties of soufhem NJ neglelta is present in early Summer (June), but

serotina and will oviposit on floral buds of this host. The great majority of larvae re; Y ^

eclose to join the bulk of the neglecta population which begins flight in late May/J

die. The venter of the s E

of ideUa. Neglectamajor is much larger (Figs. 17-18) and is found only‘in Appalachian deciduous woodlands (Fig. 27). S It

feeds solely on black cohosh Cimicifuga racemosa (L.) Nutt. The species (Fig. 12) associated with mite-induced leaf galls

aarscsr

Figs. 22-27. Celastrina distribution in the mid-Atlantic region.

22, distribution of C. idella. 23, distribution of C. ladon lucia.

24, distribution of C. /. ladon. 25, distribution of C. neglecta.

26, distribution of C. ra'gra. 27, distribution of C.

neglectamajor.

Figs. 28-29. Genitalia 3 of rf C. z'de//a from nr. Chatsworth,

Burlington Co., NJ, 13 May 1987. 28, dorsal view of valva with

apical terminal process (100X). 29, ventral view of upper half of

ring; uncal lobes with spines (100X).

i (400X).

DISCUSSION

Celastrina idella becomes the fifth North American lycaenid described from the New Jersey pine

barrens 4 . The region is a natural community with distinctive flora and a rich history of scientific study

(Forman 1979). Located on the unglaciated outer coastal plain, the barrens is an “island” of highly acidic,

sandy soil which supports large stands of undisturbed pine forests, heathlands, and bogs. Its close proximity

to major metropolitan centers and easy access to exploration account for its popularity with naturalists for

nearly two centuries. Part of the uniqueness of the pine barrens is its mixed flora. Many plant species of

both northern and southern affinity reach their range extremes in the pine barrens where they overlap

(McCormick 1970).

In similar fashion, the sibling pair of spring Celastrina in the pine barrens represent insects of

differing affinities whose ranges overlap only in southern New Jersey. The first taxon, C. ladon lucia , is

most common in northern woodlands and peatlands of Canada and New England where it feeds on many

plant families. Its southernmost coastal population occurs in the New Jersey pine barrens (Fig. 23). This

coastal lucia population is unique in that it utilizes a single host, flies very early, and is darker blue than

northern populations. It may be a remnant of a much larger lucia population that once florished on expanded

heathlands of the exposed continental shelf. Physiological constraints prevent lucia from extending further

south despite the presence of acceptable hosts.

The second taxon, C. idella , is a denizen of southeastern coastal swamps and holly forests. It occurs

from Georgia to southern New Jersey with its northernmost population in the New Jersey pine barrens.

Similarly, physiologic constraints must prevent idella from occurring further north even though holly still

remains common along the coast. (For instance, C. idella is absent from the American holly forest on the

dunes at Sandy Hook, Monmouth Co., NJ, just north of the New Jersey pine barrens, and is also absent from

the Long Island pine barrens of eastern Suffolk Co., NY, where inkberry is locally common.) Even though

these two siblings are adapted to single hosts, it should be stressed that they are also restricted by habitat

preferences. Their ranges are smaller than their respective hosts.

How often sympatric speciation creates new species is still a matter of dispute. Once thought

impossible, the concept has gained respect among evolutionary biologists (Gibbons 1996). Charles

Darwin’s ideas of the formation of species through ecological specialization have been revived (Morell,

1999). Most organisms require geographic isolation (allopatry) to form new species. However, some plant¬

feeding insects seem to be particularly adept at finding empty niches, exploiting opportunities, and

diverging from others sympatrically. Divergence is spurred by strong selection and rapid adaptation.

Specialization on host floral buds is the niche of Celastrina and each separate host represents a

potentially different niche. Wherever C. I lucia and idella co-occur, host specialization prevents their

contact. Adults of the two species fly at different times coupled to the bud stage of their respective hosts.

The degree of interbreeding (if any) between these two taxa is unknown. Population genetics models have

shown that it takes very little interbreeding to erase differences. The small percent (3-6%) of New Jersey

idella individuals with imperfect “lucia” disc on VHW (Fig. 6) could be taken as evidence of interbreeding

between C. /. lucia and idella. However, all other phenotypic and biologic characteristics of the taxa

remain distinct with no introgression. The “lucia” disc is most likely genetically controlled, but instead of

acquisition through recent interbreeding it can be equally explained as an expression of an ancestral gene

induced or unmasked by local cold temperatures (especially soil temperatures where pupae rest).

We are not aware of any evidence of interbreeding between C. /. lucia and C. idella in New Jersey.

Further south within its range, idella flies parapatrically to inland C. /. ladon. C. I ladon is common in the

Piedmont and reaches the inner coastal plain (Fig. 24) where there is evidence of partial hybridization with

idella. The flights of the two taxa overlap chronologically in Virginia and the Carolinas, and contact is

inevitable. The width of the hybridizing suture zone is presently unknown, but is suspected to be small.

) 8

On the outer coastal plain and barrier islands C. idella is the exclusive spring Celastrina taxon and

C. /. ladon is absent. Should the hybridization zone on the inner coastal plain prove to be substantially

large, then the two taxa are not separated at the species level in the south. In that case, southern New Jersey

would harbor endemic populations of C. idella.

The origins of C. idella are unknown. We suspect that it originated in the Pleistocene during glacial

stages of low sea level when the Atlantic coastal plain was enormously widened and constituted a

significantly larger belt of land. During these intervals the exposed outer coastal plain extended 100-200 km

eastward from today’s shoreline and was covered with similar vegetation (Emery et.al. 1967). Given the

abundance of Ilex and the relative sparcity of flowering dogwood Cornus florida on today’s outer coastal

plain, idella was most likely the only spring Celastrina flying in this expanded biome during low sea levels

from Delaware Bay southward. It may have evolved quickly by a host shift from a closely related ancestor.

The hypothetical ancestor presumably had a light venter phenotype similar to present day neglecta ,

neglectamajor , or the undescribed cherry gall species. Curiously, Ilex flowers are toxic to the larvae of

these three species in the lab - they generally do not survive beyond the first instar stage. Selection would

have favored a founder with larval resistance to Ilex toxins. Allozyme studies are underway to investigate

the closest relative(s) of idella and its systematic relationship with other taxa.

ACKNOWLEDGMENTS

We acknowledge our deep gratitude to David Iftner, Gordon Pratt, and Dale Schweitzer who provided

many stimulating discussions regarding this new taxon and Celastrina biology. These individuals also

reviewed the manuscript draft and offered valuble critical comments. We greatly appreciate the unselfish

assistance given by F. Martin Brown and John Eliot regarding nomenclatural issues. We also acknowledge

and extend our warmest thanks to the following individuals and their respective institutions for providing

access to reference collections: The Academy of Natural Sciences of Philadelphia (Don Azuma, Jon

Gelhaus, and Jason Weintraub), Allyn Museum of Entomology (Jackie and Lee Miller), American Museum

of Natural History (Fred Rindge and Eric Quinter), Carnegie Museum (John Rawlins), Cleveland Museum

of Natural History (Sonja Teraguchi), Dayton Museum of Natural History (Reed Watkins), Delaware

County Institute of Science (John Hallahan), Eastern College (David Unander), Field Museum (Phil

Parillo), Florida State Collection of Arthropods (John Heppner), Illinois Natural History Survey (Kathleen

Zieders), Los Angeles County Museum of Natural History (Julian Donahue), Michigan State University (Mo

Nielsen), Museum of Comparative Zoology (David Furth and Stephan Cover), New Jersey State Museum

(David Parris), Newark Museum (John Michalski), The Ohio State University Museum of Biological

Diversity (Eric Metzler), Pennsylvania Department of Agriculture (Karl Valley), Pennsylvania Department

of Forestry (John Quimby), Pennsylvania Natural Diversity Inventory (Barbara Barton), Purdue University

(Arvin Provonsha), Reading Museum (Michael Feyers), Royal Ontario Museum (Chris Darling), Rutgers

University (Timothy Casey), Staten Island Institute of Arts & Sciences (Edward Johnson), United States

National Museum (Robert Robbins), University of California Riverside (Greg Ballmer and Gordon Pratt),

University of Delaware (Dale Bray and Tom Wood), University of Guelph (Steve Marshall), University of

Louisville (Charles V. Coveil), University of Michigan Museum of Zoology (Mark O’Brien), University of

Pennsylvania (Andrew Binns), Wagner Institute (Eugene Bolt and Susan Glassman), Yale Peabody Museum

(Charles Remington). We also sincerely thank the following individuals for allowing us the opportunity to

examine their private collections and/or forward specimens to us: Chris Adams, Tom Allen, George

Balogh, Andrew Beck, Richard Boscoe, John Bowe, John Calhoun, Tom Carr, Charles V. Coveil, Harry N.

Darrow, Jim Dowdell, David Eiler, Karl Gardner, Robert Gardner, Ron Gatrelle, William Grooms, David

Iftner, Phil Kean, Ron King, Marc Minno, Mo Nielsen, David Parshall, John Peacock, Jim Popelka, Joe

Riddlebarger, Frank Rutkowski, Dale Schweitzer, John Shuey, Ernest Shull, Don Simonds, Richard Smith,

Reed Watkins, William Wright, Joe Zeligs, Ben Ziegler, and Harry Zirlin. A special thanks is extended to

Carol Spawn (Chief Librarian of The Academy of Natural Sciences of Philadelphia) and Pamela Gilbert

(Entomology Librarian, British Museum of Natural History) for aiding in retrieval of the Abbot references;

9

also to Phil Ackery and Pamela Gilbert (BMNH) for obtaining color transparencies of the original Cramer

and Abbot paintings. We are grateful to Jim Springer (North Jersey Butterfly Club) and Steve Walter (New

York City Butterfly Club) who kindly shared their databases. Steve Walter also supplied helpful

information regarding the Long Island pine barrens. We sincerely appreciate the expertise and efforts of

Paul Marsh and Richard D. Smith, Research Entomologists (USDA), who identified the larval parasitoids

and ant attendants.

LITERATURE CITED

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MS. descriptions.] vol. 16, no. 212, British Museum (Natural History), London.

ALLEN, T.J. 1997. The Butterflies of West Virginia and their Caterpillars. Univ. of Pittsburgh Press,

Pittsburgh. 388 pp.

BOISDUVAL, J.A.B.D., and J.E. LE CONTE. 1833. Historic Generale et Iconographie des Lepidopteres

et des Chenilles de l’Amerique Septentrionale. Roret, Paris. 228 pp.

BROWN, F.M. 1970. The Types of Lycaenid Butterflies Named by William Henry Edwards Part HI.

Plebejinae. Trans. American Ent. Soc. 96: 423-431.

EDWARDS, W.H. 1883. On the Polymorphism of Lycaena Pseudargiolus. Papilio 3:85-97.

CLENCH, H.K., and L.D. MILLER. 1980. Papilio ladon Cramer vs. Argus pseudargiolus

Boisduval and Le Conte (Lycaenidae): A Nomenclatorial Nightmare. J. Lepid. Soc. 34(2): 103-119.

COMSTOCK, W.P. 1940. Butterflies of New Jersey. J. New York Ent. Soc. 48: 47-84.

EMERY, K.O. et.al. 1967. Freshwater Peat on the Continental Shelf. Science 158:1301-1307.

FORMAN, R.T.T. 1979. Pine Barrens: Ecosystem and Landscape. Academic Press, New York. 601 pp.

GATRELLE, R.R. 1986. The Papilionoidea of South Coastal Area of South Carolina. Southern. Lepid.

Bull. no. 2. 16 pp.

_. 1999. An Evolutionary Subspecific Assessment of Deciduphagus henrici (Lycaenidae)

Based On Its Utilization of Ilex and Non -Ilex Hosts: Description of a Third Ilex Associated

Subspecies. Designation of a Neotype and Type Locality for Deciduphagus irus. The Taxonomic

Report, Vol. 1:6, 14 pp. The Int. Lepid. Survey, Goose Creek, SC.

GIBBONS, A. 1996. On the Many Origins of Species. Special News Report: Speciation. Science 273:

1496-1498.

GLASSBERG, J. 1993. Butterflies Through Binoculars. A Field Guide to Butterflies in the Boston-New

York-Washington Region. Oxford Univ. Press, New York. 160 pp.

_. 1999. Butterflies Through Binoculars. The East. A Field Guide to the Butterflies of

Eastern North America. Oxford Univ. Press, New York. 242 pp.

GOCHFELD, M. and J. BURGER. 1997. Butterflies of New Jersey. Rutgers Univ. Press, New Brunswick,

NJ. 327 pp.

IFTNER, D. C., and D. M. WRIGHT. 1996. Atlas of New Jersey Butterflies. Special Private Publication,

Sparta, NJ. 28 pp.

LAYBERRY, R.A., P.W. Hall, and J.D. Lafontaine. 1998. The Butterflies of Canada. Univ. of Toronto

Press, Toronto. 280 pp.

LEGRAND, H.E., Jr., and T.E. Howard, Jr. 1999. Notes on the Butterflies of North Carolina. Sixth

Approximation. 183 pp. [Online] Available: http://www.ncsparks. net/butterfly/nbnc.html

MCCORMICK, J. 1970. The Pine Barrens. A Preliminary Ecological Inventory. NJ St. Mus., Res. Report

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MORELL, V. 1999. Ecology Returns to Speciation Studies. Science 284: 2106-2108.

PAVULAAN, H., and D.M. Wright. 1994. Out of the Azure and Into the Lab: The Current State of

Celastrina research. The Mulberry Wing 10 (1): 406.

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10

PRATT, G.F., D.M. WRIGHT, and H. PAVULAAN. 1994. The Various Taxa and Hosts of the North

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SHAPIRO, A.M. 1966. Butterflies of the Delaware Valley. Amer. Ent. Soc., Philadelphia, PA. 79 pp.

SMITH, D.S. 1988. Heritable Divergence of Rhagoletispomonella Host Races by Seasonal Asynchrony.

Nature 336: 66-67.

SMITH, J.B. 1910. The Insects of New Jersey. Ann. Report of the NJ St. Mus. 1909. MacCrellish and

Quigley, Trenton. 888 pp.

SMITH, J.E. 1797. A Natural History of the Rarer Lepidopterous Insects of Georgia. 2 vols., London.

WALTER, S. 1993. Out of the Blue (?) and Into the Azure. Mulberry Wing 9(1): 5-6.

WILLIAMS, T. 1998. Reports. Earth Calendar. Audubon 100 (3): 20.

WOOD, T.K., and M.C. KEESE. 1990. Host Induced Assortative Mating in Enchenopa Treehoppers.

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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.

volume 1 is $65. US. The subscription year for volume 1 began in August of 1998 and will run to December 31 of 1999.

i CD

> (ie. a

se Creek, SC 29445.

11

Volume 1

24 December 1999

Number 10

The Taxonomic Report

II y/r OF THE INTERNA TIONAL LEPIDOPTERA SURVEY ^F

THREE NEW HESPERIOIDAE (HESPERIINAE) FROM SOUTH CAROLINA:

NEW SUBSPECIES OF

EUPHYES BIMACULA , POANES AARONI, AND HESPERIA ATTALUS

RONALD R. GATRELLE 1 ’ 2

126 Wells Road, Goose Creek, South Carolina 29445

ABSTRACT. Euphyes bimacula arbogasti is described as a new subspecies from Berkeley County, South

Carolina. It is known from only a few widely scattered colonies in the coastal swamp forests of the southeastern United

States from Georgia to southeastern North Carolina. It is darker then E. b. bimacula and E. b. illinois. Poanes aaroni

minimus is described as a new subspecies from Bull Swamp, Orangeburg County, South Carolina. This unique inland

subspecies is presently known only from the type locality. It is darker then P. a. aaroni and P. a. howardi. Hesperia attalus

nigrescens is described as a new subspecies from the relict dunes of Sandy Island National Wildlife Sanctuary, Horry

County, South Carolina. This isolated subspecies is much darker than H. a. attalus and H. a. slossonae. The Sandy Island

colony of H. a. nigrescens is believed to be the only remaining colony of this subspecies. All three subspecies are similarly

Additional keywords: Threatened species, original descriptions.

INTRODUCTION

At least 33 species/subspecies of butterflies were originally described from populations inhabiting

east coastal Georgia or south coastal South Carolina by the earliest workers on American Lepidoptera in

the 1700’s and early 1800’s. Since then however, very little taxonomic attention has been given to the

Lepidopterian fauna of the mid-Atlantic area of the United States between Florida and New Jersey. This has

been especially true for the last half of the 1900’s when very few lepidopterists, and even fewer butterfly

taxonomists, have been residents of the mid-Atlantic area.

A result of this long term scarcity of collectors is that few specimens from this region are available

for study. This informational void has given rise to taxonomic oversimplification and misrepresentation in

the popular literature of the taxa occupying the area between Florida and New York and from the Atlantic

coast to the Mississippi River. Most modern butterfly books recognize few species as occurring in more

than one subspecies throughout this vast area of the United States.

This is in stark contrast with the west coastal region of the United States with its abundance of

lepidopterists and subspecies. In this western area it is generally expected that each mountain range and

valley system will harbor different subspecies - and indeed they often do. In California, for example, some

(supposed) subspecies are separated by only a few hundred yards. Yet, the consistent impression given in

the popular literature is that the species of South Carolina (from its coast to its mountains) are not expected

to differ subspecifically from those of New Jersey, Missouri, or Louisiana.

1 Curator, Museum of the Hemispheres, Goose Creek, South Carolina.

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

All figures actual size.

Figs. 1-40. 1/6 (d/v), 3 E. b. bimacula, 29 June 1986, Passadumkeag, Penobscot Co., ME, leg. Grey. 2/7 (d/v), ? E. b. bimacula, 29 June 1986, and

same as 1. 3/8 (d/v), 3 P. a. aaroni, 20 August 1976, nr. New Gretna, Burlington Co., NJ, leg. Adelberg. 4/9 (d/v), $ P. a. aaroni, same data as 3.

5/10 (d/v), 3 P. a. howardi, 24 April 1971, 10 mi. w. Daytona, Volusia Co., FL, leg. Roman. 11/16 (d/v), 3 neotype E. b. illinois, 22 June 1968,

Denine Crossing, Grundy Co., IL, leg. N.G. Seaborg. 12/17 (d/v), ? E. b. illinois, 28 June 1975, William’s Prairie, Johnson Co., IA, leg. Gatrelle. 13/18

(d/v), 3 holotype P. a. minimus, 1 June 1992, Bull Swamp, Orangeburg Co., SC, leg. Gatrelle. 14/19 (d/v), 9 allotype P. a. minimus, 6 June 1992,

and same as 13. 15/20 (d/v), 9 P. a. howardi, 25 July 1976, 5 mi. w. of 1-95 on Hwy. 92, Volusia Co., FL, leg. Gatrelle. 21/26 (d/v), 3 holotype E.

b. arbogasti, 12 July 1970, nr. jet. of 1-26 and 17-A, Berkeley Co., SC, leg. Gatrelle. 22/27 (d/v), 9 allotype E. b. arbogasti, 21 May 1962, nr.

Blichton, Bryan Co., GA. 23/28 (d/v), 3 holotype H. a. nigrescens, 10 October 1995, Horry Co., SC, leg. Gatrelle. 24/29 (d/v), 9 allotype H. a.

nigrescens, 3 October 1995, and same as 23. 25/30 (d/v), 3 P. aaroni ssp., 27 May 1995, Sabine Pass, Jefferson Co., TX, leg. Slotten. 31/36 (d/v),

3 H. a. attalus, 2 June 1974, 8 mi. west of Medicine Lodge on 160, Barber Co., KS, leg. Heitzman. 32/37 (d/v), 9 H. a. attalus, 28 May 1974, and

same as 31. 33/38 (d/v), 3 H. a. slossonae, 21 June 1994, Chesterfield Co., SC, leg. Gatrelle. 34/39 (d/v), 9 H. a. slossonae, 5 September 1994,

Hwy. 19, Marion Co., FL, leg. Gatrelle. 35/40 (d/v), $ P. aaroni ssp., same data as 25 _

The states of Georgia, South Carolina, and North Carolina compose the most ecologically diverse

area in the eastern United States. This diversity occurs in all points of the compass, but is delineated most

notably from the coastline to the mountains. South Carolina, being in the heart of this area, has five very

definitive life zones: 1) semi-subtropical south coastal islands, 2) maritime swamp forests, 3) arid

sandhills, 4) piedmont, and 5) the edge of the Appalachian Mountains. As one traverses these zones, from

the south-east to the northwest, South Carolina’s species changes greatly. On a given fall morning one could

observe/collect such locally common tropical taxa as Heliconius charitonius tuckeri W. P. Comstock and

F. M. Brown, 1950; Danaus gilippus berenice (Cramer, [1775]); Hemiargus ceranus antibubastus

Hubner, [1818]; Pterourus palamedes (Drury, [1773]); Urbanus dorantes (Stoll, [1790]); and Calpodes

ethlius (Stoll, [1782]) on Hunting Island in Beaufort County, then drive just five hours to Oconee County

and in the afternoon observe/collect such locally common temperate species as Boloria bellona (Fabricius,

1775); Speyeria cybele (Fabricius, 1775); Lycaena phlaeas americana Harris, 1862; Polites peckius (W.

Kirby, 1837); and Enodia anthedon A. H. Clark, 1936.

As with its species, South Carolina’s subspecific diversity is more pronounced from east to west

(southeast to northwest) than from south to north. In this respect, South Carolina is similar to southern

California where the basic life zones and their accompanying subspecies are encountered east/west as they

correspond to the mountain ranges and valleys. This similar geographical subspecific pattern should be

expected because the subspecific distribution of the taxa in both South Carolina and California arose in

concert with the geological evolution of these states coastal areas, mountains, valleys, and arid regions.

Some of South Carolina’s species are represented in the state by one subspecies which is endemic

to the coastal area and another which is endemic from the midlands to the mountains: Anthocharis midea

midea (Hubner, 1809) (coastal islands) and A. m. annickae dos Passos and Klots, 1969 (remainder of

state); Asterocampa celtis reinthali Friedlander, 1988 (south coast) and A. c. celtis (Boisduval and

Leconte, [1834]) (remainder of state). Some do not occur in the coastal area but are represented by one

subspecies in the sandhills and another in the mountains: Satyrium edwardsii edwardsii (Grote and

Robinson, 1867) (mountains) and S. edwardsii undescribed (southern sandhills); Chlosyne gorgone

gorgone (Hubner, 1810) (sandhills) and C. g. carlota (Reakirt, 1866) (upper piedmont and mountain

foothills). Occasionally a species may have three subspecies within the state: Satyrium calanus cal anus

(Hubner, 1809) (south coastal), S. c. falacer (Godart, 1824) (mountains), and S. calanus undescribed

(sandhills).

There are at least three undescribed butterfly species in South Carolina and several undescribed

subspecies. Three of these heretofore undescribed subspecies are described as new Hesperiinae in this

paper. Two of these are apparently geographically unique and are thus limited to single populations (one is

in the Sandhills region and the other in a coastal dune area). The third is widely distributed from Georgia to

North Carolina, but is rare within this range. Each of these subspecies should be considered as a taxon in

need of special environmental protection. One, Hesperia attalus nigrescens Gatrelle, 1999 n. ssp., is

already protected because most of its population is expected to be found within the Sandy Island National

Wildlife Sanctuary in Horry County, SC.

A NEW SUBSPECIES OF EUPHYES BIMACULA

Review of bimacula subspecies.

Euphyes bimacula bimacula (Grote and Robinson, 1867) was described from the female - the

male being unknown at the time. There was no figure. The original description makes no mention of how

many specimens were involved. I believe it is most probable that this species was described from a single

specimen. Miller and Brown (1981) were unaware of the location of the type, but suggested that it might be

in the AMHN or NY State Museum. I have not tried to locate the type. (Any Information regarding the

3

existence of the female holotype would be greatly appreciated.) It is possible that the type is no longer

extant. Further, if bimacula was described from a single specimen, then there are no syntypes. In which

case, a neotype should be designated from the vicinity of Philadelphia, Pennsylvania.

The Annals of the Lyceum of the New York Natural History Society is a rather rare work. It was

only published in 10 volumes from 1860 to 1869. Because the original description is so rare, and also very

detailed and accurate, I think it a good idea to reproduce it here:

Hesperia bimacula, n. s. Female. Head, above, and prothorax, covered with dark brassy yellow scales; the tips of

the palpi are stained with the same shade. Thorax and abdomen, above, blackish, clothed with lateral, longer, mixed

olivaceous and aeneous hair. Beneath, the palpi are white; the eyes are also margined narrowly with white. Thoracic

parts, beneath, clothed with longer, mixed griseous and whitish hair. Abdomen, beneath, whitish, with longitudinal

darker streaks, laterally clothed with mixed griseous scales. Legs, obscure aeneous; femora with white scales

inwardly. Antennae, rather short and stout; above, sub-annulate; “club,” blackish; beneath, more plainly annulate with

yellowish white, the apices tinged with dark fulvous.

Wings, broad, somewhat longitudinally wrinkled; above, lustrous olivaceous brown, with dark olivaceous longer

scales. Anterior wings, above, without markings, except two, unequal, pale yellowish, obliquely placed maculations at

the base of the interspaces between the upper median nervules. Of these, the upper is much the smaller, and is brought

nearer the external margin than the lower; beneath this latter there are faint indications of a third, paler spot, above the

internal nervure. Base of the wing costal region, sparsely clothed with obscure aeneous-yellow scales; fringes,

prominent, whitish, with a faint darker basal shade. Posterior wings, immaculate, with longer, aeneous olivaceous and

yellowish hairs spreading over the disc and along internal margin, this latter as long as the abdomen, anal angle slightly

and rather broadly exserted; fringes as on anterior wings. Beneath, the anterior wings are evenly covered with obscure

aenrous or golden yellow scales, which are very prominent along costa, leaving the lower portion of the wings, above

internal margin, free; the two yellow maculations of the upper surface are here repeated. The posterior wings are

immaculate, concolorous with anterior pair, being almost everywhere evenly covered with golden yellow scales, least

prominently so before internal margin.

Expanse, 1.50 inches. Length of body, 0.70 inches.

Habitat. - Atlantic District. (Philadelphia!)

A little larger than Hesperia pocohontas, Scudder, and resembling this species in the shape of the wings. It will be

readily recognized by the whitish fringes and the simplicity of its markings.

Because we are dealing with bimacula subspecies in this paper, the primary character to be noted in

the original description is the repeated reference to “yellow” or “golden yellow” scaling (or hairs) on both

the fore and hind wings (both dorsally and ventrally). Nominotypical bimacula is bright Dijon mustard

orange (with slightly lighter veins) on the ventral hind wings (Figs. 6 & 7). The dorsal fore wings of male E.

b. bimacula are broadly fulvous in the cell and distal of the stigma (Fig. 1).

This subspecies ranges from the Atlantic coast (between Virginia and New Brunswick, Canada)

west to Ontario, in Canada and to Indiana in the US where it transitions to E. b. illinois (Dodge).

I have examined 102 examples of this subspecies. The following is a summary listing of sites (by

county) from which I have seen specimens. PENNSYLVANIA: Clinton, Huntingdon, Centre, Tioga,

Montgomery; MAINE: Penobscot; MASSACHUSETTS: Middlesex; NEW JERSEY: Burlington, Ocean;

WEST VIRGINIA: Hampshire; MICHIGAN: Crawford, Newaygo, Jackson (this southeastern Mich,

population is near illinois).

Euphyes bimacula illinois (Dodge, 1872) was described from Bureau Co., Illinois where Dodge

recorded it as being “.. .quite abundant upon grassy slopes on the high rolling prairie that forms the divide

between the Illinois and Rock Rivers.” He took a series of over forty specimens, nine of which were

females. None of these were designated as types. In fact, he offered them in exchange to anyone who could

provide him with specimens of “...North American butterflies not of common occurrence in Northern

Illinois.” He gave no illustration of his new skipper.

Ray Stanford (in Ferris and Brown, 1981) recognized Illinois as a valid subspecies. I concur with

this position based on my personal experience with this subspecies during the 1960’s and 70’s in my home

state of Iowa. It differs from nominate bimacula in that it is larger, more fulvous on the dorsal forewings

(especially in females), and most significantly, grayish (or olive) orange rather than yellowish orange on the

ventral hind wings (Figs. 16 & 17).

This prairie subspecies was once abundant from Colorado to Indiana. Today it is found only in a

few remnant prairies of that region. I encountered it in such prairies in Iowa. Its range is very similar to that

of Boloria selene nebraskensis (Holland, 1928). Nebraskensis and Oarisma powesheik (Parker, 1870)

are often found with it.

I have examined 63 specimens of E. b. illinois from the following states (by county). ILLINOIS:

Cook, Grundy; IOWA: Johnson, Howard, Guthrie; MINNESOTA: Pine; WISCONSIN: Jackson;

COLORADO: Yuma.

I have not been able to locate any of Dodge’s syntypes. The best lead I had was that some of these

specimens may have been deposited in the Field Museum in Chicago and from there to the Allyn Museum. I

received no reply from my inquiry to the Allyn Museum about the possibility of any of Dodge’s specimens

being there. In the apparent absence of a type, I herein designate a male (Figs. 11 & 16) taken 22 June 1968,

Denine Crossing, Grundy County, Illinois, leg. N. G. Seaborg as NEOTYPE of Hesperia illinois Dodge,

1872. This location is only 45 km east of Bureau County. This specimen in deposited in the Florida State

Collection of Arthropods, Gainesville, Florida. If any syntypes are found, I withdraw this specimen as

neotype only on condition that such syntype is designated as lectotype. I here redefine the type locality more

scientifically as a geoecological type locality as follows: high prairies of north-central Illinois.

E. bimacula bimacula is a descendant of E. b. illinois and not the southeastern population. This is

evidenced by the continuous range and similar size and phenotype of these two subspecies. If we had to

recognize only two subspecies, they would be the smaller and much darker southeastern subspecies I

describe below and a northern one, bimacula , which would range from New England to Colorado.

One of the most erroneous assumptions that persists among many taxonomists is the general idea that

the primary line of evolution for the taxa in the northeastern US ascends from the Florida refugium. The fact

is that nearly all the northeastern taxa which now inhabit the formerly glaciated areas of the northeast are

derived from western sources. This is especially true for species whose larvae feed on grasses and sedges.

Undescribed Southeastern bimacula subspecies. I have seldom encountered Euphyes bimacula

here in the Southeast, and when I have, it has never been in any numbers. I have collected it in Chatham

County, Georgia with Dr. R. T. Arbogast and in Berkeley County, South Carolina near Summerville. I

believe it to be extinct at the Summerville location. It appears this extinction was caused by drainage of the

area by Westvaco (lumber/paper) for the purpose of municipal/industrial development and pine tree

farming. Hopefully, this taxon is still extant in as yet undiscovered populations in Berkeley County.

It has long been the general opinion of skipper collectors that the small, dark southeastern

populations of E. bimacula from the Green Swamp of southeastern North Carolina southward represent an

undescribed subspecies (Figs. 21,22,26 & 27). Throughout this region, specimens are markedly darker

above and below in comparison to the two northern subspecies and average smaller. Evolutionally, it has

long been separated from the northern/westem subspecies.

The most probable reason for this taxon having remained undescribed for so long is that few people

have very many of them and usually from only one location. Its scarcity is evidenced by the bimacula

distribution map on page 254 in Opler and Krizek (1984). I know of no one familiar with it who has ever

questioned its subspecific distinctness. Mr. Harry Pavulaan was also interested in describing this taxon, but

has graciously deferred to me because I have been working on it longer. I appreciate his input and

encouragement to proceed.

Euphyes bimacula arbogasti Gatrelle, new subspecies.

A NEW SUBSPECIES OF POANES AARONI

Review of 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.

However, Holland (1931) gives a figure of a male “paratype” on PL. XLVI, fig.37. This specimen is in the

Carnegie Museum NH, Pittsburgh and is probably actually the type. It is a typical aaroni male.

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

(Figs. 3,4,8 & 9). Unfortunately, some researchers continue to follow Clark and Clark (1951) who

introduced the idea that all populations from New Jersey to Miami, Florida are but one subspecies. The

Clark’s clearly acknowledged that the populations in New Jersey were recognizable, then offhandedly

dismissed this by attributing the differences of aaroni to “nothing more than a depauperate light and

somewhat undersized variety of the species.” New Jersey aaroni are not undernourished faded dwarfs.

They are an easily recognizable, valid subspecific taxon.

Both P. a. aaroni and P. a. howardi (Skinner, 1896) are broadly fulvous dorsally. However, the

fulvous is more extensive and lighter in nominotypical aaroni - especially in females. When atypically

large dark males of aaroni and atypically small light males of howardi are encountered they indeed look

very much alike dorsally. However, New Jersey aaroni and Floridian howardi are quite different ventrally

(as attested to by Holland, page 391). On the ventral hindwing, the vast majority of howardi are marked like

Poanes viator zizianae Shapiro (1971) - with a long light central dash and three to four small light spots

(see the description of howardi below). The ventral hindwings of nearly all aaroni are marked more like

those of Euphyes dion (W.H. Edwards, 1867) with a faint central light streak and no spots.

I consider true P. a. aaroni to be confined to New Jersey and Delaware. However, it is possible

that it ranges westward to the eastern and western shores of the upper Chesapeake Bay in Maryland (Fales,

1974). It is not found in Virginia by the Clark’s own determination. (They state that many VA specimens are

noticeably darker then either Floridian or New Jersey specimens. We will examine this more later.)

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. All but two of these

were simply labeled as being from Florida. The two with site specific labels are stated as being from

Georgiana on the Indian River. This is the eastern coastal area of Florida. The taxon was not figured and no

specific specimen was designated as the type. However, Holland (1931) figures the male type on PI. XLVI,

fig.38. This specimen is absolutely typical of this subspecies and thus represents it well. The original

description is as follows (key elements differentiating aaroni, howardi, and minimus are in bold):

Pamphila howardi, n. sp. Male. - Expands 1.50 inches. Upper side: Superiors tawny with a fuscous border a

little more than one-eighth inch in width; there are from one to four small subapical tawny spots in the fuscous

border; at end of cell a dark spot which may or may not be connected with the stigma; stigma rather more than an

eighth inch in length, very narrow and unbroken, and extending to inner margin. Inferiors have the same fuscous

border and tawny central area. Under side: Superiors with tawny central area and border same as upper side; there is

a large triangular spot extending into the wing from the base. The tawny colour above this spot is of a darker hue than

that below and outside of it. Inferiors very light brown, generally with four or five very faint tawny spots in the

The females are larger, without the stigma and have the underside of the inferiors immaculate. Described from

eight specimens in my own collection and four in that of the U.S. National Museum, through the courtesy of Prof.

L.O. Howard. They are all from Florida; two being from Georgiana, on the Indian River; exact locality of others

unknown. This species belongs to the arpa, palatka, Aaroni, viator group. It is a much larger species than Aaroni;

it has not the bright immaculate inferiors below like arpa and differs from palatka in the stigma, which in that

species is in two short sections. The superiors in viator above are fuscous, covered with tawny spots.

I have examined over 300 spread howardi and observed hundreds more in the field over the last 30

years. The mounted specimens I have examined are from the following states (by county). FLORIDA:

Baker, Dade, Duval, Levy, Putnam, Seminole, Volusia; GEORGIA: Bryan, Chatham, Glynn; SOUTH

CAROLINA: Beaufort, Colleton, Charleston, Jasper; NORTH CAROLINA: Currituck; VIRGINIA: Princess

The populations from Miami, Florida to Beaufort, South Carolina are very similar in size and

markings, dorsally and ventrally. The populations in this area are broadly fulvous above and the vast

majority of specimens (especially males) have the viator- like streak and spots on the ventral hindwings (as

highlighted in the original description).

From Beaufort, South Carolina through Virginia most specimens are still broadly fulvous above but

many specimens are encountered without the v/stfor-spotting of the ventral hindwings. In some areas the

ventrally unspotted phenotype dominates. These are the darker specimens mentioned by the Clarks in the

Butterflies of Virginia on page 170. Despite the somewhat atypical appearance of some of these mid-

Atlantic populations, they are still part of subspecies howardi and should be referred to as such. I believe

the tendency for specimens in this area to be darker and to lose the ventral HW spots is due to past genetic

contact with the freshwater inland subspecies P. aaroni minimus described below.

Undescribed Texas P. aaroni subspecies. It is my opinion that the aaroni population in east

coastal Texas is most likely an undescribed subspecies (Figs. 25,30,35 & 40). I only have four worn

specimens of this in my collection but they are unlike anything I have seen from the eastern US. I would

want to examine several more before describing it. They are very large and orange fulvous rather than

yellow fulvous. Their wings are quite rectangularly elongate and pointed at the apex (the wings of the three

eastern subspecies are more square). The viator- spotting on the ventral hindwings is also very prominent. I

strongly encourage someone in Texas who has a good series of these to describe it.

Undescribed inland South Carolina P. aaroni subspecies. On 17 May 1991 I was amazed to

discover an inland fresh water population of Poanes aaroni at Bull Swamp in Orangeburg County, South

Carolina. I found it common and by far, the dominant skipper at this site in May, June and September of

1991, 1992, and 1993. (I have not visited the site since.)

North of Florida, aaroni is only known from coastal salt or brackish marshes. According to Opler

and Krizek (1984), this skipper is “...restricted to coastal salt marshes, usually in association with smooth

cordgrass ( Spartina alternifolia ).” Gochfeld and Burger (1997) state that P. aaroni aaroni is restricted to

salt marshes on both coasts of southern New Jersey. The fresh water South Carolina colony is surrounded

by dry sandhill habitat and is located 120 km. inland and at least 100 km. from the nearest populations in

Charleston and Colleton counties, South Carolina.

I know of only one other fresh water record from this region. This is found in Harris (1972) and is

by H. W. Eustis for Richmond County, Georgia in the late 1940’s. Harris figures a male from that

population on plate 10. This specimen looks like a good P. aaroni howardi as it is brightly colored and

quite unlike the dark Bull Swamp population. It is likely that Eustis’ specimens were transients from the

sizable howardi population that inhabits the vast swamps along the Savannah River from the coast inland

into at least Screven County.

The Bull Swamp population is distinctive in several aspects. First, its sandhills location, fresh

water ecology, and dark phenotype indicates that it is evolutionally distant from all coastal populations from

Florida to New Jersey, and as such, has long been isolated from the other subspecies in time and space.

Second, its phenotype is consistently distinct from both aaroni and howardi.

Therefore, I describe this population as a new subspecies, Poanes aaroni minimus.

Gatrelle, new subspecies.

A NEW SUBSPECIES OF HESPERIA ATTALUS

Review of attalus subspecies.

Hesperia attalus attalus (W.H. Edwards, 1871) was described from two females collected by

G.W. Belfrage near Waco, Texas. One of these specimens was figured (ventral aspect) by Holland (1931),

PI. LII, fig. 24. This same specimen was designated as lectotype by F.M Brown and L.D. Miller (1977).

H. a. attalus is a bright, colorful Hesperia as can be seen from figs. 31,32,36 & 37. This western

prairie subspecies averages slightly larger than the Eastern US populations. This western subspecies has

been widely isolated from its eastern counterparts for thousands of years. There are no blend zone

populations between western and eastern attalus.

I have no personal experience with this subspecies. However I have examined 47 specimens from

the following states (by county). OKLAHOMA: Cleveland; KANSAS: Barber; TEXAS: Smith. The

majority of these are from the J.R. Heitzman collection which is now housed in the FSCA, Gainesville.

Hesperia attalus slossonae (Skinner, 1890) was described from one male and one female

collected by Mrs. A.T. Slosson. The origin of the specimens is simply given as Florida. The original

description is as follows (I have highlighted key phrases in bold):

Pamphila slossonae n. sp. - In size and markings this species comes nearest to P. leonardus Harris. The male

expands rather less than an inch and a half. Ground color of inferior [superior] wings dark brown, with basal half of

wing thickly covered with tawny scales which, beyond the stigma, form three spots, and just above these are two

small square ones. The three subcostal spots are represented as in leonardus. Stigma narrow, black, and concave

posteriorly. Inferior wings same color as superior, with tawny scales scattered over the basal half. There are four

small, square, tawny spots close together, with a fifth elongated one at right angles with the four, on outer half of wing

running nearly parallel with the outer margin, only the angle made by the spots is more acute. Fringes of all wings

dingy white. The maculation on underside of superiors is nearly the same as in leonardus, but in color very different,

the ground color in this species being made up of yellowish and greenish scales; underside of inferiors olive-green,

with the spots of the upperside repeated, except there is a sixth one near the centre of the wing, and all are dingy white.

Fringes on underside same as above. Head and thorax above covered with greenish hair, beneath grayish yellow. Palpi

almost white.

The female expands a little more than an inch and a half. Superior wings very dark brown, with a broken band of

yellow consisting of five spots commencing at the first nerve above the interior margin and extending to within one-

eighth inch of apex. The subcostal spots are present, as in the male, with spots fainter; underside of wings also same

as in male, but white spots on inferiors smaller. Fringes in female dark on superiors, but gradually getting lighter on

inferiors as the anal angle is approached.

Over the last 30 years, I have encountered this subspecies at several locations throughout the

southeast. My personal records are from (by county): FLORIDA: Escambia, Levy, Marion; GEORGIA:

Burke; SOUTH CAROLINA: Aiken, Orangeburg, Chesterfield; NORTH CAROLINA: Hoke. The great

majority of the several hundred individuals I have observed or collected from these widely separated areas

are of the same well marked phenotype described in the original description (Figs. 33,34,38 & 39). In

females there is moderate ventral variation toward brownish, non-greenish-yellow individuals.

I have also examined 73 slossonae in institutional and private collections from the following states

(by county): FLORIDA: Levy, St. Johns, Bradford, Putnam; NEW JERSEY: Ocean; GEORGIA: Crawford.

Throughout their range, all slossonae males, and most females, exhibit at least some golden tawny

scaling on the ventral hindwings. The ventral hindwings of most males have a great deal of golden over¬

scaling (unless they are worn) and a prominent row of cream spots in the upper postmedian area. The

ventral hindwing varies quite a bit in female slossonae , but possesses at least some cream spotting in the

postmedian area - with many females having a well developed contrasting row of creamy white spots. The

dorsal forewings of males are usually brightly tawny immediately distad of the stigma with conspicuous

tawny scaling in the upper part of the cell and prominent apical spots. On their dorsal hindwings, male

10

slossonae usually have a conspicuous amount of golden fulvous hairs, especially near the inner angle, and a

well developed row of upper postmedian spots. The dorsal forewings of female slossonae are variable.

Some have a well developed row of postmedian spots, including the apical area, while others only have a

few spots in the postmedian area. The dorsal hindwings of females are also variable with the postmedian

spotting ranging from prominent to faint. However, almost all females have some dorsal hindwing spotting.

All of these key traits are mentioned in the original description.

MacNeill (1964), in his classic work on the western American Hesperia , briefly delves into the

origins of the eastern species of this genus. His position is that the eastern species, and particularly the

southeastern taxa, represent the oldest and thus most phylogenetically stable assemblage of species in the

genus (which originated in the Nearctic). In other words, the southeastern taxa are all very old and highly

evolved. On page 10 he states: “In [the] eastern United States, and particularly the southeast, a small group

of relatively unrelated species occurs, none of which shows close relationships to any other known species.

Apparently little geographic diversity is expressed by any of these.”

I concur with MacNeill. This is evidenced by the great stability of the slossonae phenotype from

central Florida to New Jersey - where the only difference is that individuals become smaller northward.

This homogony is the backdrop that manifests the evolutionary importance of a newly discovered, isolated,

phenotypically unique population of Hesperia attalus.

Undescribed coastal South Carolina H. attalus subspecies. On 3 October 1995 I discovered a

population of Hesperia attalus near the South Carolina Coast at the southern tip of Horry County. This site

is on private property near the newly designated Sandy Island National Wildlife Sanctuary. The elevation at

the site is about 100 ft and is part of the Sandy Island remnant dune uplift. Prior to the advent of Europeans

and deforestation, this population was separated and isolated from the inland sandhill populations of

slossonae (Figs. 33 & 36) by approximately 50 to 80 km of (often swampy) forest. Even today, it remains

separated from the inland subspecies by at least 35 km.

This population is distinctive on two levels.

First, MacNeill’s determination that all southeastern Hesperia are evolutionarily old,

phenotypically stable taxa, requires that this uniquely distinct and isolated segregate has 1) been in

existence a very long time, and 2) is evolutionarily distant (subspecifically distinct) from slossonae.

Second, it is just as distinct phenotypically from H. attalus slossonae as H. attalus slossonae is from H

attalus attalus. It is very dark and melanistic, especially in the male.

Its geographic location dictates that it most likely came into being in only one of two ways. It most

likely is a relict of an attalus that once inhabited the coastal area when ocean levels were much lower. Its

range then extending out, many miles in some areas onto what is now the Continental Shelf, up and down the

mid-Atlantic coast in semi-barren sandy habitats (Emery et. al., 1967; Remington, 1968). Thousands of

years of shore line compression have pinned this taxon between the Atlantic Ocean and the Maritime

Forests of the eastern seaboard. If this is so, it is an isolated taxon in the process of geologically induced

extinction. This may be the only population still extant.

The only other plausible course of evolution is that it is a descendent of inland ancestors whose

range was once connected to the coastal area. When this connection was severed by increased forestation,

this subspecies evolved in situ. This scenario, on the surface, seems supported by the presence of other

species at this site which are also natural parts of the inland sandhill community - most notably Hesperia

meskei straton (W.H. Edwards). However, this course is not the most likely because the other members of

the biota at this site, which are common to the sandhills, remain subspecifically the same. The uniqueness of

the attalus at this site indicates that it had already evolved to its present taxonomic state before it arrived

(from an area now offshore). MacNeill concluded that the evolution of all southeastern Hesperia was

settled millennia ago. For these reasons, I now describe this population as a new subspecies.

11

Hesperia attalus nigrescens Gatrelle, new subspecies.

,'Sirs*”

i tiny patch at the distal end, in the one exception the tawny is in the form of

range of this subspecies and its relation to slossonaet

, in light of the rapid destruction of the world’s living

My thanks go to the following persons. To Dr. J.D. Lafontaine (Canada), FSCA library, and the U. of Florida

Science Library, for reprints of original descriptions; my son Ben Gatrelle for photography of specimens, Dr. John

Heppner FSCA for access to the FSCA collections, and to the congregation of my church for affording their pastor

(me) the time to research bugs.

LITERATURE CITED

BROWN, F.M. & L.D. MILLER. 1977. Trans. Amer. Ent. Soc., 103:281-282. (In Part. Series on the Types

Named by W.H. Edwards)

CLARK, A.H. & L.F. CLARK, 1951. Butterflies of Virginia. Smithsonian Misc. Collect. 116 (7) 239 pp.

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