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LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


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22 April 1976 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


RonaLp W. Honcess, President 


S. S. Nicouay, President-elect Oxar H. H. Mie.xe, Vice President 
Donatp R. Davis, Ist Vice President Joun M. Sniper, Treasurer 
Witu1aM Hovanirz, Vice President Lee D. MILLER, Secretary 


Members at large (three year terms): 


D. C. Fercuson (1975) J. T. Brewer (1976) C. V. CovELL, Jr. (1976) 
R. O. KENDALL (1975) K. S. Brown (1976) D. F. Harpwick (1977) 
J. A. Powetxi (1975) K. W. Puiie (1976) J. B. Zecier (1977) 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and 
formally constituted in December, 1950, is “to promote the science of lepidopterology 
in all its branches, .... to issue a periodical and other publications on Lepidoptera, 
to facilitate the exchange of specimens and ideas by both the professional worker and 
the amateur in the field; to secure cooperation in all measures” directed towards 
these aims. 

Membership in the Society is open to all persons interested in the study of 
Lepidoptera. All members receive the Journal and the News of the Lepidopterists’ 
Society. Institutions may subscribe to the Journal but may not become members. 
Prospective members should send to the Treasurer full dues for the current year, 
together with their full name, address, and special lepidopterological interests. In 
alternate years a list of members of the Society is issued, with addresses and special 
interests. There are four numbers in each volume of the Journal, scheduled for 
February, May, August and November, and six numbers of the News each year. 


Active members—annual dues $13.00 
Student members—annual dues $10.00 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $18.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
John M. Snider, 3520 Mulldae Ave., San Pedro, Calif. 90732 U.S.A. 


Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA 


by Cyrit F. pos Passos 


Price: Society members, $5.00 U.S.; non-members, $7.50 U.S. Paper covers, revisions 
of the Melitaeinae and Lycaenidae supplied separately. 


Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni- 
versity of Louisville, Louisville, KY 40208, U.S.A. 


The Lepidopterists’ Society is a non-profit, scientific organization. The known 
office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second 
class postage paid at Lawrence, Kansas, U.S.A. 66044. 


JOURNAL OF 


Tue LeEpPIDOPTERISTS’ SOCIETY 


t Volume 30 1976 Number 1 


PRESIDENTIAL ADDRESS 1975— 
TO MY FELLOW AMATEURS 


ANDRE BLANCHARD 
P. O. Box 20304, Houston, Texas 77025 


The title of Dr. Rindge’s Presidential Address to the twelfth annual 
meeting of the Pacific Coast Section at San Diego, California (1965) was: 
“The Importance of Collecting—Now’. In it he explained that we were 
—we still are of course—in a losing race against the spread of civilization 
and concomitant destruction of breeding grounds for Lepidoptera. His 
address was a plea for immediate and intensive collecting of butterflies 
and moths. I am sure that few professionals needed such a reminder but 
I hope that many amateurs listened and followed his advice. Yet, and 
in full agreement with everything that he said, I would like to add my 
two cents worth of comments to Dr. Rindge’s appeal. 

The number of amateurs interested in moths is disproportionately 
smaller than that of those interested in butterflies, yet there are many 
times more species of moths than of butterflies. If there are, say, ten 
times more species of moths and ten times less people interested in them, 
this is a deficiency factor of one hundred against the moths. In such a 
situation it is indeed very likely that many, many species of moths will 
disappear from the earth before they are seen by a single taxonomist. 
How can we hope to redress such an imbalance? By showing that study- 
ing moths can be more rewarding because there is a lot more to be 
discovered about them. 

Before I go any further, let me tell you that I am very much aware 
of the fact that we do not have enough collectors even of butterflies, 
that we would like to have several times more of them. Nothing that I 
am going to say should be interpreted as an attempt to dissuade anybody 
from collecting butterflies: my ambition is only to convince some ama- 
teurs to work on moths in parallel with butterflies and to kindle an inter- 
est in moths in some neophytes. 


JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


bo 


Most of us come to lepidopterology for the pleasure of assembling a 
beautiful collection. We may differ as to how geographically extensive 
it will be or as to which groups of Lepidoptera will be included, but I 
think that nearly all of us start that way. Let me tell you, between 
parentheses that, from my contacts with many professionals, I have come 
to the conclusion that most of them are dyed in the wool amateurs who 
have found a way of getting paid for doing what they would gladly do, 
at their own expense, if they could afford to do so. 

However, many of us amateurs can not be happy for long, with collect- 
ing Lepidoptera the way others collect postage stamps. Mere collecting 
of course is all right and quite satisfying for many, but those who gradu- 
ate to the studying stage find their hobby considerably more satisfying. 
The key words are studying, doing research, and as there is immensely 
more that is unknown and unexplored among the Heterocera, the chances 
of hitting on something new are considerably better with moths than 
with butterflies. Where is the amateur who is not thrilled when he 
discovers something new like a wide extension of range, a new food- 
plant, a new detail of a life history or a species new to Science? Moths, 
just because they are not overall as pretty as most butterflies, reserve 
their rewards for those who study them a little more deeply. 

Lepidopterology is a wonderful science in which you do not need 
to be an expert—as you would have to be in Mathematics, Physics or 
Biology for instance—to do some simple, simple but useful research. 

A good point of moths is that they are rather easy to collect: this is 
not the place to explain how, but I rather like my method which con- 
sists of setting the traps at sunset, going to sleep, and gathering the 
loot at daybreak. The only disadvantage of this method is that you 
miss the day flying species and those which come only at bait. 

Moths, of course, are divided in Macroheterocera and Microheterocera. 
Both groups are replete with discoveries waiting to be made, but the 
micros are even more so. 

One of the most promising avenues to discovery is through rearing of 
larvae: rearing and careful field observations are research at its best, 
and this type of activity requires only very inexpensive equipment. When 
you come to think that in some good size genera, the larva or the food- 
plant of not a single species is known, you realize that by rearing all the 
larvae that you can procure, you stand a more than even chance of dis- 
covering from time to time something well worth your trouble. Good 
records of everything should of course be maintained, some larvae and 
pupae preserved by pickling or dehydrating. 

There was a time, late in the nineteenth and early in this twentieth 
centuries when entomologists vied for the description of new species. 


VoLUME 30, NuMBER 1 3 


Today, it sometimes seems that they need an excuse to do so, unless it 
is in the course of a group revision. These group revisions, however, 
which have become their main activity, bring forth amazing results: 
hardly any genus is revised without recognizing several new species 
which had been overlooked, or splitting the genus into two or more 
genera. In any case, these group revisions necessitate bringing together, 
and making available to the reviser, all the material available in all the 
museums and the private collections. If you have any material available 
in the genus being revised, this is your chance of contributing something 
valuable which will, of course be fully acknowledged. 

I have already mentioned that the micros are even less well under- 
stood than the macros. This, really is an understatement: some groups 
of micros are so poorly known that it is almost useless to request identi- 
fication of specimens: more than half return with at most a genus name. 
These are orphan groups and you do not have to go very far down the 
check list of micros to find such groups: the Epipaschiines—a subfamily 
of the Pyralids—are a good example. I have assiduously collected them, 
prepared male and female genitalia and discovered that they are too 
similar to be of much help, on top of that several species look alike and 
there is some sexual dimorphism. I may be wrong but I am inclined to 
think that this is, in part at least, why these groups have been left alone 
and that, in cases like that, the success of a revision and the willingness 
of a potential reviser to tackle the job depend on the availability of 
good, long reared series of specimens with preserved larvae, pupae, eggs 
and careful records of everything including foodplants. Professionals will 
take care of all this for economically important species, but we, amateurs, 
could and should do it for all kinds of species. These would be wonderful 
projects for young amateurs. I say young, because the fruition of their 
efforts may come only after several years of routine and not immediately 
rewarding work. But would not the result be worth the effort? 

Will my plea bring forth the hoped for response? Not if I merely urge 
the amateurs to go ahead blindly into a world unfamiliar to most of 
them. This is where I turn to the professionals and tell them: we want. 
we need your help! I wonder if one of you “Pros” would come forth, 
prepare for us and publish in the Journal a clear and easy guide to the 
moths and particularly the micros. I do not mean something as complete 
and forbidding as Forbes’ key in Volume One of his “Lepidoptera of New 
York and Neighboring States,” but a summary of the most easily observed 
characters of the different families and—where possible—subfamilies, 
well illustrated with sketches showing palpi, antennae, wing shapes, types 
of maculation, etc. This, in my opinion, is the most helpful tool that the 
professional could give the amateur. 


4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


I directed my address to “My Fellow Amateurs” because I meant it as 
a plea to some of them, who may not realize that the study of the moth, 
less glamorous perhaps than the study of the butterfly, is in the long 
run more rewarding. But, to the enthusiasm of the amateur, let the 
professional add the support of his knowledge. Let him give us the guide 
we need, let him stake out the road to lead us into the strange but 
wonderful world of the humble moth. 


CAPTURE OF A HYBRID LIMENITIS ARTHEMIS ASTYANAX) X L. 
ARCHIPPUS (NYMPHALIDAE) IN SOUTHERN GEORGIA 


On 22 September 1974 a recently emerged male hybrid between Limenitis arthe- 
mis astyanax (Fabricius) and L. archippus (Cramer) (form rudidus Strecker) was 
captured near Fort Jackson, Savannah, Chatham Co., Georgia. Limenitis archippus 
was abundant in the area, but L. a. astyanax was not observed. The specimen (Fig. 
1) is presently in the author's collection. 

Platt & Greenfield (1971, J. Lepid. Soc. 25: 278-284) reported the capture of a 
similar interspecific hybrid in North Carolina and listed 12 previously known records 
of such hybrids. An additional North Carolina specimen was reported by Green- 
field & Platt (1974, J. Lepid. Soc. 28: 72-75). The individual captured at Savannah 
and the two from North Carolina are apparently the only known records of this form 
from the southeastern U.S.A. 


RicuHArp T. Arsocast, 114 Monica Blvd., Savannah, Georgia 31406. 


Fig. 1. Male L. arthemis astyanax ~% L. archippus collected near Fort Jackson, 


Savannah, Georgia, on 22 September 1974 Jobin In, 1h Arbogast. Left: dorsal surface; 
right: ventral surface. 


VoLUME 30, NuMBER 1 5 


NEW HESPERIIDAE RECORDS FOR TEXAS AND THE 
UNITED STATES 


WitutiamM W. McGuire 
P.O. Box 29884, San Antonio, Texas 78229 


AND 


MicHAEL A. RICKARD 
4628 Oakdale, Bellaire, Texas 77401 


The Rio Grande Valley of Texas, located in the extreme southern 
section of the state and encompassing areas of essentially neotropical 
habitat, offers a unique opportunity for the study of Lepidoptera in the 
U.S.A. The authors have had the good fortune to collect this area rather 
frequently during the past several years and during that time some inter- 
esting and important new records of Hesperiidae have been obtained. 

During 1972-1974 specimens of several rare Hesperiidae, previously 
known in the U.S.A. from only a few examples, were taken: Aguna 
asander (Hewitson), Aguna claxon Evans, Typhedanus undulatus (Hew- 
itson), Polythrix mexicanus Freeman, Proteides mercurius (Fabricius), 
Urbanus doryssus Swainson, Panoquina fusina evansi (Freeman), As- 
traptes gilberti Freeman, Carrhenes canescens (R. Felder), Gorgythion 
begga pyralina (Moschler) and Lerema ancillaris liris Evans. In addition 
12 species of Hesperiidae were taken that represent apparent new U.S.A. 
records, another that is at least a new Texas record, and two others that 
substantiate previous but litle known Texas records. Nomenclature and 
arrangement follows that of fvans (1952, 1953, 1955) and determina- 
tions, unless otherwise indicated, were made by Rickard. 


Epargyreus exadeus cruza Evans. Fig. 1. McGuire collected 1 2 in McAllen, 
Hidalgo County on 18 October 1973 as it fed on blossoms of Queen’s Crown, 
Antigonon leptopus H. and A. (Polygonaceae). Previous records for this species are 
confusing: Holland (1931), p. 330, listed E. exadeus (Cramer) as “a straggler in 
southern California, New Mexico and Arizona.’ However, Evans (1952) stated 
that what Holland illustrated was E. exadeus cruza rather than exadeus exadeus 
(Cramer), and noted other distribution for cruza as Mexico, Guatemala, Nicaragua, 
Salvador, Costa Rica, and Panama (transitional to exadeus exadeus). This is the 
first known record of this skipper for Texas. 

Aguna metophis (Latreille). Fig. 2. Rickard took a worn @ in Bentsen-Rio 
Grande Valley State Park, Hidalgo County, 6 September 1969, det. H. A. Freeman. 
Other Texas records include Mission, Hidalgo County, 10 September 1972, 1 Q 
(Roy O. Kendall); Santa Ana National Wildlife Refuge, Hidalgo County, 10 and 
27 October 1973, 1 ¢ each date (Rickard); Loop 374 west of Mission, Hidalgo 
County, 18 October 1973, 1 &, and 19 October 1973, 1 @ (McGuire) and 26 


6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


S 


Figs. 1-5. 1, Epargyreus exadeus cruza Evans, 9, ventral, McAllen, Texas, 18 
October 1973; F 28 mm. 2, Aguna metophis (Latreille), 9, ventral, Loop 374, 
Mission, Texas, 19 October 1973; F 23 mm, tail 10 mm. 3, Codatractus alcaeus 
alcaeus (Godman & Salvin), 2, ventral, McAllen, 19 October 1973; F 26 mm. 4, 
Urbanus pronta Evans, 6, ventral, Ciudad Victoria, Tamaulipas, Mexico, 24 De- 
cember 1973; F 23 mm, tail 5 mm. 5, Urbanus esmeraldus Butler, ¢, ventral, 
McAllen, 18 August 1972; F 20 mm, tail 10 mm. 


October 1973, 1 ¢@ (Rickard); Galveston, Galveston County, 7 August 1973, a 
worn @ (McGuire). Distribution noted by Evans (1952) includes Mexico, Nicar- 
agua, Panama, Venezuala, Matto Grosso and South Brazil. 

Codatractus alceaus alceaus (Godman & Salvin). Fig. 3. The first record for 
this species in Texas was given by Freeman (1951) as a single specimen from 
the Davis Mountains, no data. A single worn specimen was collected and reported 


VoLUME 30, NUMBER | 74 


10 


Figs. 6-10. Astraptes egregius egregius Butler, 2, dorsal, Bentsen-Rio Grande 
Valley State Park, Texas, 18 October 1973; F 24 mm. 7, Astraptes alardus latia 
Evans, ¢, ventral, San Fernando, Tamaulipas, Mexico, 23 October 1973; F 28 mm. 
8, Achalarus jalapus (Plotz), ¢, ventral, Sullivan City, Texas, 31 July 1972; F 
25 mm. 9, Bolla clytius (Godman & Salvin), ¢, dorsal, Abrams, Texas, 18 October 
1973; F 15 mm. 10, Sostrata bifasciata nordica Evans, é&, dorsal, Bentsen-Rio 
Grande State Park, 26 October 1973; F 15 mm. 


by J. Richard Heitzman (1970) from Boca Chica, Cameron County, 27 June 1968. 
On 19 October 1973 McGuire collected 1 ¢ at Penitas and 1 @ at McAllen, both 
Hidalgo County. Obviously uncommon in Texas, it is at times abundant at Ciudad 
Valles, San Luis Potosi, Mexico (H. A. Freeman, pers. comm.) which is about 500 


8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


mi south of the Texas border and was recorded by Evans (1952) from Honduras, 
Nicaragua and Costa Rica, as well as Mexico. 

Urbanus pronta Evans. Fig. 4. Rickard took a @ along a railroad track near 
Madero, Hidalgo County on 19 October 1969. It was visiting blossoms of Lantana 
horrida H. B. K. It was papered as U. proteus (Linnaeus) and the true identity 
not discovered by the author until September 1973. Previous distribution records 
are from throughout Mexico and Central America south to Paraguay (Evans, 1952). 

Urbanus esmeraldus Butler. Fig. 5. A single specimen of this species was found 
by McGuire among his Urbanus material following Rickard’s discovery of U. pronta. 
The specimen is a fresh 6, collected 18 August 1972 at McAllen. This species is 
rather widespread from Mexico south to Paraguay (Evans, 1952) and at times 
rather common in southern Tamaulipas, Mexico, within 350 mi of the Texas border. 

Astraptes egregius egregius Butler. Fig. 6. A @ was taken by McGuire, 18 
October 1973 in Bentsen State Park. The usual habitat of this species is central 
Mexico south to the Amazons (Evans, 1952) and this record, as confirmed by Mr. 
H. A. Freeman, represents a significant range extension as well as a new United 
States record. 

Astraptes alardus latia Evans. Fig. 7. Rickard took three specimens in 1973: 
Bentsen State Park, 16 June (1 worn 2) and 10 October (1 fair 9), and Santa 
Ana Refuge, 1 September (1 fresh ¢). All were collected in wooded areas as they 
rested under large leaves. This distinctive Astraptes has been recorded from Central 
America and Colombia by Evans (1952), and found rarely by McGuire in Ta- 
maulipas, Mexico, within 200 mi of Texas. 

Achalarus jalapus (Plotz). Fig. 8. A fresh ¢ was taken by McGuire, 31 July 
1972 near Sullivan City in western Hidalgo County. Roy O. Kendall collected 
(and det.) 1 2 on 8 September 1972 and 1. 4 on 9 September 1972 in Mission, 
Hidalgo County. In 1973, Rickard took 2 ¢ 6 at McAllen, 23 September and 20 
October, and John B. Vernon collected 1 @ at the same location on the latter date. 
This species has probably been often confused with the more common A. toxeus 
(Plotz), from which it can be distinguished by the presence of the lobed hindwing 
in jalapus. Recorded by Evans (1952) from Mexico (Jalapa; Granahl; Guada- 
lajara), Guatemala, Honduras and Columbia, it is not uncommon 300 mi south of 
Texas (McGuire). 

Bolla clytius (Godman & Salvin). Fig. 9. On 18 October 1973 McGuire collected 
2 64 and 1 2 southwest of Mission near the village of Abrams, Hidalgo County; 
these were all taken along a wooded canal as they visited Aster blossoms. Determi- 
nation of these specimens prompted the authors to contact Mr. J. W. Tilden, who 
had previously reported Bolla brennus from the Rio Grande Valley (Tilden, 1971) 
and ask that he recheck his Bolla specimens. This was done and Tilden confirmed 
his earlier determination of B. brennus, thus establishing the presence of two 
species of Bolla in Texas and the U.S.A. Evans (1953) records this species from 
only Mexico and Honduras. 

Sostrata bifasciata nordica Evans. Fig. 10. Rickard took 2 ¢ 8 in Bentsen-Rio 
Grande Valley State Park on 26 October 1973. They were patrolling small patches 
of light in a wooded area late in the afternoon. This species has also been found 
rather commonly by McGuire in the Ciudad Victoria area of Mexico, and is recorded 
from Mexico, Guatemala, Honduras, Nicaragua, and Costa Rica by Evans (1953). 

Heliopetes arsalte arsalte (Linnaeus). Fig. 11. McGuire collected a pair of fresh 
specimens at Boca Chica, east of Brownsville, Cameron County, late in the evening 
of 20 October 1973; both were flying about in open chaparral in company with 
Heliopetes laviana (Hewitson) and H. macaira (Reakirt). McGuire had previously 
collected this species within 200 mi of the Texas border, near Ciudad Victoria; 
vans (1953) lists distribution throughout Mexico, Central and South America, 
and Trinidad. 


VoLuME 30, NuMBER 1 9 


15 


Figs. 11-15. 11, Heliopetes arsalte arsalte (Linnaeus), 2, ventral, Boca Chica, 
Texas, 20 October 1973; F 16 mm. 12, Pirwna microsticta (Godman), 2, ventral, 
Sullivan City, 20 October 1973; F 10 mm. 13, Corticea corticea corticea (Pl6tz), 
¢, dorsal, Bentsen-Rio Grande Valley State Park, 3 September 1972; F 12 mm. 14, 
Rhinthon osca (Plotz), ¢, dorsal, Loop 374, Mission, 20 October 1973; F 21 mm. 15, 
Conga chydaea (Butler), ¢, dorsal, Bentsen-Rio Grande Valley State Park, 15 July 
1972; F 14 mm. 


Piruna microsticta (Godman). Fig. 12. Evans (1955) recorded 1 ¢ from Texas 
as well as specimens from Mexico. Holland (1931), p. 361 noted the species as 
occurring in northern Mexico and “reported as having been taken in Arizona near 
the Mexican border.” After finding specimens in arrid terrain in southern Tamau- 
lipas, Mexico, a search of similar habitat near Sullivan City, Hidalgo County, 
led to McGuire’s capture of 1 2 on 20 October 1973, which reinforces Evans’ 
earlier record for Texas. 

Corticea corticea corticea (Plétz). Fig. 13. We have taken a number of examples 
of this rather common Mexican species to date. Rickard collected 1 ¢ at Madero 
on 4 November 1973 and 2 @@ in Bentsen-Rio Grande Valley State Park, 16 
December 1973. A subsequent search of the authors’ collections turned up addi- 
tional records: Bentsen State Park, 3 September 1972, McGuire (1 ¢) and Santa 


10 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Ana National Wildlife Refuge, 9 November 1968, Rickard (1 ¢, 1 9). These speci- 
mens had been mislabeled Nastra julia (Freeman). The obscure appearance and 
superficial resemblance to N. julia has doubtless caused other collectors to overlook 
or mislabel corticea. The species in widely distributed throughout Mexico, Central 
America and South America as far south as Argentina (Evans, 1955). 

Rhinthon osca (Plétz). Fig. 14. Rickard captured a fresh Q in a wooded area 
along Loop 374 west of Mission, Hidalgo County, on 20 October 1973; he collected 
a second @ in good condition south of Mission near the village of Madera, Hidalgo 
County, 26 October 1973 as it visited blossoms of Eupatorium odoratum L. Pre- 
viously, R. osca has been considered a subspecies of the Antillean R. cubana 
Herrich-Schaffer), but the authors are advised by H. A. Freeman (pers. comm.) 
that it should be accorded specific status. McGuire has previously collected 
specimens as far north as Ciudad Mante, Tamaulipas, Mexico, and Evans (1955) 
notes distribution from Mexico south to Ecuador. 

Conga chydaea (Butler). Fig. 15. McGuire collected specimens in Bentsen-Rio 
Grande Valley State Park on 15 July 1972 (1 ¢, 1 @) and 2 September 1972 (1 
6); Rickard collected 2 92 on 25 October 1973 as they visited blossoms of 
Queen’s Crown along a canal in McAllen. This species is relatively common in 
Mexico to the south of Ciudad Mante, and noted by Evans (1955) to be recorded 
as far south as Argentina. 


ACKNOWLEDGMENTS 


The authors would like to thank the United States Department of 
Interior, Bureau of Sport Fisheries and Wildlife, and the Texas Parks 
and Wildlife Department, Interpretations and Exhibits Branch, for per- 
mits making possible Lepidoptera research in Santa Ana National Wild- 
life Refuge and Bentsen-Ric Grande Valley State Park, respectively. 
Additionally, the authors wish to thank the personnel at Santa Ana and 
Bentsen for their assistance and cooperation; Mr. H. A. Freeman for 
confirming determinations, rendering advice and clarifying certain points 
relative to these Hesperiidae; Mr. Roy O. Kendall for his continuing 
support and aid in the ongoing study of Texas Rhopalocera and _ his 
critical review of this manuscript; and to Dr. C. E. Hall of the University 
of Texas Medical Branch, Galveston, Texas for the photographs used in 
this article. 


LITERATURE CITED 


Evans, W. H. 1952. A catalogue of the American Hesperiidae indicating the classi- 
fication and nomenclature adopted in the British Museum. Part II. Pyrginae, 
Sec. 1, London: British Museum, 178 p., pls. 10-25. 

- 1953. A catalogue of the American Hesperiidae indicating the classifica- 

tion and nomenclature adopted in the British Museum. Part ITI. Pyrginae, 

Sec. 2, London: British Museum, 246 p., pls. 26-53. 

= 1955. A catalogue of the American Hesperiidae indicating the classifica- 

tion and nomenclature adopted in the British Museum. Part IV. Hesperiinae 

and Megathyminae. London: British Museum, 449 p., pls. 54-88. 


FREEMAN, H. A. 1951. Ecological and systematic study of the Hesperiidae of Texas. 
So. Meth. Univ. Studies, 6: 1-64. 


VoLuME 30, NuMBER 1 LE 


HeirzMan, J. R. 1970. A new U.S. butterfly record and a migration of Eunica 
monima in Texas (Nymphalidae). Mid-Continent Lepid. Series, 12: 10-11. 

Ho.tuanp, W. J. 1931. The butterfly book. Rev. ed. New York: Doubleday Doran 
and Co., Inc., 424 p., 77 pls. 


TitpEn, J. W. 1971. Aguna claxon (Hesperiidae) new to the United States. J. Lepid. 
SOG.2 25 PABST 


HAPALIA NIGRISTRIATALIS A SYNONYM OF UDEA ANGUSTALIS 
(PYRALIDAE: PYRAUSTINAE) 


In my recent paper on the Udea angustalis group (Munroe, 1974, Can. Ent. 106: 
139-142), I did not consider Hapalia nigristriatalis Hampson (1918, Ann. Mag. 
Nat. Hist. 9(2): 395), described from a single male from San Antonio, Colombia, 
collected by Palmer. Examination of a photograph of the holotype, made by me at the 
British Museum (Natural History) in 1958 (Fig. 1), shows that H. nigristriatalis 
should be transferred to Udea Guenée, 1844, where it becomes Udea nigristriatalis 
(Hampson), new combination, and falls as a synonym of Udea angustalis (Dognin, 
1905), already known to range from southern Mexico to Bolivia. 


EucENE Munroe. Biosystematics Research Institute, Agriculture Canada, Ottawa, 
Ontario, Canada. 5 


Fig. 1. Hapalia nigristriatalis Hampson, 1918, holotype, male, San Antonio, West 
Colombia, Palmer, British Museum (Natural History). Black and white print from 
Kodachrome transparency. Wingspan of specimen 22 mm. The type-label was 
made by Hampson before he began to use the name Hapalia Hiibner for the collective 
genus he had for many years called Pionea Guenée. 


12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


A NEW SUBSPECIES OF ARGYREUS HYPERBIUS 
(NYMPHALIDAE) FROM NEW GUINEA 


Curis SAMSON 
Saruman, St. Giles in the Wood, Beckley, E. Sussex, England 


Argyreus hyperbius niugini Samson, subsp. nov. 


Male. Forewing length: 37-45 mm. 

Dorsal surface: Margins well defined, particularly those of the hindwings which, 
when combined with broad submarginals, produce a wide band. In addition to 
black bar of hindwing discal cell, there are remnants of another on discocellulars. 

Ventral surface: Constant character appears to be the absence of silver spot 
which, in nominate subspecies and majority of others, is present in dark postbasal 
area of hindwing cell. Absence of such spot also characterised in the Australian 
subspecies: inconstans Butler. 

Female. Forewing length: 35-48 mm. 

Dorsal surface: Extensive charcoal-blue band, bisecting forewing; black bars 
in cell are pronounced. White bar in space 4, plus spot therein usually inferior. 
Hindwing submarginal bands broad, moreso than those of male. 

Ventral surface: Markings well defined on fore- and hindwings; broad hindwing 
submarginals. As in male, silver spot absent in dark area of hindwing cell. 

Holotype. Male: New Guinea, Nondug] (Central Highlands), 5,500 ft., October 
16, 1950. Coll. by Wm. Brandt (E. J. L. Hallstrom). Forewing length, 43 mm. 

Allotype. Female: same as holotype, but November 1950. Forewing length, 
45 mm. 

The above primary types are in the Australian National Insect Collection, Can- 
berna vA. Cor: 

Paratypes. 6 ¢ 4,5 29, Daulo Pass, Eastern Highlands District, New Guinea, 
8,000 ft., August 1971 (2 646, 3 @@ to British Museum [Natural History]; 2 
é é to American Museum of Natural History; 2 ¢ 6, 2 2 retained by author). 
Also in the British Museum (Natural History): 1 9, Br. New Guinea, Foothills 
between Kikori R. & Purari R. (J. P. de Verteuil); 3 ¢ 4, Watut R. to Buiang, west- 
side of Herzog Mts., 3,200-5,400 ft., early 1928 (A. F. Eichhorn); 4 ¢ 6, Saiko, 
Bubu R., Upp. Waria R. Sept., Beg. October, 5,500 ft., 1936; 2 4 en es) oe 
Zageheme, Cromwell Mts., East Finisterre Range, 20 VI to 7 VIL31, 4800’ (F. Shaw 
Meyer); 1 8, same as above, but 16 VII.31. Retained by author: 1 3, 1 2, New 
ae Western Highlands, Kandep, 8,000-8,500 ft., 23.1.61 to 14.2.62, W. W. 
Brandt. 

Further material of this subspecies collected by Wm. Brandt in New Guinea, 
now in the Australian National Insect Collection: 5 @ é, 2 99, Western High- 
lands, Kandep, 8,000-8,500 ft., 23.1.61 to 14.2.62; 6 646, 1 9, Kedama Range, 
Mt. Kaindi, 6,500 ft., 24.2.1952, Sir Edward Hallstrom; 8 ¢ 6, 4 92, Nondugl 
(Central Highlands), 5,500 ft., 1950, Sir Edward Hallstrom (dates range from 
Sept. 24 to Dec. 12); 3 44, Western Highlands, Mt. Hagen Valley, Keltiga, 
5,600 ft., 28.9 to 25.10.1961; 1 2, Western Highlands, Jimi River, 4,700 ft., 16.7 
to 21.9.1961; 1 6, Telefomin (Eliptamin), 4,500-5,500 ft., 19 June to 14 Sept. 
1958. 


The Rijksmuseum at Leiden, Nederlands, possess at least 2 males of 
A. hyperbius niugini from Irian Jaya (formerly Dutch New Guinea and 


VoLUME 30, NuMBER 1 13 


Fig. 1. Argyreus hyperbius niugini. A, holotype (male) and allotype (female); 
B, ventral surfaces of same. 


West Irian), i.e., Paniai, 14 & 15.X1I.1939. This locality is now known as 
Enaratoli and lies to the east of Wissel Lakes, at approximately 5700’. 
In the British Museum (Natural History) there is a female A. hyperbius 
subspecies labelled: Dutch N. Guinea, Kobotil, O. Kaba (BM 1922- 
165). This appears to be within the known variation of A. hyperbius 
javanica Oberthur, and is a dark, well marked example. I am unable to 
locate any examples from the Moluccas or intermediate islands, but 
A. hyperbius subspecies reappear in the west in the Sunda Islands, Java 
and Sumatra, Sulawesi (formerly Celebes) and through India to Abys- 
sinia. 

I have seen males and females of A. h. niugini from the Central and 
Northern Districts of Papua, and according to Ray Straatman (pers. 
comm., 1973): “. . . the species flies in open areas (grassland) at alti- 
tudes from 1500 to 3000 metres and is most common at about 2000 to 
3000 metres.” D’Abrera (1971, p. 210) records the Australian subspecies, 
A. h. inconstans as occurring also in New Guinea and possibly Papua. All 
specimens that I have observed from Papua New Guinea are quite 


14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


~~ DJAYAPURA 
Sar VANIMO 
Sie. = we ALEXANDER 

= @® NTS 

Ss 

- _S 


RABAUL 


< 

Bae 

c= ee 
‘ 

2 

= 

= 


MARKHAM R. 
WATUT R. & HERZOG MTS. 


Be sak 


GOODENOUGH 


6% FERGUSSON 


Fig. 2. Map of Papua New Guinea (excluding Manus, New Ireland, Woodlark 
and Bougainville) indicating main ranges and localities for paratypes and primary 
type material. 


separable from those in Australia which, to the best of my knowledge, 
are restricted to that area. Thus, we record the new subspecies from 
mainland Papua New Guinea and Irian Jaya. 

The life-history of A, h. niugini is probably similar to that of the 
nominate subspecies from North India, the larvae feeding on Violaceae; 
indeed, the early stages of A. hyperbius from Japan are well illustrated 
and documented in Shir6zu & Hara (1960, p. 31). 

As noted from the material examined, many specimens cf A. h. niugini 
were collected by William Brandt, including the primary types; thus, I 
think it only fitting to include some notes on this accomplished naturalist: 

William Brandt came to Australia from Sweden about 1949 and was 
employed by the late Sir Edward Hallstrom to collect butterflies for him 
in New Guinea. For five years Brandt built up an impressive collection 
of Lepidoptera, primarily from Papua and New Guinea, and mainly of 
the larger species. In 1955 Hallstrom lost interest in his collection of 
butterflies and donated it to the Australian Government, whereupon it 
became part of the Division of Entomology, C.S.I.R.O. at Canberra. 
From 1955 until his retirement about 1969, Brandt continued to collect 


VoLuME 30, NuMBER 1 15 


mainly Lepidoptera in New Guinea, New Britain, New Ireland, New 
Hebrides, Solomons and many of the smaller New Guinea islands. In 
this work he was financed largely by the Bishop Museum, Hawaii; 
during the latter period the Lepidoptera continued to come to the Aus- 
tralian National Insect Collection, while the other insects went to the 
Bishop Museum. 

Brandt collected for a total of about 15 years in New Guinea, the last 
three or four years before his retirement being spent working on the 
Collection in Australia or, for about two years, working on New Guinea 
Lepidoptera at the British Museum ( Natural History ). During the period 
when Brandt was collecting for Hallstrom, the latter insisted that the 
data labels bore his name, as well as the actual collector. Hallstrom was 
very interested in New Guinea and set up an Experimental Livestock 
Station at Nondugl, the type locality for the new subspecies herein 
described. 


ACKNOWLEDGMENTS 


I am grateful for the help afforded me by Paul E. Smart (Saruman 
Museum) and T. G. Howarth (British Museum), both of whom allowed 
me access to useful material and much valued advice; also to Dr. R. de 
Jong (Rijksmuseum, Nederlands) and H. J. Balcliffe (Royal Geographical 
Society, England). Special thanks are extended to Dr. Ian F. B. Com- 
mon (C.S.I.R.O., Canberra, Australia) for supplying much useful data, 
plus specimen material for primary and paratypic designation. 


LITERATURE CITED 


D’Asrera, B. 1971. Butterflies of the Australian Region. Lansdowne, Melboume. 
415 p. 

Hemmiunc, F. 1967. The generic names of the butterflies and their type-species 
(Lepidoptera: Rhopalocera). Bull. British Mus. (Nat. Hist.), Ent., Suppl. 
9. 509 p. 

McCussyy, C. 1971. Australian butterflies. Nelson, Melbourne. 206 p. 

Serrz, A. 1927. The macrolepidoptera of the world. Vol. 9. Stuttgart. 

SHmrozu, T. & A. Hara. 1960. Early Stages of Japanese Butterflies in Colour. 
Vol. 2. Hoikuscha, Osaka. 148 p. 


16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


CALLOPHRYS ERYPHON (LYCAENIDAE) IN MAINE 


WakrkREN J. KIEL 
P. O. Box #2, Whitefield, New Hampshire 03598 


On the basis of several specimens of a large, boldly marked Callophrys 
recently examined and determined by Mr. J. D. Lafontaine and Mr. 
Harry K. Clench, Callophrys (Incisalia) eryphon (Boisduval) is here 
reported to be established in a small acid bog located in western Maine. 

Clench (in Ehrlich & Ehrlich, 1961) summarizes the eastern range of 
C. eryphon as “eastward to Rockies, Nebraska and northern Manitoba.” 
Klots (1951) does not include C. eryphon as occurring east of the Great 
Plains, nor is it mentioned in his list of casual species. In the past 
decade however, the species has been discovered to occur much farther 
east. Recent eastward records include Port Hope, Ontario ( Riotte, 1967); 
Pine Co. (Masters, 1972) and Cook Co. (Huber, 1966), Minnesota; and 
Chippewa and Luce counties, eastern upper peninsula of Michigan 
(Nielsen, 1966). The recent Maine checklist (Brower, 1974) contains 
no mention of C. eryphon. 

The bog, Black Spruce—sphagnum heath, is roughly 3-5 acres in 
extent, and is located adjacent to state Rt. 16, east of the Maine-New 
Hampshire border, approximately 6 mi. S of Wilsons Mills, near the 
confluence of the Diamond and Magalloway rivers. The area is approxi- 
mately 1,260 ft. in elevation and surrounded by low mountains. The 
bog supports a rather dense growth of Black Spruce (Picea mariana), 
Tamarack (Larix laricina) and other typical associate plants, e.g., Labra- 
dor Tea (Ledum groenlandicum), Rhodora (Rhododendron canadense ), 
laurel (Kalmia sp.), Bog Rosemary (Andromeda glaucophylla), various 
other heaths, grasses and sedges. From early to mid-June OEneis jutta 
Hubner occurs here, along with an unusually productive colony of 
Callophrys augustinus augustinus (Westwood), and the proliferation 
of flowering plants attract a variety of other Canadian Zone butterflies 
in the late spring. Ease of access has made the area a popular collecting 
spot with numerous New England entomologists. 

During two trips to the bog on 8 and 9 June 1974, a good series 
of C. eryphon was taken, both sexes being fairly common. A number of 
these were later positively identified by Mr. Lafontaine and Mr. Clench. 
A small series of the same catch was also deposited in the Dartmouth 
College Museum Collection. 


Phe presence of Callophrys eryphon this far east raises a number of 


VoLUME 30, NuMBER | LZ 


questions concerning (1) sympatry with C. niphon clarki Freeman, (2) 
significance of the bog environment and (3) larval food plant. 

I recently re-examined the series of C. eryphon now in the Dartmouth 
College Collection to check the possible inclusion of C. niphon. All 
specimens in the series (7 males, 4 females in fresh to slightly worn con- 
dition) appear to be eryphon, suggesting that at least within the bog 
contines, niphon does not occur. Locally, however, C. niphon is generally 
common, usually in association with pine woods and occasionally exhibit- 
ing local outbreaks (Grey, 1967). 

The observed association of this C. eryphon colony with a bog needs 
further clarification. It may be merely apparent, being presently known 
only from this one locality, and may display other habitat preferences if 
and when other colonies are uncovered. It should be noted that the 
Bog Elfin, C. lanoraieensis Sheppard, well known from large bogs of 
north central Maine, does not occur here. 

Various endemic pines are listed as food plants for C. eryphon in the 
Western montane regions. Other pines are recorded in the east for C. 
niphon: Klots (1951) suggests “probably only ‘hard’ pines, i.e. vir- 
giniana, rigida, etc., not White Pine (P. strobus)”, although Ferguson 
(1954) does not rule out the possibility of this latter species. The com- 
mon native pines appear to be almost wholly lacking at the Wilsons 
Mills locale, therefore, it appears possible that something entirely differ- 
ent, perhaps one of the spruces could serve as the host. Nearly all of 
the C. eryphon collected were taken on or near young Black Spruce, the 
butterflies often alighting on the fresh terminal growth. McGugan (1958) 
includes White Spruce (Picea glauca) as a larval collection source for 
C. niphon clarki, suggesting spruce as an alternate choice for both but- 
terflies. Clearly, however, the matter will remain in question until fur- 
ther observations and life history work can be conducted. 

The discovery of Callophrys eryphon in the east will doubtless generate 
continued intensive searches for additional colonies of this “western” 
butterfly. Collectors having specimens of northern New England niphon 
should check their material carefully and forward any suspect examples 


along with data to Mr. Clench for determination. 


ACKNOWLEDGMENTS 


Sincere thanks are extended to Mr. J. Donald Lafontaine, Biosys- 
tematics Research Institute, Ottawa, and to Mr. Harry K. Clench, Carne- 
gie Museum of Natural History, Pittsburgh, for species determination, 
and, especially to the latter, for helpful comments and suggestions and 
criticism of the manuscript. Thanks are also due Mr. Paul S. Miliotis, 


18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Dunstable, Mass., for originally raising the question of the true identity 
of the Maine specimens and for checking the botanical names; to Mr. 
Donald J. Lennox, Jefferson, N. H., for initial introduction to the bog 
and helpful discussions relating to host plant possibilities; and to Mr. 
Richard E. Gray, Dartmouth College Museum, Hanover, Nae. for 
permitting me to examine material in the collection of that institution. 


LITERATURE CITED 


Brower, A. E. 1974. A list of the Lepidoptera of Maine—Part 1, the Macrolepi- 
doptera. Life Sci. Agric. Exp. Sta., Univ. Maine at Orono. 136 p., Oemligs.; 
1 map. 

EHRLICH, ». R., & A. H. Eueticu. 1961. How to know the butterflies. William C. 
Brown Co., Dubuque, Iowa. 262 p., 525 figs. 

Fercuson, D. C. 1954. The Lepidoptera of Nova Scotia. Proc. Nova Scotian Inst. 
Sci, 23(3). 37> pnt ses. 16) pls, 1 map: 

Grey, L. P. 1967. Maine in Season Summary. NEWS Lepid. Soc. No. 3, p. 14. 

Huser, R. L. (compiler). [1966.] Minnesota 1965 annual butterfly summary. ILI p., 
hectographed. 

Knots, A. B. 1951. A field guide to the butterflies of North America, East of the 
Great Plains. Houghton Mifflin Co., Boston. 349 p., 40 pls. 

Masters, J. H. 1972. Butterfly records for three northeast Minnesota counties 
Aitkin, Carlton, and Pine Counties. Bull. Assoc. Minn. Ent. 5(2): 19-26. 

McGucan, B. M. (compiler). 1958. Forest Lepidoptera of Canada recorded by the 
Forest Insect Survey, Vol. I—Papilionidae to Arctiidae. Forest Biology Division, 
Canada Dept. of Agriculture. 76 p., 46 figs. 

NietsEN, M. C. 1966. Occurrence of Callophrys eryphon (Lycaenidae) in Michigan. 
J. Lepid. Soc. 20: 41-42. 

Riorre, J. C. E. 1967. New and corrected butterfly records for Ontario and for 
Canada. J. Lepid. Soc. 21: 135-137. 


MATHILDANA NEWMANELLA (OECOPHORIDAE) IN ARKANSAS 


Through the courtesy of H. N. Greenbaum (Department of Entomology, University 
of Arkansas), I recently have been able to examine moths collected with a Malaise 
trap set up 24 hours a day in Fayetteville, Washington Co., Arkansas. Among the 
moths collected 22-26 May 1975 were two females of Mathildana newmanella 
(Clemens). Hodges in his recent revision of the North American Oecophoridae 
(1974, Moths Amer. North of Mex., Fasc. 6.2: 122), reported this moth, originally 
described from Virginia, as occurring from Quebec to North Carolina and extending 
west only to southern Ohio. The new record from western Arkansas considerably 
extends the known range of M. newmanella, and, with the range of the deciduous 
forests ending only a little farther west, this may be near the western limits of its 
distribution. M. newmanella may be a diurnal flier, as are related species such as 
Esperia sulphurella (Fabricius ). 


Joun B. Heppner, Department of Entomology and Nematology, University of 
Florida, Gainesville, Florida 32611. (Florida Agricultural Experiment Station Journal 
Series No. 5954. ) 


VoLuME 30, NuMBER | 19 


NOTES OF MARYLAND LEPIDOPTERA. 5. 
A NEW SUBSPECIES OF POANES MASSASOIT (HESPERIIDAE) 


WILLIAM A. ANDERSEN 
220 Melanchton Avenue, Lutherville, Maryland 21093 


AND 


ROBERT S. SIMMONS 
1305 Light St., Baltimore, Maryland 21230 


On 19 June 1962, one of us (WAA) captured two unusual specimens 
of Poanes massasoit Scudder in New Bridge, Dorchester County, Mary- 
land. Upon seeing these specimens and learning of their origin, Sim- 
mons suggested that they might be representatives of a new subspecies, 
and further collecting trips were planned. On 12 July 1962, 28 additional 
specimens were collected from the same locality, and all were noted to 
differ considerably from both P. m. massasoit Scudder and P. m. hughi 
Clark, the previously described northeastern subspecies. 

Our collections contain many examples of P. m. hughi Clark from 
northcentral Maryland including its type locality. Morphological com- 
parisons of our eastern shore specimens with those of P. m. hughi and 
P. m. massasoit indicate that important taxonomic differences exist be- 
tween the three entities. A distributional study reveals that our new 
specimens (Figs. 1-10) represent the most southeastern end of a cline, 
in which P. m. massasoit is the most northerly taxon, with hughi repre- 
senting an intergrade between our new subspecies and P. m. massasoit. 

The apparent differences noted as one studies the cline from north to 
south are that the specimens become somewhat larger and there is a 
progressive loss of areas of yellow scales, especially on the underside of 
the hindwing of both sexes and on the upper surface of the wings of the 
female. In this study we will make comparisons with P. m. hughi alone 
as Clark (1932) has already very adequately compared P. m. massasoit 
and P. m. hughi. 

We name this new subspecies after our late, good friend Franklyn H. 
Chermock, who plied us with specimens, good humor and many inter- 
esting stories about collecting and collectors. 


Poanes massasoit chermocki Andersen and Simmons new subspecies 
Figs. 1-4, 9-10 
Holotype. Male: Forewing length 14.7 mm. Upper surface, forewing: plain, 


unmarked, color dark, blackish brown with faint mahogany irridescence. 
Upper surface, hindwing: same as forewing. 


20) JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Under surface, forewing: dark brown generally with slightly lighter brown 
scales at apical area and just inside costal and outer margins. Three small, sub- 
apical, yellow-brown spots at costal margin in spaces between veins R-3 and R-4, 
R-4 and R-5, and R-5 and M-1. 

Under surface, hindwing: tan at outer margins, brown centrally. Five yellow, 
submarginal spots arranged in a rough semicircle paralleling outer margin. A small 
yellow spot also in discal cell. Between this spot and two of the submarginal spots 
is a well-defined area of tan scales. 

Allotype. Female: Forewing length 16.0 mm. Upper surface, forewing: ground 
color is the same as in male. Three subapical yellow spots extending in a line 
inward from costal margin. In postmedian area between veins M-2 and M-3, and 
and M-3 and Cu-1 are two larger yellow spots, the lower one squarish. 

Upper surface, hindwing: same as in male. In rare specimen, only a faint 
suggestion of one yellow spot in postmedian area. 

Under surface, forewing: as in male except with addition of two postmedian spots 
corresponding to those of upper surface. 

Under surface, hindwing: same as in male. 

Type localities. Holotype: New Bridge, Dorchester County, Maryland, June 
19, 1962. Allotype: same locality, July 6, 1963. 

The types are deposited in the U. S. National Museum, Washington, D.C. 
Male and female paratypes will be deposited in the American Museum of Natural 
History, New York and in the Carnegie Museum, Pittsburgh, Pennsylvania. 


Differences between P. m. chermocki and P. m. hughi 


1. The most striking difference is in the under surface of the hindwing where 
the extensive yellow marking (extending from the discal cell outward to 
include the submarginal yellow band) of hughi is reduced to the narrow 
yellow submarginal, roughly semi-circular band of individual spots of 
chermocki. 

2. The maculation of the chermocki female is much reduced on the upper 
surfaces, so that in half the specimens the forewing is immaculate on the upper 
surface. 

3. The size of chermocki is somewhat larger, averaging about 0.5 mm larger 
per forewing. 


Discussion 


The locality of New Bridge, Dorchester County, is in the southernmost 
section of the eastern shore of the Chesapeake Bay, 80 miles due south 
of the nearest known colony of P. m. hughi in Cecil County. Cecil County 
is the northernmost county on the eastern shore of Maryland. It is 
divided between coastal plain in its southern portion and piedmont in 
its northern. The specimens of P. m. hughi from Cecil County were 


> 


_ Figs. I-10. Poanes massasoit subspecies from Maryland. Left side dorsal sur- 
faces; right, ventral surfaces. 1-4, P. m. chermocki (subsp. nov.), New Bridge, 
Dorchester County: (1 & 2) holotype, male, 19 June 1962; (3 & 4) allotype, female, 
6 July 1963. 5-8, P. m. hughi: (5 & 6) male, Towson, Baltimore County, 3 July 
1954; (7 & 8) female, Beltsville, Prince Georges County, 20 July 1967. 9-10, P. m. 
chermocki, form “suffusa”, New Bridge, Dorchester County, 6 July 1963. 


21 


VoLUME 30, NUMBER 1 


Tcm 


22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


collected in the piedmont area. Much collecting on Maryland's eastern 
shore coastal plain between the two areas has not produced any closer 
colonies. This deme in Dorchester County is thus geographically some- 
what isolated from the nearest population of P. m. hughi and at the 
present time this population represents the only one known to us from 
this region. 

The specimens of P. m. chermocki are rather homogeneous in their 
maculation and size as compared with P. m. hughi populations. The 
under surface of the hindwings of specimens of chermocki are strikingly 
similar and there is only slight variation in the spots of the upper surfaces 
in the female. In our hughi specimens, on the other hand, the upper 
surfaces of the female vary from being spotted on both wings to some 
having none on either wing, this being in agreement with Clark’s 
description of his subspecies. The underside of the hind wing is similarly 
varied. Clark (1932) himself described one specimen from Beltsville, 
Maryland, which is very like chermocki and he pictured another such 
specimen in the frontispiece of his Butterflies of Virginia (Clark & Clark, 
1951). We note that in our collections of hughi from north central Mary- 
land forms similar to chermocki occur at a rate of approximately 4 per 
cent. The form “suffusa” also occurs in this new subspecies. In our 
series its incidence is about the same as in hughi, i.e., 1 in 10. 


ACKNOWLEDGMENTS 


The authors are greatly indebted to Dr. Austin P. Platt of the Univer- 
sity of Maryland, Baltimore County, who not only photographed the 
specimens and helped make up the plate but also read the manuscript 
and made helpful criticisms and suggestions. We also thank William F. 
Andersen who developed and printed the photographs. 


LITERATURE CITED 


Crark, A. H. 1932. The butterflies of the District of Columbia and vicinity. U.S. 
Natl. Mus. Bull. 157. 337 + ix p. 

Crark, A. H. & L. F. Ciarx. 1951. The butterflies of Virginia. Smithsonian Misc. 
Coll. 116, No. 7. 239 + vii p. 


VoLUME 30, NuMBER 1 23 


NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF 
BUTTERFLIES OF EL SALVADOR. VII. ARCHAEOPREPONA 
DEMOPHON CENTRALIS (NYMPHALIDAE) 


ALBERTO MuysHONDT 
101 Avenida Norte #322, San Salvador, El Salvador 


During August 1971, one of my sons found a strange-looking larva 
on a very small unknown shrub in a ravine near the city of San Salvador. 
The larva unfortunately died before pupating due to the lack of food. We 
could not identify the shrub because it was not in flower, and our efforts 
to substitute other similar plants for food were unsuccessful. Two years 
later, we found a female Archaeoprepona demophon centralis Fruhstor- 
fer (Charaxinae) ovipositing on a larger flowering shrub and were able 
to rear the species to adult. The larvae were the same as the single 
specimen collected in 1971. Since that time we have reared the species 
several times. 

The rearing was done inside transparent plastic bags. The larvae were 
kept supplied with fresh leaves of the foodplant. The plastic bags were 
cleaned daily of excess moisture and excreta. Shortly before pupation, 
the larvae were transferred to a wooden box with mosquito-net windows. 
The adults emerged in the same box. Measurements of the different 
instars were recorded, and photographs were taken of all developmental 
stages. Some specimens of the early stages and exuvia were preserved in 
alcohol. These will be sent to the American Museum of Natural History, 
New York. The adults were determined by Dr. A. H. B. Rydon, the 
foodplant by Dr. D. Burch, University of South Florida. 


lane GyYcLEe 


Egg (Figs. 1, 2). White, spherical but for slightly flattened base, smooth, no 
visible sculpturing at 15 magnification, 2.5 mm diameter. When ready to hatch 
in 6 days, head and body marks visible through eggshell. 

First instar larva (Fig. 4). General color pale brown. Head roundish, naked, 
with a dark brown M mark on front. Naked body thickens from head to 2nd 
abdominal segment, then narrows caudad, terminates in two short tails. Promi- 
nence on dorsal meson of 3rd thoracic segment, another subdorsally on each side 
of 2nd abdominal segment directly above corresponding spiraculum. Thoracic 
segments with fine, dark brown lines. Abdominal segments darker brown dorsally. 
Anal prolegs smaller than others. Thoracic spiraculum larger than abdominals. Second 
abdominal spiraculum very high, completely out of line from others, except that 
8th abdominal spiraculum also slightly higher. Grows from 4-10 mm in 13 days. 

Second instar larva (Fig. 5). Head with same aspect as in first instar except 
for one short horn on each epicranial apex. Body also as in first instar except dorsal 
hump more noticeable and tails longer and straight. Grows to 14 mm in § days. 


24 JourRNAL OF THE LEPIDOPTERISTS SOCIETY 


Rae 


Figs. 1-6. Archaeoprepona demophon centralis. 1, Egg showing characteristic M 
on larval head, width about 2.5 mm; 2, egg showing dorsal markings of the larval 
body; 3, egg parasitied by larvae of an unknown Chalcididae; 4, first instar larva, 
about 6 mm long, on resting perch; 5, second instar larva, about 14 mm long, 
baring a vein; 6, third instar larva, about 27 mm long. 


Third instar larva (Fig. 6). Same general colors as earlier. Head with longer 
horns; appearance of small, blunt, lateral projections below and behind horns. Mesal 
projection on 3rd thoracic segment and subdorsal ones on 2nd abdominal segment 
very pronounced. Dark brown line separating dorsal darker area from paler sub- 
spiracular area which is criss-crossed by faint brown lines as is the thoracic zone. 
Caudal projections longer, still straight. Grows to 30 mm in 8 days. 

Fourth instar larva (Fig. 7). Head and horns covered by small tubercles, pro- 
ducing a coarse aspect; lateral projections more noticeable. Body color darker, 
mostly over supraspiracular and dorsal areas, darker slanting lines. Appearance of 
tiny, bright blue spots along spiracular zone, around mesal prominence on 3rd 
thoracic segment and on longer, slightly crooked tails. Grows to 48 mm in 13 days. 

Fifth instar larva (Figs. 8, 9). Head and horns much rougher, lateral projections 
quite noticeable. Body color brown on thoracic segments and subspiracularly on 


VoLuME 30, NuMBER 1 D5 


Figs. 7-11. Archaeoprepona demophon centralis. 7, Fourth instar larva crawling, 
about 45 mm long; 8, fifth instar larva recently moulted, about 50 mm long; 9 
detail of head of fifth instar larva; 10, prepupa in typical resting position, about 
72 mm long; 11, prepupa ready to pupate, notice placement of tails. 


9 


abdominal segments. Dark brown triangle dorsally covering area limited by 3 
very prominent projections on 3rd thoracic and 2nd abdominal segments. Dorsa 
of remaining segments from spiracular line to meson, paler brown with darker 
brown slanting bands. Tails dark brown with yellow spots, very long and crooked. 
Profusion of tiny, bright blue spots on tails, along spiracular line and around 
thoracic prominence. Prolegs very thick. Grows to 70-80 mm in 18 days. 

Prepupa (Figs. 10, 11). Drastic color change. Head and body green, darker on 
head, thoracic prominence and along spiracular line from 6th—9th abdominal seg- 
ments. Blue spots still visible. 

Pupa (Figs. 12-14). Bluish-green with scattered whitish, irregular spots; head 
bifid with points yellowish. Spiracula light brown surrounded by white ovals. Prior 
to adult emergence becomes very dark gray, some adult colors visible through 
cuticle. Lateral view: ventral profile only slightly convex; dorsal profile widens 
gradually from green cremaster to 5th abdominal segment, then the 4th forms a 
distinct hump, then narorws gradually to bifid head. Dorsal and ventral aspect: 
lateral profile widens smoothly from cremaster to wingcases, at thoracic level, then 


26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 12-18. Archaeoprepona demophon centralis. 12, Pupa, ventral aspect, about 
12 mm long; 13, pupa, lateral aspect; 14, pupa, dorsal aspect; 15-18, sequence show- 
ing emergence of adult male. 


narrows abruptly to head. Measurements: 42 mm long, 18 mm laterally, and 20 
mm dorso-ventrally at widest points. 

Adult (Figs. 19-22). Males and females shaped similarly. Forewing with 
faintly convex costal margin forming a rounded apex, concave outer margin to 
rounded tornus, straight inner margin. Hindwing with convex costal margin, strong 
humeral lobe, rounded outer angle continuing with rounded outer margin to 


VoLUME 30, NUMBER 1 27 


* * £5 Ries % # 


Figs. 19-22. Archaeoprepona demophon centralis. 19, Adult male, dorsal view, 
wingspan about 82 mm; 20, adult female, dorsal view, wingspan about 103 mm; 
21, adult male, ventral view; 22, adult female, ventral view. 


rounded anal angle, rounded inner margin with a fold. Dorsal ground color of 
forewing dull black in males, dark brown in females, with a greenish-blue iridescent 
isosceles triangle from median area of wing with base on mid-inner margin; 2 small 
subapical, bluish spots. Hindwing with small basal, black triangle and _ broad, 
greenish-blue iridescent band from mid-costal margin ending in a point near 
inner margin, close to anal angle; this band with definite pale blue tinge along 
basal border. Inner fold and thin border along ccstal margin, pale gray. Males 
with black brush of hairs near base of hindwing. Ventrally, dominant color pale 
brown with complicated pattern of darker brown areas and thin black lines; along 
outer margin of hindwing a row of small “eyes” with light blue pupils. Antennae 
and eyes black, proboscis orange. Dorsal and ventral surfaces of body concolorous 
with corresponding wing surfaces. Wing span 80 mm in males, up to 105 mm in 
females. 
Total developmental time for this species 85 days. 


NATURAL HISTORY 


The females of Archaeoprepona demophon centralis deposit their eggs 
singly on the lower surfaces of the mature leaves of a shrub determined 
by Dr. D. Burch to be at least very close to Malpighia glabra L. (Mal- 
pighiaceae). The pure white egg is quite visible against the dark green 


leaf. 


JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


i) 
(©) 


The newly hatched larva completely consumes the eggshell, leaving 
only the shiny base on the leaf, and stays for one day without further 
feeding. It then moves to the tip of the leaf and nibbles around the 
central vein, baring it and affixing to it small, frass pellets woven with 
silk until the vein is prolonged beyond the leaf limits. Pieces of leaf 
tissue hung from short lengths of silk are added to the resultant perch. 
This is used as a resting perch during the day where the small larva is 
very inconspicuous among the dried bits of leaf. It is abandoned by the 
larva only momentarily to feed at dawn or dusk. First, second and third 
instar larvae use a perch. If the leaf is consumed during this period, 
another perch is made on a second leaf, and in some cases, a third. From 
the fourth through fifth instars, the larva wanders about the plant, but 
usually stays motionless during the day resting on a twig, head and tail 
hanging at the sides, giving the dorsal anterior part of the body a rep- 
tilian appearance (i.e., head of small snake or lizard) with the promi- 
nences on the second abdominal segment resembling the eyes and the 
mesal prominence on the third thoracic segment, the snout. The effect 
is enhanced by the crawl of the larva which imitates an inquiring snake 
head. The mesal prominence on the third thoracic segment is retractile; 
when touched it almost disappears. 

The prepupal larva becomes green and very inconspicuous, blending 
with the green foliage. At this time, the larva wanders for two days 
searching for a suitable pupation site which may be on the same food- 
plant. Once the site has been chosen, usually a twig, the larva weaves 
a silk pad girdling the twig, reinforcing the twig’s attachment to the 
shrub. The anal prolegs are then applied to the pad, the tails are 
positioned on either side of the twig, and the larva hangs incurved ven- 
trally with its head almost touching the anal prolegs. It hangs thus for 
two additional days. 

The pupa is also very cryptic due to its green color which matches 
the foliage. It is very passive, but when roughly handled it reacts by 
swinging laterally and then returning to the vertical position. The pupal 
color changes to very dark gray shortly before the adult emerges. Some 
of the adult colors are actually visible through the pupal cuticle before 
emergence occurs. 

The adult emerges rapidly and hangs from the smoky gray pupal 
cuticle while expanding its wings and ejecting the meconium (Figs. 15- 
18). The adult is ready to fly in about 25 minutes. 

The adults of A. demophon centralis, like most Charaxinae, are strong, 
swift fliers, producing an audible rustling noise at short distance. They 
usually keep to tree-tops and come to the ground only to feed on fer- 


VoLUME 30, NUMBER 1 29 


menting fruits (e.g., mangoes and guayabas) or vertebrate excrement. 
Both sexes are lured easily to baits of fermented banana. They also feed 
on sap oozing from tree wounds. The male exhibits a strong territorial 
behavior and perches on its chosen tree, chasing vigorously any tres- 
passing butterfly. The female flies at lower levels, mostly when ready 
to oviposit. We have observed A. demophon centralis most frequently 
around wooded coffee plantations from 600-1200 m elevation. 

The foodplant, Malpighia glabra L. (?), is found most commonly in 
wooded ravines within the flight range of the adult. It is a small shrub 
with perennial, opposite, stipulate, ovate leaves and pinkish flowers with 
characteristic dentate petals arranged in cymes. The fruit is a red drupe 
when mature with up to 3 carpels. 

It was observed that the eggs of A. demophon centralis are very often 
parasitized by Chalcididae wasps (Fig. 3), and the very young larvae are 
preyed upon by spiders. 


Discussion 


Species belonging to the genus Archaeoprepona, prior to its establish- 
ment by Fruhstorfer in 1916, were placed in the genus Prepona Boisduval 
(Hemming, 1967). The type species for Archaeoprepona was designated 
as demophon Linneaus. Le Moult (1932) chose to ignore Fruhstorfer’s 
genus and established the invalid synonym Pseudoprepona with demo- 
phon again as the type species. Some modern authors (e.g., Descimon, 
et al., 1973) consider Archaeoprepona to be a subgenus of Prepona. 

Basic differences in the adults: black hair tufts on the hindwing of 
Archaeoprepona, against the honey-colored tufts of Prepona and the male 
genitalic differences and antennal colorations as noted by Fruhstorfer 
(1916) consistently correspond to morphological differences during the 
larval and pupal stages. The data pertaining to the immature stages are 
found in a number of sources. Included are the descriptions of the early 
stages made by Miller (1886) of Archaeoprepona amphimachus Fabri- 
cius, A. catachlora Staudinger, A. demophon extincta Staudinger (all un- 
der the generic name Prepona) and Prepona laertes Hiibner. Lichy’s 
(1933) description of the early stages of Prepona omphale guatemalensis 
Le Moult and that of P. omphale octavia Fruhstorfer by Muyshondt 
(1973a) also are important sources. In addition to the above life history 
of Archaeoprepona demophon centralis, another study that will soon be 
published on a yet, undetermined species of Archaeoprepona is taken into 
consideration. 

All the descriptions of the mentioned Prepona species show that the 
two horns on the head are fused and appear as a single epicranial pro- 


30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


jection. The Archaeoprepona species have two distinct horns, one on 
each epicranial apex plus a shorter horn below and behind each large epi- 
cranial horn. The general habitus of the larvae of both groups is similar 
except Prepona species lack the median dorsal prominence on the third 
thoracic segment which is present in all known Archaeoprepona larvae. 
Prepona larvae do not show the drastic color change when entering the 
prepupal stage which we noticed in rearing Archaeoprepona demophon 
centralis and the undetermined A. sp. Miiller’s (1886) descriptions make 
no mention of prepupal color changes. The pupae of Prepona gradually 
are humped dorsally while Archaeoprepona species show a very pro- 
nounced hump on the fourth abdominal segment. Also the pupae of 
Prepona have orange, irregular spots; these are white in Archaeoprepona 
pupae. 

All these differences favor the concept of two genera, but the many 
similarities, including behavioral patterns, indicate that both genera 
belong under the same immediate higher classification. On the same 
basis it also is very evident that these genera are closely related to the 
Zaretis-Siderone complex (Rydon, 1971; Muyshondt, 1973b and ms. in 
prep.). These facts seem to be in agreement with Rydon’s treatment of 
the Charaxidae (with family status) which, in addition to other sub- 
families that include Old World genera, separates the American genera 
into three subfamilies: Preponinae (Prepona, Archaeoprepona, Agrias, 
Anaeomorpha and Noreppa), Zaretidinae (Coenophlebia, Zaretis and 
Siderone) and Anaeinae (Fountainea, Hypna, Anaea, Polygrapha, Con- 
sul, Cymatogramma and Memphis) with Zaretidinae being the link be- 
tween Preponinae and Anaeinae. 

Some authors have tried to prove close phylogenetic relations between 
the Charaxinae and other groups, e.g., Limenitinae (Reuter, 1896; Miiller, 
1886). According to Fruhstorfer (1924), Hahnel thought there was an 
affinity between Prepona and the Apaturinae. If the consensus is fol- 
lowed that the more reliable morphological characters to determine 
phylogenetic relationships are those which resist, to a greater degree, 
changes induced by selection, and these are the characters of the early 
stages (Brower, et al., 1963), it appears that neither the Limenitinae nor 
the Apaturinae show much in common with the Charaxinae, nor with any 
other group of the Nymphalidae. Rydon (1971) may be right in assign- 
ing family status, as other authors have also done, to the Charaxinae. The 
smooth, spherical eggs (flattened base and micropylar area or not), the 
spineless (Dornenlossen of Miiller [1886]) larvae and the peculiar 
pupae of this group have nothing in common with either the very sculp- 
tured (pineapple-like) eggs, the odd-looking, spine-covered larvae or the 


VoLuME 30, NuMBER 1 31 


pupae with characteristic projections of the Limenitinae. In a series of 
future articles observations on the local Adelpha (Limenitinae) will 
emphasize this point. The spherically ribbed eggs and the flattened pupae 
of the Apaturinae do not have much in common with the Charaxinae 
either. It is true that the Charaxinae show some behavioral similarities 
with other nymphalids (e.g., the making of resting perches during the 
larval stage, adult feeding of fermenting fruit, etc.), but this might very 
well be the result of selective forces producing independently convergent 
successful strategies in species otherwise distantly related as is the case 
in many color similarities between species forming Mullerian mimicry 
complexes. 

It frequently happens that during the larval stage of a given species 
(e.g., Morpho peleides Kollar, Dione moneta Hiibner, various Apatura 
spp. and Smyrna blomfildia [Fabricius] in particular), or during the 
pupal stage (e.g., Opsiphanes tamarindi Felder, O. cassina Felder, 
Zaretis itys [Cramer], and Anaea eurypyle [C. & R. Felder] ), there are 
very noticeable differences in coloration even though during the early 
stages there should be less chance for diversification. There are also dif- 
ferences of this sort in the adults of the same species, even coming from 
the same brood. We once obtained three adult males of Morpho peleides 
hyacinthus Butler, with four “eyes” on the ventral side of the forewing 
while another 18 specimens, males and females, only had three as is 
common. All 21 specimens were from eggs deposited by the same female 
on the same date in our insectary. They were fed their normal foodplants 
and were kept under identical conditions until adult emergence. Le 
Moult’s (1932) opinion was that species of Prepona and Archaeoprepona 
are less variable than their near relatives, the Agrias, and have stable 
external characteristics. According to Dr. Descimon (pers. comm.), the 
numerous new species and subspecies that Le Moult named on the 
basis of external characteristics has caused great confusion. Vane-Wright 
(1974) thought many of Le Moult’s taxonomic conclusions were unsound 
because he “split” many previously accepted species on little evidence. 

Much “lumping” and “splitting” has been done in the past without 
having enough data to form sound conclusions. It is important that more 
investigations on the early stages of the neotropical Rhopalocera be 
conducted so that systematists can use the results to establish a more 
accurate scheme of classification. 


ACKNOWLEDGMENTS 


We wish to express our gratitude to Dr. A. B. Klots for encouraging us 
to publish the results of our studies and to Dr. G. L. Godfrey for read- 


32 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


ing, criticising and suggesting improvements for our article. We espe- 
cially thank Dr. A. H. B. Rydon and Dr. H. Descimon, Ecole Normale 
Supérieure, Paris, for supplying us with a wealth of reference material 
and Dr. L. D. Miller for sending Vane-Wright’s publication on Prepona. 
Special mention is due to my children: Orlando, who first found the larva 
of the species reported in this study; Marilynn, who first saw the ovi- 
positing female; and the youngest member of the team, Pierre, who 
obtained many other eggs and carefully fed the larvae during their long 
developmental time. 


LITERATURE CITED 


Brower, L. P., J. Vz. Brower, & C. T. Cortins. 1963. Experimental studies of 
mimicry. 7. Relative palatability and Miullerian mimicry among neotropical 
butterflies of the subfamily Heliconiinae. Zoologica (New York) 48: 65-84. 

Descimon, H., J. M. bE Matcut, & J. R. StorFEL. 1973. Contribution a l’étude des 
Nymphalides neotropicales. Description de trois nouveaux Prepona mexicains. 
Alexanor 8: 155-159. 

FruustorFer, H. 1924. Genus Prepona Byd., p. 550-566. In A. Seitz (ed.), The 
macrolepidoptera of the world. Vol. 5. Stuttgart. 

Hemminc, F. 1967. The generic names of the butterflies and their type-species. 
(Lepidoptera: Rhopalocera). Bull. British Mus. (Nat. Hist.), Entomology. 
Suppl. 9. 509 p. 

Le Mou tt, E. 1932. Etudes sur les Prepona. Nov. Entomol., fasc. 2. 

Licny, R. 1933. Observations biologiques sur les différents états de Prepona 
omphale s/sp. guatemalensis Le Moult. (Lép., Nymph.). Nov. Entomol., 
fasc. 3. 

Mutter, W. 1886. Sidamerikanische Nymphalidenraupen. Versuch eines natiir- 
lichen Systems der Nymphliden. Zool. Jahrb. Zeitschr. Syst., Geogr., Biol. der 
Thiere 1: 417-678. 

Muysuonpt, A. 1973a. Notes on the life cycle and natural history of butterflies 
of El] Salvador. I. Prepona omphale octavia (Nymphalidae). J. Lepid. Soc. 
27: 210-219. 

. 1973b. Notes on the life cycle and natural history of butterflies of El 
Salvador. II. Anaea (Zaretis) itys. (Nymphalidae). J. Lepid. Soc. 27: 294-302. 

Reuter, E. 1896. Uber die Palpen der Rhopalocera. Helsingfors. 

Rypon, A. H. B. 1971. The systematics of the Charaxidae (Lepidoptera: Nympha- 
loidea). Ent. Rec. J. Var. 83: 219-233, 283-287, 310-316, 336-341, 384-388. 

VaNE-Wricut, R. I. 1974. Eugene Le Moult’s Prepona types (Lepidoptera: 
Nymphalidea, Charaxinae). Bull. Allyn Mus. 21: 1-10. 


VoLUME 30, NUMBER | 33 


NEW CATOCALA OF NORTH AMERICA (NOCTUIDAE) 


A. E. BROWER 
8 Hospital Street, Augusta, Maine 04330 


All but one of these species fall in our two most difficult groups of 
the small Catocala—the crataegi group in the East and the andromache 
group in the West, though when we really know the early stages of all 
species, further division may be indicated. I have spent a great amount 
of time and work mounting specimens, preparing genitalia and studying 
these genitalia to arrive at only indefinite characters based on structure 
on which to separate the species. The adults vary throughout their 
range and in some local areas noticeably different specimens occur. The 
great need is for adequate material of eggs, larvae and pupae of all 
species to be available for comparison, and in the adults, a series from 
many scattered places in its range. Larval descriptions in the crataegi 
group are tenuous or non-existent, and nothing seems to have been pub- 
lished on the early stages of any of the andromache group, except cheli- 
donia Grote. With probably the greatest series in existence of these 
species and closely related ones before me, I believe all of the following 
proposed species warrant the title new species, when a series is closely 
studied. For many years specimens of the andromache group have been 
sought by two experienced workers, J. W. Johnson and Erich Walter, 
and several females of each of several species have been confined for 
eggs, without securing an egg. The series of specimens have been 
possible only through the generous help of collectors and the institution 
named in connection with types. Unless otherwise stated all types remain 
in the author's collection at present. 


Catocala texarkana Brower new species 
(Fig. 1) 


The forewing is of a light uniform gray, somewhat darker basally, and along the 
inner margin, the basal area in the fold toward the transverse anterior line with 
definite ribbing. The median area from costa to inner margin or to fold is conspicu- 
ously whitish, with a segment of a median line to the reniform usually present, and 
some darker shading between the reniform and transverse posterior line. The lines 
are narrow, brownish black, with a few short teeth or rounded bows; the _ basal 
half-line is faint with an outward angle and a rounded bow to its end; the t.a. line 
is heavy black, with additional black shading basally, of short zig-zags to the 
fold, thence toothed inward and then somewhat outwardly bowed to inner margin; 
the t.p. line is fine, scarcely toothed except for the short two large teeth, thence 
an inward arc to the blacker horizontal segment in the fold, thence bowed outward 
to the inner margin; and subreniform usually separate, irregular, small; the grayer 
reniform rather small, upright, set out by an irregular, conspicuous white border: 


34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


beyond the t.p. line a variable brown band is present, followed by a faint white 
subterminal line; a terminal line of shallow crescents; from the large teeth of the 
t.p. a dark dash to below the apex. The hindwing is orangish yellow; the outer 
band moderate, leaving a small yellowish to orangish apex, and is usually broken 
before anal end, swollen toward the upper end and again in the M:Cu area; 
this bends and extends to the base of the wing. Beneath, the hindwing repeats 
the upperside, except paler; the forewing is as usual. The species texarkana is 
most readily separated from other species of the crataegi group, when, with its 
form bridwelli, it is compared with them in series. I name this form bridwelli 
(Fig. 2) after L. H. Bridwell who reared from Crataegus many specimens of 
both forms. The type 6 of bridwelli, Forestburg, Texas, 1200 ft., 13 May 1939. 
In addition 32 specimens have been set aside as this form. It is characterized 
by a much blacker basal area including a heavy black t.a. line, some becoming 
browner and grayer toward the base. The types have the lighter gray median 
area continued to the inner margin, but many of this form have the inner margin 
darker, in an extension of the basal area. 

Holotype. ¢, Forestburg, Texas, 1200 ft., 10-12 May, 1940, L. H. Bridwell, 
probably reared from larvae beaten from Crataegus. 

Allotype. @ ditto, 13 May, 1939, reared from Crataegus. 

Paratypes. 40 ¢ and 42 9; Forestburg, Texas, 12 ¢, 15 9, May 1939, mostly 
reared; Forestburg, May 1940, 28 ¢ and 25 9; Lincoln Co., Arkansas, May 3 
and 4, 1938, 29; all L. H. Bridwell. From other localities I have the following: 
a ¢ labelled “Texas”, W. N. Tallant collection; a 2 College Station, Texas, April 
19, 1929; a ¢ Churchill Bridge, Brazoria Co., Texas, 3-V-1968 (A. and M. E. 
Blanchard); all three too poor to include in the types. The Blanchard’s showed 
me the Churchill Bridge collecting locality when I visited Houston. 

Expanse (part reared) of a series of each: ¢ 41 mm.; 2 43 mm. 


Catocala lincolnana Brower new species 
(Fig. 3) 


Lincolnana is a simply marked species, brownish gray on basal, apical, and 
anal areas, and along inner margin beyond transverse anterior line; the remainder 
light gray with some brown. The basal half-line is evident, the transverse anterior 
line is heavy, black, of short zig-zags, nearly straight and oblique to the inner 
margin; the reniform, a small upright oval, fringed with white; the subreniform 
irregular, rather large, and separate; the transverse posterior line is fine, shortly 
toothed, largely transverse to the fold, when a long inward segment carries it 
to near the t.a. line. The subterminal line of lighter short hastae is evident, the 
terminal line of short bars scarcely evident. The hindwing above is as usual in 
the Crataegus-feeders, with a larger orangey apical area, a large oval black spot 
before the anal angle; the median band a bit narrow and extending to the base; 
this pattern repeated below on the hind wing and the usual pattern beneath on 
the forewing. 

Expanse: ¢ 47 mm. 

Holotype. ¢, Lincoln Co., Arkansas, 1 June 1937, L. H. Bridwell. Type in 


author’s collection. 


Catocala johnsoniana Brower new species 
(Fig. 6) 
The ground color of the forewings is ashy gray with prominent black lines. 
Beyond the transverse anterior line, from the costa a darker oblique shade passes 
across the reniform and on to the transverse posterior line above the subreniform; 


VotumMeE 30, NuMBER 1 35 


Figs. 1-6. Catocala species. 1, Catocala texarkana, Forestburg, Texas, 10-12 
May 1940, ¢, holotype; 2, C. texarkana form bridwelli, Forestburg, Texas, 13 
May 1939, §; 3, C. lincolnana, Lincoln Co., Ark., 1 June 1937, ¢, holotype; 4, 
C. erichi, Green Valley Creek, San Bermardino Mts., Calif., ex ovum, 19 May 
1966, ¢, holotype; 5, C. californiensis, Valyermo, Los Angeles Co., Calif., 27 
June 1957, 3. holotype; 6, C. johnsonana, Kernville, Kem Co., Calif., 17 June 
1965, ¢, holotype. 


beyond and above the reniform a less defined paler blotch extends to the t.p. line. 
The basal half-line is black and transverse; the t.a. line is heavy and black, nearly 
straight to Cus, then curved and tapered to A:, thence obliquely outward to inner 
margin. The basal dash is present. The t.p. line extends outward on R, then 
barely toothed to most outward point, thence with three short acute teeth to the 
loop forming the small pale subreniform (open, closed, or disconnected), thence 
strongly curved to A:, and nearly direct to the inner margin. The subterminal 
line is whiter gray, the terminal line of connected crescents. The fringes are gray. 
The medium-sized inconspicuous reniform is commonly darker-centered and out- 
lined by paler scales. The hindwing above is orangish yellow, the outer black 
band well developed with an apical patch of ground color, usually a small spot 
broken off before the anal angle; median black band narrowed, with an outward 
bulge on R and a second one Mz to Cuz, little tapered, usually but little turned 
inward and ending bluntly. Beneath, the forewing is pale yellow with the usual 
dark and light areas; the hind wing orangish yellow; the outer black band medium 


36 JoURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


width, with small apical area of ground color, usually unbroken. The median 
band rather narrow, oblique, nearly even, until Mz, when set out more than width 
of band, soon narrowed, the very end of the terminal portion turned toward inner 
margin, and ending abruptly. 

Expanse: ¢ 49.5 mm.; 9 52 mm. 

Holotype. ¢, Kernville, Kern Co., Calif., 17 June 1965 (Erich Walter). 

Allotype. ¢@, Kern River Canyon, Kern Co., Calif., 29 May, 1954 (Wm. A. Rees). 

Paratypes. 5 &, 1 @: Kern Canyon, Kern Co., Calif., elevation 2800 ft., 29 
May, 1954 ¢ (C. A. Hill); Kernville, Kern Co., Calif., 2500 ft., 16 June 1965 
4, 21 June 1965 2 ¢ (Erich Walter); 18 June 1966 ¢ (J. W. Johnson and E. 
Walter); Hughes Lake, Los Angeles Co., Calif., 23 June 1971 2. Holotypes 
and paratypes collection A. E. Brower. Allotype and part of paratypes returned 
to Natural History Museum, Los Angeles. 


Catocala californiensis Brower new species 
(Fig. 5) 


The ground color of the forewings is ashy gray, varying from light to brownish 
gray, with a broad lighter band from costa beyond the transverse anterior line to 
the subreniform, and including a still whiter gray subreniform; with a black shade 
from mid-costa over reniform to strongly inbowed transverse posterior line, which 
has short stout teeth on M; and M2 with a short tooth inward on the fold. The 
lines are heavy black, especially the transverse anterior line, which is nearly 
straight to the fold; the t.a. set out by a paler gray basal edging, the t.p. less so by 
an outer paler line; the basal half-line black, somewhat jagged; the basal dash 
absent; a more or less evident subterminal band of lighter sagittate marks; the 
terminal black scalloped band is usually continuous; the fringes gray. The hind- 
wing above has the inner black band narrow, nearly straight, swollen outward 
from Ms to Cur, then greatly attenuated and sharply angled inward toward the 
inner margin; the outer band of moderate width, usually broken before anal end, 
and with a yellow patch before apex. Beneath, both wings are much paler, the 
forewing with the usual pattern, and the hind wing a duplication of the upper side. 

Expanse: ¢ 46 mm.; 2 47.5 mm. 

Holotype. ¢, Ranch two and one half miles south-southwest of Valyermo, 
Los Angeles Co., Calif., 4800 ft., 27 June 1957 (Noel McFarland). © 

Allotype. 92, Pinyon Flats, Santa Rosa Mts., Calif., 10 July, 1967, ultra-violet 
light, (J. W. Johnson and Erich Walter). 

Paratypes. 10 ¢, 11 9: near Acton, Mint Canyon, Los Angeles Co., Calif., 
3 June 1950 g (Chas. A. Hill); Tujunga, Los Angeles Co., Calif., 26 June 1940 9 
(C. Henne); ranch two and one half miles south-southwest of Valyermo, Los 
Angeles Co., Calif., 4800 ft., 26 June 1957, 3 ¢, 3 9, and 1 July 1964 @ at 
black light (Noel McFarland); nine miles southeast of Pearblossom, Los Angeles 
Co., Calif., 27 June 1947 g (Noel McFarland); Pinyon Flats, Calif. [Santa Rosa 
Mts.], 5 July 1970 6, 16 July 1967 ? (the last two J. W. Johnson handwriting ); 
Pinyon Flats, Santa Rosa Mts., Calif., 4000 ft., 3 July 1970 6, 10 July 1967-2 
6,1 9, 11 July 1967 1 ¢,3 Q, ultra-violet light, 16 July 1969 6, 18 July 1968 2 
?, ultra-violet light (J. W. Johnson and Erich Walter ) 


Catocala erichi Brower new species 
(Fig. 4) 


The forewings are black, tinged with brown, apearing more or less overscaled 
with somewhat smaller white scales with a bluish sheen, best developed in the 
median area. Basal half-line of black inner, and white outer lines; transverse anterior 
line, strongly zig-zag of white inner and black outer portions, appearing black on 


VoLuME 30, NuMBER | 37 


both sides; before and somewhat below the reniform is a striking white patch, and 
another in the median area between the inward bulge of the transverse posterior 
line and the subterminal line; the apical area is brownish black, and from it to 
the outer angle the margin is variably gray and white; the reniform is rather large, 
upright, irregular, vague, partially outlined with black and may be more so outside 
with white; the subreniform is usually separate, lighter with brownish overshading; 
the transverse posterior line is short-toothed, largely transverse, black inside, white 
outside; the subterminal line, of short broad white chevrons, is prominent; the 
terminal line is of short bars or shallow crescents; terminal dark crescents form 
a more or less continuous line at the base of the fringe. On some the black 
obscures all except the two white patches and the subterminal line. The hind- 
wings are deep red with strong black bands, with white to pink margined apices; 
the median black band more or less abruptly ending or greatly contracted with 
the narrowed end turned upward, with long black hair near base of wing. Beneath, 
on forewing the wings are much paler with the bands as usual, and on _ hind- 
wing a paler reproduction of the upper side. During separated years broods of 
this species were reared from eggs and with caught specimens form an unusually 
similar series of specimens, well separated from francisca Hy. Edwards, and from 
the more northern complex of mariana, Hy. Edwards, edwardsi Kusnezov and, eldo- 
radensis Beutenmiiller. Larvae of erichi (named for the chief collector), two 
broods, lost in the last instar the dark brown patch on the abdominal hump, 
while larvae of francisca kept their patch. 

Expanse (nearly all reared); ¢ 66 mm.; 2 68 mm. 

Holotype. ¢, emerged 19 May 1966, reared by J. W. Johnson, ova Green Valley 
Creek, San Bernardino Mts., Calif., 7000 ft., Aug. 1965 (E. Walter). 

Allotype. @, San Bernardino Mts., Calif., 7000 ft., reared by J. W. Johnson, 
emerged May 1971, ova by E. Walter summer of 1970. 

Paratypes. 11 ¢, 8 @: Green Valley Creek, San Bemardino Mts., Calif., 
7000 ft., ova from female August 1965 by Erich Walter, reared by J. W. Johnson 
and Erich Walter, adults emerged May 18 until June 21, 1966, 4 6, 3 @. Of a 
second lot of ova, summer of 1970, by Erich Walter, reared by both Johnson 
and Walter in 1971, 4 ¢, 3 2 emerged late June to July 27; Hathaway Creek, 
near Barton Flats, San Bernardino Mts., Calif., 2 August 1940 ¢ (C. Henne); 
Camp O-ongo, near Running Springs, San Bernardino Mts., Calif., 28-31 August 
1967 @ (C. L. Hogue). Types at present in my collection, paratypes returned 
to Natural History Museum, Los Angeles, J. W. Johnson and Erich Walter. 


38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


A CHECKLIST OF THE BUTTERFLIES OF GRANT COUNTY, 
NEW MEXICO AND VICINITY? 


CiirForD D. FERRIS” 
College of Engineering, University of Wyoming, Laramie, Wyoming 82071 


Little has been published on the butterflies of New Mexico, although 
the type localities for several species lie in the northern and western 
portions of the state. Early collecting by H. Skinner and J. Woodgate 
and later collecting by A. B. Klots forms the basis for much of what we 
know about its butterfly fauna. Collecting in recent years by D. Cowper, 
R. Holland and M. E. Toliver has expanded our knowledge. Williams 
(1914), Hubbard (1965), Toliver (1971) and Holland (1974) have 
published papers which treat limited geographic areas within New 
Mexico. 

Since 1965, the author has collected actively in the southwestern por- 
tion of the state, primarily in Grant County, with some additional collect- 
ing in Catron, northern Hidalgo, Luna and western Sierra counties. 
The area so defined comprises a fairly uniform faunistic region. Although 
records are available from the Peloncillo Mts. in Hidalgo Co., they are 
not included as another faunistic region is involved. Hubbard’s paper 
lists butterflies from the Pinos Altos Mts. which lie in Grant Co. His 
list of 52 species, which excludes the Hesperioidea, represents only a 
part of the 157 confirmed species in the current study. A checklist of the 
presently known species appears in a subsequent section of this paper. 


Localities 


Most of the collecting was in various areas of the Gila National Forest, 
on several ranches and with scattered collecting in the desert areas south 
and southwest of Silver City. The primary forest collecting sites in- 
clude: the Burro Mts. south of Silver City; the Black Range east of 
Silver City; Cherry Creek Canyon (Pinos Altos Mts.), Signal Peak and 
Lake Roberts north of Silver City; and the Mogollon Mts., especially the 
Willow Creek area, in Catron Co. Sonoran desert localities include: 
along the Gila River, especially in the vicinity of Red Rock; along State 
Road 90 between Lordsburg and Silver City; vicinity of Faywood Hot 
Springs and City of Rocks State Park; and Hachita. 


| Published with the approval of the Director, Wyoming Agricultural Experiment Station, as 
Journal Article no. JA 713. ; 
* Research Associate, Allyn Museum of Entomology, Sarasota, Florida; Museum Associate, Los 


Angeles County Museum of Natural History, Los Angeles, Calif. 


VoLUME 30, NuMBER Il 39 


Vegetation types of the region are quite varied. Collecting sites in 
the Mogollon Mts. are generally open Canadian Zone meadows associ- 
ated with live streams, usually a ponderosa pine (Pinus ponderosa Laws. ) 
and willow (Salix sp.) association containing a variety of grasses, herbs 
and shrubs. The Black Range and sites north of Silver City are generally 
Transition Zone associations with ponderosa pine, pinyon pine (Pinus 
monophylla Torr. & Frem.), Douglas fir (Pseudotsuga taxifolia var. 
glauca [Mayr] Sudw.) and aspen (Populus tremuloides Michx.) at the 
higher elevations, and at lower elevations a variety of deciduous trees 
and shrubs including Acer, Alnus, Celtis, Cercocarpus, Crataegus, Prunus, 
Populus sp. (cottonwood), Rhus, Salix and Platanus wrightii S. Wats. 
Grasses and herbs abound, especially near permanent water. Canadian 
Zone vegetation occurs at higher elevations, especially on Signal Peak 
9000’. The Burros Mts. are covered primarily with pinyon pine, alligator 
juniper (Juniperus deppeana Steud.) and Emory Oak (Quercus emoryi 
Torr.); several other species of oak are also present. Acacia greggii 
Gray, Chilopsis linearis (Cav.) Sweet. and Prosopis juliflora (Swartz) 
DC. occur at lower elevations. Principal vegetation types in the desert 
areas include grasses, Acacia, Agave, Chilopsis, Prosopis, Yucca and 
various cacti. Cottonwood, hackberry (Celtis sp.) and other deciduous 
trees are associated with riparian areas, and outcroppings of oak are 
common. 

Annual rainfall in the Silver City area averages 0.406 meters. The 
principal moisture occurs in the months January—March, July—October, 
with the heaviest precipitation during the summer months. July is 
usually the wettest month and May the driest. The mean temperature 
is 12.67°C with average highs and lows of 28.55°C and -4.83°C respec- 
tively. 

Specific collecting sites with alphabetic codes are given below. In 
several cases where only one or two records are from a locality, no code 
is used and the localities are clearly shown on the maps (Figs. 1 & 2). 
Either the code letters or the locality name appears on the maps. 

CATRON COUNTY. Whitewater Creek and Catwalk area near Glenwood 4800’ 
(CA); Datil ca. 7000’ (D); Gila Cliff Dwellings and EE Canyon 5600-6100’ 
(EE); Frisco Hot Springs 4500’ (FS); Glenwood 4700’ (G@); Gila Wilderness 
Area (varying elevations) (GW); Gilita Creek 7800-8000’ (GK); Little Creek 
and Gila River Junction 5750 (LC); Luna 7000’ (LN); Mogollon 6600’ (M); 
Pleasanton 4600’ (P); Pie Town ca. 8100’ (PT); Quemado Lake 7200’ (QL); 
Reserve ca. 5800’ (R); Silver Creek, crossing at State Road 78 8560’ (SK); SS 
Basin T10S, RI5W, S21 7600’ (SS); Willow Creek (camp area) 1800’ (WC). 

GRANT COUNTY. Ancheta Canyon 5800-6400’ (AC); Ash Spring Canyon 
6700-6900’ (ASC); Bayard 5880’ (B); Black Canyon Camp 7000’ (BC); Bill 
Evans Lake 4700’ (BE): Burro Mountains (Homestead Road and USFS 851) 
ca. 6650’ (BM); Bear Mountain 6500-7000’ and Bear Mtn. Ranch 6250’ (BR); 


40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Cliff 4500’ (C); Copperas Canyon 6000-7400’ (CC); Cherry Creek Canyon and 
McMillen Canyon 6700-7300’ (CCC); Cottonwood Canyon 6300’ (CN); Dry 
Gallinas Canyon 6800’ (DGC); East Canyon, Black Range 7400’ (EC); USFS 
152, Black Range (varying elevations) (F); Ft. Bayard, E. of Silver City 6000- 
6200’ (FB); Faywood Hot Springs 5000’ and City of Rocks State Park 5200’ (FH); 
Fierro 6600’ (FI); Gallinas Canyon area, Black Range 6800-7000’ (GC); Gold 
Gultch 5800-5900’ (GG); Gila River near Gila 4250’ (GR); East Fork of Gila 
River ca. 6000’ (GRE); Hachita 4100’ (H); Hachita Mountains 4800-6000’ (HM); 
Iron Creek Camp, Black Range 7100’ (IC); Kneeling Nun Vista, Black Range 6990’ 
(KN); Kwilleylekia Ruins 4550’ (KR); Lower Gallinas Canyon 6400-6800’ (LGC); 
Lake Roberts 6000’ (LR); L-S Mesa, north of Silver City, USFS 853 6200-6500’ 
(LS); Little Walnut Creek 6600’ (LW); Mule Creek area, Jupe Means Ranch 
5800-6300’ (MC); Mill Creek Canyon 6800-7100’ (MI); Mogollon Creek 4640’ 
(MK); Moon Ranch 5200’ and Buckhorn 4800’ (MR); Pinos Altos 7000’ (PA); 
Pine Cienega 6500’ (PC); Ricolite Canyon near Red Rock 4200’ (RC); Red Rock 
4000’ (RR); Sherman 5600’ (S); SA Canyon, Gila Wilderness Area 6800-7400’ 
(SA); Silver City 5900-6100’ (SC); Sapillo Creek 5800-6200’ (SE); San Lorenzo 
5800’ (SL); Skate’s Canyon 6600’ (SN); Soldier Canyon 6300’ (SO); Signal 
Peak 8000-9000’ (SP); State Road 90, milepost xx between Silver City and Lords- 
burg (SR-xx); Santa Rita 6500’ (ST); Tyrone 5700-5800’ (T); Thunderbird 
Camp (1 mi. N) 6500’ (TC); Upper Gallinas Canyon 7000’ (UGC); Vanadium 
5950’ (V); White Signal (vicinity) 6000’ (WS). 

HIDALGO COUNTY. Animas 4300’ (A); Animas Valley 4700-5100’ (north- 
south) (AN); Deming ca. 3000’ (DM); Lordsburg (vicinity) 4240’ (L). 

LUNA COUNTY. Columbus (vicinity ) 4000’ (CO). 

SIERRA COUNTY. Emory Pass area 8200’ (EP); Kingston 6250’ (K). 


Checklist 


In the list of species which follows, the counties and localities are noted 
as well as the months in which specimens have been taken. Additional 
comments are appended when applicable. Excepting the Hesperioidea, 
the order of families generally follows that proposed by Ehrlich & Ehrlich 
(1961). Subfamilies are not designated (Hesperioidea excepted) and 
genera are listed alphabetically according to recent revisionary work. 
Relative abundance is not noted, as this is strongly seasonal ( year-to- 
year variation) and is affected by climatic conditions. Single specimen 
records are noted. Several collectors have kindly supplied records which 
are so designated: (HA) Bruce Harris, (H) Richard Holland, (R) 
Kilian Roever, (T) Michael Toliver, (Z) Dale A. Zimmerman. William 
Baltosser provided the Gila Wilderness records (GW). 


MEGATHYMIDAE 


Agathymus aryxna (Dyar). Grant Co.: HM; CCC, WS (Z); T (R, T). Hidalgo 
Co.: AN (T). July, immatures in Agave palmeri Engelm.; salle in October. 
Agathymus neumoegeni (Edwards). Catron Co.: M (R). Grant Co.: PA; CCC 
ies (R). August—-October, immatures in Agave parryi Engelm.; adults a 
ctoper. 
Megathymus coloradensis navajo Skinner. Catron Co.: GW; D, M, Pi RGRoe 
Grant Co.: UGC; 7 mi. N of SC (Z); CG IG MG, ST, Ts Ws (R). Aoaleye 


VOLUME 30, NUMBER 1] 


ARIZONA 


Road 


County Line 
sta Nat! Forest Boundary 
ee Ruiverr 


Fig. 1. Collecting sites in Catron Co., New Mexico. 


HESPERIDAE-PYRGINAE 


4] 


CATRON 


Miles 
-#—_____++—_____ 
O /O 20 


Autochton cellus (Boisduval & LeConte). Catron Co.: EE (Z) Grant Co. CCC, 


SP. July. 


Celotes nessus (Edwards). Catron Co.: CA (Z). Grant Co.; RR. May. 
Cogia caicus moschus (Edwards). Catron Co.: G. April. One specimen. 


42 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


C. hippalus hippalus (Edwards). Hidalgo Co.: AN (T). August. 

Epargyreus clarus huachuca (Dixon). Catron Co.; GW Grant Co.: CCC, EC, 
IC; SC (Z). May-July. 

Erynnis brizo burgessi (Skinner). Grant Co.: ASC, CCC, EC, F, FB, IC, LS, LW. 
March—May, July. 

E. funeralis (Scudder & Burgess). Grant Co.: BM, CCC, F, MC, SP, SR-26. April— 
May, August-September. 

E. icelus (Scudder & Burgess). Catron Co.: SK, WC and vicinity (R). June. 

E. meridanus meridanus Bell. Catron Co.: G, W of M. Grant Co.: FB, MC. April, 
August. 

E. pacuvius pacuvius (Lintner). Catron Co.: GW. Grant Co.: CCC, LW, PA. 
March, May—June, August. 

E. (persius) fredericki H. A. Freeman. Catron Co.: G, LN. Grant Co.: CCC, 
MC, SA, UGC. March—May, August. 

E. telemachus Burns. Catron Co.: G, GW Grant Co.: ASC, BM, BR, CCC, EC, F, 
GC, IC, LGC, LW, SP. March—June. 

E. tristus tatius (Edwards). Catron Co.: G, GW. Grant Co.: BM, MC. July— 
August. 

Pholisora catullus (Fabricius). Catron Co.: G. Grant Co.: B, FH, T. August. 

Pyrgus communis complex. P. c. communis and P. c. albescens are synchronic and 
sympatric in some areas. They are widespread in all life zones and are found 
wherever members of the Malvaceae grow. Males may be positively identified 
by their genitalia. Only partial records are listed below. 

P. c. albescens Plotz. Catron Co.: G, LC, P, WC. Grant Co.: AC, BC, BM, CCC, 
MC, SL, SO, TC, WC. June—October. 

P. c. communis (Grote). Catron Co.: WC. Grant Co.: EC, IC, LS, MC, UGC. 
May-June, August, October. 

P. scriptura (Boisduval). Catron Co.: G. March. This species is bivoltine in many 
areas. I have not yet found the summer brood in Catron/Grant Co. 

P. xanthus Edwards. Catron Co.: 3 m. E of M (H); WC area (R). April, June. 

Staphylos ceos (Edwards). Catron Co.: G. Grant Co.: GR, RR. Hidalgo Co.: AN 
(T). April-May, July—August. 

Thorybes pylades (Scudder). Grant Co.: AC, BM, CCC, F, LS, MC, TC. Sierra 
Co.: EP. May—August. 

Timochares ruptifasciatus (Plétz). Grant Co.: LW (Z). One specimen on 
20-ix-63. 

Zestusa dorus (Edwards). Grant Co.: CCC, EC, IC. April-May. 


HESPERUDAE-HESPERIINAE 


Amblyscirtes aenus ssp. Catron Co.: CA (R). Grant Co.: CC: CCC, GR, Mimbres 
Canyon (Mimbres River) (R). May-July. The southwestern race of A. aenus, 
which probably represents an undescribed subspecies, is easily confused with 
A. cassus. Both occur in Grant and Catron Co. They are genitalic distinct 
entities. In aenus, the uncus has two distinct processes on each side and the 
valva is relatively blunt with the distal end straight up; in cassus, the uncus 
has one distinct and finely tapered process on each side with only a suggestion 
of the second process, and the distal end of the valva is quite pointed and 
somewhat recurved. The ventral surface of the forewings exhibits considerably 
more orange-fulvous color in cassus than in aenus. The dorsal surface spots 
in cassus are entirely orange-fulvous while they are cream to barely fulvous in 
aenus. 

A. cassus Edwards. Catron Co.: GW. Grant Co.: CC, CCC, SP, UGG win 
August. 

A. eos (Edwards). Grant Co.: LS, MC. August. 


VoLuME 30, NuMBER | 43 


CATRON 


=o) yo}|0004 


x 


\ 


- WC... 
SNe 
Pleasanton \3. 


SIERRA 


e Cook's Pk. 
8408 


ARIZONA 


Reeiebirg = 


Columbus 


MEXICO 


Fig. 2. Collecting sites in Grant Co. and surrounding areas, New Mexico 


. Note: 
Fig. 2 is to the same scale and uses the same legends as Fig. 1. 
A. exoteria ( Herrich-Schaffer ). Catron Co.: GW Grant Co.: ASC. July. 
A. nereus (Edwards). Grant Co.: CCC, GRE, (R). July—August. 
A. oslari (Skinner). Catron Co.: CA (R). Grant Co.: MC (R). May—June. 
A. phylace (Edwards). Grant Co.: TC. July. 
A. 


simius Edwards. Grant Co.: LS, MC, SR-26, TC. July—August. 


44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Atalopedes campestris (Boisduval). Grant Co.: IC. May. 

Atrytonopsis deva (Edwards). Catron Co.: G. Grant Co.: CC, CCC. April-June. 

A. lunus (Edwards). Grant Co.: LS. August. 

A. pittacus (Edwards). Catron Co.: G. Grant Co.: BR, CCC. March—June. 

A. python (Edwards). Grant Co.: CC; SC (Z). June. 

A. vierecki (Skinner). Grant Co.: BM, CCC, GG, LS, MC; SC (Z). May—June. 

Copaeodes aurantica (Hewitson). Catron Co.: GW. Grant Co.: BE, BM, CN, 
MC, PC, RR; C, SC, T (Z). April—October. 

Euphyes vestris vestris (Boisduval). Catron Co.: EE. Grant Co.: CCC. June-July. 
The correct subspecific name to apply to Rocky Mountain vestris is in doubt. 
Coastal California vestris are larger and lighter in color than inland western 
vestris. 

Hesperia harpalus susanae Miller. Catron Co.: GW, WC. July—August. 

H. pahaska pahaska Leussler. Catron Co.: FS. Grant Co.: BM, LS, MR, SE, SP, 
V. May-June, September—October. Specimens from SW Hidalgo Co. are 
referable to H. p. williamsi Lindsay. Grant Co. specimens seem to be nominate 
pahaska. 

H. uncas uncas Edwards. Grant Co.: AC, LS, MC, MR, SC, SR-26. July—Septem- 
ber. Intergrading with H. u. lasus (Edwards) occurs. 

H. viridis (Edwards). Catron Co.: WC Grant Co.: BR, CC. June-August. 

H. woodgatei (Williams). Catron Co.: FS. Grant Co.: MR, TC. July, September-— 
October. 

Oarisma edwardsii (Barnes). Grant Co.: DGC. August. 

O. garita (Reakirt). Catron Co.: WC and vicinity, LN (R). June-July. 

Ochlodes snowi (Edwards). Catron Co.: GW, WC. July—August. 

Piruna pirus (Edwards). Catron Co.: SK (R). Grant Co.: GR (R). June. 

P. polingii (Barnes). Catron Co.: GW. Grant Co.: CCC, IC. Early July, August. 
Possibly bivoltine. 

Poanes taxiles taxiles (Edwards). Catron Co.: GW, LC, SK. Grant Co.: CCC, 
DGC, LGC, IC, SP. Sierra Co.: EP. July-August. 

Polites themistocles (Latreille). Catron Co.: LN, GK (R). June-July. 

Stinga morrisom (Edwards). Grant Co.: LGC. April. 

Yvretta carus (Edwards). Grant Co.: MC. August. 

Two species have been collected in the Peloncillo Mountains which may occur 
elsewhere in Hidalgo Co. or in Grant Co.: Amblyscirtes texanae Bell; Hesperia 
pahaska williamsi Lindsay (T). Hylephila phyleus (Drury) ought to be common in 
gardens in the Silver City area, but I know of only one record of a specimen ex- 
tracted from a car radiator on 30-viii-64 (HA). The vehicle was reported to 
have come from the Gila Wilderness Area, which means that the specimen came 
from northern Grant or southern Catron Co. In August, 1970, I saw a large pyrgine- 
like skipper nectaring at desert willow along the banks of the Gila River south of 
Red Rock, Grant Co. Because of its position, it was not possible to net it. It 
exhibited large white hyaline spots on the forewings and appeared to be one of 
the following: Polygonus leo arizonensis (Skinner), Codatractus arizonensis ( Skin- 
ner), Pyrrhopyge araxes arizonae (Godman & Salvin). The latter was quite common 
at the time in southern Arizona. P. 1. arizonensis has been taken in the vicinity of 
Alamagordo, Otero Co. (H). 


PAPILIONIDAE 


Battus philenor philenor (Linnaeus). Catron Co.: GW, P. Grant Co.: FH, MC, 
SC, SO, RR. June-September. I have sighted numerous adults flying across 
desert roads in many areas of Grant Co. 

Papilio bairdii bairdii Kdwards. Catron Co.: G. A single female of typical Cali- 


VoLuME 30, NUMBER 1 45 


fornia bairdii phenotype, 8—vi-66. Hubbard lists a questionable record from 
the Pinos Altos Mtns., Grant Co. See comments under P. polyxenes. 

P. cresphontes cresphontes Cramer. Grant Co.: BM, CCC, SC, WS vicinity (Z). 
Tune—August. 

P. multicaudata (Peale MS.) Kirby. Catron Co.: GW, SK; G (Z). Grant Co.: 
CCC, EC, IC, LGC, LS, SP; BM, RR (Z). Grant-Hidalgo Co. line on State 
Road 90. March, May—August. 

P. polyxenes complex. Possibly all polyxenes-like specimens from the area should 
be referred to this species. Three specimens have been taken which in facies 
resemble P. rudkini clarki Chermock & Chermock, based upon comparison 
with paratypes of clarki. These are from Grant Co.: LR, 11—viii-68, a fresh 
male; RR, 3—vi-73, two females. 

P. polyxenes asterius Stoll. Grant Co.: H, LR, LGC, LS; SC, SP (Z). May, August. 
The subspecific name asterius has been applied, but specimens from Grant Co. 
are rather different from eastern asterius. 

P. rutulus arizonensis Edwards. Catron Co.: G, GW. Grant Co.: CCC, EC, SP. 
May-—June. 


PIERIDAE 


Appias drusilla poeyi (Butler). Grant Co.: SC (Z). One specimen on 24-vi-71. 
Anthocharis sara inghami Cunder. Catron Co.: CA, G, GW, LC. Grant Co.: ASC, 
CCG, GR, LGC, LW, RR, UGG; C, SC (Z). March-April. 

Colias alexandra ssp. Catron Co.: GW, SK, WC. June, August. See discussion of 
this subspecies in Ferris (1973). 

C. eurytheme Boisduval. Catron Co.: G, GW, LC, P, SK, WC. Grant Co.: BM, 
CCC, H, IC, LS, MC, RR, SP. Hidalgo Co.: L; 20 mi. S of H (Z). March— 
September. 

C. philodice eriphyle Edwards. Grant Co.: C, GR, SC (Z). April, September, 
November. Es 

Colias (Zerene) cesonia (Stoll). Catron Co.: GW, LC. Grant Co.: AC, MK, SC; 
5 mi. S of C, 8 mi. W of SC (Z); CCC, IC (T). March, June—November. 

Euchloe hyantis lotta (Beutenmiiller). Grant Co.; CCC, GR, SP. March, May. 

Eurema mexicana (Boisduval). Grant Co.: BR, CCC, CN, FB, LGC, SC, SP. 
Hidalgo Co.: L. April—October. 

E. nicippe (Cramer). Catron Co.: G, P. Grant Co.: CCC, FH, H, LGC, MC, 
RR, SR-26. Hidalgo Co.: L. March-September. Common and widespread. 
Nathalis iole Boisduval. Catron Co.: GW. Grant Co.: B, BM, CCC, IC, LGC, LS; 

6 mi. E of SC (Z). April—September. 

Neophasia menapia menapia (Felder & Felder). Catron Co.: GW; WC (Z). Grant 
Co.: CCC, SP; PA (Z). July. Associated with ponderosa pine. 

Phoebis agarithe agarithe (Boisduval). Catron Co.: P. Single female on 22-viii-71. 

P. sennae eubule (Linnaeus). Catron Co.: G, GW, P. Grant Co.: C, FH, LS, 
RR, SC, ST. Hidalgo Co.: L. August-September. 

P. sennae marcellina (Cramer). Grant Co.: A, AN, UGC; BM (Z); CCC, IC (T). 
Luna Co.: CO. April, August-September. 

Pieris napi mogollon Burdick. Catron Co.: GW, SK, WC. May, August. 

P. protodice protodice (Boisduval & LeConte). Catron Co.: GW, LC, WC. Grant 
oie COC ECVE. EB. 1G: sLS 2MC. PC; RR, SP: EW;.T,, SG (4). 
Hidalgo Co.: L. Luna Co.: 20 mi. S of DM (Z). March-April, form vernalis 
Edwards; June—September. 

P. rapae (Linnaeus). Catron Co.: GW. Grant Co.: B; SC (Z). March-April, June, 
October. 

P. sisymbrii elivata (Barnes & Benjamin). Catron Co.: CA. Grant Co.: CCC, EC, 
F, GC, GR, IC, LW, UGC; SC (Z). March—May. 


46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


RIODINIDAE 


Apodemia mormo nr. mejicanus (Behr). Catron Co.: CA (Z). Grant Co.: BE, BM, 
LS: CCC, SC (Z). Hidalgo Co.: SR-11. May-June, August. 

A. nais (Edwards). Catron Co.: GW. Grant Co.: CCC, MI, PA. June-July. 

A. palmerii palmerii (Edwards). Catron Co.: FS. Grant Co.: GR, RR. Hidalgo 
Co.: L. Luna Co.: DM. June, August-September. 

Calephelis nemesis nemesis (Edwards). Grant Co.: RR. Single fresh male on 18- 
vi-72. 


Emesis zela cleis (Edwards). Catron Co.: CA, GW. May. 


LYCAENIDAE 


Atlides halesus halesus (Cramer). Grant Co.: AC, BM, BR, S, SR-26; 9 mi. S of 
C, SC (Z). Luna Co.: Cook’s Peak summit (Z). March-April, August— 
September. 

Brephidium exilis exilis (Boisduval). Grant Co.: FH, KR, RR, SP, T; MK, SC (Z). 
June—October. 

Callophrys (Callophrys) apama apama (Edwards). Grant Co.: CCC. June-July. 

Callophrys (Incisalia) augustinus annetteae (dos Passos). Grant Co.: LGC, LS; 
SC (Z). April-June. 

C. eryphon eryphon (Boisduval). Catron Co.: LN, SK, WC (R). Grant Co.: 
IC (R). May-June. 

Callophrys (Mitoura) siva siva (Edwards). Catron Co.: EE, GW. Grant Co.: BM, 
CC, CCC, FB, FI, GC, LS; SC (Z). April-July. 

C. spinetorum (Hewitson). Catron Co.: GW, WC. Grant Co.: CCC, F, IC; SP 
(Z). May-June. 

Celastrina argiolus cinerea (Edwards). Catron Co.: G, GW, SK, WC. Grant Co.: 
BM, CCC, IC, PC, RR, SN, SP; SC (Z). March—October. 

Erora quaderna sanfordi dos Passos. Grant Co.: BR, CCC, LGC. February—May. 

Euristrymon ontario ilavia (Beutenmiiller). Grant Co.: BM, MC. May-June. 

Everes amyntula herri (Grinnell). Catron Co.: EE, GW, SK. Grant Co.: CCC, 
EC, F, IC, SP; SC (Z). Hidalgo Co.: AN. April-May, August. 

Glaucopsyche lygdamus arizonensis McDunnough. Catron Co.: GW. Grant Co.: 
CCC, F, SP. April—May. 

Hemiargus ceraunus gyas (Edwards). Catron Co.; EE, G. Grant Co.: AC, SR-26. 
Hidalgo Co.: L. June—August. 

H. isola alce (Edwards). Catron Co.: GW, LC. Grant Co.: AC, BC, BM, CCC, 
EC, FH, LS, MC, RR, SC, SE, T. Hidalgo Co.: L. Sierra Co.: EP. April— 
December. 

Hypaurotis crysalus (Edwards). Catron Co.: GW; CA (Z). Grant Co.: SP. July, 
September. 

Leptotes marina ( Reakirt). Catron Co.: GW. Grant Co.: BM, FH, IC, RR, SR-20; 
CCC, SC (Z). Hidalgo Co.: L. April-September. A widely distributed species. 

Lycaeides melissa nr. pseudosamuelis Nabokov. Catron Co.: GW, LC, LN. Grant 
Co.: LR, SA. August-September. Specimens are phenotypic pseudosamuelis. 

Lycaena arota schellbachi Tilden. Catron Co.: SK. August. 

Ministrymon ines (Edwards). Grant Co.: BM. One female on 24—v-75. 

M. leda (Edwards). Grant Co.: BM, RR; CCC (Z). May-June. 

Phacostrymon alcestis oslari (Dyar). Grant Co.: SR—20. July. 

Plebejus (Icaricia) acmon texanus Goodpasture. Catron Co.: G, GW, LC, WC. 
Grant Co.: AC, BM, CCC, F, FH, GC, IC, LS, MC, PC, SN, SP, SR=26.5¢; 
SC (Z). Hidalgo Co.: L. Sierra Co.: EP. April—October. 

P. icarioides buchholzi dos Passos. Grant Co.: CCEeSP itme: 

Plebejus (Plebejus) saepiolus nr. gertschi dos Passos. Catron Co.: SK, WC and 
vicinity (R). June. 


VoLUME 30, NUMBER 1 | 


Shijimiaeoides battoides centralis (Barnes & McDunnough). Grant Co.: LGC. One 
fresh male on 8—viii-75. Genitalia checked to verify species. See Shields (1974) 
for generic discussion. 

S. rita rita (Barnes & McDunnough). Grant Co.: AC, BM, SC, SR-26, T. Hidalgo 
Co.: AN, SR-13 (T). August. 

Strymon melinus franki Field. Catron Ce.: GW, LN. Grant Co.: BC, BM, CCC, 
F, GR, MC, RR, SC, SE, SN, T. Hidalgo Co.: L. April—October. 


LIBYTHEIDAE 


Libytheana bachmanii ssp. Catron Co.: FS. Grant Co.: FH, GR, MK, RR; SC (Z). 
March, August-September, November. In a series of specimens from this region, 
individuals can be assigned to both bachmanii (Kirtland) and _ larvata 
(Strecker). 


NYMPHALIDAE 


Agraulis vanillae incarnata (Riley). Grant Co.: SC. July-August. Scarce. 

Anaea andria andria Scudder. Catron Co.: EE, G. Grant Co.: BM, CCC, GR, RR, 
SR-26; MK, SC, 6 mi. W of SC, T (Z). Grant-Hidalgo Co. line along State 
Road 90. March—April, August-September. 

Asterocampa celtis montis (Edwards). Catron Co.: GW. Grant Co.: FH, GR. 
Sierra Co.: K. June, August-September. 

Chlosyne gabbii sabina (Wright). Catron Co.: G. Grant Co.: CCC, IC, LGC, SA; 
SC (Z). March—May, July. 

C. lacinia crocale (Edwards). Catron Co.: G, GW. Grant Co.: B, BR, CCC, FB, 
FH, GR, KR, MC; SC, WS (Z). Hidalgo Co.: L. Sierra Co.: K. April, June, 
August-September. All specimens are referred to crocale, although individuals 
may be selected from a large series which are phenotypic adjutrix Scudder, 
nigrescens (Cockerell) and rufescens (Cockerell). This region probably forms 
a blend-zone for the Texas adjutrix and the Arizona White Mtns. crocale. 

C. nycteis drusius (Edwards). Catron Co.: SK. June. 

Cynthia annabella Field. Catron Co.: GW. Grant Co.: CCC, V; SC, T (Z). 
August—October. 

C. cardui (Linnaeus). Catron Co.: WC. Grant Co.: BM, CCC, F, LS, MC, SC, 
WS; 5 mi. S of C (Z). Hidalgo Co.: AN (T). March—September. 

C. virginiensis (Drury). Catron Co.: GW, SK. Grant Co.: BM, CCC, F, IC, LR, 
SP; SC (Z). Hidalgo Co.: AN (T). April-June, August—October. 

Danaus gilippus strigosus (Bates). Catron Co.: G, GW. Grant Co.: CCC, FH, 
GR, MC, RR; 5 mi. S of C, SC (Z). Hidalgo Co.: L. May—September. 

D. plexippus plexippus (Linnaeus). Catron Co.: GW, LC. Grant Co.: C, FB, 
BEE MGs SC; R26, S: BM. lOvmi Sof C,.5 mi W of SG, LT CZ). June— 
September. 

Dymasia dymas dymas (Edwards). Grant Co.: RR. May-June, September. 

Euphydryas anicia alena Barnes & Benjamin. Catron Co.: CA. Grant Co.: 5-6 mi. 
NW of SC (Z). March—April. 

Euptoieta claudia (Cramer). Catron Co.: GW, P, SK, WC. Grant Co.: AC, BM, 
Beco He. it KH. LS MC RRO SP. I: SC (7%). Hidalgo Co.:A. April 
October. Widespread and common in all life zones. 

Limenitis archippus obsoleta Edwards. Grant Co.: C (Z). Hidalgo Co.: L. July- 
September. 

L. astyanax arizonensis Edwards. Catron Co.: GW, LC, SK. Grant Co.: CCC, 
LGC, MI; SC (Z). June—-September, November. 

L. weidemeyerii angustifascia Perkins & Perkins. Catron Co.: GW, Sk, WC. Grant 
Co.: BC, CCC, IC, LGC; C (Z). June-August. 


48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Limenitis (Adelpha) bredowii eulalia (Doubleday). Catron Co.: GW. Grant Co.: 
CCC, LS, MI, SC; 6 mi. SW of T (Z). May—September, November. 

Nymphalis antiopa (Linnaeus). Catron Co.: GW, WC. Grant Co.: B, BM, CCC, 
LGC; SC (Z). April—-September. 

N. californica californica (Boisduval). Catron Co.: GW. Grant Co.: CCC; SP (Z). 
March, June—July, October. 

N. milberti furcillata (Say). Catron Co.: GW, WC. August. 

Phyciodes campestris camillus Edwards. Catron Co.: G, GW, LC, WC. Grant Co.: 
H. Hidalgo Co.: L. August-September. 

P. mylitta arizonensis Bauer. Catron Co.: LN, SK. Grant Co.: BM, C, CCC, DGC, 
SA, UGC. Sierra Co.: EP. March-September. See Howe (1975) for nomencla- 
ture. 

P. picta picta Edwards. Catron Co.: G, LC, LN. Grant Co.: H, SA. Hidalgo Co.: 
L. May, August-September. 

P. tharos (Drury). Catron Co.: G. Grant Co.: C, GR, KR. August-September. 
Several distinct forms are commonly collected, including distincta Bauer. 
Phyciodes (Anthanassa) texana texana (Edwards). Grant Co.:; CCC, RR. May-— 

June. 

Poladryas minuta arachne (Edwards). Catron Co.: WC. Grant Co.: CCC, IC, SP. 
June-August. See Scott (1974) for nomenclature. 

Polygonia interrogationis (Fabricius). Grant Co.: SC. One specimen on 3-xi-62; 
6 mi. S of C. One specimen on 10-x-63 (Z). 

P. satyrus (Edwards). Catron Co.: WC. Grant Co.: CCC, IC, SP; SC (Z). May— 
August. 

P. zephyrus (Edwards). Catron Co.: GW, SK, WC. Grant Co.: SP (Z). One 
specimen on 30—vi-74; May, August in Catron Co. 

Precis lavinia coenia (Hibner). Catron Co.: GW. Grant Co.: BM, GR, KR, LGC; 
CCC, SC (Z). June-August, October. One male was taken at the Gila River 
locality on 10-viii-75 which is very close to nigrosuffusa (Barnes & McDun- 
nough ). 

Speyeria atlantis nausicaa (Edwards). Catron Co.: GW, SK, WC. Grant Co.: CC, 
CCC, IC, SP. May—August. 

S. nokomis nitocris (Edwards). Catron Co.: LC, WC. August-September. 

Texola elada perse (Edwards). Grant Co.: BE, GR, RR. May-June, August. 

Thessalia alma (Strecker). Catron Co.: GW, SS. Grant Co.: BM; 3 mi. N of WS 
(Z). June-July, September. 

Thessalia theona thekla (Edwards). Grant Co.: BM, BR, CCG) “ECReSGawvs: 
Hidalgo Co.: AN (T). May—June, August, October. 

Vanessa atalanta rubria (Fruhstorfer). Catron Co.: GW. Grant Co.: F B: SG. (7): 
Hidalgo Co.: A. June—September. 


SATYRIDAE 


Cercyonis meadii ssp. Catron Co.: M (R) Grant Co.: MC (R). August. Intensive 
searching for this species during August, 1975 in both Catron and Grant Cos. 
failed to produce a single specimen. 

Cercyonis oetus charon (Edwards). Catron Co.: GW, WC. July—August. 

Cyllopsis pertepida dorothea (Nabokov). Grant Co.: BM, BR, FB) SNe e@ars: 
SC; CCC, 3 mi. N of WS (Z). Sierra Co.: EP. June—September. See Miller 
(1974) for nomenclature and additional comments. 

Gyrocheilus patrobas tritonia (Edwards). Catron Co.: M (Z). Grant Co.: CCC, 
IB, UGC, Sierra Co.: EP. September—October. 

Megisto rubricata cheneyorum (R. L. Chermock). Catron Co.: GW. Grant Co.: 
CCC, FB, LGC; SC (Z). June-August. See Miller (1976). 

Neominois ridingsii ssp. Catron Co.: SS. One fresh male on 17—vi-72: 


VoLuME 30, NuMBER | 49 


Oeneis alberta daura (Strecker). Catron Co.: Crest Trail off State Road 78 (R). 
June. 

Asterocampa leilia leilia (Edwards) and Eurema proterpia (Fabricius) have been 
taken in the Peloncillo Mountains and may stray into the Silver City area. On 
23-viii-73, D. A. Zimmerman saw a specimen of Heliconius charitonius vasquezae 
Comstock & Brown at the north edge of the Western New Mexico University campus. 
The specimen was not captured, but Zimmerman is quite familiar with the species as 
he has collected it in Florida and the Neotropics. 


ACKNOWLEDGMENTS 


The author wishes to thank Michael E. Toliver and Richard Holland, 
both of Albuquerque, New Mexico, for providing records and comment- 
ing critically concerning the first draft of this paper. Bruce Harris, Clear 
Lake, South Dakota, Kilian Roever, Phoenix, Arizona, and Dr. Dale A. 
Zimmerman, Silver City, New Mexico kindly provided many additional 
records. Dr. John P. Hubbard, New Mexico Department of Game and 
Fish, Santa Fe and William Baltosser, Silver City, generously provided 
records from the latter’s collecting trips into the Gila Wilderness Area 
during 1975. Special thanks are due the author's cousin, Ralph A. Fisher, 
Jr., who lives in the Silver City area. He made a major contribution to 
the paper by collecting specimens over a ten-year period. Many of the 
early spring and late fall records are a result of his efforts. He also 
provided elevation data and other information used in the paper. 


LITERATURE CITED 


EuRuicH, P. R., & A. H. Exriicu. 1961. How to know the butterflies. W. C. 
Brown, Dubuque. 262 p. + vii. 

Ferris, C. D. 1973. A revision of the Colias alexandra complex (Pieridae) aided by 
ultraviolet reflectance photography with designation of a new subspecies. J. 
Lepid. Soc. 27: 57-73. 

Ho.uanp, R. 1974. Butterflies of six central New Mexico mountains, with notes on 
Callophrys (Sandia) macfarlandi (Lycaenidae). J. Lepid. Soc. 28: 38-52. 
Howe, W. H. (Eprror). 1975. The butterflies of North America. Doubleday & 

Co., New York. 633 + xiii p., 97 plates. 

Husparp, J. P. 1965. Some butterflies of the Pinos Altos Mountains, New Mexico. 
J. Lepid. Soc. 19: 231-232. 

Minter, L. D. 1974. Revision of the Euptychiini (Satyridae), 2. Cyllopsis R. Felder. 
Bull. Allyn Mus. Ent. No. 20. 98 p. 

. 1976. Revision of the Euptychiini (Satyridae), 3. Megisto Hiibner. Bull. 
Allyn Mus. Ent. No. 33. 23 p. 

Scott, J. A. 1974. Adult behavior and population biology of Poladryas minuta, 
and the relationship of the Texas and Colorado populations. Pan Pacif. Ent. 50: 
9-22. 

SHIELDS, O. 1974. Studies on North American Philotes (Lycaenidae). Bull. Allyn 
Mus. Ent. No. 19. 10 p. 

Touriver, M. E. 1971. Preliminary notes on the butterflies of Roosevelt County, New 
Mexico. J. Lepid. Soc. 25: 213-214. 

WituraMs, R. C., Jr. 1914. One hundred butterflies from the Jamez [sic] Moun- 
tains New Mexico (Lepid.), with notes and descriptions of a new species. 


Ent. News 25: 263-268. 


50 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


BIOLOGY AND TAXONOMY OF THREE GALL 
FORMING SPECIES OF EPIBLEMA (OLETHREUTIDAE) 


WILLIAM E. MILLER 


North Central Forest Experiment Station, Forest Service, USDA, 
St. Paul, Minnesota 55108. 


Epiblema Hibn. in North America comprises 39 species, mostly named 
over a half century ago (Brown, 1973). Natural history information is 
available for fewer than 10 species and consists chiefly of host and para- 
site records. Studies of several species received impetus during 1920-50 
because of their superficial resemblance to introduced lepidopterans and 
their role as alternate hosts of parasites. These interests are exemplified 
respectively by Thompson (1928) and Bobb (1942), the latter citing 
related literature. As far as known, Epiblema feed on Compositae; the 
late instar larvae bore in the stems. The three species treated in this 
paper produce rudimentary galls. 

Besides reviewing taxonomy, this paper augments natural history 
knowledge of scudderianum (Clem.) and gives new information about 
desertanum (Zell.) and discretivanum (Heinr.). It reports hosts, maps 
geographic records, describes feeding patterns and galls, outlines seasonal 
life histories, and integrates the literature on the genus. 


Taxonomy 


The following review identifies the species treated. It is abbreviated to 
primary citations. I examined all types. 


Epiblema scudderianum (Clemens) (Fig. 1) 


Hedya scudderiana Clemens (1860, p. 358) (Type in Academy of Natural Sciences 
of Philadelphia, illustrated by Miller (1973) ). 

Euryptychia saligneana Clemens (1865, p. 141) (Possible type in British Museum 
(Natural History) (Miller, 1973) ). Paedisca affusana Zeller (1876, p. 307) 
(Lectotype designated here, “Zeller Coll. Walsingham Collection . . .; Paedisca 
affusana Z. Il, 307 fig. 38 Am. Sept. Rssl . . .; Type; B. M. 2 Genitalia slide 
No. 5738,” British Museum (Natural History), left forewing and distal part of 
right forewing missing, hindwing length 8.0 mm). 


Epiblema desertanum (Zeller) (Fig. 2) 


Paedisca desertana Zeller (1876, p. 306) (Lectotype designated here, “Dallas, Tex. 
Boll; Type 14338; Paedisca desertana Z.; . . . Lectotype des. W. E. Miller,” 
Museum of Comparative Zoology, 4, forewing length 8.0 mm). 


VoLUME 30, NUMBER 1 Bik 


an 


Figs. 1-3. Wings of Epiblema. 1, scudderianum ¢, Ottawa Co., Michigan, fore- 
wing 8.0 mm; 2, desertanum 6, Ingham Co., Michigan, forewing 8.0 mm; 3, discre- 
tivanum 9, Chatham Co., Georgia, forewing 6.5 mm. 


Epiblema discretivanum (Heinrich) (Fig. 3) 


Eucosma discretivana Heinrich (1921, p. 823) (Type No. 23743, National Museum 
of Natural History ). 

Forewing patterns of scudderianum (Fig. 1) and desertanum (Fig. 
2) scarcely vary while that of discretivanum (Fig. 3) varies without 
regard to sex in degree of shading, particularly in basal and mid-dorsal 
areas. Size of adults is shown by the following forewing length ranges: 
scudderianum, 7.0-10.5 mm (136 examples not sexed); desertanum 7.0- 
§.5 mm (25); and discretivanum, 5.5-7.5 mm (37). Male genitalia are 


illustrated by Heinrich (1923) and female genitalia and wings by Brown 
(O73 


Hosts 


Except as noted, host records refer to identified adults that developed 
naturally on the indicated plant species. Plant specimens were diagnosed 
or verified by E. C. Leonard, National Herbarium; J. H. Beaman, Michi- 
gan State University; and Harmon Runnels, Ohio Agricultural Research 
and Development Center. 

Hosts of scudderianum in decreasing order of observation frequency 
were the Canada goldenrod complex, Solidago altissima L.-canadensis L.; 
tall goldenrod, S. gigantea Ait.; early blooming goldenrod, S. juncea Ait.; 
elm leaved goldenrod, S. ulmifolia Muhl.; and prairie goldenrod, S. 
nemoralis Ait. I reared moths from an unidentified host in Florida which 
was likely camphor weed, Heterotheca subaxillaris (Lam.) Britt. & 
Rusby, a host noted on museum specimens from Florida and Texas. I 
observed typical galls on Aster ericoides L. in northern Ohio but did not 
succeed in rearing adults. 

I found desertanum only on the grass leaved goldenrod, Solidago 


52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY » 


sraminifolia (.) Salisb. This goldenrod often occurred on the same 
sites as one or more of those above; both scudderianum and desertanum 
sometimes occurred at such sites. Epiblema discretivanum occurred 
mostly on groundsel-tree, Baccharis halimifolia L., but I reared adults 
also from narrow leaved groundsel, B. angustifolia Michx., and B. 
glomeruliflora Pers. 


Geographic Distribution 


Three types of locality records appear in Fig. 4: (1) where I reared 
adults that were subsequently identified, (2) where museum specimens 
that I verified were collected (museums included National Museum of 
Natural History, Canadian National Collection, and American Museum of 
Natural History), and (3) where I observed galls only. The map shows 
gall-only records where there were gaps in the first two types of records. 
Only one map symbol appears where two or more for the same species 
were close enough to overlap. 

Records for scudderianum occur from Maine south to Florida and 
west to North Dakota and Texas. Those for desertanum occur through 
practically the same area while those for discretivanum are confined to 
the coastal plain from Georgia and Florida to Texas (Fig. 4). 


Larval Feeding Pattern 


Between hatching and entering stems, scudderianum larvae evidently 
fed at host tips. This was inferred from several series of observations 
typified by the following example. In mid-July, I examined 25 Solidago 
altissima-canadensis plants with incipient galls in a field in southern 
Michigan. The tip of every plant had been mined by a small insect no 
longer present. In the same field on the same date, I examined another 
25 plants that had mined tips. Of this group, 16 had incipient scud- 
derianum galls; larvae on the remaining nine plants probably did not 
survive to start galls. Limited observations suggest similar pre-gall feed- 
ing by desertanum. I did not observe discretivanum for pre-gall feeding. 

A total of 85 incipient scudderianum galls which I examined in June 
and July in Ohio, Maryland, and Michigan had one and usually two small 
openings between the gall chamber and outside. One Opening was 
gradually enlarged throughout the summer whereas the other usually 
was not. I assume the latter to be the passage by which the larva entered 
the stem. It was often located just above a leaf attachment. The en- 
larged opening served as a hatch through which the larva periodically 
ejected debris, mostly frass. The debris hatch was covered with silk 
when not in use. Some entry passages may have been converted to debris 


VoLUME 30, NUMBER 1 53 


100 200 300 mi 


== ——— = 
161 322 483 km 


Fig. 4. Distribution of records for Epiblema scudderianum (circles), desertanum 
(squares), and discretivanum (triangles). Solid symbols signify reared adults; half 
open symbols, museum specimens; full-open, galls. 


hatches. I observed ejection of debris from galls brought indoors; frass 
intercepted by leaves beneath galls is a common sight in the field. 
Kellicott (1878) also reported ejection of debris by scudderianum. 
Debris hatches were usually located in the lower half of scudderianum 
galls. After larvae became full-grown and ceased to feed and excrete, 
they permanently sealed debris hatches with a dark brown noncellular 
material probably of larval origin. Such plugs were closely fitted and 
when removed looked somewhat like train wheels. 

Debris hatches and plugs occurred in desertanum and discretivanum 
galls but were located in the upper half of galls. No entry passages 


54 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 5. Galls of Epiblema discretivanum on Baccharis halimifolia. Left, sectioned 
to show larval tunnel and moth exit. Right, intact. 


Fig. 6. Upper stem of flowering Solidago altissima-canadensis. Left, normal 
plant. Right, branched plant infested by Epiblema scudderianum. Gall is visible 
near center of the segment of stem shown. 


separate from debris hatches were identified in either species, the entry 
passages probably having been converted to debris hatches. 

Apparently due to host reaction, scudderianum and desertanum galls 
occasionally ruptured, exposing the larval feeding chamber. Counts in 
Maryland in mid-July showed 7 of 25 scudderianum and 9 of 48 desert- 
anum galls in various stages of rupture. Several larvae were continuing 
their feeding in adjacent parts of the same stems. 

Stem tunnels of larvae that pupated ranged in length as follows: 
scudderianum, 3.2-5.0 cm (16 observations); desertanum, 4.5-4.7 cm 
(2); and discretivanum, 2.1-3.3 cm (24). 

Mature larvae excavated moth exit tunnels in the upper half of galls, 
leaving only a thin layer of plant tissue. They spun silken funnels that 
guided the emerging insect into the exit. Moth exits were completed by 
all three species before winter. The gall of discretivanum (Fig. 5) is 
similar in appearance and gross structure to the other two galls; that of 
scudderianum has been illustrated often and is shown together with 
desertanum galls by Miller (1963). 


VoLUME 30, NUMBER 1 55 


Infestation by scudderianum was often accompanied by branching 
of host plants late in the summer (Fig. 6). 


Seasonal History 


Scudderianum flew in May and June in northern localities. At a light 
near an old field in southern Michigan, I caught 30 moths between May 
25 and June 21 during two years of observations. In northern Ohio 
during one year of observation, 6 moths emerged within the above dates 
from galls held in an outdoor insectary. In Maryland during one year of 
observation, 4 moths emerged between April 30 and May 12 from galls 
in an outdoor insectary. In studying galls of Gnorimoschema gallaesoli- 
daginis (Riley) (Miller, 1963), I obtained one moth of Epiblema 
scudderianum during the above periods. 

I made one observation concerning the flight period of desertanum: 
on June 26 in Maryland, 13 galls had protruding empty pupal cases and 
two had live pupae. I made no comparable observations on discretiv- 
anum. 

The earliest dates I observed incipient scudderianum galls were June 
20 in Maryland and June 24 in northern Ohio; desertanum galls, July 
19 in Maryland. All three species overwinter in galls as mature larvae. 

The earliest dates I observed scudderianum pupae were April 11 in 
Maryland, May 7 in northern Ohio, and May 22 in southern Michigan; 
desertanum pupae, May 24 in southern Michigan. 


DiscussION AND CONCLUSIONS 


Type examination and fixation confirms and formalizes identities and 
synonymies of the three species. The synonyms were proposed by Fer- 
nald (1882) as well as Heinrich (1923); it is uncertain whether Fernald 
saw all types but certain that Heinrich did not. Adults of the three 
species are recognizable from forewing pattern despite the variability in 
discretivanum. Larvae of some Epiblema are characterized by MacKay 
(1959) and pupae of two are included in Mosher’s (1916) classification. 
Eggs of Epiblema strenuanum (Wlkr.) E. carolinanum (Wlshm.), and 
E. otiosanum (Clem.) are known (Peterson, 1965; Thompson, 1928; 
Decker, 1932). 

Scudderianum, associated with four genera of hosts, has more known 
hosts than any North American Epiblema. It has often been reported 
from the Canada goldenrod complex. The five host species reported 
here, representing two genera, appear to be new records. Ellis (1925) 
listed Bidens frondosa as a frequent host, and two other genera observed 
once by him as hosts, referring to scudderianum as the bidens borer. One 


56 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


wonders if he confused scudderianum with the true bidens borer, otio- 
sanum, but he stated correctly that his insect was univoltine whereas otio- 
sanum is multivoltine (Decker, 1932). Desertanum has only one known 
host in the North; its host or hosts in the South, where Solidago gramini- 
folia does not occur (Fermald, 1950), are unknown. Thus far, discreti- 
vanum is known only from the three species of Baccharis listed in this 
paper. Heinrich (1921) said it formed a gall on “wild myrtle” which 
Bottimer (1926) identified as B. halimifolia. 

All three Epiblema studied appear confined to eastern North America. 
Distributions of scudderianum and discretivanum and their most frequent 
hosts closely coincide; the same is true for desertanum in the North. 
Discretivanum likely occurs in the West Indies and other Caribbean 
areas because its hosts are there (Small, 1933). 

Riley (1883) stated that scudderianum, in one of several alternate 
feeding patterns, fed first on tips and later formed galls. Kellicott (1882) 
confirmed gall forming but not tip feeding. Riley mixed species and 
even genera of goldenrod-feeding olethreutids in his observations. 
Whether he observed true scudderianum tip feeding is uncertain. Pre- 
gall tip mining by scudderianum and desertanum may represent an early 
feeding mode characteristic of other olethreutid larvae (MacKay, 1963). 
Similar feeding has been reported in otiosanum (Decker, 1932), caro- 
linanum (Thompson, 1928) and tripartitanum (Zell.) (Bottimer, 1926). 

Early debate abcut whether scudderianum induced gall formation 
arose in part by scudderianum moths apparently emerging from Gnori- 
moschema galls (Kellicott, 1882). Judd (1951) as well as I observed the 
same phenomenon. Old galls of these two gall makers can be confused. 
On the other hand, scudderianum larvae whose galls rupture might find 
their way into Gnorimoschema galls just as some otiosanum larvae 
wander to new sites before overwintering (Decker, 1932). 

In contrast to linear stem boring, scudderianum, desertanum and dis- 
cretivanum concentrate their feeding within a short length of stem. 
Maximum stem swelling seems to be associated with the point of maxi- 
mum internal feeding. After scudderianum larvae form their chambers, 
cambial activity is greatly accelerated. Tissues internal to the cambium 
are grazed (Blum, 1953). Ejection of debris is essential where feeding 
is intensified in a small area. Branch proliferation due to scudderianum 
has counterparts in hosts of strenuanum (Crawford, 1933) and caro- 
linanum (Thompson, 1928). 


North American Epiblema with known biologies overwinter as mature 
larvae. Scudderianum and desertanum are apparently univoltine in the 
North. In Ontario over several years, Brodie (1909) observed scudder- 


VoLUME 30, NUMBER 1 BF 


ianum moths flying between June 12 and July 1, two weeks later than I 
observed in southern Michigan. Moth flight, gall formation, and pupa- 
tion appear to be earlier in Maryland than in the Midwest. 

Galls of all three species observed in this study persist for a year or 
longer after their makers leave and many are used as homes by other 
arthropods (Miller, 1966). 


LITERATURE CITED 


Bium, J. L. 1953. Vascular development in three common goldenrod galls. Pap. 
Mich. Acad. Sci. Arts Lett. 38: 23-34. 

Boss, M. L. 1942. Parasites of the oriental fruit moth and of certain weed-infesting 
larvae. Va. Agr. Exp. Sta. Tech. Bull. 79. 23 p. 

Borrmenr, L. J. 1926. Notes on some Lepidoptera from eastern Texas. J. Agr. Res. 
33: 797-819. 

Bropiz, W. 1909. Lepidopterous galls collected in the vicinity of Toronto—No. 2. 
Can. Ent. 41: 73-76. 

Brown, R. S. 1973. Phylogenetic systematics: Its application to the genus Epiblema 
(Lepidoptera). M.S. Thesis, Univ. of Arkansas. 179 p. 

CLEMENS, B. 1860. Contributions to American lepidopterology. No. 6. Proc. Acad. 
Nat. Sci. Philadelphia 1860: 345-362. 

1865. North American micro-lepidoptera. Proc. Ent. Soc. Philadelphia 5: 
133-147. 

Crawrorp, A. W. 1933. Glypta rufiscutellaris Cresson, an ichneumonid larval para- 
site of the oriental fruit moth. New York Agr. Exp. Sta. Tech. Bull. 217. 29 p. 

Decker, G. C. 1932. Biology of the bidens borer, Epiblema otiosana (Clemens ) 
(Lepidoptera, Olethreutidae). J. New York Ent. Soc. 40: 503-509. 

Exuis, W. O. 1925. Some lepidopterous larvae resembling the European corn borer. 
WeeNerokes, o0: 117-192. 

FERNALD, C. H. 1882. A synonymical catalog of the described Tortricidae of North 
America north of Mexico. Trans. Amer. Ent. Soc. 10: 1-64. 

FERNALD, M. L. 1950. Gray’s manual of botany. 8th ed. American Book Co., New 
York. 1632 p. 

Hernricu, C. 1921. Some Lepidoptera likely to be confused with the pink bollworm. 
J. Agr. Res. 20: 807-836. 

. 1923. Revision of the North American moths of the subfamily Eucosminae 
of the family Olethreutidae. U.S. Nat. Mus. Bull. 123. 298 p. 

Jupp, W. W. 1951. Hymenoptera and an inquiline moth reared from the goldenrod 
gall caused by Gnorimoschema gallaesolidaginis Riley (Lepidoptera). Proc. 
Nova Scotia Inst. Sci. 22(4): 1-7. 

Ketuicotr, D. S. 1878. <A new gall moth, and notes on larvae of other gall moths. 
Can. Ent. 10: 201-204. 

. 1882. Is Paedisca scudderiana a gall-maker? Can. Ent. 14: 161-163. 

MacKay, M. R. 1959. Larvae of the North American Olethreutidae (Lepidoptera ). 
Can. Ent. 91, Suppl. 10. 338 p. 

1963. Evolution and adaptation of larval characters in the Tortricidae. 
Gan. Ent: 95: 1321-1344. 

Mitter, W. E. 1963. The goldenrod gall moth Gnorimoschema_ gallaesolidaginis 
(Riley) and its parasites in Ohio. Ohio J. Sci. 63: 65-75. 

1966. Spiders in old insect galls on goldenrod. Ohio J. Sci. 66: 618-619. 

Mirter, W. E. 1973. Clemens types of Olethreutinae (Lepidoptera, Tortricidae ). 
Trans. Amer. Ent. Soc. 99: 205-234. 


58 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Mosuer, E. 1916. A classification of the Lepidoptera based on characters of the 
pupa. Bull. Ill. Lab. Nat. Hist. 12(2): 17-160. 

Peterson, A. 1965. Some eggs of moths among the Olethreutidae and Tortricidae 
(Lepidoptera). Fla. Ent. 48: 1-8. 

Ritey, C. V. 1883. Notes on Paedisca scudderiana. Amer. Nat. 17: 1069-1070. 

SMALL, J. K. 1933. Manual of the southeastern flora. Published by the author, 
New York. 1554 p. 

Tuompson, R. W. 1928. The golden-glow borer (Epiblema carolinana Walsingham ). 
Ann. Rep. Ent. Soc. Ontario, 58, 1927, p. 73-75. 

ZeLLER, P. C. 1876. Beitrage zur Kenntniss der nordamericanischen Nachtfalter, 
besonders der Microlepidopteren. III. Verh. Zool.-bot. Ges. Wien 25: 205-360. 


A POSSIBLE SOURCE OF MORTALITY IN PAPILIO TROILUS 
(PAPILIONIDAE) POPULATIONS IN EASTERN TEXAS 


While collecting Papilio troilus (L.) eggs in the vicinity of the Hardin town dump 
in Hardin Co., Texas, on 21 March 1972, several trails of leaf-cutter worker ants 
were observed carrying pieces of leaves to their nests presumably for culture medium 
for their fungal “gardens” (Creighton, 1950, Bull. Mus. Comp. Zool. Harvard Univ. 
104: 325-329). These ants were kindly determined by Dr. Wm. L. Brown, Jr. 
to be Atta texana (Buckley). One of the small shrubs which was rapidly being 
defoliated of its succulent spring growth of leaves was Sassafras albidum (Nutt.), 
which was also the main plant species being utilized for oviposition by P. troilus. 
While looking at one particular sassafras plant I was able to rescue a troilus egg 
on an excised leaf section that was actually being carted off down the stem by an 
ant and three more eggs on leaves certainly destined for the same outcome. The 
probable fate would have been death, but the proximate source of mortality could 
have been any of several, including desiccation, predation by ants, or fungal attack 
of either eggs or larvae. These eggs and the resulting larvae were instead reared 
through to P. troilus adults on Sassafras albidum in Ithaca, New York. Specimens of 
P. troilus and Atta texana from these Hardin County populations have been deposited 
in The Cornell University Collection: Lot 1023, sublot 13b. 


J. Mark Scriser, Department of Entomology, Cornell University, Ithaca, New 
York 14853. 


VoLUME 30, NUMBER 1 59 


MIGRATION OF BUTTERFLIES ALONG THE GULF COAST OF 
NORTHERN FLORIDA 


F, A. URQUHART AND N. R. URQUHART 


Life Sciences, Scarborough College, University of Toronto, 
West Hill, Ontario, Canada MIC 1A7 


For the past five years (1970-75) we have been making an intensive 
study of the Gulf Coast migrations of the monarch butterfly (Danaus 
plexippus plexippus [L.]) with special reference to the main migration 
route along the coast of northern Florida. Most of our observations and 
alar tagging have been conducted in the vicinity of Apalachee Bay at 
St. Marks Lighthouse, Wakulla Beach, Shell Point, and Live Oak Point. 
Although the migrants pass through this area during the latter part of 
October, the peak period of abundance has been consistently between 
October 20-25. Prior to October 20 and after October 25, relatively few 
migrants were recorded, although stragglers continued to pass through 
the area as late as mid-January. On one or two days during this brief 
period, a peak movement occurs involving countless thousands of indi- 
viduals. The results of this study, as they pertain specifically to the 
monarch butterfly, will be presented at a later date. 

In the present paper we report that along with the monarchs were 
three other species of migrating Lepidoptera, namely, Agraulis vanillae 
(L.), Urbanus proteus (L.) and Phoebis sennae eubule (L.). Each of 
these species occurred in great numbers during the migrating period, 
and, during the free-flight period (non-feeding) they all moved in a 
definite westerly direction, following the coastline. 

The cloudless Sulphur, Phoebis sennae eubule, a well documented 
migratory species ( Williams, 1930, 1958), was observed migrating north- 
ward from the open ocean toward land at Live Oak Point. On reaching 
land, these migrants altered direction abruptly and followed a west 
direction. This band of migrants, extending 200’ across, flew in an end- 
less procession within a few centimeters of the water and, on land, close 
to the vegetation; only an occasional specimen moved toward the flower- 
ing goldenrod to feed. An average of 480 migrants per minute passed 
the observation point. 

The gulf fritillary, Agraulis vanillae, has been reported as a migrant by 
many authors, as listed by Williams (1930, 1958). As this species moved 
along the coast, they fed upon the nectar of Baccharis halimifolia and 
Solidago puberula. An average of 32 specimens was recorded feeding 
upon the nectar of each of five B. halimifolia bushes, a plant that grows 


60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-4. Gulf Coast migrants. 1, Danaus p. plexippus; 2, Agraulis vanillae; 
3, Phoebis sennae eubule; 4, Urbanus proteus. Many specimens collected were travel- 
worn, indicating long flights. 


extensively along the sand dune area of the coast and flowers at the 
time of the peak migration. On being disturbed, these migrants con- 
tinued westward, following the coastline. The reports of flight direc- 
tions, as given by Williams and others for this species, are rather con- 
fusing since, depending upon the geographic location of the observation, 
flights might be recorded in every compass direction. This is due, in 
part, to the southerly flight of migrants, aberrant flights due to topo- 
graphical influences, motion toward flowering plants, and the presence 
of non-migrating nomadic breeding populations. We believe that the 
movement of these migrants is southerly, as mentioned by Harris (1972), 
but, like the monarch butterfly, has a westward direction on reaching the 
coastal area. 

The long-tailed skipper, Urbanus proteus, has been reported by Scud- 
der (1889) as occurring in numbers, but with no record of flight direc- 
tions. Williams (1958) records sight observations of a migration to the 
north in the spring and to the south in the autumn in Florida, but the 
actual locality in Florida where the observations were made is not given. 


VoLuME 30, NuMBER 1 61 


Although general migration patterns for a given geographic area can 
be indicated by careful observations for a given geographic area, it is 
necessary to have accurately recorded observations for many localities 
to establish a definite flight path. The most definitive method is that 
of following a marked individual of a moving population by means of 
individual designations, as shown on the cover of Insektenmigrationem 
(Annual Report for 1974). In this manner release-recapture lines can 
be accurately plotted over long distances. For small species, such as 
Urbanus proteus, one must rely on observations, such as contained in the 
present paper, and reports from other, preferably widely distributed, 
localities. 


LITERATURE CITED 


Harris, L. H. 1972. Butterflies of Georgia. Univ. of Oklahoma Press. 326 p. 

INSEKTENMIGRATIONEM. 1974. Germany (53 Bonn, Adenauerallee 150-164). 

ScuppER, S. H. 1889. Butterflies of eastern U.S.A. and Canada with special refer- 
ence to the New England States. Cambridge, U.S. 

WituraMs, C. B. 1930. The migration of butterflies. Oliver & Boyd, Edinburgh. 
A473 p. 

. 1958. Insect migration. Collins, London. 235 p. 


OCCURRENCE OF LEPTOTES CASSIUS THEONUS (LYCAENIDAE) 
IN GEORGIA 


On 29-30 September 1974, I collected 20 males and 6 females of Leptotes cassius 
theonus (Lucas) on Skidaway Island, Chatham Co., Georgia. This species was not 
mentioned by Harris (1972, Butterflies of Georgia. University of Oklahoma Press ) 
and has apparently not heretofore been recorded from Georgia. All the specimens 
were taken from what appeared to be a well-established colony situated along a short 
stretch of dirt road just northwest of Priest Landing on the Wilmington River side 
of the island. Although it was quite abundant in this particular locality, L. c. theonus 
was nearly overlooked because of its superficial resemblance to Hemiargus ceraunus 
antibubastus Hiibner, which was equally abundant. It appears likely that careful 
exploration of Georgia’s coastal islands will reveal the existence of additional L. c. 
theonus colonies. 


Ricuarp T. Arsocast, 114 Monica Blvd., Savannah, Georgia 31406. 


62 JouRNAL OF THE LEPIDOPTERISTS’ SOCIETY 


NEW HESPERIIDAE RECORDS FOR MEXICO 


Hucu Avery FREEMAN 
1605 Lewis Drive, Garland, Texas 75041 


During the past year I have been making concentrated efforts to com- 
plete a study that I have been making on the Hesperiidae of Mexico. 
Through research conducted in the American Museum of Natural His- 
tory, collecting widely over Mexico and determining specimens for 
various museums and collectors I have found 12 new species and 33 new 
records of hesperiids from Mexico during that period of time. In this 
article I will deal only with the new records and in a separate article 
the new species. 


PYRGINAE 


Tarsoctenus praecia (Hewitson, 1857) 


Type locality. Santarem, Brazil. 
Distribution. Lower Amazon River, Para, Brazil. There is a specimen in the 
American Museum of Natural History labelled, Mexico. 


Drephalys dumeril (Uatreille, 1824) 


Type locality. Uncertain. 

Distribution. The British Museum contains specimens of this species from the 
following locations: Panama, Venezuela, Br. Guiana, Surinam, Fr. Guiana, Ecuador 
(La Chima) and Brazil (Bahia, Maranham, Joinville). I have seven males of this 
species in my collection from Candelaria Loxicha, Oaxaca, Mexico, collected during 
July, August, September and October, 1968-71, all obtained from E. C. Welling. 


Drephalys oria Evans, 1952 


Type locality. Honduras. 

Distribution. There are two males and a female from Honduras in the British 
Museum. There is a male in the American Museum of Natural History from Rancho 
San Carlos, Oaxaca, Mexico, collected March 12, 1969 by Peter Hubbell, and there 
is a male in my collection with the same data. 


Aguna aurunce (Hewitson, 1867) 


Type locality. Amazons. 

Distribution. The British Museum contains specimens of this species from the 
following locations: Guatemala, Nicaragua, Panama, Colombia, Venezuela, Trini- 
dad, Br. Guiana, Fr. Guiana, Peru (Tarapote), Upper Amazons (Teffe, Itaituba, 


Obidos ) and Para, Brazil. I have a male in my collection from Tenosique, Tabasco, 
Mexico, August 24, 1962, collected by E. C. Welling. 


Zestusa staudingeri (Mabille, 1888 ) 
Type locality. Guatemala. 
Distribution. There are no specimens of this species in the British Museum or 


VoLuUME 30, NuMBER | 63 


American Museum of Natural History. I have 21 specimens in my collection from 
Rancho Belen, 20 km south of San Christobal, Chiapas, Mexico, collected during 
March by Robert Wind. 


Ridens philia Evans, 1952 


Type locality. Colombia, Cauca, Juntas: M. de Mathan: end 1897-8. 

Distribution. Colombia (Cauca, R. Dagua). I have 21 males and three females 
in my collection from Sta. Rosa, Comitan, Chiapas and Candelaria Loxicha, Oaxaca, 
Mexico, collected by E. C. Welling and T. Escalante during March, August, Sep- 
tember and October, 1965-69. 


Astraptes fulgor Hayward, 1938 


Type locality. Argentina. 

Distribution. The British Museum has one male with no locality data. I have in 
my collection two males and a female from Rancho San Carlos, Oaxaca, Mexico, 
August 1968. I have examined two males of this species from Campeche, Campeche, 
Mexico, November 15, 1967, collected by H. L. King. 


Astraptes tucuti Williams, 1927 


Type locality. Panama. 

Distribution. Guatemala, Nicaragua, Costa Rica, Colombia, Peru (Chanchamayo, 
Ucayali), Bolivia and Brazil. I have a male from Tenosique, Tabasco, Mexico, 
September 13, 1962, collected by E. C. Welling, and a male from Rancho Dos 
Amatos, Veracruz, Mexico, August 1969, collected by Peter Hubbell. 


Astraptes samson Evans, 1952 


Type locality. Colombia. 

Distribution. Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, 
Br. Guiana, Fr. Guiana, Ecuador (Paramba), Peru (Inambari, Huancabamba), 
Bolivia (Reyes) and Amazons (St. Paulo d’Olivenca to Para). There is a male in 
the American Museum of Natural History from Presidio, Veracruz, Mexico, Sep- 
tember 1940. I have two males in my collection from Muste, Chiapas, Mexico, 
August and September, 1968, collected by E. C. Welling. 


Calliades zeutus (Moschler, 1878) 


Type locality. Colombia. 

Distribution. Panama, Br. Guiana, Upper Amazons (Iquitos) and Bolivia ( Buena- 
vista). There is a male in my collection and one in the collection of E. C. Welling 
from X-Can, Quintana Roo, Mexico, June 25, 1967, collected by E. C. Welling. 


Thorybes albosuffusa Freeman, 1943 


Type locality. Fort Davis, Texas. 

Distribution. This species is found over the Big Bend section of Texas. I have 
a female from 1 mile south of Cedritos, Coahuila, Mexico, June 23, 1957, and there 
is a specimen in the American Museum of Natural History from Colima, Mexico. 


Bungalotis erythus (Cramer, 1775) 


Type locality. Surinam. 
Distribution. Honduras, Nicaragua, Colombia, Fr. Guiana, Br. Guiana and Brazil 


64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(Para, Maranham). I have examined a specimen from E. C. Welling that he 
collected at X-Can, Quintana Roo, Mexico, July 10, 1967. 


Bungalotis astylos (Cramer, 1780) 


Type locality. Surinam. 

Distribution. Honduras, Costa Rica, Panama, Colombia, Venezuela, Trinidad, Br. 
Guiana, Fr. Guiana, Ecuador, Peru (Amazones, Carabaya, Chanchamayo, Rentema 
Falls), Upper Amazons (Ega) and Brazil (Para, Manaos, Matto Grosso, Rio, Petro- 
polis). There are two males in the American Museum of Natural History from Presi- 
dio, Veracruz, Mexico, July 1939 and July 1941 (C. C. Hoffman). 


Dyscophellus ramon Evans, 1952 


Type locality. Bugaba, Panama. 

Distribution. Honduras, Panama and “Peru.” There is a female in the American 
Museum of Natural History from Presidio, Veracruz, Mexico, July 1941 (C. C. 
Hoffman ). 


Nascus broteas (Cramer, 1780) 


Type locality. Surinam. 

Distribution. Panama, Colombia, Venezuela, Trinidad, Fr. Guiana, Ecuador (La 
Chima, Paramba), Peru (Amazons), Upper Amazons (Ega, St. Paulo d’Olivenca), 
Brazil (Para, Espirito Santo) and Paraguay. I have a specimen from Tenosique, 
Tabasco, Mexico, August 24, 1962, collected by E. C. Welling. 


Gindanes brontinus Godman & Salvin, 1895 


Type locality. Nicaragua. 
Distribution. Nicaragua. E. C. Welling collected two males of this species at 
Piste, Yucatan, Mexico, July. I have one specimen and he has the other. 


Pythonides assecla Mabille, 1883 


Type locality. “E. Brazil.” 

Distribution. Honduras, Nicaragua, Colombia, Br. Guiana, Fr. Guiana, Upper 
Amazon, Peru (Chanchamayo, Tarapote). In material received from E. C. Welling 
for determination, there were three females of this species from Mexico, two from 


Tenosique, Tabasco, August, and September 1962 and one from X-Can, Quintana 
Roo, July 2, 1965. 


Ebrietas livius (Mabille, 1897) 
Type locality. Bolivia. 
Distribution. Ecuador (Loja, Rio Numbala), Peru (La Merced, Rio Tabaconas), 


Bolivia (Yungas) and Bahia. There is a specimen in my collection from Tenosique, 
Tabasco, Mexico, September 17, 1962, collected by E. C. Welling. 


Pyrgus oileus orcus (Stoll, 1780) 
Type locality. Surinam. 
Distribution. Common from Costa Rica to South America. There are three males 
of this apparent subspecies from Tapachula, Chiapas, Mexico, August 21, 1949 
(Ik. G. Smyth) in the Los Angeles County Museum. 


VoLUME 30, NUMBER 1] 65 


HESPERINAE 


Synapte puma Evans, 1955 


Type locality. Balboa, Panama. 

Distribution. Panama. I have in my collection 19 males and two females of this 
species from Candelaria Loxicha, Oaxaca, Mexico, collected from March to October, 
1968-69 (E. C. Welling). 


Thargella caura (Plotz, 1882 ) 


Type locality. Surinam. 

Distribution. Nicaragua, Costa Rica, Venezuela, Trinidad, Br. Guiana, Fr. Guiana, 
Bolivia (Santa Cruz), Amazons (Nauta to Para) and Bahia. In specimens received 
for identification from E. C. Welling there was a male of this species from Tenosique, 
Tabasco, Mexico, August 12, 1962, in poor condition, and I have a female from 
Orizaba, Veracruz, Mexico, May 1907 (C. C. Hoffman) which belongs to the 
American Museum of Natural History. 


Lerema lumina ( Herrich-Schaffer, 1869 ) 


Type locality. Uncertain. 

Distribution. Guatamala, Costa Rica, Panama and Colombia. I have three males 
of this species in my collection collected by Robert Wind at San Carlos, Chiapas, 
Mexico, July 2, 1969. 


Onophas columbaria ( Herrich-Schaffer, 1870) 


Type locality. Brazil. 

Distribution. Panama, Colombia, Trinidad, Br. Guiana, Surinam, Fr. Guiana, 
Ecuador (La Chima, Paramba), Bolivia (Buenavista), Amazons (Teffe-Para) and 
Bahia. I examined a female specimen of this species collected by H. L. King at 
Kabah Ruins, Yucatan, Mexico, November 17, 1967. 


Turesis theste Godman, 1901 


Type locality. Costa Rica. 

Distribution. Apparently very rare as the British Museum has only the type, and 
there are no specimens in the American Museum of Natural History. E. C. Welling 
sent me two males of this species for determination. He collected them at Tenosique, 
Tabasco, Mexico, August 17, 1962. 


Vertica grandipuncta (Mabille, 1883) 


Type locality. Para, Brazil. 

Distribution. Br. Guiana, Ecuador (Ambato), Upper Amazons (Pebas, Tonantins, 
Santarem, Tapajos), Brazil (Para). I received two males of this species from 
E. C. Welling for determination. They were collected at Tenosique, Tabasco, Mexico, 
August 24 and 27, 1962. 


Talides alternata Bell, 1941 


Type locality. Santa Catharina. 

Distribution. Guatemala, Costa Rica, Colombia, Venezuela, Surinam, Ecuador 
(Balzapamba), Peru (Chanchamayo, Chiriamayo), Para and Novo Friborgo. I 
have in my collection three males and two females of this species collected at 


66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Candelaria Loxicha, Oaxaca, Mexico, during August and October, 1968-69, by E. C. 
Welling. 


Telles arcalaus (Stoll, 1782 ) 


Type locality. Surinam. 

Distribution. Honduras, Panama, Colombia, Venezuela, Trinidad, Br. Guiana, Fr. 
Guiana, Peru (Rio Colorado) and Amazons (Para, St. Paulo d’Olivenca). There is 
a male specimen of this species in the American Museum of Natural History labelled 
“Ochls north Mexico,” with no other data. 


Carystoides orbius Godman, 1901 


Type locality. Nicaragua. 

Distribution. Nicaragua, Trinidad, Br. Guiana, Fr. Guiana, Surinam, Ecuador 
(Canalos, Rio Bobonaza), Upper Amazons and Para. In specimens received from 
E. C. Welling there was a male of this species collected at Tenosique, Tabasco, 
Mexico, September 4, 1962. 


Mellana balsa freemani Steinhauser, 1974 


Type locality. Santa Tecla, El Salvador. 

Distribution. Santa Tecla, El Salvador, June, July and October. Sayaaxche, El 
Peten, Guatemala, August 10, 1963 (E. C. Welling). I have in my collection four 
males from Candelaria Loxicha, Oaxaca, Mexico, July and August, 1969 and 1971 
(E. C. Welling ). 


Mellana tecla Steinhauser, 1974 


Type locality. Santa Tecla, E] Salvador. 

Distribution. Santa Tecla, Tamanique, E] Salvador. I have in my collection 18 
males of this species from Muste, Chiapas, Mexico, July 1968 (E. C. Welling) and 
five males from Candelaria Loxicha, Oaxaca, Mexico, July and August, 1969 and 
1971 (E. C. Welling). 


Panoquina panoquinoides (Skinner, 1891) 


Type locality. Florida. 
Distribution. Florida, Texas, Cayman Island, Tobago and Pernambuco. I have 


a male and a female of this species in my collection from Progreso, Yucatan, Mexico, 
October 6, 1971 (E. C. Welling). 


Xeniades pteras Godman, 1900 
Type locality. Panama. 
Distribution. Costa Rica, Panama, Colombia (St. Marta) and Venezuela. I have 


a male and female of this species in my collection from Muste, Chiapas, Mexico, 
collected October 22, 1968, by E. C. Welling. 


Saliana hewitsoni Riley, 1926 


Type locality. Upper Amazons. 

Distribution. Panama, Colombia, Ecuador (Paramba, La Chima, Zamora), Peru 
(Moyabamba), Bolivia (Reyes, Buenavista) and Amazons (Nauta, Para). Dr. 
W. W. McGuire brought a female specimen of this species here for me to determine 


VoLUME 30, NuMBER 1] 67 


which he had received from Peter Hubbell. The specimen was collected at Cate- 
maco, Veracruz, Mexico, November 15, 1973 by Sr. Abraham Ramirez. 


ACKNOWLEDGMENT 


This research was made possible by a grant from the National Geo- 
graphic Society. 


68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


GENERAL NOTES 


RANGE EXTENSIONS OF 
CALLOPHRYS FOTIS, C. POLIOS, COLIAS ALEXANDRA AND 
EREBIA CALLIAS (LYCAENIDAE, PIERIDAE AND SATYRIDAE) 


Until 1971, Callophrys (Incisalia) fotis schryveri (Cross) was known only from 
Colorado and southern Wyoming (Ferris & Stanford, J. Lepid. Soc. 24: 256-266). 
In 1971 and 1972, it was collected in numbers by S. Kohler in the vicinity of 
Missoula, Missoula Co., Montana. Voucher specimens are deposited in the collection 
of S. Kohler and the author. 

Ferris & Fisher (1973, J. Lepid. Soc. 27: 112-118) discussed the distribution of 
Callophrys (Incisalia) polios (Cook & Watson) in North America. At that time, 
polios was unknown from the Great Plains region. It was recorded from eastern 
Minnesota and Wisconsin, and northeastern Illinois. Eastern limits for the subspecies 
obscurus Ferris & Fisher included Missoula Co., Montana; the Black Hills, South 
Dakota; Albany Co., Wyoming; and south along the Colorado Front Range. In 
May 1973, T. McCabe collected C. polios obscurus in the Killdeer Mts., Dunn Co., 
North Dakota, a locality several hundred miles north and east of the nearest known 
locality of this subspecies. Voucher specimens are deposited in the collection of 
T. McCabe and the author. 

In his revision of the Colias alexandra complex, Ferris (1973, J. Lepid. Soc. 27: 
57-73) listed no records south of the United States-Mexico border. Colias alexandra 
harfordii H. Edwards has now been collected at several localities in Baja California 
(state) in the Sierra San Pedro Martir (Holland, 1972, J. Res. Lepid. 11: 147-160; 
H. Real, in litt.). 

Since my paper on Erebia callias Edwards appeared (Ferris, 1974, J. Lepid. Soc. 
28: 230-236), a number of new Colorado records for this insect have been sent to 
me by R. E. Stanford and D. Cowper from the following counties: Boulder, Gilpin, 
Gunnison, Ouray, Pitkin, Rio Grande and San Juan. 


ACKNOWLEDGMENTS 


I wish to thank the following for providing distribution records: D. Cowper, 
J. D. Eff, R. Holland, S. Kohler, T. L. McCabe, H. G. Real and R. E. Stanford. 

This note is published with the approval of the Director, Wyoming Agricultural 
Experiment Station, as Journal Article JA 722. 


Cuirrorp D. Ferris, College of Engineering, University of Wyoming, Laramie, 
Wyoming 82071. (Research Associate, Allyn Museum of Entomology, Sarasota, 
Florida; Museum Associate, Los Angeles County Museum of Natural History, Los 
Angeles, California. ) 


AMBLYSCIRTES BELLI (HESPERIIDAE): A NEW RECORD FOR KENTUCKY 


On July 26, 1969, between 1500 and 1600 I collected a single Amblyscirtes belli 
lreeman (@) that was resting on a leaf in a wooded area of Chickasaw Park in 
Louisville, Jefferson Co., Kentucky. Determination was tentatively made by C. V. 
Covell, Jr. of the University of Louisville and confirmed by Mr. H. A. Freeman of 
Garland, Texas. This species has not been reported for Kentucky (Covell, 1974, 
J. Lepid. Soc. 28: 253-256), the nearest known record being in Pope Co., Illinois 
(Irwin & Downey, 1973, Ill. Nat. Hist. Surv. Biol. Notes No. 81: 1-60). 


Ricard A, Henperson, 2609 Welsford Way, Louisville, Kentucky 40222. 


VoLUME 30, NUMBER 1 69 


FACTORS AFFECTING THE OCCURRENCE OF 
MELANIS PIXE (RIODINIDAE) IN EXTREME SOUTHERN TEXAS 


The lower Rio Grande Valley of Texas has a varied lepidopteran fauna including 
many neotropical forms that reach their northernmost points in this area. One of 
the least known of these is Melanis (= Lemnax = Lymnas) pixe pixe (Boisduval), 
Riodinidae. This medium-size butterfly (wingspan about 3.5 cm) has a velvet-black 
ground color with an apical yellow spot on each forewing. A single basal spot on 
each wing and six marginal spots on the hindwings are an intense scarlet color. 

This species is neither illustrated nor mentioned in any of the standard American 
butterfly guides, i.e., Holland, Klots, Ehrlich & Ehrlich, or dos Passos. An illus- 
tration of M. gynaeceas Godm. & Salv., which may be an extreme version of M. 
pixe, may be found in Biologia-Centrali-Americana (Godman & Salvin, 1882:  v. 
38, pl. 110, fig. 3). Despite previous collections of this M. pixe in the Brownsville 
(Cameron Co., Texas) area (summarized below), the present report is believed to be 
the first published note on U.S. records of this species. The northernmost published 
record of this butterfly that I could find was in the state of Veracruz (Hoffman, 
1940, Am. Inst. Biol. Mex. 11(2): 639~-739), at least 420 km south of Brownsville. 
L. E. Gilbert (pers. comm.) has observed pixe at Cd. Mante, Tamaulipas, 380 km 
south of Brownsville. 

In June, July and August of 1961, I observed this species in Brownsville. Larvae 
were found and reared to imago feeding on an introduced ornamental, guamuchil, 
Pithecellobium dulce (Roxb.) Benth. This leguminous shrub is native to Mexico 
and through Central America to Colombia and Venezuela (Little & Wadsworth, 1964, 
Common Trees of Puerto Rico and the Virgin Islands, Agric. Handbook 249, 
U.S.D.A.). Adult M. pixe are very poor fliers and are often seen hovering around 
the host plant. Their flight pattern is suggestive of many of the warning-colored 
day-flying moths. 

Melanis pixe was first reported from the U.S. on 21 March 1956 by O. O. Stout 
in Brownsville. (This and the following records are from the files of R. O. Kendall.) 
Field collections of M. pixe occurred from 1957-1961; none were subsequently 
reported until the late 1960’s, when C. A. Kendall sighted one. All U.S. records to 
date are from Brownsville. I saw no further specimens after the superabundance in 
1961. 

Possibly, the absence of M. pixe in 1962 and subsequent years resulted from the 
effects of the catastrophic freeze of 9-12 January 1962. Brownsville reported a 
low temperature of 19°F (12 January), the coldest since 1899. Massive damage to 
citrus groves and substantial damage to native vegetation resulted. This freeze 
undoubtedly was the most destructive to native wildlife and vegetation recorded 
(Heitzman, 1962, J. Lepid. Soc. 16: 249-150; James, 1963, Southwest. Nat. 8: 
45—46.). 

The first reported occurrence of M. pixe in the Brownsville area corresponded 
with one of the warmest periods on record in the same area. No severe freezes were 
recorded from 1952-1961 (lowest temperature 29°F); no freezing temperatures were 
recorded from 1954-1958, inclusive. The warm vears of the mid-1960’s again per- 
mitted the occurrence of M. pixe in the lower Rio Grande Valley. 

Many butterflies of northern Mexico are found in southern Texas, but only under 
certain environmental conditions (Gilbert, 1969, J. Lepid. Soc. 23: 177-185). Popu- 
lations of M. pixe may be completely eliminated by these severe freezes and must 
disperse from Mexico; on the other hand, populations may simply be so low in 
numbers and local in nature that detection is unlikely. 

A puzzle yet to be solved is determination of the native larval foodplant of 
M. pixe. Adults flying around Pithecellobium dulce showed no attraction behavior 
toward Texas ebony, P. flexicaule (Benth.) Coult., a common tree in southern 
Texas; M. pixe larvae have never been found on this species. A specimen of Texas 


70 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


ebony was less than 10 m from the smaller P. dulce that supported M. pixe larvae. 
Pithecellobium dulce has been reported common around Cd. Victoria, Tamaulipas 
(Robert Runyon, botanical voucher sheet 777, Univ. Texas at Austin Herbarium), 
320 km south of Brownsville. There possibly has never been a native foodplant for 
M. pixe in the Brownsville area. Thus, M. pixe may have occurred in southern Texas 
only since the introduction of guamuchil. 


ACKNOWLEDGMENT 


I wish to thank R. O. Kendall for information on previous sightings of this species. 


RaymMonp W. Neck, Texas Parks and Wildlife Department, John H. Reagan Build- 
ing, Austin, Texas 78701. 


NEW FOODPLANT AND OVIPOSITION RECORDS FOR BATTUS PHILENOR 
(PAPILIONIDAE) 


Battus philenor (1..) feeds on several species in the plant family Aristolochiaceae. 
In the central and southern Appalachian regions of the eastern U.S.A., Aristolochia 
serpentaria L, is a predominant native foodplant species (Scudder, 1889, The butter- 
flies of the eastern U.S. and Canada, 2: 1219-1364; Holland, 1898, The butterfly 
book, Doubleday, Doran & Co. New York; Forbes, 1960, Comell Univ. Agr. Expt. 
Sta. Memoir #371). The northern range limits of Battus philenor probably are 
extended by feeding upon the introduced ornamental Aristolochia sipho L’Her or 
Asarum spp. In Texas, Kendall (1964, J. Lepid. Soc. 18: 129-157, and pers. comm. ) 
reports only Aristolochia longiflora (Engelm. & Gray). Here we report feeding and 
oviposition by B. philesor on Aristolochia serpentaria and A. reticulata (Nutt.) in 
eastern Texas. These observations apparently represent new foodplant records for 
the state of Texas and the U.S.A., respectively. 

During a collecting trip to the “Big Thicket” region of eastern Texas in late 
March, 1972, eggs and larvae of first through third instars of the pipevine swallow- 
tail, Battus philenor, were found upon the small perennial Aristolochia reticulata in 
a longleaf pine forest near Camp Waluta, approximately 6 mi. NW of Silsbee between 
routes 92 and 69 in Hardin County. In addition, several females were seen ovi- 
positing on these plants. Another trip during the following spring to the same area 
yielded similar observations. From 8 April 1973 through 13 April 1973, eggs, larvae 
of various instars and ovipositing females were abundant upon the A. reticulata 
(Fig. 1). Eggs were laid most frequently in groups of two, three or four per plant, 
though the number laid by an individual female on any one occasion ranged from 
one to seven per plant. 

While following one particular female engaged in the characteristic ‘ovipositional 
searching’ flight between 12:25 and 13:00 hours on 12 April 1973, oviposition was 
observed upon Aristolochia serpentaria. Although A. reticulata plants were more 
abundant, none were selected by this female for oviposition, or even approached. 
Three A. serpentaria plants were supplied with one, three and two eggs respectively. 

Although A. serpentaria is not apparently a widespread hostplant for Battus 
philenor in Texas, it is more common in other states to the northeast. Aristolochia 
reticulata, however, has not to our knowledge ever been reported as a foodplant 
of B. philenor. The explanation for the intensive use of A. reticulata in the Waluta 
site and the absence of records elsewhere probably stems from the fact that A. 
reticulata has a relatively restricted range. It is found in the humus of sandy soils 
of pine-hardwoods or pine savannahs only in eastern Texas, southwestern Arkansas 


VoLUME 30, NuMBER 1 ck 


Fig. 1. Battus philenor female ovipositing upon Aristolochia reticulata 12 April 
1973 in a long leaf pine savannah near Silsbee, Hardin County, Texas. 


and northwestern Louisiana (Pfeifer, 1966, Ann. Mo. Bot. Gdn. 53: 115-196; 
Correll & Johnston, 1970, Manual of the vascular plants of Texas, Texas Research 
Foundation). 

Specimens of Aristolochia reticulata have been deposited in the Bailey Hortorium 
at Cornell University. Battus philenor individuals which were found and reared 
upon the A. reticulata have been deposited in the Cornell Collection (Lot 1023; 
sublot 18). 


ACKNOWLEDGMENTS 


Thanks are extended to Geraldine Watson for identification of the two plant 
species, but most especially for her enthusiastic and valuable guidance in locating 
the many beautiful and priceless portions of the Big Thicket region of eastern 
Texas which were appropriate for our studies. The travel expenses were borne by 


N. S. F. Grant # GB 33398. 


J. Mark ScripeR AND Pau P. FEENy. Department of Entomology, Cornell Uni- 
versity, Ithaca, New York 14853. 


—~l 
bo 


OURNAL OF THE 3 E 
LEPIDOPTERISTS SOCIETY 


BOOK REVIEW 


THE SWALLOWTAIL BUTTERFLIES OF NortH AMmeEricA, by Hamilton A. Tyler. 1975. 
Naturegraph Publishers, Healdsburg, Ca. vii + 192 p., illus. + 16 plates. Price 
$5.95 (U.S.) paperback, $9.95 cloth. 

This book is a pleasant blending of the scientific and popular knowledge of the 
North American swallowtail butterflies. The geographic region encompassed extends 
from the arctic into the North American faunal region of Mexico and to the southern 
political boundary of that country. The author has chosen to include neotropical 
species as well as an aid to visitors to Mexico. The format is of a convenient, almost 
field guide size. There is a wealth of information presented in a rather informal 
manner. The usual taxonomic treatment appears with keys, wing venation diagrams, 
etc., but also included are biogeographical data, evolutionary speculations, mimicry, 
hilltopping, hybridization experiments and other interesting items. There is extensive 
information on distribution, food plants and immatures. A nice feature, not seen 
in many scientific books, is a discussion of the origin of the scientific names used to 
describe the members of the supergenus Papilio. 

The author divides the swallowtails into the four genera: Papilio, Eurytides, 
Parides and Battus. These are then subdivided into species groups within each 
genus forming the basis for the book chapters. The genera Parnassius and Baronia 
are given only brief treatment as they are not true swallowtails. For each species, 
the various subspecies and forms are discussed, and information is provided such 
that similar appearing species may be differentiated. Data are then presented for 
habitat, flight period, food plants, early stages and distribution. 

Interspersed throughout the text are line drawings of ova, larva and pupa. There 
are two sets of color plates which are composed of water color renditions of the 
principal species and many of the subspecies discussed in the text. The plates are 
certainly adequate and many present rather natural portrayals of the imagines. A 
“color errata” noting color registration shifts to be corrected in subsequent printings 
is included. There are three indices: a general index, an index of swallowtail names 
and an index of larval food plants. A “selected” bibliography appears at the end 
of the book which, while perhaps selected, presents in four pages a reasonably 
comprehensive treatment. 

The informal style may aggrevate some workers, but I found it quite refreshing. 
The book is certainly current and cites ongoing and incompleted taxonomic and 
breeding studies. Generally it appeared free from typographical errors. The only 
criticism that I would offer is the lack of inclusion of authority names when the 
scientific names of plants are listed. A few food plant omissions were noted, but 
these are of minor consequence. It would have been helpful if the undersides of 
all of the members of the Old World swallowtail group (machaon-polyxenes-bairdii, 
etc.) had been figured, as in many cases, the ventral surfaces contain the necessary 
diagnostic characters for reliable species separation. 

This book is a useful addition to the library and should appeal to the amateur 
and serious collector alike as an informative and ready reference. 


Currorp D. Ferris, Bioengineering Program, University of Wyoming, Laramie, 
Wyoming 82071. 


EDITORIAL STAFF OF THE JOURNAL 
GeorceE L. Goprrey, Editor 


Illinois Natural History Survey, Natural Resources Building 
Urbana, Illinois 61801 U.S.A. 


Witu1AM H. ALxen, Associate Editor James G. STERNBURG, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study 
of Lepidoptera. Contributors should prepare manuscripts according to the following 
instructions. 

Text: Manuscripts should be submitted in duplicate, and must be typewritten, 
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11 
inch paper. Titles should be explicit and descriptive of the article’s: content, including 
the family name of the subject, but must be kept as short as possible. The first men- 
tion of a plant or animal in the text should include the full scientific name, with 
authors of zoological names. Insect measurements should be given in metric units; 
times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). 
Underline only where italics are intended. References to footnotes should be num- 
bered consecutively, and the footnotes typed on a separate sheet. 

Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically 
under the heading LireRaTurE Crrep, in the following format: 


SHepparD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, 
London. 209 p. 

196la. Some contributions to population genetics resulting from the 
study of the Lepidoptera. Adv. Genet. 10: 165-216. 


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

Illustrations: All photographs and drawings should be mounted on stiff, white 
backing, arranged in the desired format, allowing (with particular regard to lettering ) 
for reduction to their final width (usually 4% inches). Illustrations larger than 81% 
x 11 inches are not acceptable and should be reduced photographically to that size 
or smaller. The author’s name, figure numbers as cited in the text, and an indication 
of the aricle’s title should be printed on the back of each mounted plate. Figures, 
both line drawings and halftones (photographs), should be numbered consecutively 
in Arabic numerals. The term “plate” should not be employed. Figure legends must 
be typewritten, double-spaced, on a separate sheet (not attached to the illustrations ), 
headed ExpLANATION OF FicurEs, with a separate paragraph devoted to each page of 
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for tables should not be capitalized. Tabular material should be kept to a minimum 
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Proofs: The edited manuscript and galley proofs will be mailed to the author for 
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Page Charges: Authors with grant or institutional funds are requested to pay a 
charge of $27.50 per printed page (including tabular and black-and-white illustrative 
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case of authors who have no grant or institutional funding, as it is not intended that 
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be requested to pay this charge for material in excess of 20 printed pages. 

Address all correspondence relating to the Journal to the editor. Material not 
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the editor of the News: Ron Leuschner, 1900 John Street, Manhattan Beach, 
California 90266. 


ALLEN PRESS, INC. eg rso LAWRENCE, KANSAS 
USA 


CONTENTS 


PRESIDENTAL AppRESS 1975—To My FrLttow Amateurs. André 
Blanchard) 003 BO ONO SRS Seen 


New HEsPeRUDAE RECORDS FOR TEXAS AND THE UNITED STATES. 
William W. McGuire and Michael A. Rickard 


A New SupspEcies OF ARGYREUS HYPERBIUS (NYMPHALIDAE) FROM 
New Guinea. Chris Samson 


CALLOPHRYS ERYPHON ( LYCAENIDAE) IN Marne. Warren J. Kiel 


Nores OF MARYLAND LEPIDOPTERA. 5. A NEw SuBSPECIES OF POANES 
MASSASOIT (HeEspERuDAE). William A. Andersen and Robert 
S. Simmons 


NOTES ON THE LIFE CYCLE AND NATURAL History OF BUTTERFLIES 
OF Ex SALvapor. VII. ARCHAEOPREPONA DEMOPHON CENTRALIS 
(NymMpHALDAE). Alberto Muyshondt 


New CATOCALA oF NortH AMERICA (Noctumae). A. E. Brower __ 


A CHECKLIST OF THE BUTTERFLIES OF GRANT County, NEw Mexico 
AND Vicinity. Clifford D. Ferris 


BioLoGy AND TAXONOMY OF THREE GALL FORMING SPECIES OF 
EPIBLEMA (OLETHREUTIDAE). William E. Miller 


MIGRATION OF BUTTERFLIES ALONG THE GULF COAST OF NORTHERN 
Fioriwa. F. A. Urquhart and N. R. Urquhart 


New HEspreruparE Recorps ror Mexico. Hugh Avery Freeman 


GENERAL NOTES 


Capture of a hybrid Limenitis arthemis astyanax < L. archippus (Nym- 
phalidae) in southern Georgia. Richard T. Arbogast _._________ 


Hapalia nigristriatalis a synonym of Udea angustalis (Pyralidae: Pyraus- 
tinae).: Eugene Maaro@: i010 


Mathildana newmanella (Oecophoridae ) in Arkansas. John B. Heppner __. 


A possible source of mortality in Papilio troilus (Papilionidae) populations 
in eastern Texas,’ J. Mark Seriber02 0 ee 


Occurrence of Leptotes cassius theonus (Lycaenidae) in Georgia. Richard 
Ts APOC dsG NN SA UU 


Range extensions of Callophrys fotis, C. polios, Colias alexandra and Erebia 
callias (Lycaenidae, Pieridae and Satyridae). Clifford D. Ferris __ 


Amblyscirtes belli (Hesperiidae): A new record for Kentucky. Richard 
Aly SHR TSOR OL NI i Ce UNO ML 


Factors affecting the occurrence of Melanis pixe (Riodinidae) in extreme 
southern Texas, Raymond W. Neck 0k 


New foodplant and oviposition records for Battus philenor ( Papilionidae). 
J: Mark Scriber and Paul P. Péeny 0 ee 


BOOwK Pee ee a een ea 1 CI OL A 


12 


19 


i 


Volume 30 1976 Number 2 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


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


¢. hodges 


12 July 1976 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


RonaLp W. Honcss, President 


S. S. Nicoxay, President-elect Ouar H. H. Mic.kz, Vice President 
Donatp R. Davis, Ist Vice President Joun M. SnwwER, Treasurer 
WiLL1AM Hovanitz, Vice President Ler D. MILLER, Secretary 


Members at large (three year terms): 


D. C. Fercuson (1975) J. T. BREwer (1976) C. V. CovELL, Jr. (1976) 
R. O. KENDALL (1975) K. S. Brown (1976) D. F. Harpwick (1977) 
J. A. Powrett (1975) K. W. Purp (1976) J. B. Zrecier - (1977) 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and 
formally constituted in December, 1950, is “to promote the science of lepidopterology 
in all its branches, . . . . to issue a periodical and other publications on Lepidoptera, 
to facilitate the exchange of specimens and ideas by both the professional worker and 
the amateur in the field; to secure cooperation in all measures” directed towards 
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Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA 


by Cyrit F. pos Passos 


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JouRNAL OF 
Tue LeprIpopreRIstTs’ SOCIETY 


Volume 30 1976 Number 2 


A STUDY OF THE PENINSULAR FLORIDA POPULATIONS 
OF THE MONARCH BUTTERFLY 
(DANAUS P. PLEXIPPUS; DANAIDAE) 


F. A. URQUHART AND N. R. URQUHART 
Scarborough College, University of Toronto, West Hill, Ontario, Canada M1C 1A7 


A number of contradictory reports have been published concerning 
the presence or absence of the monarch butterfly (Danaus p. plexippus 
(L.) ) in the peninsular part of Florida (hereafter referred to as Peninsular 
Florida and being that portion of the State between Latitude 30°N-24°N) 
throughout the year. Some published reports indicate that the monarch 
butterfly is absent during the summer months while others report the 
presence of this species throughout the year (Urquhart, 1960). Brower 
(1961) reported breeding populations in January and throughout the 
spring and summer months (March through August). 

To clarify this situation, since it forms an integral part of our complete 
investigation of the movements of this species throughout North America, 
a continuous study of the populations in Peninsular Florida was com- 
menced in 1968 and carried out through 1973. The results of this six 
year study program are here presented. 


PROCEDURE 


With the active cooperation of our research associates (Urquhart, 
1960) living in Florida, together with a number of field expeditions, a 
program of field observations, collecting of larvae, tagging of imagoes 
and recording of larval stages of development was set up in the following 
areas (Fig. 1): Area I (Orlando); Area II (Temple Terrace, Lithia, 
Palmetto, Bradenton, Sarasota); Area III (Fort Lauderdale, North Miami 
Beach, Homestead ). Observations were made on the presence of imagoes 
and species of milkweed (Asclepias) at Area IV (Key Vaca and adjacent 


74 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Research areas: (I (Orlando); II (Temple Terrace, Lithia, Palmetto, 
Bradenton, Sarasota); III (Fort Lauderdale, North Miami Beach, Homestead); IV 
(Key Vaca); BR (area studied by Brower at Highland Hammock State Park, Lake 
Childs, Old Venus, 22 mi. SW Clewiston ). 


Keys). Larvae collected were reared (outdoors) to the imago stage and 
alar tagged (Urquhart, 1960) and released. 

A specimen alar tagged and liberated at a particular time and later 
recaptured at some other place gave information as to the direction flown, 
distance travelled and the time interval involved. In addition, the total 
number of specimens tagged was recorded as well as the total of recap- 
tures, in order to ascertain whether or not, in terms of percentage of 
recaptures, there was any significant movement within a given population. 


VOLUME 30, NUMBER 2 15 


RESULTS 


Flight of Imagoes 


During the autumnal migration, the majority of females in a moving 
population are in a state of ovarian dormancy (Urquhart, 1960). A small 
percentage of females, however, mate prior to moving southward or, 
in some cases, during the movement, and are capable of laying fertilized 
eggs on the milkweed plants of various species along the route of migra- 
tion ( Urquhart, 1960). It has been found that if these ovipositing females 
are kept under laboratory conditions, they will continue to lay their full 
complement of eggs. The female offspring, if maintained in the labora- 
tory, do not enter a state of ovarian dormancy and hence will mate and 
lay viable eggs. Thus, a population of monarchs may be maintained in 
the laboratory throughout the year ( Urquhart & Stegner, 1966). If, on 
the other hand, a female that is in the state of ovarian dormancy is held 
in the laboratory under ambient conditions, ovipositing does not take 
place nor do eggs develop in the ovaries, even after a period of five 
months. However, females in ovarian dormancy that were collected from 
the overwintering sites in California (Monterey) in January and held 
under ambient laboratory conditions developed eggs in the last week of 
February and laid viable eggs in March (Urquhart, 1960). As yet no 
significant research has been carried out as to the factors involved in 
initiating or depressing ovarian dormancy, although light period is sus- 
pected. Thus, a migrating population of monarch butterflies includes 
some gravid females and some in ovarian dormancy. In a particular area, 
such as Florida, one will encounter a population in which some of the 
individuals are part of a migrating population and capable of laying 
viable eggs; others may be in a state of ovarian dormancy and will con- 
tinue their migration southward without ovipositing; while still others 
may be the offspring of ovipositing migrants, thus giving rise to what 
might be considered a “resident population.” 

As a result of the alar tagging program, which has been carried out in 
North America over the past 24 years, the release-recapture records indi- 
cate that a few migrants from eastern Canada and the eastern United 
States travel through Peninsular Florida (Fig. 2). One alar tagged 
migrant released in Grafton, Ontario, Canada was recaptured in Miami, 
Florida having flown south, a distance of 1223 mi. Another migrant alar 
tagged in Binghamton, New York was recaptured at Miami, Florida 
having flown south southwest, a distance of 1114 mi. A third migrant 
tagged at Port Hope, Ontario, Canada was recaptured in Havana, Cuba 
having flown south a distance of 1403 mi. A specimen tagged at Anna 
Maria, Florida in research Area II was recaptured in Miami, having flown 


76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


— =_— 


| 
| e 
ee 
| 
| | 
| i] 
| 
| vy | | 
he 
r | 
32 See ee | I 
| | 
\ | 
aS C) \| 
SK \| 
a | 
<< | 
“Sd 
~ 


SQ 


¢- 


| 

| 

| 

| 

| 

| 

| . 
1 oe a 


Fig. 2. Long distance migration records (release-recapture lines): (A) Miller 
Key to Kingsville, Texas; (B) Port Hope, Ontario, Canada to Havana, Cuba; (C) 


Anna Maria to Miami; (D) Grafton, Ontario, Canada to Miami; (E.) Binghamton, 
New York to Miami. 


southeast a distance of 195 mi. A migrant alar tagged at Miller Key in 
Area II was recaptured in Kingsville, Texas having flown a distance west- 
ward of 860 mi. All of these distances are measured as straight lines 
between the point of release and that of recapture. The actual route 
taken is, of course, not known. 

Although two long flight records are reported for specimens alar tagged 
in Area II, most of the recaptures were for shorter distances and flights 
were in various directions (Fig. 3; Table 1). 


Of a total of 4146 specimens tagged in Peninsular Florida, only 39 


VoLuME 30, NuMBER 2 77 


Fig. 3. Release-recapture lines for specimens tagged in Area II showing move- 
ment away from the area at various times of the year and the number of tagged 
specimens that were recaptured close to the site of tagging—indicated by a circle 
with enclosed number. 


78 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Taste 1. Tagging results showing distances travelled, direction of flight and 
number of recaptures in each case. 


Distance 


Travelled Flight Number of 
Tagged at (Km ) Direction Recaptures 
Bradenton 0 0 22, 
Bradenton 28 WNW 1 
Bradenton 13 SSE ) 
Sarasota 16 N 1 
Sarasota 0 0 2 
Myakka State Park 0 1) 1 
Ft. Lauderdale 0 0 1 
Coquina Beach 228 SE 1 
Mullett Key 1003 W 1 
Anna Maria 24 NNE 1 
Palmetto 0 0) 1 
Boca Grande Dall NNW IL 


were recaptured (.94%) even though the tagged specimens were released 
in well populated areas where the chances of recapture should have been 
much greater, as has been the case in other parts of North America dur- 
ing the migrating period (Urquhart, 1960). 


Presence of Larvae 


Brower (1961) recorded breeding populations of the monarch butter- 
fly in the area marked BR (Fig. 1). These observations were made at: 
Highland Hammock State Park; east side of Lake Childs; Old Venus; 
and 22 mi. SW of Clewiston, given as the junction of highways 846 and 
833. The observations were made during the months of March (25-31), 
April (2-30), June (15-30), July (7-31), and August (1-4). Brower 
also reported a breeding populatioin in the same area in January. 

During the years 1958-1973 inclusive, larvae were collected in the 
field and reared outdoors, the resulting imagoes alar tagged and released. 
The number of larvae collected each day was recorded and summated 
for a particular month. The number of larvae recorded varied for a 
particular month from one year to another. This was due to the variable 
activity of the field assistants, the size of the population in any one year 
and the restricted oviposition of a particular female or females with 
respect to the number of milkweed plants present. Thus, although a total 
of 116 larvae were collected in June of 1969, fewer were collected in 
June of 1970 and 1971. Hence, the data presented in Table 2 and the 
summation with imagoes in the histogram (Fig. 4) only indicate trends 
and not accurate field population densities for each year. 


VoLUME 30, NuMBER 2 79 


760} 


680 


510 


360 


NUMBER OF SPECIMENS COLLECTED 
Py 
fe 
i—) 


280 
200 
120 


40 


MONTHS 


Fig. 4. Number of specimens collected during the entire research period for each 
month. Broken line in January indicating what was possibly the true population 
figure as explained in the text. Two peaks of abundance are noted. 


Foodplant 


The presence of a breeding population of monarch butterflies depends 
upon the presence of various species of the genus Asclepias, upon which 
the larvae feed. 

Of the 21 species of Asclepias listed for Florida (Woodson, 1954), 15 


occur in Peninsular Florida, the remainder being reported for the north- 


TaBLE 2. Number of specimens (larvae and imagoes) collected per month for all 
years (1968-1973). 


Jan. Feb. March April May June July Aug. Sept. Oct. Nov. Dec. 


220 16 17 48 202 122 58 169 198 158 59 92 
(208) (88) (78) (177) (269) (155) (112) (87) (142) (358) (706) (118) 
Total 428 104 95 225 471 277 170 256 340 516 765 210 


80 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


ern portion of Florida here referred to as the Panhandle area. Brower 
(1961) recorded larvae feeding upon 4 species of Asclepias, namely: 
tuberosa rolfsii, humistrata, tomentosa and curtisii. Of these the most 
abundant were tuberosa rolfsii and tomentosa, the others recorded as 
uncommon. In addition to Brower’s records, verticillata, which occurs in 
rather dry, wooded areas, and lanceolata, found in wet, marshy areas, 
are of common occurrence throughout Peninsular Florida. The remain- 
ing species occur as widely scattered plants and are recorded from only 
a few localities. 

Unlike some species of Asclepias, such as A. syriaca, which occurs in 
dense growths, the plants of species found in Peninsular Florida tend to 
be widely separated from one another. There are some species, such as 
humistrata and tuberosa, found as flowering plants in home gardens, and 
much of the data recorded in this paper are based on larvae collected 
from such plants. 

Whether or not the larvae of the monarch butterfly will feed on the 
foliage of all species of Asclepias is not known. Brower (1961) has 
recorded 4 species upon which larvae were found feeding. Urquhart 
(1960) has recorded 10 species of which 4 are found in Peninsular 
Florida (incarnata, tuberosa, tomentosa, curassavica), as well as 3 
species of related genera. 

From the above xecords one may assume, subject to further study, 
that the larvae of the monarch butterfly will feed on all species of 
Asclepias. Hence, breeding populations may occur in many areas of 
Peninsular Florida in addition to those recorded here, so long as tempera- 
ture permits the survival of both the host plant and the larvae. 


Discussion 
Areas of Concentration and Flight Directions 


It has been well documented that the autumnal migrants follow a 
northeast to southwest flight direction (Urquhart, 1960). It has also been 
shown that although small breeding populations may occur south of Lat. 
39°N, by far the greatest concentration of summer population build up 
takes place between Lat. 45°N and Lat. 35°N and east of Long. 100°. 
This is specifically correlated with the distribution of Asclepias syriaca, 
a species which, by the nature of its rhizomatous root system, grows in 
dense clusters often covering many acres of marginal land as well as 
appearing in considerable abundance along gravel roadways and rail- 
way lines (Fig. 5). Other species occur in this area of syriaca abundance, 
such as amplexicaulis, exaltata, tuberosa and verticellata, thus adding 
to the great abundance of host plants throughout the area indicated. 


VOLUME 30, NUMBER 2 81 


a een SS AN \ 


ie | we 
( | vi a se We aN 
| cae i ieee eaten N5e- ee 


Fig. 5. Area of Asclepias abundance (stipled) and direction of monarch migration 
from this area to the southwest (solid arrow) accounting for the relatively meager 
migration through Peninsular Florida. 


Since the line of autumnal migration is from northeast to southwest, as 
indicated by the arrow in Fig. 5, the majority of the autumnal migrants 
will tend to by-pass Peninsular Florida. Observations made on a number 
of field trips to northern Florida, as well as the eastern section of the 
United States (east of and including the Appalachian Highlands) have 
definitely indicated this trend. Therefore, relatively few autumnal 
migrants from the area of most abundant host plants travel down Penin- 
sular Florida. There will be a certain small number of migrants emanating 


82 JourNAL OF THE LEPIDOPTERISTS SOCIETY 


from those areas where species of Asclepias occur in smaller numbers in 
the south and southeast parts of the continent; together, this constitutes a 
minority of the autumnal migrants. 

If the above concept is correct, namely that Peninsular Florida is by- 
passed, then one would expect to find large numbers of migrants along 
the Florida Panhandle Gulf coast. This has been found to be the case 
(Urquhart, 1966). During the autumnal migration countless numbers of 
monarchs move west along the Gulf coast with great concentrations in the 
Apalachee Bay area at a time when relatively few specimens can be ob- 
served in the Gulf coast area of Peninsular Florida. Of those that do 
enter Peninsular Florida, field observations together with alar tag returns 
indicate that the movement is south and for the most part along the coast 
of the Gulf of Mexico. 

If the Peninsular Florida populations move southward one would ex- 
pect to find migrants in some of the islands of the Gulf of Mexico and, 
perhaps, in parts of southern Mexico and Central America. As a result 
of field expeditions to the Yucatan Peninsula of Mexico and Guatemala, 
both larvae and imagoes of the monarch butterfly were collected in the 
autumnal migration period (Fig. 6). Of the latter, many were definitely 
migrants as indicated by the colour change and scale loss ( Urquhart, 
1960). The situation in these areas is similar in all respects to that found 
in Florida. 

It would therefore appear that the small, migrating population of 
monarch butterflies moving down Peninsular Florida, together with 
generations resulting from ovipositing migrants, travel via the offshore 
islands of Florida in the Gulf of Mexico and along the Florida Keys, 
eventually reaching Cuba. From Cuba, the flight continues from the 
area of Pinard el Rio to Yucatan, and from there to various parts of 
southern Mexico and Central America in which migrants in ovarian 
dormancy, gravid migrants, and generations of larvae and imagoes result- 
ing from the latter occur (Fig. 6). 

No large over-wintering concentrations of migrants in dormancy, as 
reported for California and observed in at least one area of Mexico (ms. 
in preparation), were located. It is assumed, however, that individuals 
from Yucatan and Guatemala, as well as other adjacent areas, will, during 
the vernal migration, travel northeastward, entering Cuba and eventually 
Florida in the spring and early summer, thus increasing the population 
of Peninsular Florida during this period. 


Correlation Between Larval Population and Migrants 


As pointed out in this paper and previously documented (Urquhart, 
1960, 1966), migrant females may be in a physiological state which is 


VoLUME 30, NuMBER 2 


83 


Fig. 6. Migration route from Peninsular Florida to Cuba and Yucatan. Migrant 
monarchs and larvae found at Chetumal, Flores, Morales and Guatemala as indicated. 
In addition to localities indicated in the figure, migrants and larvae were found at 
Villahermosa, Palenque, Zinacantan, Tuxtla and Oaxaca in the isthmus area of 
southern Mexico. 


84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


referred to as ovarian dormancy; or they may be capable of ovipositing 
wherever species of milkweed occur along the southward migratory 
route. Hence, larvae will occur from late summer (August) to late fall 
(November) along the migratory route so long as freezing temperatures 
do not occur, limiting the growth of milkweed (Urquhart, 1960). Ovi- 
positing migrants reaching Peninsular Florida will oviposit on various 
species of milkweed and, so long as the temperature is suitable for larval 
growth and the host plant available, larvae and imagoes may be found 
throughout the year (Urquhart & Stegner, 1966). 

Two peaks of monarch abundance (Fig. 4) occur throughout the year, 
a small peak in April-June and a larger one in August-November. The 
first peak is correlated with the advent of ovipositing vernal migrants, as 
discussed previously, and the second and larger peak with the advent 
of a much larger population of ovipositing autumnal migrants. Correlated 
with the occurrence of migrants, and hence a larger population of 
imagoes, is the presence of more abundant food plants as compared to 
conditions during the winter period. Decreased populations occur in 
January—March and July-August as the result of the decrease in the num- 
ber of migrants. The meager population is, however, maintained as a 
result of the lingering presence of a few migrant stragglers as well as 
first and second generations resulting from gravid female migrants. 

The rate of growth of the larvae during the winter period is reduced 
due in part to the lack of available host plants and partly to the effect 
of low temperatures lowering the growth rate. Concerning the latter, 
observations on various field expeditions during periods of low tempera- 
tures showed that larvae will drop from the host plant and remain on 
the ground in an inactive state until the return of higher temperatures. 


Factors Limiting a Large Population in Peninsular Florida 


Larval response in seeking host plants. Urquhart (1960) indicated, 
by movement tracings of larvae with respect to the ability of the larvae 
to locate the host plant when in close proximity to it, that they were 
unable to locate the host plant except by random movement. Experiments 
carried out in the field indicated that many larvae, when once dislodged 
from the host plant, were unable to locate the same plant or a substitute 
one when the plants were widely separated. As a result of numerous 
observations on various field trips through Peninsular Florida it is con- 
cluded that the various species of milkweed plants are widely separated 


in any growth area. Hence, many larvae would not complete the life 
cycle in most parts of F pied 


~ VotuME 30, NuMBER 2 85 


15 


13 


11 


NO. OF SPECIMENS TAGGED 
it~) 


1 2 3 4 5 6 7 
RECAPTURES—WEEKLY INTERVALS 


Fig. 7. Showing the rapid decrease in recapture of tagged specimens due to 
emigration from the area of release. 


Insufficient food supply and competition with other species. 
Although there are a number of species of milkweed found in Peninsular 
Florida, there are not sufficient numbers of any one species, or of all 
species, to support a large population such as occurs in northeastern 
United States. Also, with the advent of low temperatures, particularly 
in areas north of the frost line (Lat. 28°N), many species of milkweed 
will drop their leaves and not resume growth until the following spring: 
in addition, many species die off during the winter period and resume 
growth the following spring (Woodson, 1954). 


86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Brower (1962) has hypothesized competition for host plants with 
Danaus gilippus berenice (Cramer). However, in our six years of inten- 
sive field studies, during which time many hundreds of monarch larvae 
were collected and reared for migratory purposes, that were carried out 
through the entire calendar year in many localities throughout Florida 
(both Peninsular and Panhandle), there was no evidence of such com- 
petition. Throughout most of Florida, the larvae of D. g. berenice were 
scarce, only a few specimens being collected by our field assistants and 
ourselves, while monarch larvae, in many areas, were quite abundant. 
It is suggested that the observations of Brower were for a limited area 
and during a short period of the year during which place and time, D. g. 
berenice larvae were locally abundant. 

Imago migration. It has been shown in this paper that relatively few 
alar tagged specimens were recaptured, even though all were liberated 
in areas of high human population, as compared to returns for other parts 
of the continent (Urquhart, 1960); further, that the majority of recap- 
tures were made during the first two weeks following release with a 
marked drop in recaptures in the third and fourth weeks and none after 
six weeks (Fig. 7). From this it is concluded that the imago population 
does not remain in a given area but like the true migrants move away 
from the area to the south in the fall and winter and north in spring and 
early summer. 


SUMMARY 


1. Owing to the southwest pattern of the autumnal migration and the 
relationship of larvae to the presence of the abundant supply of milk- 
weed plants in eastern North America, relatively few migrants pass 
through Peninsular Florida. 

2. Female migrants that are in ovarian dormancy together with migrant 
males move southward or southwestward to Cuba and thence to the 
Yucatan Peninsula. 

3. Gravid migrant females deposit eggs on species of milkweed along 
the migratory route. 

4. Generations resulting from eggs deposited by gravid females move 
southward and southwestward, as in 2 above; the females in gravid state. 

5. As a result of the paucity and scattered nature of the milkweed 
plants, limiting survival of larvae, together with the advent of low 
temperatures which prolongs larval growth period and reduces available 
food plants, and the movement southward of the imagoes, the monarch 


population in Peninsular Florida remains relatively low throughout the 
year, 


VOLUME 30, NUMBER 2 87 


6.A definite correlation exists between the number of larvae and 
imagoes and the autumnal and vernal migrations. 


ACKNOWLEDGMENTS 


The present investigation is part of a larger research program dealing 
with the migrations and ecology of the monarch butterfly and sponsored 
by research grants from the National Research Council of Canada, the 
National Geographic Society, and the Insect Migration Research 
Association. 


LITERATURE CITED 


Brower, L. P. 1961. Studies on the migration of the monarch butterfly I. Breed- 
ing populations of Danaus plexippus and D. gilippus berenice in south central 
Florida. Ecology 42: 76-83. 

1962. Evidence for interspecific competition in natural populations of the 
monarch and queen butterflies, Danaus plexippus and D. gilippus berenice in 
south central Florida. Ecology 43: 549-552. 

Urounart, F. A. 1960. The Monarch Butterfly. Univ. Toronto Press, Toronto. 
361 p. 

1966. A study of the migrations of the Gulf Coast populations of the 
monarch butterfly (Danaus plexippus L.) in North America. Ann. Zool. Fennici. 
3: 82-87. 

Urouuart, F. A. & R. W. Stecner. 1966. Laboratory techniques for maintaining 
cultures of the monarch butterfly. J. Res. Lep. 5: 129-136. 

Woopson, R. E. 1954. The North American species of Asclepias L. Ann. Missouri 
Bot. Garden 41: 1-211. 


88 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


FUGITIVE COLOR IN THE MALES OF CERTAIN PIERIDAE 


Harry K. CLENCH 
Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15213 


In late June 1973 I collected a series of Nathalis iole Boisduval at Nicolls 
Town, North Andros Island, in the Bahamas. They were spread shortly 
after my return home and a curious fact was then immediately evident. 
The small, oval, androconial patch of each of the males, near the base 
of the costa on the upperside of the hindwing, was bright, vivid orange. 
In all our other males of N. iole this patch is dull yellow, regardless of 
whether they came from the West Indies (including New Providence 
Island in the Bahamas), the United States or Central America. 

A month or so later Mr. Allan Watson of the British Museum was 
visiting us and I told him about this odd population. I showed him the 
series, the first time in several weeks that I had seen the specimens myself. 
The orange color of the androconial patches seemed to be much duller 
and less striking at this time than I had remembered it earlier. In late 
August, two months from when they had been captured, I examined the 
series once again. The orange color was completely gone, and the speci- 
mens were all perfectly ordinary-looking N. iole, with the androconial 
patches the usual dull yellow. 

I then wrote to two colleagues, Dr. Lee D. Miller and Mr. F. Martin 
Brown, both of whom live in “iole country,” and described the situation, 
asking them if they could catch and examine fresh N. iole males to see 
what color the patches were in their areas. They both reported them to 
be orange, as they had been in the Andros specimens. 

In early June 1974 I took additional N. iole in the Bahamas, this time 
on Grand Bahama Island, at Pine Ridge, 4 mi. NE of Freeport. In July, 
just over a month since they were captured, the scent patches of the 
males were orange (although not as bright as I recall them in the Andros 
series); in August, two months after capture, the orange was gone (a 
slight trace in one specimen may have remained). Examination of speci- 
mens under a dissecting microscope in July showed nothing unusual 
other than a uniform orange tinge to the patch. 

This color change in the androconial patch of male N. iole had been 
observed long ago by Gundlach (1881:100). 

On North Andros in June 1973 I also caught a pair, male and female, 
of Eurema (Pyrisitia) messalina blakei Maynard. After they had been 
spread an examination showed that the male (but not the female) had 
a distinct pink bar on the underside of the forewing, running from near 


VoLuME 30, NuMBER 2 89 


the base distad for about 2/3 of the inner margin, in interspace Cuz-2A. 
Other E. messalina males in the collection, from the Bahamas, Cuba and 
and Jamaica, had no pink on any of them. 

I kept this curiosity in mind when in the Bahamas again in June 1974. 
On South Andros Island, near Driggs Hill, E. messalina was not uncom- 
mon and I obtained a series, examining each of them as it was caught. 
The males all had the pink bar. 

On my return to Pittsburgh I examined again the now year-old male 
from North Andros. To my surprise the pink bar had disappeared com- 
pletely. I had examined this male in April 1974 and am reasonably sure 
that the pink bar was still present, some 10 months after capture. Exam- 
ination of my June 1974 specimens in August, two months after capture, 
showed that of eight males, the patch was absent in two, extremely faint 
in five, clearly visible in one. 

When the newer specimens had been spread I examined the males 
under the dissecting microscope. The scales in the pink area were white 
and all seemed identical in structure to the scales in the white areas 
nearby on the wing, except that each one had its distal edge narrowly and 
sharply bright red, as though it had been dipped in red ink. 

The situations in the two species are probably related. In both species 
only males are involved; and in both species the color is fugitive. It may 
also be significant that the ground color of Eurema messalina is white and 
the fugitive color of the bar is pink, whereas in Nathalis iole the ground 
color of the patch is yellow, which would appear orange if combined 
with pink. In other words, the transient pigment may be red in both, but 
appear as pink in E. messalina on the white ground, and as orange in 
N. iole because the patch substrate is yellow. 

The transient color in N. iole is clearly associated with an androconial 
patch, a readily seen and morphologically distinct structure. The under- 
side of the forewing near the inner margin, where the pink bar occurs in 
E. messalina, is also a common site of androconial structures. It seems at 
least possible that the pink color transiently indicates an androconial area 
in that species that otherwise is not characterized by any visible structural 
modification. 

If this is indeed the case, then androconial patches or areas may be 
present in more species than we know. Ordinary visible scent patches 
are useful taxonomic structures, their presence, location and configuration 
providing valuable clues to relationships. If less easily visible patches 
also exist, their attributes could be just as useful once they are made 
known. Some of them may be transiently visible because of an evanes- 
cent pigment, as is presumably true of E. messalina. In some species 


90 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


such pigments might be even more fugitive than in either of those re- 
ported here, so examination of males should be made when they are 
captured or as soon as possible thereafter. 

It is even conceivable that androconial patches may exist without any 
ordinarily visible clue whatever, should the transient substance(s) associ- 
ated with them be unpigmented. Perhaps such patches could be “de- 
veloped,” by prompt treatment with a suitable reagent that would render 
the substance visible; or perhaps submicroscopic structural differences 
exist in the scales, invisible under conventional, magnification but show- 
ing up, say, in the scanning electron microscope. 

Transient pigment need not always be associated with androconial 
areas. Clark & Clark (1951: 87) describe fugitive pink color in the 
North American pierid, Euchloe olympia Edwards, that may well be 
non-androconial. They make no mention of the sex of the individuals 
that show it, but the manner of their description suggests that it may be 
in both sexes; and the pink is found on wing areas where androconial 
patches rarely occur (anterior part of the hindwing underside; costal 
area of the forewing on both surfaces). From the Clarks’ description we 
cannot yet rule out completely the possibility that the evanescent pink 
of E. olympia is androconially associated, but it seems unlikely. 


LITERATURE CITED 


Criark, A. H. & L. F. Cuarx. 1951. The butterflies of Virginia. Smithsonian Misc. 
Coll. 116 (7), vii + 239 p., ill. 

Gunpbiacu, J. 1881. Contribucion a la entomologia Cubana. Habana (G. Montiel). 
445 + xii p. 


VoLUME 30, NUMBER 2 91 


LARVAL MORPH VARIATION IN CHLOSYNE LACINIA 
(NYMPHALIDAE) 


RayMonp W. NEcK* 
Department of Zoology, University of Texas, Austin, Texas 78712 


Mature larvae of the patch butterfly, Chlosyne lacinia (Geyer), exhibit 
striking color polymorphism. First described by T. D. A. Cockerell in 
Edwards (1893) and illustrated by Neck et al. (1971), the three morphs 
are as follows: rufa, an all orange or orange-red form; nigra, an all black 
form; and bicolor, a form basically black as in nigra but with a prominent 
mid-dorsal row of orange to orange-red blotches whose proximity to each 
other gives the impression of a stripe. The black pigment is present in 
the cuticle while the orange pigment is in a deeper layer (hypodermis ) 
that shows through window-like areas of the cuticle devoid of pigment. 

Laboratory crosses utilizing adults of known larval phenotypes revealed 
that the larval color polymorphism involves two unlinked autosomal loci 
(Neck et al., 1971). At one locus bicolor (B) is dominant to nigra (b). 
At the second locus rufa (R) is dominant to non-rufa (r). The ruta 
allele is epistatic to the first locus; no phenotypic expression of the first 
locus occurs when the rufa allele is present at the second locus. 

Although there are only three major morphs in this species, a variety 
of subtypes exists which indicate the influence of genetic modifiers 
and/or environmental factors. The most common variant is a nigra form 
with circular light yellow-orange spots of varying intensity at the base 
of each mid-dorsal spine. This phenotype, named dot, is pictured in Plate 
1D of Neck et al. (1971). It may be the same form reported by Cockerell 
(in Edwards, 1893) and Gorodenski (1969) although the description by 
Cockerell (“with numerous yellow-white spots”) could possibly refer to 
the super-dot form described below. 

Nearly all nigra have a small spot at the base of each mid-dorsal spine. 
(42 of 50 or 84.0% nigra larvae collected 10 September 1972 at Austin 
were definitely dot; others had very small dot.) In some individuals it 
is inconspicuous enough to be unnoticed unless critically examined under 
a dissecting microscope. In other individuals the dot is so large as to 
create confusion with the bicolor morph. In most cases the dot of the 
nigra tends towards yellow whereas the block of the stripe in bicolor 
is more orange. In addition, a black mid-dorsal line present in those 
larvae is seen in the bicolor larvae only between the orange spots and 


1 Present address: Texas Parks and Wildlife Department, John H. Reagan Building, Austin, 
Texas 78701. 


92 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Parental larval phenotypes: black x black. 


Phenotypes of Progeny 


brood number black dot total 
129 84 86 170 
DAP) 194 8 . 202 — 
273 120 jLIL Val 
DATA 92 Dil 119 
301 50 Dall 71 


never in them. In dot nigra this line is seen between and entering the 
dot, continuing almost to the base of the spine. 

Super-dot is also a variant of nigra with a yellow-orange spot at the 
base of each mid-dorsal, medio-dorsal, supra-spiracular and_ infra- 
spiracular spine. Super-dot also has a spot at the base of paired spines 
just above each proleg, a spot also present in bicolor and nigra. The more 
heavily yellowish-pigmented dot larvae also have yellow spots at the 
base of supra-spiracular spines, but they lack yellow at the base of medio- 
dorsal spines. A relationship between the dot and super-dot varieties 
may exist. One larva, collected as fourth instar was an extreme form of 
dot, i.e., very large spots at the bases of the mid-dorsal spines (but not 
others). This larva subsequently molted into a fifth instar super-dot 
form. Although dot larvae would thus appear to have a tendency to 
produce an abundance of yellowish spots, many dot phenotypes occur 
in fifth instar larvae. 

Variation exists in the amount of melanin present in the cuticle. Nigra 
exhibits definite black and gray forms although intermediates occur 
that are difficult to place in either category. Dot larvae may be either 
gray or black, but all super-dot forms observed have been gray. The 
more evident dot larvae, however, are grays. The more intense pigmenta- 
tion in the black form tends to cover up the dot, but it can still be 
observed in most specimens. Therefore, unless the genetic basis involves 
incomplete dominance or incomplete penetrance of a normally dominant 
allele, two separate genetic systems are involved. A form of. bicolor 
exists with extensive melanization which obscures most of the orange. 
This form is sometimes difficult to distinguish from nigra, particularly 
in third and, sometimes, fourth instar. Inspection of less heavily mela- 
nized thoracic segments where the yellow-orange pigment can be seen 
more readily reveals such larvae as bicolor. 

Intense melanization is also observed in rufa. Black pigmentation 
which is normally restricted to the base of various spines and inter-seg- 
mental membranes may be so intense that it is difficult to differentiate 


“VoLUME 30, NuMBER 2 93 


TABLE 2. Parental larval phenotypes: dot x dot. 


Phenotypes of Progeny 


brood number black dot total 
en 455 74 0 74 
507B 288 13 301 
516 44 15 59 
517 72 36 108 
559 70 24 94 
570 62 0 62 


it from nigra. Heavily pigmented rufa may be correctly determined as 
rufa by orange pigmentation which can be detected in areas around and 
ventral to spiracles. Also, rufa lacks the mid-dorsal black line present 
in the non-rufa morphs. A similar darkening appears in Chlosyne harrisii 
(Scudder) as the normally orange larvae of this species have been ob- 
served to be “nearly black at maturity, the fulvous being represented 
merely by a few dots and small spots” (Edwards, 1877). In this case, 
however, the imago was also strongly melanic; this has not been observed 
in C. lacinia. 

Environmental conditions may exert a significant force in production 
of black, hidden-stripe and heavily melanized rufa forms. These forms 
are readily seen in laboratory cultures; they are quite rare in field popu- 
lations. Caterpillars raised under crowded laboratory conditions have 
been noticed to be generally darker than caterpillars occurring in the 
wild (see Long, 1952, 1953, and references therein). Cooler temperatures 
are also known to produce larvae which are more heavily pigmented 
than larvae reared at warmer temperatures (David & Gardiner, 1962). 
On the other hand, dot are often seen and super-dot forms have been 
seen in natural populations. The occurrence of super-dot forms appears 
to be correlated with large population sizes. Although observations are 
limited, this may indicate a relaxation of selection pressures as reported 
for adult phenotype of Melitaea aurinia Rott. (Ford & Ford, 1930). 

Attempts to determine the genetic basis of these forms have been in- 
conclusive (Tables 1 and 2). Certain black x black crosses yielded black 
and dot larvae in 1:1 and 3:1 ratios in addition to some broods highly 
skewed for black, i.e., very few dot. However, several dot X dot crosses 
yielded broods with both black and dot larvae and some broods with 
only black larvae! The genetic basis of these forms is vague and may be 
highly influenced by environmental conditions as previously reported in 
the larvae of the arctiid moth Utetheisa pulchella L. (Kettlewell, 1964). 
Kettlewell (1944) reported results which he interpreted as being due 


94 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


to the expression of a gene which is normally buffered in natural con- 
ditions but is released under artificial laboratory conditions. Major 
variation within morph classes in some amphibians (Volpe, 1961; Resnick 
& Jameson, 1963) has been shown to be the result of incomplete pene- 
trance (variable expressivity) of various genes. 


CONCLUSION 


In Chlosyne lacinia genetic modifiers and environmental influences 
apparently act upon a basic three-phenotype polymorphic system. The 
result is to modify a genetically discontinuous system of variation into a 
system of nearly continuous variation. Such continuous variation was 
described for the larvae of this species by two workers (Koehler, 1927; 
Comstock, 1927, 1946). 


LITERATURE CITED 


Comstock, J. A. 1927. Butterflies of California. Published by author, Los Angeles. 
334 p. 

. 1946. A few pests of sunflower in California. Bull. South. Calif. Acad. 
Sci. 45: 141-144. 

Davin, W. A. L. & B. O. C. Garpiner. 1962. Observations on the larvae and 
pupae of Pieris brassicae (L.) in a laboratory culture. Bull. Ent. Res. 53: 417— 
436. 

Epwarps, W. H. 1877. Description of the preparatory stages of Phyciodes harrisii, 
Scudder. Can. Ent. 9: 165-168. 

1893. Notes on a polymorphic butterfly, Synchloe lacinia, Geyer (in Hub. 
Zutr.), with a description of its preparatory stages. Can. Ent. 25: 286-291. 

Foro, H. D. & E. B. Forp. 1930. Fluctuations in numbers, and its influence on 
variation, in Melitaea aurinia, Rott. (Lepidoptera). Trans. Roy. Ent. Soc. 
London 78: 345-351. 

Goropenski, S. A. 1969. The genetics of three polymorphic larval color forms 
of Chlosyne lacinia (Lepidoptera, Nymphalidae). Genet. Res. 14: 332-336. 
KETTLEWELL, H. B. D. 1944. Temperature effects on the pupae of Panaxia domi- 

nula Linn. Proc. S. London Ent. Nat. Hist. Soc. 1943-4: 79-81. 

1964. The inherited and environmental contributions to the patterning 
of Utethesia pulchella L. (Lep.). Entomologist (London) 97: 169-172. 
KOEHLER, P. 1927. Biologia de Chlosyne Saundersi Pbl. & Hew. Revista Soc. Ent. 

Argentina 1-2: 3-4. 

Lone, D. B. 1952. Some problems of polymorphism in insects. Proc. Roy. Ent. 

Soc. London 27: 99-110. 


1953. Effects of population density on larvae of Lepidoptera. Trans. Roy. 

Ent. Soc. London 104: 543-585. 

Neck, R. W., G. L. Bus & B. A. DrummMonp. 1971. Epistasis, associated lethals 
and brood effect in larval color polymorphism of the patch butterfly, Chlosyne 
lacinia. Heredity 26: 73-84. 

Resnick, L. EK. & D. L. JAMeson. 1963. 
Science 142: 1081-1083. 

VoLrE, E. P.. 1961. 
134: 102-104. 


Color polymorphism in Pacific tree frogs. 


Variable expressivity of a mutant gene in leopard frog. Science 


VoLUME 30, NuMBER 2 95 


PORTABLE OUTDOOR CAGES FOR MATING FEMALE GIANT 
SILKWORM MOTHS (SATURNIIDAE )' 


THOMAS A. MILLER * AND WILLIAM J. COOPER ” 


U. S. Army Medical Bioengineering Research and Development Laboratory, 
USAMBRDL BLDG 568, Fort Detrick, Maryland 21701 


Like many lepidopterists who colonize and study the saturniids, we 
must find time apart from professional duties and other daily activities 
to pursue our interest in this family of moths. Therefore, our rearing 
methods must be both efficient and reliable if we are to accomplish any- 
thing other than the mere maintenance of colonies. Although most of 
the saturniid species we have reared can be colonized for many genera- 
tions without difficulty, we have at times had a particular problem obtain- 
ing fertile eggs to perpetuate existing colonies or to begin new ones. This 
problem has been a function largely of limited available time to ensure 
the mating of females either by hand mating or by the use of indoor 
cages. For this reason we undertook an evaluation of methods for the 
routine, unattended mating of female saturniids indigenous to our area. 
We decided beforehand that the tying out of virgin females and the use 
of various-size stationary cages or traps (Collins & Weast, 1961; Quesada, 
1967; Villiard, 1969) were not suitable to meet our requirements. Our 
approach, therefore, was to construct and test a series of portable out- 
door cages that could be substituted locally for tying out or stationary 
cages and could also be easily transported for use in remote field areas. 
These cages were designed to minimize or prevent the escape of females 
placed therein while permitting pheromone-seeking males access for the 
purpose of copulation. 


MATERIALS AND METHODS 


1973 Studies. The cages used during 1973 were fabricated from heavy- 
gauge galvanized metal screen (1.27 cm openings). Each cage was 
cylindrical and contained a series of interior and exterior baffles as 
shown in Fig. 1. All components were joined by soldering. The cages 
were spray painted inside and out, in a pattern of brown, green, and 
black, to camouflage them and minimize the probability of human tamper- 
ing. We constructed cages of two sizes: (1) small cages with the 
cylindrical portion 23 cm in height and 23 cm in diameter and (2) 

1 The opinions contained herein are the private views of the authors and should not be con- 


strued as official or reflecting the views of the Department of the Army. 
2 Entomologist and Research Chemist, respectively. 


96 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


TOP VIEW 


SIDE VIEW 


i tease 


Fig. 1. Diagrammatic wire-screen mating cage: interior baffles (A, B, B’, C); 


exterior baffles (D, D’); and dead space (E). Side view without exterior baffles 
shown. 


large cages with the cylindrical portion 46 cm in height and 38 cm 
in diameter. The entrance and baffles were approximately proportional 
to the size of the cage. Additional details of construction are shown in 
Figs. 2 and 3. The studies during 1973 were conducted at Edgewood 
Arsenal, Harford Co., Maryland. The saturniid species we used were 
obtained as diapausing pupae from various sources (Table 1). All 
pupae were held over winter (1972-1973), during which they were 
exposed to outdoor temperatures from the time they were obtained until 
the emergence of adults. Newly emerged virgin females were placed 


VoLuME 30, NuMBER 2 | 97 


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ages 
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Figs. 2-3. Wire screen cages used during 1973: 2, front view of cage containing 


C. promethea female no. 1; 3, angular view of cage containing C. promethea female 
no. 4. 


outdoors in mating cages on the day of emergence or on the following 
day. Small cages containing females of species in which males are day- 
flying [e.g., Callosamia promethea (Drury )] were placed outdoors in the 
late afternoon (after 1630 EST) but were not left overnight. Large cages 
containing females of species in which males are night-flying (i.e., all 
others in Table 1) were placed outdoors in the late afternoon or early 
evening, left overnight, and checked the next day between 0700 and 
0800 EST or 1200 and 1300 EST. This schedule was used because work 
requirements precluded activities at other times. Cages were hung from 
tree branches such that they were about 1.5 m above ground. All 
females used in the cages were held indoors afterwards and allowed to 
oviposit so that the fertility of the eggs could be verified. 


98 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Taste l. Sources of saturniid moths used in mating-cage studies during 1973 and 
1974. 


Species Source 


1973 Studies - 
Antheraea polyphemus Diapausing pupae purchased from a dealer in fall of 1972. 


Actias luna Diapausing pupae reared in Harford Co. during summer of 
1972. 

Hyalophora cecropia Diapausing pupae collected in Harford Co. during winter 
of 1972-1973. 

Callosamia promethea Diapausing pupae collected in Harford Co. and Portage Co. 


(Ohio) during winter of 1972-1973. 
1974 Studies 


Hyalophora cecropia Diapausing pupae reared in Harford Co. during summer of 
1973. 
Callosamia promethea Diapausing pupae reared in Harford Co. during summer of 


1973; and diapausing pupae collected in Harford Co. and 
Portage Co. (Ohio) during winter of 1973-1974. 


1974 Studies. The design of the cages used during 1974 was based on 
the previous year’s results, as will be discussed later. We used metal cans 
to construct tubular cages of two sizes: (1) small cages with a 14 cm 
length and 10 cm diameter and (2) large cages with a 17 cm length and 
15.5 cm diameter. For both sizes, the metal ends of the can were re- 
moved. One of the open ends was covered with a plastic lid that allowed 
us to remove and repiace female moths. The other open end was covered 
with galvanized metal screen, which was soldered into place. For the 
small cages the screen had diamond-shape openings measuring 1.27 
cm on a side (Fig. 4). For the large cages the screen had hexagonal- 
shape openings measuring 1.5 cm on a side (Fig. 5). To minimize human 
tampering, these cages were also spray painted in a camouflage pattern 
of brown, green, and black. The studies during 1974 were conducted at 
Edgewood Arsenal and Havre de Grace, both in Harford Co., Maryland. 
The saturniid species we used and their sources are shown in Table 1. 
The large cages were used for female Hyalophora cecropia (Linnaeus), 
and the small cages were used for female C. promethea. The procedures 
for handling the pupae, the schedule for setting out the females in the 
cages and rechecking them, and verification of the fertility of the eggs 
were all the same as in the 1973 studies. 


RESULTS 


1973 Studies. During 1973 we used a total of 12 virgin females, repre- 
senting four saturniid species, to evaluate the wire-screen mating cages 


VOLUME 30, NuMBER 2 99 


ed 


Figs. 4-5. Tubular cages used during 1974: 4, small cage showing C. promethea 
female no. 8 on diamond-shape screen; 5, large cage showing H. cecropia female 
no. 2 on hexagonal-shape screen. 


(Tables 2 and 3). Each of the two Antheraea polyphemus (Cramer) 
females attracted a male and mated with no apparent difficulty. Neither 
of the females moved from their original positions in the cages, even 
though the first of these did not attract a male until the third night. The 
first two female Actias luna (Linnaeus) also attracted males and mated 
without apparent difficulty. The third female of this species escaped 
from the cage during the first night. This was not totally unexpected, 
since this particular female was a very excitable individual and had 
escaped from the cage the previous day while being held indoors. The 
single female H. cecropia did not attract, or at least did not mate with, 


100 JouURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Results of studies using various saturniid females in large, wire-screen 
mating cages. Edgewood Arsenal, Harford Co., Maryland, 1973. 


Night 
Date 
21 I ] Mal Moths Eggs 
Reto r En ae nC nee etaered _ Mated Fertile 
Antheraea polyphemus 
1 9-V 10-V 12-V de At, 
2 19-V 20-V 20-V + + 
Actias luna 

i 15-V 15-V 16-V ae + 

2 18-V 19-V 19- 4 bie 

3 20-V 21-V — = jo 


Hyalophora cecropia 
1 30-V 31-V 1-Vi? + + 


1Female escaped from cage during night of 21-V. ; 
2No male attracted outdoors; female mated with reared male that emerged indoors on 31-V. 


any males during the first night. The cage, containing this female, was 
brought indoors the following morning and left there until noon. By 
noon, a male H. cecropia, which had emerged indoors the previous day, 
entered the mating cage and copulated with the female. With the excep- 
tion of the one female A. luna that escaped, we obtained large numbers 
of fertile eggs from each of the foregoing females. The six female C. 
promethea all attracted males and successfully mated (Table 3). In two 
cases the males did not enter the cage but mated with the females 
through the screen. In one case, two males found their way into the 
cage, although only one of them mated with the female. In another 
instance we were unable to bring the cage indoors after mating had 
occurred, and the moths evidently escaped during the night. We obtained 


Taste 3. Results of studies using Callosamia promethea females in small, wire- 
screen mating cages. Edgewood Arsenal, Harford Co.., Maryland, 1973. 


Afternoon 

F ] Fond F ] M 

‘emale Hemale Female 1 Moth 

Number Emerged in Cage Naa Morea ee 
| 2-VI 2-VI 3-VI ai i 
2 4-VI 4-VI 4-V}! a a 
3 4-VI 4-VI 5-VI ae Bi 
4 7-VI 8-VI 8-VI +. + 
5 7-VI 8-VI 8-VI' + ae 
6 8-VI 8-VI 8-VI dk — 

‘Male did not enter cage; mated with tarale through screen. 


*Mated moths left in cage overnight (8-9-VI); both escaped. 


VoLUME 30, NuMBER 2 101 


TABLE 4. Results of studies using Hyalophora cecropia females in large tubular 
mating cages. Edgewood Arsenal (EA) and Havre de Grace (HDG), Harford Co., 
Maryland, 1974. 


Night 
Date HS 
Female Female Female Male Cage Moths Eggs 
Number Emerged in Cage Attracted Location Mated Fertile 

I 8-VI 8-VI —! EA ae ata 

p) 9-VI 9-VI 10-VI EA i st 

o 9-VI 9-VI 11-VI HDG + + 

4 10-VI 10-VI 11 & 12-VI? EA +. + 

5 11-VI 11-VI Ls EA ie zi 

6 13-VI 13-VI 14-VI EA + + 

vl 13-VI 13-VI 13-VI EA ae 4. 

8 13-VI 13-VI 14-VI HDG + + 

9 15-VI 15-VI 15-VI HDG ++ -- 

10 16-VI 16-VI 17-VI HDG -- + 

GR 17-VI 18-VI 18-VI HDG + + 

12, 18-VI 19-VI 19-VI HDG + + 

13} 22-VI 22-VI 22-VI HDG 4+ + 

14 22-VI 22-VI 22-VI HDG + + 


1No male observed; female oviposited night of 11-VI; eggs not fertile. ‘ 
2 Female mated with a different male each night; deposited fertile eggs after each mating. 
3. No male observed; female oviposited night of 14-VI; eggs fertile. 


a large number of eggs from five of the female C. promethea. Since the 
male C. promethea are day-flying, we were able to make observations 
as they arrived in the area of a caged female. Two or three males were 
usually attracted to each female, and they flew in apparent random direc- 
tions until contacting the cage. Once they encountered the cage, they 
maintained contact and moved in a circular manner around it. Then 
they either found the entrance or contacted the female as it rested on 
the outer portion of the cage. On two occasions, as mentioned earlier, 
the males mated with the females directly through the wire screen. Those 
males that found the entrance were not particularly aided by the exterior 
baffle (Fig. 1) because their persistent contact with the cage caused 
them to fly into the entrance without actually contacting the exterior 
baffles. 

1974 Studies. During 1974 we used 14 female H. cecropia and 10 
female C. promethea to evaluate the tubular mating cages (Tables 4 
and 5). Thirteen of the female H. cecropia successfully attracted males, 
mated and deposited fertile eggs (Fig. 6). One female H. cecropia 
deposited eggs on the wire screen of the cage during the fourth night. 
No male was observed mated with this female, and the eggs were not 
fertile. Another female H. cecropia also deposited eggs on the wire 
screen of the cage during the fourth night, but these eggs proved to be 


102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


— 


TaBLeE 5. Results of studies using Callosamia promethea females in small 
tubular mating cages. Edgewood Arsenal (EA) and Havre de Grace (HDG), 
Harford Co., Maryland, 1974. 


Afternoon 
Date ———— 
Female Female Female Male Cage Moths Eggs 
Number Emerged in Cage Attracted Location Mated Fertile 
1 27-V 28-V 28-V EA + — 
2 27-V 28-V 29-V EA + + 
3 8-VI 8-VI 8-VI EA a Bie 
4 8-VI 8-VI 8-VI EA a Ei 
5 9-VI 9-VI 9-VI EA aly Je 
6 10-VI 10-VI 10-VI HDG -- “+ 
Ff 11-VI 11-VI 12-VI EA + + 
8 11-VI 11-VI 12-VI EA — + 
9 11-VI 12-VI 12-VI HDG + + 
10 16-VI 16-VI 16-VI HDG — + 


fertile even though no male was observed copulating with this female. 
A third female H. cecropia copulated with two separate males (Fig. 7) 
during the second and third nights in the cage and deposited fertile eggs 
after each mating. Each of the 10 female C. promethea (Table 5) 
attracted a male, mated, and deposited fertile eggs. 


DIscussION AND CONCLUSIONS 


The studies conducted during 1973 demonstrated to us that it was 
possible, through the use of the wire-screen mating cages, to obtain 
fertile eggs of A. luna, A. polyphemus and C. promethea. The information 
on the single female H. cecropia, although obtained under indoor con- 
ditions, demonstrated that a male of this species will enter the cage and 
mate with a female. The time and effort required during the studies 
were minimal, even though we used several females of some species to 
collect data on efficiency. Where a single fertile female would suffice 
to provide eggs for a colony, the time required would be even less. We 
found that the schedule required for setting out the cages and rechecking 
them could be made compatible with working hours. 

Despite these successes, we recognized certain shortcomings in the 
design of the wire-screen cages and the requirements for their use. 
lirst, the virgin female moths can escape, although the frequency of 
this occurrence appears to be very low because of their general quiescence 
prior to mating. Second, the cages had to be checked daily because after 
mating the female moths become active enough: to eventually find their 
way out of the cage. Because of these shortcomings and the observed 
ability of C. promethea to copulate through the wire screen, we designed 


VoLUME 30, NuMBER 2 103 


Figs. 6-7. Hyalophora cecropia mated pairs during 1974: 6, female no. 6 showing 
typical copulation through screen; 7, female no. 4 showing copulation with second 
male and eggs deposited on screen after previous mating. 


and successfully tested the tubular mating cages the following year. The 
wire screens used in the tubular mating cages appear to present no bar- 
rier to copulation when a male arrives at the cage. The tubular mating 
cages precluded the escape of females and obviated the requirement to 
recheck them daily. This constituted a definite advantage because our 
professional duties sometimes required us to be away for several days 
at a time. It would appear to be possible to set out a caged virgin female 
and have no requirement to recheck the cage for a week or more. Within 
this time period, even if the female mated on the first night or day and 
deposited eggs on the wire screen, we could still return and remove the 


104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


eggs before they hatched. We did not evaluate this concept during 1974, 
but plan to do so during a subsequent season. In addition, we plan to 
evaluate the concept of placing developing female pupae (in their 
original cocoons) in tubular mating cages and leaving them unattended 
while they emerge, attract males, mate, and deposit eggs on the wire 
screen. If this technique proves successful, it will permit an even longer 
period (perhaps two weeks ) between setting out and returning to collect 
the eggs. With one exception, we did not use these tubular mating cages 
operationally during 1974, because we had obtained sufficient numbers 
of fertile eggs during the experiments with H. cecropia and C. promethea. 
In subsequent seasons, however, we plan to incorporate the use of these 
cages into our routine system for rearing saturniids. 

One of the large tubular mating cages was used operationally in south 
Texas (5 mi. S George West, Live Oak Co., 28-September-74, 0845 CST), 
where one of us (TAM) set out a virgin female Eupackardia calleta 
(Westwood). The female was left unattended in the cage, although 
observations were made from nearby. Seven E. calleta males, which are 
morning fliers, were attracted within 10 min, and one of them copulated 
through the wire screen without difficulty. The use of this cage made it 
possible to easily obtain fertile eggs to colonize this species, and also 
provided us with data on an additional saturniid species. 


LITERATURE CITED 


Cotiins, M. M. & R. D. Weasr. 1961. Wild Silk Moths of the United States. 
Collins Radio Co., Cedar Rapids, Iowa. 138 p. 

Quezapa, J. R. 1967. Notes on the biology of Rothschilda Paroma (Lepidoptera: 
Saturniidae ), with special reference to its control by pupal parasites in El] 
Salvador. Ann. Ent. Soc. Amer. 60: 595-599. 


Vituiarp, P. 1969. Moths and How to Rear Them. Funk and Wagnalls, New 
York: 242 \p: 


VoLUME 30, NuMBER 2 105 


LARVAL FOODPLANTS, SPATIAL AND TEMPORAL 
DISTRIBUTION FOR FIVE SKIPPERS (HESPERIIDAE ) 
FROM TEXAS! 


Roy O. KENDALL? 
Route 4, Box 104-EB, San Antonio, Texas 78228 


AND 


M. A. RickarpD 
7003 Bissonnet # 1602, Houston, Texas 77036 


Although fairly common at times, Decinea percosius (Godman), 
Chiomara asychis georgina (Reakirt), Timochares ruptifasciatus (Plotz), 
Xenophanes trixus (Stoll) and Cabares potrillo (Lucas) are among the 
most uncommon skippers in collections from Texas. Local larval food- 
plants in the Gramineae, Malpighiaceae, Malvaceae, and Verbenaceae 
are given for the first time. Records of specimens collected by others, 
unless otherwise stated, are from personal correspondence or communica- 
tion. Biological arrangement follows dos Passos (1964), and botanical 
arrangement, Gould (1969). 


Decinea percosius (Godman) 1900 


First recorded from the United States by Freeman (1948) who col- 
lected a ¢ at Brownsville, Cameron Co., Texas, 28 Nov. 47. The 
following year he and Otto Buchholz collected 50 examples (42 6,8 2) 
between Brownsville and Southmost, 4 & 7 April 48. So far as we can 
determine, it was not collected again in Texas until 1969. 

Based on available data, this species appears to be only a periodic 
resident of extreme south Texas. In nature, it most likely feeds on broad- 
bladed grasses. It may have a larval or pupal diapause, or perhaps only 
a retarded immature development during winter months. Three broods 
are indicated. First adults emerge in late March and early April. The 
2nd brood appears from late June to early July, and the 3rd from mid- 
October to late November. Peak emergences are in early April, late 
June, and early November. 

Rearing records. At a spot near Southmost, SE of Brownsville, Rickard col- 


lected 19, 8 Nov. 69. The following day at the same spot, he collected 1 ¢, 
1 9, and Kendall took 3 2. On 10 Nov. at this location Kendall collected + more 


1 Contribution No. 313. Bureau of Entomology, Division of Plant Industry, Florida Department 
of Agriculture and Consumer Services, Gainesville 32602. 

2 Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, Florida 
Department of Agriculture and Consumer Services. 


106 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


2, one of which was demonstating oviposition behavior, and it was kept alive 
for possible egg production in the lab. Fifty-three eggs were deposited on Steno- 
taphrum secundatum ( Walt.) Kuntze, GRAMINEAE: 10 Noy. (8), 14 Nov. (13), 15 
Nov. (10), 16 Nov. (3), and 17 Nov. (19). Some eggs were preserved, and the 
remaining eggs started hatching the morning of 21 Nov. A few hatched each 
morning thereafter for about 10 days. First instar larvae readily ate S. secundatum. 
Most of the larvae died, however, in 2nd instar and the remainder in the 3rd instar. 
The exact cause of death was undetermined, but climatic conditions were suspect. 
Humidity in the lower Rio Grande Valley is very high, especially at night. In the 
rearing facility at San Antonio it was relatively low. 

At the same Southmost location on 4 April 70, Rickard collected 7 additional 
specimens (5 ¢, 2 @). From one confined female, about 20 eggs were obtained 
on ryegrass, Lolium perenne L., GRAMINEAE. After one day the 2 was placed in 
a killing jar and kept as a cabinet specimen. Eggs hatched 15-16 April and readily 
ate L. perenne. On 25 April 70 Rickard gave Kendall two of these larvae, one of 
which was in process of molting. Because the grass blade had dried around the 
larva, some difficulty was experienced in completing the molt; it died later and 
was preserved. The remaining larva, 2nd instar when received, ate L. perenne, 
S. secundatum and Cynodon dactylon (L.) Pers. It molted on 8, 15 & 25 May 
and pupated 16 June. A 2 emerged about 0645 CDT 26 June 70. It was interesting 
to note that a formal tent or shelter was not constructed by the larva until it was 
fully mature. At this time it remained head down in the shelter pending final 
defecation after which it pupated head up. It was also noted that feeding was 
largely at night. 

The remaining larvae kept by Rickard were switched to Johnsongrass, Sorghum 
halepense (1) Pers., GRAMINEAE, and all died during pupation, probably from 
malnutrition. These were reared in Houston, Texas where the relative humidity is 
comparable to that of the lower Rio Grande Valley area. 

Other Texas records. Cameron Co.: Brownsville and vicinity: J. R. Heitzman 
found it fairly common during the period 19 June-1 July 69; Joseph F. Doyle III, 
collected 1 6, 18 June 70; Rickard collected 3 6, 1 9, 21 March and 4 4, 5 April 
70; Kendall saw one 22 Oct. 72. Hipatco Co.: Relampago, Rickard collected 2 ¢, 
18 Oct., 2 6,1 9, 19 Oct. and 1 4, 30 Oct. 71. 


Chiomara asychis georgina (Reakirt) 1868 


First recorded for the United States by Skinner (1901); collected by 
Irvin Runyeon from southern California to the Arizona border. It is 
well established in Cameron and Hidalgo counties of southern Texas 
and ranges up the coast to Harris Co. and inward to Bexar Co. Three 
and possibly 4 broods are indicated. There appears to be an immature 
semi-diapause during January and February even though adults have 


been collected in each month of the year except February; the known 
larval foodplant is deciduous. 


Rearing records. While collecting in Bentsen-Rio Grande Valley State Park 
(B-RGVSP), Hidalgo Co., 29 Aug. 69, Rickard observed a female ovipositing on 
the foliage of Malpighia glabra L., MaLpicHIACKAE. Examination of the plant dis- 
closed 2 larvae in leaf shelters, one was lost, the other produced a @ on 25 Sept 
69. Kendall collected a larva feeding on this plant in Santa Ana National Wildlife 
Refuge (SANWR), Hidalgo Co., 13 Noy. 71; it pupated 6 Dec. but died before 
emergence. On 11 Sept. 72 at McAllen, he collected another larva; it pupated 20 


VoLuME 30, NuMBER 2 107 


Sept. and a Q emerged 29 Sept. 72. The larva of this species may feed on other 
Malpighiaceae such as Thryallis angustifolia (Benth.) Kuntze found in Bexar Co., 
and Janusia gracilis Gray found in far western Texas, New Mexico and Arizona over 
which area the insect ranges. 

Other Texas records. BEXAR Co.: Roy W. Quillin, Cibolo River in NE Bexar 
Co., 1 §, 21 Aug. 60; at his home in San Antonio, 1 ¢, 19 Aug. 65. CAMERON Co.: 
H. A. Freeman (1951), Brownsville, June and Aug. Roy O. & C. A. Kendall, 
Brownsville, 1 ¢, 3 April 57; 1 9, 4 Dec. 64; 1 ¢, 29 Oct. 65; 1 9, 25 Nov. 66; 
Harlingen, 5 ¢, 2 2, 20 Oct. 65. J. W. Tilden, Brownsville and vicinity, 4, 17-19 
@ermGow 4m. WwW of Boca Chica, 2 6; 26 Oct. 63; nr. Brownsville, 1 ¢, 1 9, 
13 Nov. 63. Joseph F. Doyle III, Brownsville, 1 9, 21 Dec. 67. M. A. Rickard, 
Brownsville, 2 ¢, 2 Sept. 67; several on 17, 18 and 31 Aug. 69; 1 ¢, 7 Sept. 69; 
2mometG Ock, 69:2 9, 9 Nov. 69; | 2, 4 April 70; 1 6, 17 Aug. 70.. Harris Co:: 
John B. Vernon, Houston, 1 ¢, 23 Oct. 71. Hmatco Co.: H. A. Freeman (1951), 
Pharr, March—June and Aug.—Dec. Roy O. & C. A. Kendall, B-RGVSP, 4 ¢,1 2, 
2IeNove 62.) Madero, 1 9, 11 Jan. 72; SANWR, 1 ¢, 14 Jan. 72; Mission, 1- 3, 
12 Sept. 72; McAllen, 1 ¢, 22 Oct. 72. J. W. Tilden, B-RGVSP, 1 ¢, 10 Nov. 63; 
SANWR, 1 ¢, 11 Nov. 63. Fay H. Karpuleon, nr. McAllen, 1 ¢, ? Dec. 65. J. R. 
Heitzman, SANWR, several, 21 & 24 June 66. M. A. Rickard, SANWR, 1 ¢, 1 9, 
oaAueo7; Lic. 5 Oct. 68;-1 2, 6 Oct. 68; 2 2,4 July 69; 1 36, 26 July 69; 1 ¢, 
Se wmenO-eomo sl, May 70: 26, 1 9,18 Aug: 70: 1 6, 12\Sept. 70; 1 @, 15 
icine 29 Oct. 71; 1 6, 31 Oct. 71; B-RGVSP, several, 15;.16, 17, 24, 25, 
29 & 30 Aug. 69 and 6 Sept. 69; 1 ¢, 14 Aug. 71; Madero, several, 25 Oct. 69; 
1 2,9 Nov 69. KiLEBERG Co.: Roy O. & C. A. Kendall, nr. Kingsville, 1 ¢, 23 Aug. 
64. Live Oax Co.: E. M. & S.F. Perkins, nr. Lake Corpus Christi State Park 
(LCCSP), 1 ¢, 10 Nov. 62._M. A. Rickard, hwy. 9 nr. Mathis, 1 @, 9 March 69. 
SAN Patricio Co.: J. W. Tilden (1974), Welder Wildlife Refuge, 1 ¢, 13 Oct. 63 
(in ROK coll.); 1 ¢, 2 Nov. 63. J. R. Heitzman, LCCSP, one or more, 16 June 68. 


Timochares ruptifasciatus (Plotz) 1884 


Recorded from the United States for the lst time by Skinner (1901); 
collected by Irvin Runyeon from southern California to the Arizona 
border. Its distribution in Texas seems to be restricted to Cameron and 
Hidalgo counties where it is well established. It has been collected each 
month except December and only a few times in January and February. 
Rearing has disclosed a larval diapause. The deciduous local larval 
foodplant Malpighia glabra L., MALPIGHIACEAE, is grown as an orna- 
mental in extreme southern Texas where its fruits are prized for making 
jellies. This explains why Freeman (1951) found the skipper mostly 
in city flower gardens. Comstock (1953) observed a 2 oviposit on a 
malpighiaceous vine in Mexico. He reared it through, described and 
illustrated the egg, last instar larva and pupa. He was unable to identify 
the larval foodplant except as to family. 


Rearing records. At Brownsville, 10 Nov. 69, Kendall found a single Ist 
instar larva on juvenile leaves of Malpighia glabra. It had made a rudimentary 
shelter by cutting and folding over a portion of the tender new leaf. In the lab 
the larva continued to feed on this plant and pupated 19 Dec. 69; a 2 emerged 
11 Jan. 70. Rearing was conducted at laboratory temperatures ranging from 50°— 
75° F, a comparable temperature range for outdoor temperatures in Brownsville 


108 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


at that time of year. Humidity, however, was not comparable. Two more larvae © 
were collected on this plant 15 Oct. 71 in SANWR. These were reared in a semi- 
outdoor environment. They pupated on 7 and 12 Jan. 72, and 2 9 emerged, one 
on 29 Jan., the other 7 Feb. 72. Kendall revisited SANWR 12 Jan. 72 and found 1 
last instar larva; it was preserved. On 8 Sept. 72 he collected a @ at Mission, 
Hidalgo Co., confined it with M. glabra and obtained 7 eggs, 4 of which were 
preserved. The remaining eggs hatched 12 Sept. 72. Larvae were placed on a 
living plant in the lab garden. Only 1 larva could be found when examined 23 
Oct. 72; it was brought into the lab where it died 22 Nov. 72. Again 20 Nov. 73, 
3 mi. E of Harlingen, Cameron Co., Kendall collected 2 larvae eating M. glabra. 
These were sleeved on a living plant in the lab garden on 1 Dec. 73. Upon return- 
ing from an extended field trip 5 March 74, examination showed one larva had 
died, the other had just pupated; a ¢ emerged 18 March 74. Rickard collected a 
pupa 22 March 70 on M. glabra in B-RGVSP; a 2 emerged 30 March 70. 

Other Texas records. CAMERON Co.: H. A. Freeman (1951), Brownsville, 1 3, 
1 9, 29 Aug. 44. J. W. Tilden (1974), Brownsville, 1 ¢, 19 Oct. 72. Rickard, 
Brownsville, 2 6, 14 Aug. 69; 1 ¢, 18 Oct. 69; 1 2, 19 Oct. 69; 1 ¢, 21 March 
70; 1 6, 23 March 70; 1 6, 4 April 70; 2 9, 5 April 70; 1 Oli Amen ier 
19 Oct. 71. Roy O. & C. A. Kendall, Brownsville, 1 ¢, 19 Oct. 72; 1 9, 22 Oct. 
72. Hmatco Co.: H. A. Freeman (1951), Pharr, 1 9, 30 Aug. 44; 3 ¢, 21 Oct. 
44; 1 9, 25 Aug. 46. Rickard, B-RGVSP, 1 9, 24 Aug. 69; 1 6, 6 Sept. 69; Madero, 
1 g, 19 Oct. 69; SANWR, 1 9, 17 May 70; 1 6, 15 Aug. 71. W. W. McGuire, 
Mission, several, 3 & 4 Sept. 72. Roy O. & C. A. Kendall, Mission, 6 6, 2 9, 9 
Sept. 72; 4 6, 1 2, 10 Sept. 72; 6 6, 1 @, Il Sept. 72:53 G2 wSeoman 


Xenophanes trixus (Stoll) 1784 


Barnes & McDunnough (1913) published the Ist record for the 
United States; 2 6, 2 ?, May and June from Brownsville, Texas; year 
and collector not given. Freeman (1951) gives an uncited Skinner 
record for July in Brownsville. So far as we can determine it was not 
collected again in Texas until 1968 which tends to indicate a periodic 
residency in the state. Local collection data indicates 3 and possibly 
4 broods. Larvae probably feed through the winter months when 
temperatures are mild; a local freeze may destroy the temporary 
population. 


Rearing records. Near Southmost, SE of Brownsville, Kendall observed a @ 
demonstrating oviposition behavior around Malvaviscus drummondii Torr., MAt- 
VACEAE, 9 Nov. 69. This specimen avoided capture but another 9 was taken and 
kept alive for egg production. Confined with M. drummondii twigs it deposited 
30 eggs the following day and died 11 Nov. Young larvae soon ate holes through 
the micropile but died before escaping from the egg shell. Also on 9 Nov., 1 
3rd instar larva was found hiding in a very rudimentary shelter on M. drummondii; 
it continued to feed sparingly in the lab until 24 Jan. 70 when illness was vias: 
it died 26 Jan. and was preserved. 

Unaware of Kendall’s findings, Rickard observed and captured, 28 Feb. 70, a 
? ovipositing on juvenile leaves of M. drummondii at the same Southmost sear 
He also found one late instar larva on the plant in a poorly constructed leaf nest 
same as Kendall had observed. This larva molted 2 March and pupated 12 March. 
An adult emerged 30 March but escaped in his home and was never found. 

Rickard gave the live @ and egg collected 28 Feb. to Kendall who obtained 


VoLuME 30, NuMBER 2 109 


additional eggs: 1 March (27), 3 March (5), 4 March (1), 5 March (25). The 
2 was killed 8 March and kept as a cabinet specimen. Eggs were placed in an 
unheated room. By 22 March the embryo had completely developed and some 
larvae had eaten through the micropile, but all died before completing the hatching 
process. In a separate effort, 21 March 70, Rickard experienced the same dis- 
appointment. Low humidity is now suspect as the cause of death. 

Other Texas records. CAMERON Co.: J. R. Heitzman, nr. Brownsville, 2 4, 
21 June 69; 1 ¢, 24 June 69; 1 ¢, 25 June 69; 1 ¢, 1 9, 26 June 69. Roy O. & 
C. A. Kendall, Brownsville, 1 ¢, 7 Nov. 69; 3 6, 8 Nov. 69; 1 9, 10 Nov. 69; 
nr. Southmost, 17 ¢,2 9, 9 Nov. 69; 13 ¢, 3 9, 10 Nov. 69. M. A. & S. Rickard, 
Brownsville & vicinity, 1969: 1 $,2 9, 27 July; 1 6, 17 Aug.; 1 6, 18 Aug.; 1 2, 
IeCre oes 296 7 Sept; 10 4,4 9, 18 Oct; 2 3, 1 2, 19'Oct; 4 4, dD. 9, 25 Oct.; 
3 4,26 Oct; 9 4,8 2, 8 Nov. 2 8,1 2,9 Nov.; 2 4, 13 Dec. 1970: 4 3,2 2, 28 
Heb46 4 2, 2 March; 2 6,2 9, 4 April; 1 9, 5 April: 5 6, 8 9, 24 May; 
cpm aueelade llsept, WOnl: 2)°65)2) O19 May-"5. d2 20 May:\3 6; 6 July; 
common on 21 Aug., 18 and 19 Oct.; 2 ¢, 30 Oct. 


Cabares potrillo (Lucas) 1857 


First recorded for Texas and the United States by Weeks (1904); 
Brownsville, Texas, summer 1903, leg. [Wm. H.] Doll. This species is 
generally restricted to the lower Rio Grande Valley in Texas. It seems 
to be best established in Hidalgo Co. where its deciduous larval food- 
plant, Priva lappulacea (L.) Perse, VERBENACEAE, is most abundant. 
There appear to be 3 full broods with considerable overlapping in 
September, October and November. Based on limited rearing, it would 
seem the larvae feed through the winter months with growth and de- 
velopment greatly retarded. It has been collected each month except 
February and July. 

In Mexico, nr. Cd. Mante, Tamaulipas, Kendall observed several 2 ovi- 
positing on this same plant 20 Jan. 74. Examination disclosed first 
instar larvae on several small plants at that time. It will be noted that 
the time frames of these 2 locations are comparable even though one 
is neotropical, giving stability to residency of the species in Texas. 

Rearing records. Kendall collected a 9, 13 Oct. 68, at SANWR and kept it 
alive for eggs. Earlier, Rickard had informed Kendall of seeing a 92 oviposit on 
an unknown plant at this location. Based on his description, it was easy to locate 
the larval foodplant, P. lappulacea (det., Dr. Marshall C. Johnston, University of 
Texas, Austin). The @ deposited eggs on this plant on 14, 15 & 19 Oct. They 
were not inventoried but were estimated to be about 25, some of which were pre- 
served. Hatching started 17 Oct. Larvae were placed on a protected potted plant 
outdoors. Later, it was discovered a moth larva had eaten most of the foliage. Only 
2 Cabares potrillo larvae could be found 2 Nov.; they were glossy black and quite 
small; one soon died, the other pupated 25 Nov. and a Q emerged 27 Dec. 68. 

A return visit to SANWR 9 Nov. 68 produced 6 larvae on this plant. Larval 
shelters are constructed in such a way that they are very difficult to find and 


recognize. Insufficient foodplant was brought back to the lab and it became neces- 
sary to preserve 1 larva. The others pupated 25 Nov. (2), 26 Nov. (1), 1 Dee. 


110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(1), and 3 Dec. (1); the last one was injured and was preserved. Adults emerged: 
1 4, 30 Dec. 68; 2 9,7 Jan. 69; 1 3, 10 Jan. 69. 

Other Texas records. BEXAR Co.: Roy O. & C. A. Kendall, San Antonio, 1 4, 
4 April 59. Cameron Co.: Roy O. & C. A. Kendall, 10 mi. SE of Brownsville, 1 ¢, 
1 April 57; Santa Maria, 1 ¢, 14 Nov. 71. Tilden (1974), Brownsville, 1 2, 30 
Oct. 63; Santa Maria, 1 ¢, 18 Oct. 72 (doubtless there are other county records 
unknown to us at this time). Hmatco Co.: Freeman (1951) found it well estab- 
lished at Pharr and vicinity from 1946-1959; specimens were taken in Mar., June, 
Oct., and Nov. Joseph F. Doyle III], SANWR, 1 6, 21 Sept. 68; 3 ¢, 19 Oct. 68; 
1 g,1 92, 24 Nov. 68. J. R. Heitzman, SANWR, 1 9, 21 June 66; 1 ¢, 24 June 
66; several on 22, 24 & 25 June 68; B-RGVSP, several, 25 & 28 June 68. Roy O. 
& G. A. Kendall, SANWR, 1 6 22 June 68; 3 6; 1 9, 21 Oct) 66a Ocr 
68; 2 3,9 Nov. 68; 1 2, 11 Nov. 68; Mission, 1 ¢, 10 Oct. 72. M. A. & S. Rickard, 
1968: 2 @,2 9,5 Aug, 1 6, 6 Aug; 2 6, 21 Aug; 1 657 Seppe ieee 
21 Sept.; 2 6,1 9,5 Oct. 2 9,6 Oct; 1 6,1 9, 12 Oct; 1G) 13 Oca 
9 Nov.; 1 6, 10 Nov.; 1 6, 16 Nov.; 2 6, 24 Nov.; 1970; 1 GSS 0Wicrcae ia. 
DO Marche led Oli Mayan ween Ocha le 


ACKNOWLEDGMENTS 


We wish to thank the U. S. Department of the Interior, Fish & Wild- 
life Service, Bureau of Sport Fisheries & Wildlife, Albuquerque, New 
Mexico, and the Texas Parks and Wildlife Department, Interpretation 
& Exhibits Branch, Austin, Texas, for providing the necessary permits 
to conduct basic biological research on the Lepidoptera in the national 
refuges and state parks. We are especially grateful to the local person- 
nel of these Departments for their courtesy and assistance in conducting 
these continuing studies of Texas Lepidoptera. We are also grateful to 
those who furnished collection data for inclusion in this paper. 


LITERATURE CITED 


BarNES, WILLIAM, & J. McDuNNoucH 1913. Species of Lepidoptera new to our 
fauna with synonymical notes. Can. Ent. 45: 182-185. 

Comstock, J. A. 1953. Collecting butterflies in the coastal area of Mexico near 
Manzanillo, Colima, with notes on the life history of a rare skipper. Bull. S. 
Calif. Acad. Sci. 52: 43-45. 

pos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. 
Soc. Mem. 1, 145 p. 

FREEMAN, H. A. 1948. Notes on some North American skippers, with the descrip- 
tion of a new species from Kansas. Ent. News 59: 206. 

1951. Ecological and systematic study of the Hesperioidea of Texas (Lepi- 
doptera, Rhopalocera, Hesperioidea). Southern Methodist Univ. Stud. 6: 20. 

Goutp, F. W. 1969. Texas Plants—a checklist and ecological summary. Texas 
Agr. Exp. Sta., Texas A & M Univ. Misc. Publ. 585 rev. 

SKINNER, H. 1901. On a small collection of butterflies made in California and 
Arizona. Ent. News 12: 170-171. 

Titpen, J. W. 1974. Unusual and interesting butterfly records from Texas. J. Lepid. 
Soc. 28: 22-25. 

Weeks, A. C. 1904. Minutes of meetings of Brooklyn Ent. Soc. held at the resi- 


dence of Mr. George Franck, 1040 DeKalb Ave., Brooklyn, N. Y. Ent News 
1S: 35l. 


VoLUME 30, NUMBER 2 12 


COLLECTING COCOONS OF CALLOSAMIA SECURIFERA 
(SATURNIIDAE )* 


RICHARD S. PEIGLER 


Department of Entomology and Economic Zoology 
Clemson University, Clemson, South Carolina 29631 


Little was published on Callosamia securifera (Maassen) until Fergu- 
sons work in 1972. Skinner (1920) first synonymized the names C. 
securifera and C. carolina Jones, but the moth was long considered a 
subspecies of C. angulifera (Walker). Because the diurnal males are 
not phototaxic and the species inhabits remote coastal areas of south- 
eastern United States, specimens are still comparatively uncommon in 
collections. It is my purpose to provide some helpful advice for anyone 
wishing to collect cocoons of this moth. 


Choosing Correct Habitat 


I have searched successfully for cocoons of C. securifera in Florida, 
Georgia, South Carolina and North Carolina. However, most of what is 
said in this paper applies mainly to the habitats in South Carolina where 
I have collected most. 

The hostplant is sweetbay (Magnolia virginiana L.), but the tree 
ranges much farther north along the Atlantic Coast, west along the Gulf 
Coast and inland from both coasts than does the moth (Peigler, 1975). 
It is, therefore, advisable to go to or near localities were C. securifera is 
known to occur. Sweetbay may be easily confused with red bay (Persea 
borbonia (1L.) Sprengel), but the latter does not have white undersides 
of leaves which help to recognize sweetbay. 

There will be many areas near or between known localities of the 
moth which will appear suitable for searching because of the presence 
of the host tree, but may be unproductive. Expansive fields with stand- 
ing water throughout may have hundreds of sweetbays, but I have 
never found even an empty cocoon in such places. Even if C. securifera 
were present, it would be very hard to find cocoons unless the popula- 
tion was high because of the high density of host trees on which to look. 
Other habitats where I have not found cocoons are thin woods under 
towering pines and mixed hardwood forests. Perhaps females prefer 
to fly and oviposit in the open. The most rewarding habitats are along 


1 Published by permission of the Director of the South Carolina Agricultural Experiment Station. 
Technical Contribution No. 1250. 


112 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


roadsides in which the sweetbays are small and in the open and a lot 
of areas can be checked with an automobile. 


Choosing Correct Season 


In the fall or early winter searching may be very slow because many 
green leaves are still on the host trees. This situation exists until early 
December in the Carolinas and Georgia. On the other hand, even the 
tip leaves may be lost by March so that sweetbays are not readily seen. 
At such times I spend more time looking for the host than the cocoons 
themselves. Therefore, December and January are the best times to 
plan a collecting trip. 


Comments 


Dr. R. B. Dominick reports that he usually finds only one cocoon per 
tree (see Ferguson, 1972, p. 242). I would agree that this is true for 
the areas in South Carolina where he collects (northern Charleston 
Co. and eastern Berkeley Co.). However, in western Berkeley Co. where 
I collect most often, the population of the moth is probably much higher, 
and I seldom find isolated cocoons. The most viable number of cocoons 
I found on a single tree was six, and several times I found four and five. 
Often an empty female cocoon is found along with viable ones, indicat- 
ing that females often mate and begin ovipositing on the same tree on 
which they fed as larvae. Usually over 90 percent of the sweetbays 
examined will have no cocoons hanging on them. 

The populations may fluctuate or move. One winter I found 18 viable 
cocoons along a highway in western Berkeley Co. during three days 
of searching. The following winter I found 60 viable cocoons in four 
hours along the same stretch of road. This past winter the population 
was again high in that area. 

Cocoons may occur on a tree from 1-20 ft. above ground level, but 
most are about 5 ft. Most sweetbays in the areas where I have col- 
lected are secondary growth sprouts from cut trees, and if a cocoon 
is too high to reach, the thin tree may be easily bent over. Brown (1972 
and pers. comm.) searches tall sweetbays in central Florida with 
binoculars. 

It is important to train one’s eyes to look for hanging clumps of red- 
brown sweetbay leaves among green ones. The weight of a swinging 
cocoon detaches leaves from the stem so that cocoons seldom are 
covered with green, living leaves. The leaves on old cocoons of the 
previous season are bleached or have weathered away. A collector 


VoLUME 30, NUMBER 2 113 


looking for whitish cocoons will probably find more empty than viable 
ones. 

Few insects feed on sweetbay, but chewed leaves may not mean C. 
securifera is present. Grasshoppers or larvae of Papilio palamedes Drury 
may have done the damage. In southeastern North Carolina I have 
seen larvae of Epimecis sp. (Geometridae) feeding commonly on this 
tree in summer. Not all large cocoons on sweetbay may be C. securi- 
fera because larvae of the saturniid Antheraea polyphemus (Cramer ) 
sometimes move from adjacent red maples (Acer rubrum L.) to spin 
their cocoons on sweetbay. Often nests or egg sacs of large spiders, 
including the green lynx (Peucetia viridans (Hentz)) and a yellow 
species of Thomisidae, are attached with sweetbay leaves to stems. The 
tendrils of Smilax spp. vines commonly hold dead sweetbay leaves up 
in trees. The collector should examine possibilities closely so that a 
C. securifera cocoon is not overlooked. 


ACKNOWLEDGMENTS 


I wish to thank Dr. Richard B. Dominick and Mr. Charles R. Edwards 
for initial help in searching for cocoons of this moth. Drs. Merle 
Shepard and G. R. Carner of Clemson University helped with the 


manuscript. 


LITERATURE CITED 


Brown, L. N. 1972. Mating behavior and life habits of the sweet-bay silk moth 
(Callosamia carolina). Science 176: 73-75. 

Fercuson, D. C. 1972. Bombycoidea, Saturniidae (in part). In R. B. Dominick 
et al., The Moths of America North of Mexico, fasc. 20.2B, p. 155-269, pls. 
12-22. E. W. Classey Ltd. London. 

PeicLerR, R. S. 1975. The geographical distribution of Callosamia securifera 
(Saturniidae). J. Lepid. Soc. 29: 188-191. 

SKINNER, H. 1920. Callosamia carolina and Samia securifera (Lepid., Saturniidae ). 
Ent. News 31: 107. 


A GYANDROMORPHIC LYMANTRIA DISPAR (LYMANTRIIDAE ) 


It should not be too surprising to find a gyandromorphic Gypsy moth ( Lymantria 
dispar (L.)) among the hundred thousands that have been defoliating trees in 
New Jersey the last few years. A single gyandromorphic, adult female was caught 
6 July 1974 in Glendola, Monmouth Co., New Jersey in a light trap put up by the 
College of Agriculture, Rutgers University. The left antenna is female and the 
right, male. It is otherwise a normal specimen without any other deviations showing. 
The specimen is in the Rutgers Collection. Dr. J. P. Reed kindly loaned it to me 
for study. 


Jos—ErpH MuLLeErR, R.D. #1, Lebanon, New Jersey 08833. 


114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


WING COLOR VARIATION IN CALLOSAMIA (SATURNIIDAE)? 


RICHARD S. PEIGLER 


Department of Entomology and Economic Zoology 
Clemson University, Clemson, South Carolina 29631 


The genus Callosamia Packard is comprised of three sexually dimorphic 
species. Callosamia promethea (Drury) is melanic with males almost 
black and females a wine-red color. Callosamia securifera (Maassen) 
and C. angulifera (Walker) have brown males and orange females 
although the former species is lighter and suffused with yellow. Adults 
of the latter two species also differ seasonally in that the spring or over- 
wintering forms of both sexes are light and the summer forms are dark. 
Such seasonal variation has been oversimplified in past literature. It 
is the purpose of this paper to discuss in greater detail these wing color 
variations. 

Although C. promethea does not have seasonal forms as in the other 
two species, there is color variation. In males the area beyond the post- 
median line on the underside of the wings may be either bright red or 
blue-gray. Both color forms, and intermediates, were found among 
males collected in spring and summer from southeastern Pennsylvania. 

Orange females of C. promethea may occur among populations with 
normal dark red females. Young (1968) and Baldy (1890) reported 
such populations occurring in Ulster Co., New York and on Mt. Cata- 
wissa, Pa., respectively. The possibility of hybridization is unlikely 
because of effective temporal isolation. Orange C. promethea females 
are possibly mutations in which the gene (or genes) governing reddish- 
ness is not expressed. 

The pair of C. angulifera which Ferguson (1972, pl. 22, fig. 3, 4) uses 
to represent summer adults are scarcely darker than spring ones. In 
upper South Carolina the summer adults of both sexes are nearly as 
dark as C. promethea and the males have obsolete discal marks as in 
C. promethea. The winter of 1973-74 was very mild in South Carolina, 
and the adults of that spring varied through the entire spectrum from 
light to dark. This included specimens from wild and reared cocoons 
(kept outdoors all winter) as well as those coming to lights. Apparently, 
adults are darker when wing scale formation in the pupa occurs under 
comparatively high temperatures. In fact, C. angulifera cocoons which 
[ overwintered in the refrigerator (40°F =5°C) yielded dark adults 


' Published by permission of the Director o 


oe f the South Caroli i i i 
Technical Contribution No. 1259. yu arolina Agricultural Experiment Station. 


VoLUME 30, NUMBER 2 ~ WS 


that match the summer form. The same was true for adult hybrids of 
angulifera ¢ X promethea 2, although some of their darkness may be 
attributed to C. promethea parentage. Cocoons of the saturniid Actias 
luna (L.) which I kept at no lower than 40°F during the winter also 
produced adults similar to the summer form of that species. 

The effect of the mild winter of 1973-74 was further illustrated in 
the C. securifera population in Berkeley Co., South Carolina. Spring 
adults began flying in mid-April instead of late April, as they usually 
do, and most were dark. Although a few golden spring males and black- 
ish summer males emerged from a lot of about 20 cocoons collected in 
March 1974, most did not fit either form and were intermediate. 

Ferguson (1972) stated that females of C. securifera from Florida are 
paler than those collected from further north. However, specimens in 
my collection from Florida and both Carolinas do not follow this pattern. 
Possibly the Florida material available to Ferguson was faded. 

In conclusion, seasonal variation in Callosamia is due apparently to 
effects of temperature on pupae and is not clear-cut without inter- 
mediates. There is no constant geographical variation in any of the 
three species, although seasonal extremes in C. angulifera may be greater 
in the South. 


ACKNOWLEDGMENTS 


I wish to thank Drs. John Morse and G. R. Carner for reviewing the 
manuscript and offering helpful suggestions. 


LITERATURE CITED 


Baupy, S. 1890. [Untitled note.] Ent. News 1: 146. 

Fercuson, D. C. 1972. Bombycoidea, Saturniidae (in part). In R. B. Dominick 
et al., The Moths of America North of Mexico, fasc. 20.2B, p. 155-269, pls. 
12-22. E. W. Classey Ltd. London. 

Younc, A. M. 1968. Color variant of Callosamia promethea (Saturniidae) in New 
York. J. Lepid. Soc. 22: 185-186. 


116 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


THE GENUS COPABLEPHARON IN TEXAS, WITH DESCRIPTION 
OF THREE NEW SPECIES (NOCTUIDAE) 


ANDRE BLANCHARD 
P. O. Box 20304, Houston, Texas 77025 


Four species of the genus Copablepharon, still unrecorded from Texas, 
fly in this state. Three of these four species are new and are described 
in this paper. 


Copablepharon grandis Strecker 1875 
Fig. 1 


Lep. Rhop. Het., 129. 


My records for this species are as follows: Paducah, State owned Matador Wild- 
life Management Area, Cottle Co., Texas: 8 Sept. 1966, 4 ¢; 11 and 13 July 1967, 
3 6,3 9; 22 Sept. 1968, 1 ¢. These were identified by comparison with a pair 
of specimens in the Canadian National Collection. There are also, in the collection 
of the American Museum of Natural History, 1 ¢ and 1 @ collected at El Paso, El 
Paso Co., Texas by A. H. & S. K. Rindge. 

Fig. 1 unfortunately fails to show the definite contrast between the pale sulfur 
yellow of the forewings and the white of the hindwings. 


Copablepharon albisericea A. Blanchard, new species 
Figs) 2/05, 9013014 


Head, collar, thorax, tegulae, forewings above, hindwings above and _ beneath, 
legs, abdomen shiny silky white (Fig. 2). Forewings beneath more or less soiled 
with olivaceous in their basal halves between radius and fold. Male antennae 
weakly serrate, with hair length about half of shaft diameter. Female antennae 
simple, pubescent. Antennae of both sexes clothed with white scales above, orange 
below. Wing expanse = 40-43 mm. 

Male genitalia (Fig. 5): Uncus long, cygnate, not dilated in its middle; sides 
of tegumen broad, penicillus lobes well developed; vinculum narrow with rounded 
margin; valves long, slender, narrowest two-thirds distance from base where ventral 
margin and costa are slightly concave, clasper with broad base, tapering to a point; 
corona of 17-20 spines aligned along distal margin of valve, except near tip where 
3-5 get crowded together; clavi slender, digitate, slightly dilated distally; juxta 
broad, lightly sclerotized; sclerotization of lateral and dorsal parts of annelus at least 
equal to that of juxta; aedeagus stout, one small conical cornutus located at apex of 
small diverticulum of vesica (Fig. 9). 

Female genitalia (Figs. 13 & 14): Ovipositor cone shaped, lobes clothed near 
their apex with short, stout setae; anterior apophyses a little longer than sclerotized 
part of eighth segment collar; posterior apophyses almost twice as long as this 
collar; ductus bursae short, weakly sclerotized; bursa membranous, with an expanded 
lateral arm on the right side about half as long as left arm; ductus seminalis from 
near base of right lateral arm. 

Holotype: Male, Canadian, State owned Gene Howe Wildlife Management Area, 
Hemphill Co., Texas, 27 Sept. 1968 collected by A. & M. E. Blanchard, deposited 
in the National Museum of Natural History (No. 73431 ) 


VoLUME 30, NUMBER 2 117 


Figs. 1-8. Copablepharon spp.: 1, C. grandis 9, Paducah, Texas, 11 July 1967: 
2, C. albisericea ¢, holotype, Canadian, Texas, 27 Sept. 1968; 3, C. serraticornis &, 
holotype, Paducah, Texas, 8 Sept. 1966; 4, C. gillaspyi &, holotype, Padre Island 
National Seashore, 28 Sept. 1973. Figs. 5-7: ¢ genitalia, aedeagus omitted: 5, 
C. albisericea (A. B. 3555); 6, C. serraticornis (Slide COPA 6 in C. N. C.); 7, 
C. gillaspyi (A. B. 3612). Fig. 8: C. gillaspyi aedeagus with incompletely inflated 
vesica (A. B. 3613). 


148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 9-16. Copablepharon spp.: Figs. 9 & 10: Aedeagus with inflated vesica: 
9, C. albisericea (Slide COPA 5 in C. N. C.); 10, C. serraticornis (Slide COPA 6 
in C, N. C.), Figs. 11 & 12: Distal part of valve showing corona: 11, C. serrati- 
cornis (Slide COPA 6 in C. N. C.); 12, C. gillaspyi (A. B. 3612). Figs. 13-16: 
Female genitalia: 13, C. albisericea (A. B. 3252); 14, C. albisericea CAS Br samen: 
15, C. serraticornis (A. B. 3552); 16, C. gillaspyi (A. B. 3554). 


VoLuME 30, NuMBER 2 119 


Paratypes: All from Canadian, Hemphill Co., Texas: 27-30 Sept. 1968, 17 4, 
3 2, collected by A. & M. E. Blanchard, 8 of these ¢ are in the American Museum 
of Natural History; 27 & 28 Sept. 1968, 37 ¢ collected by D. F. Hardwick, in the 
Canadian National Collection. 


C. albisericea is readily distinguished from C. alba, the other white 
species of this genus: C. alba is the only described species of the genus 
having a very short uncus. 


Copablepharon serraticornis A. Blanchard, new species 
Figs. 3, 6, 10, 11, 15 


Head, collar, thorax, tegulae pale ochreous yellow, darker on vertex, collar and 
anterior thorax; paler on front, and metathorax (Fig. 3). Male antennae strongly 
serrate, with each segmental process developed into a thin edge extending laterally 
and supporting numerous bristles a little longer than shaft diameter. Widest por- 
tion of each serration about 2.5 times as wide as narrowest portion. Female antennae 
simple, very weakly pubescent. Antennae of both sexes clothed above with scales 
concolorous with front, beneath orange. Forewings above concolorous with thorax; 
a pale, faint grayish streak along cubitus; fringes concolorous. Hindwings above 
varying from same hue as forewings but much paler to pale buff; fringes slightly 
paler than disc. Beneath same hue as above but paler. Wing expanse: Male 36-40 
mm; female 40—45 mm. 

Male genitalia (Figs. 6, 10, 11): Very close to those of C. albisericea, differing 
from them as follows: Uncus very slightly dilated in its middle; clavi a little 
stouter and shorter; base of clasper narrower; ventral margin of valve convex all 
along; corona (Fig. 11) with 11-13 spines in a line which diverges from distal 
margin of valve as it approaches ventral margin. Vesica (Fig. 10) smaller. 

Female genitalia (Fig. 15): Similar to those of C. albisericea, differing as 
follows: Setae on ovipositor lobes stouter and longer; eighth segment collar a little 
longer; anterior apophyses a little longer than eighth segment collar; posterior 
apophyses over one and a half times as long as that collar. 

Holotype: Male, Paducah, State owned Matador Wildlife Management Area, 
Cottle Co., Texas, 8 Sept. 1966 collected by A. & M. E. Blanchard; genitalia on 
slide A. B. 505; deposited in National Museum of Natural History (No. 73432). 

Paratypes: All taken at Paducah, Cottle Co., Texas: 8 Sept. 1966, 1 2; 22 Sept. 
1968, 1 ¢, 4 @ collected by A. & M. E. Blanchard; 23 and 24 Sept. 1968, 3 é 
collected by D. F. Hardwick, in Canadian National Collection. 


Copablepharon gillaspyi A. Blanchard, new species 
Figs. 4, 7, 8, 12, 16 


Head, collar, thorax, tegulae beige (light grayish, yellowish brown) (Fig. 4). 
Male and female antennae whitish above and structurally similar to those of C. 
serraticornis. Forewings above pruinose beige with a paler whitish, poorly defined 
fascia along costa extending almost to apex and a grayish streak along cubitus, fringes 
whitish. Hindwings above concolorous with costal fascia of forewings, paler basally: 
fringes whitish. Beneath, same color as above attenuated to nearly white. Wing 
expanse = 39-44 mm. 

Male genitalia (Figs. 7, 8, 12): Similar to those of C. serraticornis, differing 
mainly in that the clasper is wider at its base and that the corona (Fig. 12) has 
over 30 spines crowded in 2—4 rows along distal margin of valve. Aedeagus similar 
to that of C. serraticornis (Fig. 8). 

Female genitalia (Fig. 16): Similar to those of C. serraticornis, differing mainly in 


120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


that the setae on the ovipositor lobes are smaller and cover only the tip of the lobes 
and that the anterior apophyses are only three fourths as long as the eighth segment 
collar and the posterior apophyses barely longer than this collar. 

Holotype: Male, Padre Island National Seashore, Kleberg Co., Texas, 28 Sept. 
1973, genitalia on slide A. B. 3245, Dr. J. E. Gillaspy collector; deposited in the 
National Museum of Natural History (No. 73433). 

Paratypes: Padre Island National Seashore, Kleberg Co., Texas, 28 Sept. 1973, 
2 4; 7 Oct. 1974, 3 6,1 9; J. E. Gillaspy collector. Same location, 29 Sept. 1975, 
1 6; North Padre Island, Nueces Co., Texas, 30 Sept. 1975, 1 ¢@; A. & M. E. 
Blanchard collectors. 


I take pleasure in naming this species for its discoverer, my friend, 
Dr. J. E. Gillaspy, Professor of Biology at A. & I. College, Kingsville, 
Texas. 

C. serraticornis and C. gillaspyi are both closely related to C. serrata 
McDunnough, but the antennae of C. serrata are not as strongly serrated, 
their serrations are scarcely more than sharp bulges to the sides; the 
widest portion of a segment is usually about 1.5 times as wide as the 
narrow portion, the male valves of C. serrata are angled near the apex 
as those of C. albisericea and not evenly convex as those of the other 
two species. 


ACKNOWLEDGMENTS 


I am deeply grateful to Mr. J. D. LaFontaine of the Biosystematics 
Research Institute, Agriculture Canada, for providing me with all the in- 
formation I needed about all the other previously described taxa in this 
genus, for many invaluable suggestions and for revising the manuscript. 


VoLUME 30, NUMBER 2 121 


NATHALIS [OLE (PIERIDAE) IN THE SOUTHEASTERN 
UNITED STATES AND THE BAHAMAS 


Harry K. CLENCH 
Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15213 


In the Lepidoptera of Florida Kimball (1965: 36) devotes most of 
his brief discussion of Nathalis iole Boisduval 1836 to an unusual sub- 
ject: the absence of early records of it from the state. He concludes, 
“Perhaps the species began to establish itself during the late 1920's,” 
and adds that a search should be made in collections for pre-1930 
records of iole from Florida. 

Kimball based his contention chiefly on two points. First, Grossbeck 
(1917, see below) cited only a single Florida record of iole. Second, the 
late W. T. M. Forbes examined the collection of Harvard’s Museum of 
Comparative Zoology and reported (in litt. to Kimball) that it con- 
tained no iole earlier than a 1924 Coconut Grove specimen that “looks 
suspicious and an authentic series dated 1933-1934. 

A survey of the Carnegie Museum Collection, which contains much 
old Florida material, shows that our oldest Florida specimens of iole 
are: a male from Dunedin [Pinellas Co.], 3 April 1921, leg. W. S. 
Blatchley, and two from Miami [Dade Co.], October 1921, leg. J. Harold 
Matteson. It is noteworthy that the W. H. Edwards Collection has none, 
although he received many butterflies from Dr. William Wittfeld, who 
lived and collected at Georgiana [Brevard Co.], on the Indian River, 
in the 1880's; nor are there any taken by G. Krautwurm, who collected 
extensively for the museum at Lutz [Hillsborough Co.] and nearby 
Stemper, notably in 1911. 

In the American Museum of Natural History are the following pre- 
1930 records of iole: a male from Jupiter [Palm Beach Co.], 1-24 
February 1920, leg. W. C. Wood; a male from Ft. Myers [Lee Co.], 22 
April 1922, leg. F. M. Jones; two females, Punta Gorda [Charlotte Co.], 
18-19 April 1922; and a female from Homestead [Dade Co.], 18 April 
1923: 

The older literature provides further interesting information. Scudder 
(1889: 1842) says of iole, “. . . not known in even southernmost Florida”; 
and farther on, “. . . does not appear to occur in the United States any- 
where east of Louisiana, although it is found in Cuba and Jamaica.” 
Skinner (1898: 63) gives the distribution of iole as “Ill., Mo. to Cala., 
N. Mex., Ariz., Tex., Mex. ...” and his supplement ( [1905]: 22) adds no 
further distributional data. Dyar (1901: 449) makes no mention of iole 


122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in a list of winter (January and February) Lepidoptera taken near 
Lake Worth [Palm Beach Co.] in 1890 and 1900. 

The earliest notice of Florida iole that I can find is the one Kimball 
mentions, Grossbeck (1917: 9), who gives a single record, “Big Pine 
Key, Sept. 20 (Ds.).” This specimen was taken by William T. Davis— 
“Ds.’—in 1913, and is conserved in the Staten Island Institute. 

Published evidence and that in available collections, therefore, sup- 
port Kimball’s basic contention completely. The fairly extensive informa- 
tion is all consistent with an hypothesis that Nathalis iole was formerly 
absent from Florida, and that it invaded the southern part of the state, 
probably from Cuba, sometime before 1913. (Davis’s capture in the 
lower Florida keys could have been within a year or two of its first 
landfall and close to the original site.) It established itself and gradually 
extended its range northward. By 1920 it had reached Jupiter on the 
east coast, by 1921 the Tampa area (Dunedin) on the west coast, and 
in 1931 it was found in Atlanta, Georgia (Harris [1950]; 1972), about 
its present northern limit. If these dates are used, iole spread northward 
at an average rate of about 42 miles per year, possibly somewhat slower 
at first, faster later on. An important point (commented on by Klots, 
1951) is that iole in the eastern part of its range is confined to the 
southernmost states (central Georgia southward). In the Mississippi 
Basin (see below) it ranges far to the north of this latitude. 

Nathalis iole may also be a relative newcomer in the Bahamas. The 
species is now known from four islands, all in the northern part of the 
archipelago: Grand Bahama (Rindge, 1955); New Providence (West, 
1966); North Bimini (Rindge, 1952) and North Andros (Nicolls Town, 
June 1973, leg. Clench; unpublished ). 

In the present connection New Providence is the most instructive 
because it is by far the most thoroughly collected island in the Bahamas, 
and collections from there date back to the 1880’s at least. Charles J. 
Maynard made a large collection there in June 1897, conserved in the 
Museum of Compartive Zoology. Nathalis iole is not represented. 
J. L. Bonhote collected extensively on New Providence in 1898; his 
specimens are now in the British Museum. Sharpe (1900) published an 
account of them, but N. iole is not mentioned. W. W. Worthington col- 
lected a large lot of Bahama butterflies for Carnegie Museum in 1909- 
1910. He visited many islands and was on New Providence in January 
1909. He took no Nathalis iole. The American Museum Collection 
contains New Providence material taken by various collectors in 1912, 
1915, 1929 and 1930 (reported in Rindge, 1952). No Nathalis iole is 
represented in this material, but these collections appear not to have 


VoLUME 30, NuMBER 2 12s} 


been extensive so no particular significance can be attached to the 
absence of iole from them. The first known captures of iole on New 
Providence were the specimens taken by West in 1945, which he re- 
ported in 1966. Interestingly, iole is also represented in a small collection 
made on New Providence for Carnegie Museum in 1946. 

In sum, N. iole was probably absent from New Providence in 1897 
and 1898 and could have been absent as late as 1930; but it was present 
in 1945. 

Nearly all species of Bahama butterflies have reached the islands 
from Hispaniola, Cuba or Florida. For the most part, each of these 
source areas generates a different, characteristic, pattern of distribution 
within the Bahamas, and most species from that source tend to follow 
the pattern. Unfortunately, the known distribution of iole is ambiguous. 
It is restricted to the northern islands, a pattern characteristic of in- 
vaders from Florida; but it also includes Andros. Invasion from Cuba 
to Andros to New Providence and other northern islands is both reason- 
able and in part substantiated by the ranges of other species. 

At present all we can say about the arrival of Nathalis iole in the 
Bahamas is that it probably reached the islands sometime between 
1898 and 1945. The chances are that it came in the period 1920-1940, 
but this is by no means certain. It may have arrived from Florida, 
where iole was by then established, or it could have come in directly 
from Cuba. 

Now back to the mainland. Nathalis iole is divisible into three dif- 
ferent geographical components which, for want of a better term, may 
be called segregates. At present, the chief discriminating attribute of 
these segregates is the northward limit of their ranges. This seems a 
slender thread on which to hang such a speculation, but two further 
points prompt me to suggest it: (1) the varied northward limits of 
their distributions suggest physiological differences in their tolerance 
to cold and a possible further difference in their migratory tendencies; 
and (2) each of the three segregates is associated with a different 
southern part of the species range. The three segregates are as follows: 

1. Southeastern. In the United States this segregate is confined to 
Florida and southern Georgia, with no evidence of repetitive northward 
migration. It is probably not cold tolerant. As detailed above, this 
segregate is a probable recent arrival in the United States from Cuba. 
The southern limits of its range include the northern Bahamas (where 
it is also a probable recent arrival), Cuba, Jamaica and possibly His- 
paniola (Brown & Heineman, 1972: 62 [“?” in table]). 

2. Central. In America north of Mexico this segregate extends from 


124 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Mississippi to Arizona, and far northward to Ontario, Manitoba, North 
Dakota, southeastern Wyoming, nearly all of Utah, and perhaps as far 
west as the eastern slopes of the Sierra Nevada in California (Warren 
Creek, 9000-9500 ft., ca. 5 mi. W Lee Vining, Mono Co.: Garth & Tilden, 
1963: 88). Whether it is resident northward or occurs there only as a 
result of repeated immigration is not known, but most authors believe 
that the latter is the correct explanation. This segregate can tolerate 
temperatures below freezing and has been found flying at Colorado 
Springs as late as early December (Brown et al., 1956: 200; Brown, in 
litt.). Regardless of the nature of its northward occurrence, it differs 
conspicuously from the Southeastern segregate in being their at all. The 
southern limits of the range of this segregate include Guatemala (1 ¢, 
Carnegie Museum) and Mexico, where it is resident to altitudes of at 
least 7000 ft. (2150 m). The range appears to be old and probably has 
not changed significantly in historic times. Early authors did not men- 
tion it as occurring much north of the latitude of Illinois, but areas north 
of there were then poorly collected and in these areas iole is sporadic 
and generally uncommon. 

3. Pacific. In the United States this segregate is confined to southern 
California (Emmel & Emmel, 1973: 22). The northernmost record that 
has come to my attention is Kennedy Meadows, Tulare Co. (anonymous 
reviewer), unless the Warren Creek record mentioned above should 
belong here. The occurrence of this segregate in California is spotty 
but it is locally common (as is true, inter alia, in Florida and the 
Bahamas), but nothing in its known range suggests that it is cold 
tolerant. Emmel & Emmel (loc. cit.) mention it as being found “occa- 
sionally . . . [at] high elevations in the mountains, especially in late 
summer, so this segregate may have a limited tendency to migrate. 
Southward the range includes the whole of Baja California. 

In summary, then, the Southeastern segregate does not range north- 
ward, is non-migratory and its southern limits are in the West Indies; 
the Central segregate ranges far northward, is probably migratory and 
its southern limits are in mainland Mexico and Guatemala; and the 
Pacific segregate does not range far northward, may be slightly migratory 
and its southern limit is in the peninsula of Baja California. 

At present the United States portions of the ranges of the three 
segregates appear to be slightly disjunct. Whether or not iole occurs 


in Alabama (where the Southeastern and Central segregates might meet) 
or in western Arizona (where the Central and Pacific segregates could 
be in contact) is of little moment in establishing the thesis of three 


segregates. 


VoLuME 30, NUMBER 2 125 


It would be satisfying to be able to delineate their common boundaries 
(if they are indeed in contact) more precisely, but even with records 
from the intervening areas is would be no easy task. Despite careful 
comparison, the only difference among these segregates that I have 
been able to find in museum specimens is that in the Central segregate 
the hind wing upperside of the winter form female is usually yellow, 
less often pale orange; whereas at least in the Southeastern segregate, 
females of the winter form are as strongly orange as those of the sum- 
mer form. In material available to me, adequate dated specimens of 
the winter forms of all three segregates are few, however, and even 
this difference is not certain. 

Curiously, both the slight facies difference and the possible physio- 
logical differences seem to concern only the Central segregate. No 
evidence as yet would indicate any real intrinsic difference between the 
Southeastern and the Pacific segregates, even though they are widely 
separated geographically and the Central segregate intervenes. 


ACKNOWLEDGMENTS 


I am grateful to the following for their help in providing information 
on specimens in their care: Dr. Frederick H. Rindge, American Museum 
of Natural History, New York; Dr. George O. Pratt, Jr., director, and 
Mr. Joseph Burke, Staten Island Institute of Arts and Sciences, Staten 
Island, New York. I also thank an anonymous reviewer of an earlier 
version of this paper. He provided a northern locality record of iole 
in California and clarified the probable status of the Warren Creek 
record. 


LITERATURE CITED 


Brown, F. M., D. Err & B. Rotcer. 1956 (1954-1956). Colorado butterflies. 
Proc. Denver Mus. Nat. Hist., nos. 3-7: viii + 368 p., ill. 

Brown, F. M. & B. HEINEMAN. 1972. Jamaica and Its Butterflies. London (Clas- 
sey): xv + 478 p., ill. 

Dyar, H. G. 1901. Notes on the winter Lepidoptera of Lake Worth, Florida. 
Proc. Ent. Soc. Washington 4: 446-485. 

EMMEL, T. C. & J. F. Emmet. 1973. The butterflies of southern California. Nat. 
Hist. Mus. Los Angeles Co., Science Ser. 26: xi + 148 p., ill. 

GartH, J. S. & J. W. TiLpEN. 1963. Yosemite butterflies. J. Res. Lepid. 2: 1-95, 
ill. 

GrossBECck, J. A. 1917. Insects of Florida. IV. Lepidoptera. Bull. Amer. Mus. Nat. 
Hist. 37: 1-147. 

Harris, L., Jr. [1950]. The butterflies of Georgia. Revised. Georgia Soc. Nat., 
Bull. 5 (mimeographed): [3] + vii + 29 + [4] p. 

. 1972. Butterflies of Georgia. Norman, Oklahoma (Univ. Oklahoma 
Press): xvi + [6] + 326 p., ill. 

Kmsa.L, C. P. 1965. The Lepidoptera of Florida, an annotated checklist. Gaines- 
ville (Florida Dept. Agr.): v + 363 p., ill. 


126 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Krots, A. B. 1951. A field guide to the butterflies. Boston (Houghton Mifflin): 
xvi + 349 p., ill. 

Rinvcr, F. H. 1952. The butterflies of the Bahama Islands. American Mus. Novit., 
sailajeev dls) joy 

_ 1955. The butterflies of the Van Voast—American Museum of Natural 
History expedition to the Bahama Islands, British West Indies. American Mus. 
Novits, 4 1715-220 po. allt 

Scupper, S. H. 1889. The butterflies of the eastern United States and Canada 
with special reference to New England. Cambridge, Mass. (author): xxiv + 
p. 1-766 (vol. 1); xi+ p. 767-1774 (vol. II); vii+p. 1775-1958 + 89 pls. 
maps (vol. III). 

SHARPE, E. M. 1900. Ona collection of butterflies of the Bahamas. Proc. Zool. Soc. 
London 1900: 197-203, ill. 

SKINNER, H. 1898. A synonymic catalogue of the North American Rhopalocera. 
Philadelphia (American Ent. Soc.): xvi + 100 + xiv p. 

. 1905. Same, supplement no. 1. [i] + 34 p. 

West, B. K. 1966. Butterflies of New Providence Island, Bahamas. Ent. Rec. 78: 
174-179, 206-210. 


MASS HIBERNATION SITE FOR NYMPHALIS VAU-ALBUM 
(NYMPHALIDAE ) 


Hibernation is cominon in several genera of butterflies and especially among 
Nymphalidae. However, in most cases the literature on such hibernating is rather 
nebulous and repetitive, without specific observational support. One takes for 
granted that hibernation spots are found beneath debris, loose bark, in hollow 
trees, etc. Therefore I was interested to see fourteen Compton tortoise shells 
(Nymphalis vau-album (Denis & Schiffermiller)) actually entering a hibernation 
site. 

The observations took place near a radio relay tower at the summit of Mohawk 
Mountain, Mohawk State Forest, near Cornwall, Connecticut on 15 October 1974 
between 1300-1400 hrs. EST. During this still, warm period my attention was 
drawn to several butterflies fluttering about near the top of the tower some 20-30’ 
about the ground. Viewing them with my binoculars I made them out to be 
N. vau-album. Several also rested on the dark green screening of the building 
windows 6’ above the ground. In each case there seemed to be a concentration at 
one common area. Watching the junction at the corner of the building and the 
roof, I could see the butterflies alight, then walk to an opening between metal 
stripping and the wall. The opening, approximately 2” long and *4” wide, was the 
focal spot for the butterflies. The butterflies approached this opening, folded their 
wings and disappeared within. Some would stay inside for a short time, then re- 
emerge, fly about in a circle, return and re-enter. According to C. L. Remington 
(pers. comm.), in Autumn, nymphalines change their phototactic responses and are 
attracted to dark areas. This opening provided such a spot, appearing black against 
the white walls. 

The greatest number of individuals seen at any one time was fourteen, and all 
seemed to be using the opening. 

Although some individuals were seen to enter and re-emerge, most stayed inside. 
Therefore, it is felt that this area was a definite hibernation spot for this species. 


Nose S. Procror. Biology Department, Southern Connecticut State College, 501 
Crescent St., New Haven, Connecticut 06515. 


-I 


VoLUME 30, NUMBER 2 12, 


OBSERVATIONS OF EUPACKARDIA CALLETA IN 
SOUTHERN TEXAS (SATURNIIDAE )! 


THOMAS A. MILLER 


U. S. Army Medical Bioengineering Research and Development Laboratory, 
USAMBRDL BLDG 568, Fort Detrick, Maryland 21701 


Eupackardia calleta (Westwood) is known to occur in southern Texas 
from the area of Victoria and Beeville south into the Rio Grande Valley 
and westward into the Big Bend Area (Collins & Weast, 1961; Ferguson, 
1971-72). During the latter half of 1974, while temporarily assigned to 
Fort Sam Houston, San Antonio (Bexar County), I was able to make 
field observations and rear this interesting and little reported saturniid. 

From 9 August-4 December, populations of E. calleta were studied 
at 13 locations in 7 southern Texas counties (Karnes, Goliad, Bee, Live 
Oak, McMullen, Jim Wells and Duval), primarily by observing immature 
stages on one of this species’ known food plants, cenizo, Leucophyllum 
frutescens (Berland. ). 

Fifty-seven cocoons were collected from 9 August-28 September. 
These contained: 18 viable pupae (15 4; 3 2); 37 pupal cases (15 4; 
15 2; 7 undetermined sex) from which the adults had emerged; 1 parasi- 
tized female pupa; and 1 dried, intact larva. All of these cocoons, with 
viable pupae or otherwise, had a very worn, smooth surface and it was 
not possible to use the appearance to estimate the cocoon age. Likewise, 
the age of the cocoon could not be estimated by determining whether 
or not the silken attachment had ligatured the stem of the food plant, 
since this condition was observed for some of the cocoons with viable 
pupae and some without. However, these observations of the appearance 
of the cocoon surface and the ligaturing of the food plant stems suggest 
that some of the cocoons with viable pupae were probably as old as 
some of those from which the adults had emerged. 

The eighteen viable pupae were kept outdoors at San Antonio and the 
adults emerged in 1974 as follows: 14 Sept. (1 é); 15 Sept. (1 ¢); 17 
Sept. (1 ¢); 18 Sept. (1 ¢ ); 24 Sept. (2 6,1 2); 26 Sept. (2 ¢); 2 Oct. 
Gere) Oct. (2 3); 9 Nov. (i ¢); 10 Oct: (1 2); HL Oct (1.3, 1 2); 
13 Oct. (1 ¢); 29 Oct. (1 ¢). Although these pupae were removed 
from their original collection locations, it is probable that the period of 
emergence at San Antonio generally coincided with the period of natural 


1 The opinions contained herein are the private views of the author and should not be construed 
as official or reflecting the views of the Department of the Army. 


128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


emergence farther south, since, as noted later, naturally-occurring males 
were observed in Jim Wells and Live Oak counties on 28 September. 

An additional 47 cocoons collected between 26 October and 29 Novem- 
ber included 27 with viable pupae (15 ¢, 12 2). The 20 cocoons with- 
out viable pupae were not examined to determine the nature of the 
contents, but all had the same smooth, worn appearance mentioned 
earlier, indicating that they were not recently formed. The cocoons with 
viable pupae were of recent origin because they were white and powdery 
in appearance, and had short, loose silken threads on the surface or 
silken networks still attached to various parts of the food plant. Observa- 
tions of the larvae reared at San Antonio during 1974 showed that they 
spin a silken network upon which they construct the typical teardrop- 
shaped cocoon. After the cocoon has taken form, the larva secretes a 
white fluid which, upon drying, imparts the powdery appearance. The 
silken threads and the powdery appearance evidently disappear with 
weathering and their presence is a reliable indicator of recent formation. 

Several naturally occurring adult males of E. calleta were observed 
in two instances in southern Texas. The first instance relates to the 
female that emerged at San Antonio on 24 September. This female was 
transported to Live Oak Co. on 28 September and set out at 0845 CST. 
Seven males were attracted from the surrounding cenizo fields within 
5 minutes. The eggs from this mated female were used to colonize this 
species at San Antonio. This method of survey for adult males would 
have been used more extensively if there had been more females among 
the viable pupae collected up to 28 September. The second instance 
was in Jim Wells Co. on 28 September where binoculars were used to 
observe males flying over the extensive cenizo fields that occur along 
U.S. Highway 281. The straight-line flights of 3 males were observed 
for some time in the hope that the moths would pause, circle, or drop 
into the vegetation and possibly indicate the presence of a naturally 
occurring female. No such behavior was observed, and the males 
eventually flew out of sight. 

Since no observations were made between 28 September and 16 
November, the dates of occurrence of early instar larvae were not 
determined. However, 4th- and 5th-instar larvae were observed in 
Goliad and Bee counties between 16 November and 4 December. Larvae 
observed on 16 and 29 November were inactive, with only a few feed- 
ing or spinning cocoons. This was probably due to the cool temperatures 
(6-7° C) prevailing at the time. The last observations in Goliad and 
Bee counties were made on 4 December. At that time a few more 
cocoons had been formed, but most of the larvae were still inactive and 


VoLUME 30, NUMBER 2 129 


not feeding. Eighteen 5Sth-instar larvae were collected on 4 December 
and transported to San Antonio where they were kept on cenizo cuttings. 
Some of these larvae began spinning in transit; all had spun cocoons 
by 7 December. The fact that these larvae began spinning within hours 
after being placed in a warm environment suggests that they might 
have remained inactive on the plants in Goliad and Bee counties 
(as they had from 16 November-4 December) until the return of warm 
weather in early spring. Unfortunately, follow-up observations of these 
natural populations could not be made. 

Although the foregoing observations provide specific records of the 
occurrence of E. calleta life stages in southern Texas, they are from 
scattered areas and chronologically incomplete. As such, they provide 
only limited insight into some aspects of the biology of this species. 
One interesting observation is that certain of the cocoons, with or 
without viable pupae, were very similar in surface appearance; and that 
cocoons of both types had ligatured the food plant stems. This suggests 
that for individuals of a single generation there is considerable varia- 
tion in the time period between cocoon formation and adult emergence. 
In this regard, Kendall (1974) has observed that some E. calleta pupae 
remain in diapause for up to 22 months. Thus it is understandable that 
confusion exists concerning generations represented by the reported 
annual bimodal emergence (March and September—November) of E. 
calleta in southern Texas. Collins & Weast (1961) reported that E. cal- 
leta emerges in southern Texas from late October through November, 
with a partial second emergence occurring the following March. Fergu- 
son (1971-72) reported that there is a partial emergence of E. calleta 
in March in southern Texas, followed by a larger emergence which takes 
place from September-November. Although these authors do not specifi- 
cally refer to these emergences as generations, this is alluded to in 
later statements concerning the fact that E. calleta is known to emerge 
in Arizona only in August, and that there may be only one generation 
of this species in Arizona. Annual bimodal emergences are known for 
other saturniids, such as Hyalophora cecropia (Linnaeus). In Illinois, 
H. cecropia has two distinct periods of emergence which are very close 
together (May-early June and late June-July). However, these two 
periods of emergence are known from an intensive 3-year study by 
Sternburg & Waldbauer (1969) to represent a single generation. In 
the case of E. calleta, the March and September-November emergences 
may represent individuals from two or even three earlier generations, 
but this situation requires considerably more detailed study to be 
understood. The observation that larvae remain inactive on the food 


130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


plants during cool periods in late November and December could repre- 
sent a mechanism whereby the development of individuals from a single 
generation becomes out of phase; with the adults eventually emerging 
during one or the other of the annual bimodal periods. Certainly, there 
was considerable variation in the progress of development among in- 
dividuals that might be considered to represent a “fall” generation; some 
had formed cocoons and pupated as early as 16 November in Karnes 
Co.; others were still in the 4th instar as late as 29 November in Bee Co. 


ACKNOWLEDGMENTS 


I wish to thank Roy O. Kendall, of San Antonio, for sharing his 
records and knowledge of this saturniid species and for taking the time 
to review this paper; and to thank Captains Alan R. Gillogly and Row- 
land N. Wilkinson, formerly of Fort Sam Houston, who accompanied 
me on some of the field trips to southern Texas. 


LITERATURE CITED 


Cotuins, M. M. & R. D. Weasr. 1961. Wild Silk Moths of the United States. 
Collins Radio Co., Cedar Rapids, Iowa. 138 p. 

Fercuson, D. C. 1971-72. Bombycoidea (Saturniidae). In R. B. Dominick 
et al., The Moths of America North of Mexico, fasc. 20.2, 269 p., 22 pls. E. W. 
Classey, Ltd. London. 

KENDALL, R. O. 1974. Personal communication. 

STERNBURG, J. G. & G. P. WALpBAUER. 1969. Bimodal emergence of adult Cecropia 
moths under natural conditions. Ann. Ent. Soc. Amer. 62: 1422-1429. 


VoLUME 30, NUMBER 2 131 


EMERGENCE OF HYALOPHORA CECROPIA (SATURNIIDAE) 
BLOCKED BY SEEDS IN THE COCOON VALVE 


G. P. WALDBAUER AND J. G. STERNBURG 
Department of Entomology, University of Illinois, Urbana, Illinois 61801 


In the fall of 1974 we collected in Champaign, Illinois, a cocoon 
of Hyalophora cecropia (L.) that, unknown to us, had the exit valve 
blocked by three unshelled sunflower seeds and half of a peanut kernel 
which had presumably been inserted there by a bird. At that time the 
cocoon was heavy and, when shaken, felt as if it contained a living 
pupa. It was chilled to break diapause and then placed in an incubator 
at 25°C with other cocoons. When it became obvious that nothing would 
emerge from this cocoon, we cut it open and discovered the seeds and a 
dead adult female that apparently had been unable to push through the 
blocked valve. We have found many cocoons with foreign objects, usually 
seeds, forced into the exit valve, but this is the first time we have seen 
direct evidence that this can result in mortality. 

Reports of seeds within the valves of cecropia cocoons are scattered 
throughout the literature. For example, Minot (1870) found beech nuts, 
and LeBaron (1870) and anonymous (1870) found corn kernels. Wald- 
bauer & Sternburg (1967) found a variety of seeds and an earthworm. 

Every fall and winter since 1965, we have found foreign objects in 
cecropia cocoons collected in or near Champaign and Urbana, Illinois, 
but we recorded data only during the first three years. The number and 
percentage of cocoons with foreign objects found each year are: 1965-66, 
15 (3.1%); 1966-67, 7 (3.5%); and 1967-68, 32 (2.9%). Twenty-two of 
these cocoons contained living pupae, one contained a dead larva, and 
31 had pupae that had been killed by a woodpecker (Waldbauer & 
Sternburg, 1970). Forty-one cocoons contained one foreign object, eight 
contained two, four contained three, and one contained four. The 54 
cocoons contained 75 different objects: 26 unshelled sunflower seeds, 
21 corn kernels, 11 unidentified pieces of nut meat, eight shelled sun- 
flower seeds, two half peanut kernels, two whole acorns, two pieces of 
shelled acorn, one entire peanut kernel, one piece of Brazil nut, and 
one dead earthworm. All of these cocoons were in exposed positions on 
a twig or branch. One was on the branch of a shrub (Cornus stolonifera) 
at a height of 1.2 m. The rest were on the twigs or branches of trees— 
mostly Acer saccharinum, Betula spp., or Malus spp.—at an average 
height of approximately 2.38 m above the ground (range: 0.3 to 6.1 m). 

With one exception, the objects were pushed through the valve of 


132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the outer envelope into the space between the outer and inner envelopes 
[see Waldbauer & Sternburg (1967) for illustration]. The exception 
was a corm kernel pushed through a hole in the outer envelope at the 
end opposite the valve. 

Several birds that occur in this vicinity are reported to insert seeds 
in crevices, knot holes, and other crannies: the bluejay (Bent, 1946), 
the white-breasted nuthatch (Bent, 1948), and the red-bellied and red- 
headed woodpeckers (Bent, 1939). However, only the bluejay is com- 
mon in the urban environment in which we collected most of the 
cocoons. 

We do not know how frequently blockage of the valve is sufficient 
to prevent the escape of the adult from the cocoon because we removed 
any foreign objects we noticed. A seed or two in a cocoon may have 
escaped our notice. Indeed, we often found sunflower seeds and corn 
kernels, apparently pushed out of the valve by emerging adults, at the 
bottom of our emergence cages. However, a blockage large enough to 
prevent the moth’s exit has obviously escaped our notice only once. 
Eleven of the 54 cocoons mentioned above appeared to be sufficiently 
blocked to prevent emergence, but we do not know if this would have 
happened, either because we removed the objects or because the cocoon 
did not contain a living pupa. 


LITERATURE CITED 


ANnonyMous. 1870. Corn kernels in cocoons of cecropia moth. Amer. Ent. 2: 370. 

Bent, A.C. 1939. Life histories of North American woodpeckers. U. S. Nat. Mus. 
Bull. 174, 334 p. 

Bent, A. C. 1946. Life histories of North American jays, crows and _ titmice. 
U.S. Nat. Mus. Bull. 191, 495 p. 

Bent, A. C. 1948. Life histories of North American nuthatches, wrens, thrashers 
and their allies. U.S. Nat. Mus. Bull. 195, 475 p. 

LeBaron, W. 1870. Corn kemels in cocoon of cecropia moth. Amer. Ent. 2: 177. 

Minor, C. S. 1870. Beech-nuts in cocoon of the cecropia. Amer. Ent. 2: 242. 

Wappavuer, G. P. & J. G. Srernpurc. 1967. Differential predation on cocoons 
of Hyalophora cecropia (Lepidoptera: Saturniidae) spun on shrubs and trees. 
Ecology 48: 312-315. 

Wavppaver, G. P. & J. G. Srernsurc. 1970. Hairy and downy woodpecker 
attacks on cocoons of urban Hyalophora cecropia and other saturniids (Lepi- 
doptera). Ann. Ent. Soc. Amer. 63: 1366-1369. 


VoLUME 30, NuMBER 2 133 


DESCRIPTIONS OF THE IMMATURE STAGES AND BIOLOGY 
OF VITULA LUGUBRELLA (RAGONOT ) 
(PYRALIDAE: PHYCITINAE )! 


H. H. NeEunzic 


Department of Entomology, North Carolina State University, 
Raleigh, North Carolina 27607 


Vitula lugubrella was described by Ragonot in 1887. In 1901, Hamp- 
son, who completed volume 8 of Ragonot’s “Mémoires sur le Lépi- 
dopteres’, included a slightly more extensive description [based in part 
on Moodna ostrinella (Clemens)] and a figure of the adult of this 
phycitine. Heinrich in 1956 described and partially illustrated the male 
genitalia. 


The present paper gives information on the appearance of the last 
stage larva, pupa and habits of the larval stage. 


Last Stage Larva 


Color.” Head yellowish brown; all groups of muscle attachments usually pale 
brown, indistinct; small brown patch at notch of postgenal region of head, with 
darker pigmentation extending dorsally along posterior margin of head capsule; 
hypostoma with brown to black markings; mandibles yellowish brown between 
articulations becoming dark brown to black distally and dark brown to black along 
lateral margins. 

Prothoracic shield yellowish brown to brown without any distinctly contrasting 
patches of color. 

Prespiracular plate yellowish brown to brown. 

Remainder of prothorax yellowish white, white or grayish white (translucent pink 
to pinkish white in living larva). 

Mesothorax and metathorax yellowish white, white or grayish white (living larva 
translucent pink to pinkish white; young last stage larva with internal organs readily 
evident, including dark purple gut and pale, whitish tracheae); mesothoracic ring 
pale brown to dark brown, usually darker along inner margin. 

Thoracic legs mostly yellowish brown. 

Abdomen similar to mesothorax and metathorax; caudal segments usually more 
yellowish (living larva with pink and dark internal color not as evident on caudal 
segments ). 

Eighth abdominal segment ring pale brown to brown, usually paler than meso- 
thoracic ring. 

Anal shield pale translucent yellowish brown, without distinct markings. 

Peritreme of spiracles pale brown to brown. 

Structural characters. Entire length 12.5-18.5 mm. Width of head 1.19-1.32 mm. 
Mandibles (Fig. 1) with 3 distal teeth. Sensilla styloconica of maxillae (Fig. 2) 
simple, tapering to a point, curved mesally. 


1 Paper No. 4769 of the Journal Series of the North Carolina Agricultural Experiment station, 
Raleigh. 


2 Of preserved larva, unless otherwise indicated. 


134 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


a 544092 9 0:5 0%0.00 599, 290 
y81000 09a 0 6 + 110 aParorrons 
5 


Seofo * of eon: 
e O Soe 
* 0 40 OD: 
* ve aPe:0 
MISS oe +o teen ot 0% tai 


4 S) 


Figs. 1-5. Vitula lugubrella. 1, mesal view of right mandible of last stage larva; 


2, mesal view of left maxilla of last stage larva; 3, lateral view of head, prothorax 
and mesothorax of last stage larva; 4, dorsal view of cephalic segments of pupa; 
5, dorsal view of caudal segments of pupa. 


VoLUME 30, NUMBER 2 135 


Most morphological characters similar to Vitula edmandsae (Packard), as described 
and figured by MacKay (1972), with at least the following exceptions: 

On the head, L1 distinctly above imaginary line between A2 and A3 (Fig. 3) 
(V. edmandsae with L1 very close to, or on, an imaginery line between A2 and A3); 
prothoracic shield with distance between XD2 and SDI only slightly less than 
distance between SD1 and SD2 (Fig. 3) (V. edmandsae with distance between 
XD2 and SD1 distinctly less than between SD1 and SD2; abdominal segments 1-5 
with D2 setae about 2 longer than D1 setae (V. edmandsae with D2 setae on 1-5 
about 3X longer than D1 setae). 


Pupa 


Color. Yellowish brown to brown. 

Structural characters. Length (excluding cremastral “spines” ) 8.3-10.0 mm. 
Width across mesothorax 2.4-2.6 mm. 

Head minutely granulate on front; vertex broadly produced anterior to prothorax 
(Fig. 4). 

Thoracic spiracles distinct, narrow, protruding (Fig. 4) dorsum of metathorax 
with 2 groups of about 16 shallow punctures. 

Cephalic 4% to %3 of abdominal segments 1-3 with numerous shallow punctures; 
punctures usually do not reach spiracles on segment 2 and barely extend to spiracles 
on segment 3; segments 4—7 with numerous shallow punctures encircling segments; 
segment 8 with 2 groups of dorsal punctures and D1 and L2 setae (D1 missing 
on some specimens); segment 9 with D1, L2, and L3 setae and sometimes vestigial 
SV2 setae; segment 10 with 12 stout, hooked setae (Fig. 5); gibba absent. 


Material Examined 


Texas: Ft. Davis, 8 larvae, 1 pupa, Quercus grisea, Liebmann 9-IX-73, Coll. H. H. 
Neunzig; 3 larvae, Quercus grisea, Liebmann 28-IV-74, Coll. H. H. Neunzig; 5 
larvae, 2 pupae, Quercus grisea, Liebmann 27-IV-75, Coll. H. H. Neunzig. 


Larval Habits 


Larvae of Vitula lugubrella live within the dried galls of Atrusca spp. 
which occur on Quercus grisea. These cynipid galls are of the “oak apple” 
type and are approximately 15-30 mm in diameter. The small larvae 
feed just inside the outer covering of the gall on the fibrous material that 
constitutes most of the internal structure (Fig. 6). Late or last stage 
larvae also feed on these peripheral fibers and, in addition, frequently 
tunnel through the center of the gall, including the central chamber in 
which the cynipid develops (Fig. 7). Large quantities of conspicuous, 
loose frass accumulate within infested galls. Pupation takes place in a 
loosely woven silk cocoon within the gall. Vitula lugubrella apparently 
is multivoltine because larvae, of various sizes, and pupae, at different 
stages of development, are present in galls in the spring and also in the 
fall. 

Vitula lugubrella is the 2nd North American phycitine species known 
to be associated with “oak apple” galls. Sosipatra thurberiae (Dyar) 


136 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Migs. 6-7. Vitula lugubrella. 6, partially grown larva in gall of Atrusca sp.; some 
of the frass in the gall was deposited by a large larva of V. lugubrella that inhabited 


the gall earlier; 7, last stage larva in tunnel made through center of gall of Atrusca 
sp. 


has previously been reported by Heinrich (1956) as an inhabitant of 
these leaf galls. It is interesting to note that Roesler (1973) has synony- 
mized Sosipatra and Vitula indicating a close relationship between 
lugubrella and thurberiae 


VoLUME 30, NUMBER 2 137 


ACKNOWLEDGMENT 


The drawings were prepared by Mrs. Lily Shen. 


LITERATURE CITED 


Herricu, C. 1956. American moths of the subfamily Phycitinae. U.S. Nat. Mus. 
Bull. 207. 581 p. 

MacKay, M. R. 1972. Larval sketches of some Microlepidoptera, chiefly North 
American. Mem. Ent. Soc. Canada 88. 83 p. 

Raconot, E. L. 1887. Diagnoses of North American Phycitidae and Galleriidae. 
Paris. 20 p. 

Raconot, E. L. & G. F. Hampson. 1901. Monographie des Phycitinae et des 
Galleriinae II. In Mém. sur les Lépidoptéres. 8. St. Petersburg. 602 p. 

Roeser, R. U. 1973. Phycitinae. Parts 1 and 2. In Microlepidoptera Palaearctica. 
A, Vienna. 752 p. & 137 p. 


SUMMER MONARCH (DANAUS PLEXIPPUS) IN SOUTHERN TEXAS 
(DANAIDAE ) 


The monarch, Danaus plexippus (L.), is by far the best-known butterfly in North 
America. During several years’ personal observation in the Brownsville, Cameron 
Co. area of southern Texas, monarchs were normally observed only from late Sep- 
tember through April. Adults were most commonly seen in the fall and early 
spring as they migrated south and north, respectively. However, in warmer years 
monarchs were observed during the winter months. Other observers (see Urquhart 
1960, The Monarch Butterfly, U. Toronto Press, pp. 165, 172-173; Yeager 1974, 
News Lep. Soc. 1974(3), p. 3) have reported similar near total absences of 
monarchs in southern Texas during summer months. Summer breeding of monarchs 
is normally limited to northern Texas and areas to the north. 

However, in June and July 1966 larvae of the monarch were found in Browns- 
ville on milkweed. Adult monarchs reared from wild larvae in the laboratory emerged 
on 17 June (2), 19 June (4), 3 July (1), 5 July (1) and 11 July (2). Climatic 
factors could have been involved in this southern extension of summer breeding 
of the monarch in 1966. Temperatures during January-May were cooler than 
normal. January was wetter than normal followed by three months of very little 
rainfall (2.16 in.) until a very wet May (6.05 in. vs. normal 2.50). 

Another possible factor needs to be considered. The monarch butterfly is subject 
to cyclical population crashes which have attributed to a cytoplasmic polyhedrosis 
virus (Urquhart 1970, Atalanta 3(2): 1-11). The year 1966 was characterized by 
generally low populations throughout most of North America, including southern 
Texas (Urquhart 1970, op. cit.). If the monarch is less prone to migrate during 
low population periods as suggested by Urquhart (1970, op. cit.), this aberrant 
summer occurrence of the monarch in the Brownsville area may have been the 
indirect result of a virus outbreak. 


RayMonp W. Neck, Texas Parks and Wildlife Department, John H. Reagan 
Building, Austin, Texas, 78701. 


138 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


BOOK REVIEWS 


Tue Burrerruies OF NortH AMERICA, by William H. Howe, Contributing Editor. 
1975. Doubleday and Co., Inc., Garden City, L. I. New York. xiii + 633 p. +97 
plates. Price $39.95 U.S. for the standard edition. 


This long awaited work, including sections by twenty contributors, edited and 
illustrated by Howe, fails to meet expectations. Some sections are very well done 
while others are not. These will be treated in specific terms subsequently. The 
general format of the book follows that of Holland’s Butterfly Book, including the 
general introduction and the sequence of presentation of families. The family 
sequence is awkward in comparison with modern books and checklists. The work 
begins with the danaids and ends with the Hesperioidea. The colored plates are 
bound in the middle of the volume. Also included are a glossary of terms, a short 
bibliography and general index plus an index of food plants. 

General criticisms of the book relate to nomenclature, literature citations, the 
plates and ease of use. It must be recognized that considerable time may elapse 
between the date when the finished manuscript is submitted to the publisher and 
the release of the bound volume. On this basis, the author can be forgiven for not 
including taxonomic work published after late 1973. On the other hand, much 
work in some families prior to that date is omitted, while in other families it is 
included. There is a basic lack of consistency in this respect. It probably relates to 
the fact that the various contributors submitted their entries over a wide time 
period, however it is the job of the editor to note additional work and insure that 
the contributors make any necessary changes. Apparently the individual authors saw 
neither the final manuscript nor the galley proofs. It also appears that the plates 
were executed without consultation with the authors and prior to the writing of 
many of the sections. The authors should have been given the opportunity to 
select the specimens illustrated. 

Despite an initial statement to the contrary on p. vii, a number of new taxa 
are introduced (which may cause problems with authority) and other taxa are 
omitted without comment. There is considerable lumping of subspecies in some 
genera without appropriate discussion. There is no reference for the new Bauer 
names in the Melitaeinae, for example, and one assumes that the book constitutes 
the first publication of these taxa. With respect to the Code of the I.C.Z.N. ( Arts. 
11, 13, 68-73), some or all of these names may represent nomina nuda. No holo- 
types have been designated; in most instances no type localities are indicated; in 
several cases the accompanying text does not clearly distinguish how the new 
subspecies differs from the related subspecies. Other problems in the text include 
incorrect and incomplete distribution information, incorrect and incomplete flight 
periods, and too many typographical errors. The first two categories could have been 
avoided by checking original descriptions of many of the species and available 
regional books and checklists, as well as the “Annual Summary’ in the News 
of The Lepidopterists’ Society. 

Although a bibliography appears at the end of the book, I question the rationale 
of its selection. A number of pertinent entries are omitted, while others of lesser 
value are included. Author and date citations included in the main text do not 
appear in the bibliography in many cases. 

Regarding ease of use, I find it awkward to have the plates bound in the middle 
of such a large volume. A more critical feature is the lack of page references in 
the plate legends, although the text descriptions cite plate and figure. 

The color quality of the plates requires some comment. Watercolors can come 
close to modern photography, and there are some practical book production 
problems that favor their use, but they definitely detract from the scientific value 


VoLUME 30, NUMBER 2 139 


of this book. There is a distinct color shift to green and blue-green in many of 
the plates. This is especially noticed in the Hesperioidea where the warm browns 
are gray-green-brown and in the lycaenids where many of the colors are off-register. 
Part of this relates to poor quality control in the printing. I had occasion to check 
several copies of the book. On given plates, the colors were not uniform throughout 
the copies. A number of plates have a pastel color background, usually green or 
blue, so that better definition is given to the specimens represented. The intensity 
of the background varied considerably from copy to copy, with concomitant color 
shifts in the specimens depicted. This situation was prominent in three copies 
examined, although not the same plates were involved. Based upon comments 
from correspondents who have seen other copies of the book, some do not exhibit 
this problem to the extent that the copies I examined did. The unrealistic blue 
and red tones throughout the book, however, appear to stem from the original 
plates. While scientific illustrators may take “artist’s license” to insure reasonable 
reproduction of their subjects in the final printed form, Howe has overdone this 
technique in many cases, producing an artificial and unrealistic portrayal. Many 
of the plates have a “washed out” aspect, which again may result from production 
difficulties, but this should not occur in a book of this scope. Generally, the colors 
are disappointing. The lycaenids, in many cases, should have been illustrated larger 
than life size. In many of the blues, it is difficult to tell just what is being repre- 
sented in the figures. 

Other problems relate to the mixing of families, genera and species over a 
number of plates. In some plates, examples from as many as six different families 
appear, when there is no valid reason for doing se. Another problem is the figuring 
of the male, female and underside of a given species over three separate plates. 
Initially some thought should have been given to the arrangement of the sexes 
and. species so that the reader would not have to leaf through several plates to find 
all the illustrations for a given insect. The species corresponding to a given genus 
should have been uniformly grouped. One wonders why Figure 32 which illustrates 
maculation in Parnassius was placed on p. 211 in the Speyeria discussion. 

On pp. 578-579, several addenda regarding distribution and taxonomy appear 
noting recent work. If some addenda could be included, why not all of the new 
works? It would have made the book both current and of more value. 

Considering the magnitude and scope of this project, it seems strange that not 
a single genitalic sketch appears, excepting a generalized drawing in the introductory 
portion. In view of the, in many cases, marginal plates, especially in the blues 
and some of the Hesperioidea, genitalic sketches of the difficult species would have 
been very valuable. These would have been extremely useful in separating look- 
alike species in such genera as Lerodea, Lerema, Nastra, Bolla, Cogia, Erynnis, 
Staphylos and Thorybes. This is also true in several other families and genera such 
as Euphydryas, Philotes, Oeneis, etc. 

In a number of instances, it was noted that the subspecies figured in a plate, 
with associated collection locality stated in the plate legend, did not agree with 
the text description of the distribution for that subspecies. The plates and associated 
text should have been carefully reviewed to prevent such inconsistencies. This situation 
is going to cause confusion and perpetuate misconceptions. There are also some 
misidentified species in the plates. These will be treated later. 

On the positive side, the introductory material by Harry Clench (butterfly 
anatomy, biology, etc.) and Alexander Klots (scientific nomenclature, collecting, 
preserving, etc.) is extremely useful and informative to the amateur and _ profes- 
sional alike. These are perhaps the best done sections in the book. The overall 
editing appears to have been carefully done to insure uniformity of format, a 
formidable task when twenty contributors are involved. 

SATYRIDAE. The text appears basically correct, although I prefer the approach 


140 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


that assigns most of the North American Coenonympha to tullia rather than the book 
approach which treats inornata, ochracea, ampelos, etc. as separate species. There 
are several recent revisions in the Satyridae that are not included. The concerned 
reader would do well to consult L. D. Miller's revisions in the Euptychiini pub- 
lished in the Allyn Museum Bulletin series. The Heitzman and dos Passos revision 
of Lethe should also be consulted (Trans. Amer. Entomol. Soc., 100: 52-99, 1974). 
Erebia debanensis is incorrectly identified as E. youngi rileyi, while discussion of 
E. debanensis, inuitica and semo is omitted. Incomplete distribution data are given 
for Erebia callias, magdalena, Oeneis alberta daura and Gyrocheilus patrobas tritonia. 
The latter is quite common in New Mexico in Grant and Sierra Counties. Records 
have appeared in published regional lists and in the Lep. Soc. News. Incomplete 
and incorrect data are given for Oeneis jutta reducta which was described from 
Montana and is found in eight Wyoming counties in addition to Colorado. The 
discussions of Cercyonis meadii mexicana and nominate meadii seem to be juxta- 
posed to some extent. It should be noted that the flight period for Paramecera 
allyni Miller (not xicaque which is Mexican) is normally from mid-July into early 
August, and not late summer as stated. The following errors were noted in the 
plates: Pl. 4, f. 22, Coenonympha ochracea brenda was described from the “Los 
Angeles area”, although Brown in his discussion of the Edwards’ types casts doubt 
upon this locality, suggesting the Great Basin. The Utah collection site is slightly 
odd, although I have taken brenda phenotypes in extreme western Nevada; Pl. 5, f. 1, 
the specimen shown is Erebia rossii gabrieli and not E. r. kuskoquima (note col- 
lection site and text discussion of distribution); Pl. 5, f. 16 is E. theano ethela and 
not demmia (note collection site, demmia is found only in one area of southern 
Colorado); Pl. 5, f. 18-19, specimens shown are Coenonympha ochracea subfusca 
and not C. o. furcae (note collection site and compare with text and Pl. 4, f. 23-24); 
Pl. 9, f. 12 is Oeneis bore/taygete and not O. melissa gibsoni. 

NYMPHALIDAE. A number of nomenclatural changes occur in this section, most 
without adequate comment. A number of new taxa are introduced and the inclusion 
of recent taxonomic work is not consistent. All of the Asterocampa taxa are treated 
as separate species. Two recently described subspecies of Euphydryas chalcedona 
are included in the text, while two earlier described subspecies of E. editha, alebarki 
Ferris and gunnisonensis Brown are omitted, although there is reference to the 
occurrence of E. editha in Wyoming and Colorado. J. Scott’s work in Poladryas 
is omitted. It is stated that Nymphalis californica herri is the Rocky Mountain race. 
This subspecies was described from the Buckhorn Mountains in Washington and 
is found in the Priest River area of Idaho. Specimens from southwestern New 
Mexico through Wyoming are typical californica. The character by which Vanessa 
atalanta rubria is separated from nominate atalanta is reversed; rubria exhibits a 
somewhat smaller subapical white bar rather than a larger bar. 

The main problem with this section is the treatment of the genus Speyeria. I 
disagree entirely with the elevation of nevadensis and electa to species status, and 
with several other nomenclatural changes. This assignment totally ignores the work 
that Grey and Moeck have done in studying the distributions and color phases of 
western Speyeria. Additionally, leto and adiaste are also elevated to species status. 
There is perhaps sound basis for the latter, but considerable further study is neces- 
sary with respect to leto. I do not accept minor variations in larval stages as the 
basis for separation at the species level. The form clio in S. mormonia eurynome 
has been given subspecific status. I can see no basis whatsoever for this action. 
Unsilvered forms are typical in all mormonia populations. I have taken mixed pairs 
in copulo. The author seems to haye ignored the Rocky Mountain populations 
of atlantis and callippe in his elevation of nevadensis and electa to species rank. 
The misconceptions are too numerous to detail here. Two other taxa are treated 
to provide resolutions to two long-standing problems. S. zerene sordida is treated as 


VoLuME 30, NuMBER 2 141 


an aberrant form of bremnerii and is probably correct. The introductory section in 
Speyeria alludes to the pfoutsi and platina problem, but the ensuing text does not 
resolve it. Skinner’s type series for S. zerene platina was a mixture of several species, 
which led Gunder to describe pfoutsi. The taxon pfoutsi should be considered a 
synonym of platina. The new species S. egleis toiyabe Howe merits some comment. 
The figure reference in the text (p. 237) is incorrect as the reader is directed to 
Plate 31, f. 13 rather than f. 14. Apparently only the holotype has been designated 
without a type series. This is unfortunate as there is no discussion concerning the 
variation of facies within this subspecies. What is shown as the holotype (Pl. 31, 
f. 14) resembles quite closely some of the pale disc specimens from the egleis cline 
found in the Pine Creek area of Elko Co., Nevada. 

Several items were noted in the plates: Pl. 34, f. 1 is a very poor rendition of 
Boloria eunomia ursadentis. This subspecies is distinctly pale and yellow-brown. 
Regarding the related text discussion of this species, it should be noted that it has 
been found to date only at high altitude on the Beartooth Plateau as cited in the 
original description (J. Res. Lep. 9(4) :243-248, 1970 [1971]). Northen Wyoming 
and southern Montana colonies of ewnomia are closer to dawsoni than to any other 
subspecies. Pl. 34, f. 3, the specimen figured is typical low altitude eunomia nr. 
dawsoni; nichollae is very dark, almost melanic, and apparently occurs above tree- 
line only at the edges of snow fields. The colors are misleading in a number of 
Boloria figures. 

LIBYTHEIDAE. There is considerable confusion in the text (p. 258) and in 
Pl. 47, f. 14-15. The description of L. carinenta applies to L. bachmanii larvata 
and vice versa. The plate figures depict /arvata and not carinenta. 

RIODINIDAE. Celephelis is combined into seven species which seems an 
improvement over McAlpine’s recent splitting of the genus. Some nomenclatural 
changes are mentioned. 

LYCAENIDAE. This section is generally well done. Several minor items are 
noted. The subspecies minnehaha is applied erroneously, as did Holland, to the 
montane Colorado population of Plebejus shasta. This situation was explained by 
the author in 1970 (Ent. News 81:203-207). Pl. 59, f. 17 does not fit at all the con- 
cept of P. shasta comstocki. The specimens shown as Plebejus icarioides pembina 
(Pl. 56, f. 27-28) are from a blend-zone region and are not representative of 
neotopo-typical western Montana specimens. The specimen shown in Pl. 50, f. 12 
is Incisalia polios obscurus Ferris and Fisher and not nominate polios. The text 
omits this taxon, described in 1973. Plebejus acmon spangelatus is assigned incor- 
rectly to lupini despite Goodpasture’s genitalic studies and revision of acmon and 
lupini (J. Kan. Ent. Soc. 46:468-485, 1973). The genus Phaedrotes is incorrectly 
retained instead of Glaucopsyche as pointed out by Brown (J. Lepid. Soc. 25:240- 
246, 1971). There are several new combinations proposed in Callophrys. In C. 
mitoura, the brood form smilacis has been elevated without comment to subspecific 
status replacing sweadneri. In the discussion of Ministrymon, it is stated that 
ines is simply the fall brood of leda. This bears further study. I have taken ines 
in May along with leda in southwestern New Mexico and typical leda in the fall. 
John A. Legge, Jr. has taken both forms from the same mesquite tree in Baja 
California del Sur in December. The distribution for Satyrium liparops fletcheri is 
listed as southern Manitoba only; it also occurs in Saskatchewan and North Dakota. 
Vaga blackburni is described as “. . . an abundant species on all of the Hawaiian 
Islands.”, which contradicts the recent proposal for inclusion of this species under 
endangered status. Pl. 60, f. 3-4 figure Lycaeides argyrognomon ricei and not anna. 
Various nomenclatural changes in Lycaena are proposed. It is stated that L. arota 
schellbachi extends into southern Wyoming. I know of no valid records for the 
state. The Everes amyntula/comyntas confusion seems to have been fairly well 
resolved in this book. The interested reader should refer to the recent revisionary 


142 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


_ 


work on the Philotes group by O. Shields and published in the Allyn Museum Bul- 
letin series, as he has proposed some substantial nomenclatural changes as well as 
describing new subspecies. 

PAPILIONIDAE. Again there are some minor items in the generally well done 
treatment of Papilio. In Pl. 62, f. 1, P. troilus as shown as P. brevicauda. In Pl. 63 
which illustrates the machaon complex, none of the undersides of the species are 
shown. Unfortunately, the undersides are diagnostic in most cases for reliable species 
separation in this group. 

The treatment of Parnassius, however, is particularly distressing. Granted that 
there has been too much “splitting’’ in this group, the authors of this section go 
too far in the opposite direction. The lumping of nearly all of the Rocky Mountain 
region phoebus under the umbrella sayii with the retention of a few questionable 
additional subspecies (hermodur for example, which is an altitudinal female form) 
is unrealistic. Even the Great Basin rubina (text misspells as rubiana) is lumped 
into sayii. It appears that the authors have recognized differences in facies when 
it suited them and ignored same when it did not. The discussions of P. clodius 
altaurus, gallatinus and menetresii are badly mixed up. PI. 78, f. 18 illustrates P. 
clodius altaurus (yellow-orange spots) and not menetresii (red spots). One should 
consult McDunnough’s paper relative to Bryk’s treatment of the North American 
Parnassius (Can. Ent. 68:216—225, 1936). There has been considerable confusion 
in the taxonomy of Parnassius in the past and the present treatment serves only to 
further rather than clarify it. 

PIERIDAE. This section, with a major contribution by A. B. Klots, appears to 
be the most carefully done in the book, and cites current research as of early 1973. 
Many of the problem areas in the Colias have been resolved, and the genus Pieris, 
noting Warren’s work, is thoughtfully handled. The illustrations accompanying this 
section are perhaps more realistic than in some of the other families. Dr. Klots is 
to be complimented on his handling of the Coliadinae. The specimen shown in 
Pl. 72, f. 17 as C. o. occidentalis from Crook Co., Oregon is probably a local 
form of C. alexandra peculiar to that region. Appias drusilla poeyi is dropped 
without comment. The subspecies mayi has been applied to all northern populations 
of Euchloe ausonides. This is not correct, as true mayi is a very distinct insect, and 
specimens from many of the colonies in the far Northwest do not qualify. Many are 
closer to nominate ausonides. The name andrewsi has been retained under ausonides 
rather than placed with hyantis as Opler suggested in his revision of the genus. 

HESPERIOIDEA. Although Freeman’s 1969 revision of the Megathymidae is 
cited, Kilian Roever, who prepared this section, has sensibly chosen a rather dif- 
ferent arrangement. Many of the Agathymus have been demoted to subspecies of 
several principal species such as neumoegeni and mariae. The western Megathymus 
have been reassigned to yuccae with coloradensis as a subspecies. 

C. D. MacNeill who authored the Hesperiidae section has revised his earlier 
work in lumping all of the harpalus/manitoba taxa under comma. Some of the 
other subspecies in Hesperia have been dropped without comment. The treatment 
of Erynnis generally follows Burns’ 1964 revision. Overall, the coverage is good 
and the various species, especially the Mexican migrants, are figured, but many 
of the plates leave much to be desired regarding color fidelity. Occurrence in the 
U.S.A. of some of the doubtful Mexican species would have been confirmed had 


several museum and private collections been examined more carefully. Several 
species cited as occurring in Wyoming should be considered doubtful and may 
relate to misidentifications as I have no confirmed records. The lack of genitalic 
sketches is a major omission. 

In summary, the book has some good points and some good sections, but on the 
whole, in this reviewers opinion, it exhibits some serious faults. Many of the 


problem areas could have been resolved had the manuscript been submitted to 


VoLUME 30, NuMBER 2 143 


several competent technical reviewers prior to publication. The selection of the 
majority of the contributors from the West Coast has lent an unfortunate parochial 
flavor to the text. One receives the distinct impression that there is nothing of 
taxonomic interest east of the Great Basin. The plates are a major disappointment, 
especially in the lycaenids and hesperiids where they are most needed. It is 
unfortunate that this book, which had the potential for being a really definitive 
treatise on the North American butterflies, has fallen so far short of the mark. I 
am sure that there will be those who will praise the book, citing its utility to 
amateurs and the fact that nearly every North American species is figured. To 
them, I would reply that while the book has its good points, the novice collector 
can be easily misled in a number of areas because of problems with the text and 
illustrations. As a consequence of the number of flaws which produce frustration in 
the reader, the work as a whole must be considered as marginally worth the price. 


CiirForD D. Ferris, Bioengineering Program, University of Wyoming, Laramie, 
Wyoming 82071. 


Mires oF Morus AND BuTTERFLIES, by Asher E. Treat. 1975. Cornell University 
Press, Ithaca, N. Y., 362 p., 150 figures (Frontispiece in color). Price $35.00 (U.S.). 


Clearly a classic work both for substance, including a remarkable amount of 
original (unpublished ) material, and for scholarly and poetic styie! The introductory 
section covers a concise history of mite-moth scientific discovery, a fine chapter 
on methods of study and a brief, thoughtful account of the important characteristics 
of mites. 

In the body of the book, major sections (Mesostigmata: Acariformes) and chap- 
ters are presented under higher taxa headings, for the mites, but within each 
chapter the arrangement is “. . . primarily ecological at familial or lower levels” 
(Author’s preface ). 

This is also a well-edited and very useful book with (1) 150 figures of excellent 
quality, (2) lists of moth-mite and mite-moth records, (3) keys to mites recorded 
from moths, (4) an extensive bibliography and (5) geographic and general indexes. 
The book should be read by not only “lepidopterists, acarologists and parasitologists” 
(vide Author) but by all amateur and professional scientists. 

Mites of Moths and Butterflies will form a solid foundation for and be a lasting 
stimulus to future studies in many fields of scientific research. Truly a basic excel- 
lent book for any lepidopterist’s library. 


WituiaM B. Nuttinc, Department of Zoology, University of Massachusetts, Amherst, 
Massachusetts 01002. 


BUTTERFLIES OF LEBANON, by Torben B. Larsen. 1974. National Council for Scien- 
tific Research, (C.N.R.S.), Beirut, Republic of Lebanon, xv + 255 p. Price: £7.50 
sterling (U.K.), $22.50 (U.S.). Distributed in the U.K. by E. W. Classey, Ltd., 
Park Road, Faringdon, Berks. SN7 7DR, England, and in the U.S. by Entomological 
Reprint Specialists, P. O. Box 77224, Dockweiler Station, Los Angeles, California 
90007, U.S.A. 


A review is a very personal thing to the reviewer. Without attempting to pass 
moral judgment, recent events in Lebanon have made the writing of this review 
especially difficult. Make no mistake about it, though, Larsen has written a very 
good book, and the Lebanese National Council for Scientific Research is to be 
congratulated for producing it in its present form. 


144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Larsen expends the first 76 pages in prefatory material: Lebanon’s history from 
the naturalist’s viewpoint, the zoogeography of the area, distributional patterns 
within the country, a comparison of the butterflies of Lebanon with those of sur- 
rounding countries, the economic importance of Rhopalocera and their classification, 
along with a Lebanese checklist. Detailed discussions of the 139 recorded species 
are given in the next 140 pages, and this discussion is followed by a few pages of 
possible, doubtfully and erroneously recorded butterflies. 

The sixteen color plates, identified by the text pages they face, range in quality 
from excellent to at least acceptable. Probably the least readable plate is that 
facing page 193, which is too dark and too red; some others are a bit too dark, 
but all are quite usable. It is unfortunate that the plates do not have figure 
numbers, but one soon becomes accustomed to the legends, and the exact details 
of the capture of each figured specimen are given, a policy that should achieve 
greater currency. 

The text, however, is where the book really excels. There is a wealth of informa- 
tion tucked into these pages, information that is so well documented that one is 
left with the impression that he could return to those spots at the appropriate 
times and be virtually assured of collecting the insects. Available foodplant informa- 
tion is given and original sources on life histories are cited, where available; when 
the early stages are unknown, informed speculation on possible foodplants is offered. 
Habitat information is detailed and based on the author’s personal experience in 
most cases; in others the best available information is cited. 

A single taxon is described as new in the book, but three others were to have 
been proposed before the present publication. These were to have been described 
in Alexanor during 1974, but publication was delayed, and the inclusion of these 
names in Butterflies of Lebanon does not constitute valid descriptions. These names, 
in the context of the book, are nomina nuda, as Larsen correctly states. By the 
time this review is printed these names will be validated, but they cannot be dated 
from 1974, as stated in the text. 

This book is a fine investment for anyone interested in Middle Eastern butter- 
flies or for one who wants a wealth of recent biological information about them. 


Lee D. Mutter, Allyn Museum of Entomology, 3701 Bayshore Road, Sarasota, 
Florida 33580. 


VOLUME 30, NUMBER 2 145 


GENERAL NOTES 


LIBYTHEANA BACHMANNII (NYMPHALIDAE) IN CONNECTICUT 


Libytheana bachmannii (Kirtland) is considered a rare vagrant in Connecticut to 
the local lepidopterists and reaches the northern terminus of its range in this area 
(Klots, 1951, A Field Guide to the Butterflies, Houghton Mifflin Co., Boston). 
Those that do occur are suspected to be immigrants from more southern territories 
and stay till they are lost to winter kill. There being only three Connecticut speci- 
mens in collections, it was unusually interesting to make several observations of 
this species during 1974. 

All specimens have taken in Guilford, Connecticut, and it is from this town that 
all present observations were made. Between 18 and 20 June three individuals were 
seen. One fed on Mock Orange (Philadelphus coronarius L.) and two rested on 
bark of Apple (Malus sp.). In each case there were considerable nearby stands of 
Hackberry (Celtis occidentalis L.), the larval hostplant of L. bachmannii, leading 
to suspicions of a possible summer brood by the immigrants. 

On 3 October three additional Snout Butterflies were encountered in an area 
with a half mile radius. Again Celtis occidentalis was quite common. However, 
in each case the butterflies were seen resting and possibly feeding on the fruits 
of Wineberry (Rubus phoenicolasius Maxim.). This fruit is very viscid to the touch, 
and its sweet juice would be a logical attractant. 

Only future observations will show whether there is indeed a summer brood of 
L. bachmannii in the Guilford area. 


Nosie S. Proctor. Biology Department, Southern Connecticut State College, 
001 Crescent St., New Haven, Connecticut 06515. 


A CLAMP FOR MARKING BUTTERFLIES IN 
CAPTURE-RECAPTURE STUDIES 


I have recently begun a study of adaptive differences in social behavior of several 
species of butterflies near Whitehall, Washington Co., New York. I initially used 
techniques described by Ehrlich & Davidson (1960, J. Lepid. Soc. 14: 227-229), 
in which two people first capture all the butterflies in a given area, then mark and 
release them en masse in a central location. Although this program was adequate 
for some species, the observed pattern of local dispersal of sedentary species was 
greatly affected for a period of several days by the spatial pattern of my collecting 
and releasing. The affected species showed the least variation in individual behavior 
when individuals were marked and released quickly at the site of capture. So I de- 
signed and built a clamp that allows speedy and efficient marking at the site of capture 
by a single person (see Fig. 1). 

The base is made of 14” Masonite, as is the anvil, which is glued to the base with 
its smooth side up. The softwood fulcrum is glued to the base. The “hinge” pins 
are # 18 brads that are driven through the base, cut short and filed smooth. The 
clip is made of 4” clear acrylic plastic with 46” holes to clear the pins and seven 
46” holes in an H pattern as shown, each chamfered from the top with a countersink. 
The pieces may be cut roughly, and the assembled clamp shaped on a vertical power- 
sander. A rubber band holds the jaws of the clamp together. A hole in the upper 
end of the base allows the clamp to be carried accessibly on a hook or string about 
the waist or neck. 

A butterfly is placed between the jaws of the clamp, and a small number is 
written on its forewing, using a permanent fiber-tipped pen (e.g., Sanford “Sharpie” 
at the suggestion of L. E. Gilbert). The several chamfered holes accommodate 


146 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


I“"HINGE" PINS 


0 CM 5 


Fig. 1. Top, side and “exploded” views of the clamp. 


butterflies of various sizes, and the pressure of the clip is adjustable by moving the 
rubber band back and forth between the fulcrum and the anvil. Since the clip is 
transparent, accurate measurements can be made of the wing length, presence and 
size of spots, etc. The butterfly is turned over and marked on the other side after 
the number on the first side is thoroughly dry. 

During a total of 21 weeks in the summers of 1974 and 1975, I used this clip to 
mark 1,612 individuals of 28 species, varying in size from tailed blue ( Everes comyntas 
Godart) to tiger swallowtail (Papilio glaucus Linnaeus). Marked individuals were 
recovered a total of 950 times. I had little success with blues (Plebejinae), which are 
simply too tiny and delicate and I had difficulty with freshly eclosed pearl crescents 
(Phyciodes tharos Drury) and bronze coppers (Lycaena hyllus Cramer), which 
combine delicate wings with hefty wing muscles. Painted ladies (Vanessa cardui 
Linnaeus; V. virginiensis Drury) and skippers (Hesperiidae) cannot be held firmly 
because their straight wings, wide bodies and strong muscles give them a mechanical 
advantage over the clip unless they are clamped close enough to the body to risk 
damage to their wings. Other species rarely caused trouble. Large species could 
escape if they could reach something to pull against with their feet. Very rarely, an 
individual would damage its wing mechanism trying to escape; such individuals ap- 
peared superficially unhurt, but were unable to fly and seldom recovered. 

The clip can also be used to capture feeding or strutting hairstreaks (Theclinae ) 
directly, with greater success and less damage than netting. 


Henry S. Horn, Department of Biology, Princeton University, Princeton, New 
Jersey 08540. 


VoLUuME 30, NUMBER 2 147 


PAPILIO TROILUS (PAPILIONIDAE) IN PURSUIT OF FISH CROW 


On 13 August 1974 while crossing a dike of an impoundment in the Barn Island 
Wildlife Management Area, Stonington, Connecticut, I noticed two fish crows 
(Corvus ossifragus Wilson) flying about 200’ over the marsh. This species is 
uncommon in that area and was therefore of interest. As I watched, a spicebush 
swallowtail (Papilio troilus L.) climbed toward the second bird, actually circled it 
and darted in on the bird. Such a response, elicited by the stimulus of a flying 
object such as a bird passing nearby has been reported by others, monarch ( Danaus 
plexippus (L.)) and red-winged blackbird (Agelaius phoeniceus L.) by Slansky 
(1971, J. Lepid. Soc. 25: 294), Lorquin’s admiral (Limenitis lorquini ( Boisduval ) ) 
and glaucous-winged gull (Larus glaucescens Naumann) by Pyle (1972, J. Lepid. 
Soc. 26: 261) and question mark (Polygonia interrogationis (Fabricius) ) with 
chimney swifts (Chaetura pelagica L.) by Hendricks (1974, J. Lepid. Soc. 28: 236). 

I was quite surprised, however, to see the crow twist in the air, grasp the butterfly 
in its bill and eat it! I know of no other records for this species of bird eating this 
type of food and certainly corvids do not normally capture prey while in flight. 
Therefore, not only were the swallowtail’s actions of interest but also the behavior 
and diet of the fish crow! 

Pre-courtship responses and aggressive territorial behavior have been listed as 
possible explanations for the butterfly’s actions. One can only conjecture what 
the butterfly was intending. 


Noste S. Procror. Biology Department, Southern Connecticut State, 501 Crescent 
St., New Haven, Connecticut 06515. 


MORE RECORDS OF BUTTERFLIES AS PREY FOR AMBUSH BUGS 
(HETEROPTERA ) 


In a recent publication (Pyle 1973, J. Lepid. Soc. 27: 305-307) it was related 
that a butterfly, Boloria selene, (Denis & Schiffermiiller), was discovered being fed 
on by two unidentified ambush bugs (Phymatidae) on a flower head in Washington 
State. It was indicated that ambush bugs were not known to capture butterflies. 
This writer has made numerous observations of attacks on butterflies by crab spiders 
and certain insects, including ambush bugs. Recorded here are observations in 
Maryland of ambush bugs capturing certain butterflies. 

A single observation of this behavior was made in 1969. However, during exten- 
sive collecting throughout Maryland in 1973-1975, 13 additional butterflies of 10 
species were seen being fed upon by ambush bugs, which evidently had captured 
the butterflies. In each case the prey and predator were collected and preserved. 
In one instance two ambush bugs were captured feeding on a single butterfly. Of 
the 15 ambush bugs collected, only 4 were nymphs. Each butterfly was on a 
flower head and probably was attacked while feeding. Except for two specimens, 
the butterflies were fresh or nearly fresh. The ambush bugs were identified as 
Phymata fasciata (Gray). The adults averaged 8.9 mm and the nymphs 5.5 mm 
in length. Data on the butterflies captured by the ambush bugs are tabulated in 
Table 1. 

From these data it can be surmised that captures of butterflies by ambush bugs 
may be quite common. Although this behavior is referred to in older texts, it has 
been generally overlooked by collectors. Comstock, Comstock & Herrick (1895, 
A Manual for the Study of Insects. 19th ed., Comstock Publ. Co., Ithaca, N.Y.), in 
discussing ambush bugs, stated, “It overpowers and captures insects like cabbage 
butterflies, honey-bees and large wasps.” Lutz (1918, Field Book of Insects. 2nd 
ed., G.P. Putnam’s Sons, New York), in a similar discussion, stated, “it conceals 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


148 


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itself in flowers, where it captures various insects including large butterflies and 
even bees.” However, the more recent text of Borror & DeLong (1954, An Intro- 
duction to the Study of Insects. 3rd ed., Holt, Rinehart, & Winston, New York) 
did not list butterflies as prey of Phymatidae. 


ACKNOWLEDGMENT 


The author thanks Dr. Jon L. Herring, Systematic Entomoiogy Laboratory, 
U.S.D.A., for identifying the ambush bugs. 


Joun H. Fates, Ridge Road, Neeld Estate, Huntingtown, Maryland 20639. 


PAPILIO XUTHUS (PAPILIONIDAE) IN HAWAII 


The Hawaiian Islands have a very limited butterfly fauna—the only large butterfly 
is the monarch (Danaus plexippus (L.)), and medium-size species are limited to 
several species of Cynthia and Pieries rapae L. (Zimmerman, 1958, Insects of Hawaii. 
Vol. 7: Macrolepidoptera, Univ. of Hawaii, Honolulu). I was very surprised, there- 
fore, on 6 February 1975 to observe a cream and white swallowtail fluttering around 
hibiscus bushes planted in the lawn of a condominium two miles north of Kaanapali 
on the island of Maui. 

Although an attempt at hand capture failed, in the process of achieving a “near 
miss” I was able to tentatively identify the butterfly as Papilio xuthus L., a native 
of Japan and East Asia. Numerous other individuals were observed in the next few 
days, and the identification was confirmed when I obtained a net and captured a 
short series of specimens. Since P. xuthus is a member of the Rutaceae-feeding group 
of Papilios, an immediate search was made of local Citrus. Eggs, young larvae and 
pupal skins were found, and two individuals were reared on young citrus leaves 
(the larvae showed no interest in the tough older leaves). 

Because of the potential importance of P. xuthus as a citrus pest, I notified the 
Hawaiian Department of Agriculture in Honolulu by telephone and was informed 
that they were aware of the introduction. In a subsequent letter, Ronald Mau, 
Survey Entomologist for the Department, kindly sent me the following information. 
Papilio xuthus was first discovered on Oahu in April 1971 and has now reached 
all the major Hawaiian Islands. It was first observed on Maui in May 1974 (I 
would guess that there was only a single introduction there since 10 months later 
the generations still appeared to be synchronized). 

A chalcoid wasp egg parasite, Trichogramma spp., according to Mau appeared to 
be giving good control, and a tachinid fly, Exorista sorbillans (Wiedmann), parasitic 
on the larvae has been introduced to supplement Trichogramma. Its successful estab- 
lishment is not certain at present. Mau also reports that P. xuthus pupae underwent 
some type of obligate diapause in 1971 and 1972, although adult and larval activity 
did not cease entirely. 

More observations on the behavior of this newly introduced species would be 
most interesting. In March 1975 on Maui, P. xuthus adults were abundant and large 
numbers of eggs and larvae were present on cultivated citrus, showing every sign 
that the population was in the “log phase” of growth. There are native Rutaceae 
in the Hawaiian flora of the genera Ptelea, Platydesma and Zanthoxylum, and the 
impact of P. xuthus on these should be carefully observed. 

It is interesting to speculate on the source of the introduction. One possibility 
that has been suggested is the accidental importation of a gravid female or two, 
but it is difficult to visualize such an “accident” with a large and attractive insect 
like P. xuthus. There is a more intriguing possibility. Papilio xuthus is known from 
the Bonin and Marianna Islands (Shirézu, 1960, Butterflies of Formosa in Colour. 


150 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Hoikusha, Osaka). The only native Hawaiian lycaenid, Vaga blackburni (Tuely), 
has its nearest relative V. ogasawaraensis (Pryer) on the Bonin Islands. Zimmerman 
(op. cit., p. 494) suggests the possibility that “blackburni is a derivative of ogasa- 
waraensis, or that they are descended from a common stock, and the original 
ancestors which populated Hawaii may have come from the Bonin Islands.” So 
it is just possible that this beautiful addition to the Hawaiian fauna arrived by 
natural dispersal, carried more than 3000 miles by the westerly winds of the 
North Pacific. 


Pau. R. Exnriicu, Professor of Biology, Stanford University, Stanford, California 
94305. 


CHARAXINAE (NYMPHALIDAE): OLD WORLD VERSUS NEW 


The life history of Anaea eurypyle confusa Hall (Muyshondt 1974, J. Lepid. Soc. 
28: 306-314) reveals some interesting differences between this species and the 
Old World Charaxinae. First, concerning the adults, is it quite certain that both 
sexes are attracted to dung and carrion? In the Old World species, only the male 
is so attracted, although both sexes are attracted to fermenting fruit and sap. 
It has been suggested that the male’s attraction to dung and carrion may have 
some connection with the male metabolism with regard to courtship and mating. 
This is comparable, perhaps, to the requirements of certain male danaids for the 
juices of withered plants of the Boraginaceae as a precursor of the male pheromones 
used in courtship. 

With regard to the larvae, I know of no Old World species in which the young 
larva constructs a perch of frass and silk on which it rests between feeds, and, as 
can be seen from Dr. van Someren’s list of foodplants (1974, J. Lepid. Soc. 28: 
315-331), the East African Charaxinae have been bred extensively. The Old World 
larvae usually spin a carpet of silk, without frass, on which they rest between feed- 
ing. I also know of no Old World species that makes a retreat by rolling or folding 
a leaf. Strangely enough, the young larvae of the genus Asterope (Eunicinae) and 
Cyrestis camillus F. (Marpesiinae) do have this habit of constructing a perch of 
silk and frass on which to rest, but none of these rolls or folds a leaf when larger. 

Generally, the pupae of the Old World Charaxinae are green and monomorphic, 
but those of Charaxes protoclea Feisth. and C. lasti Gr. Sm. are dimorphic, having 
a green and a brown form. It seems highly probable that the pupae of the closely 
allied C. boueti Feisth. and C. cynthia Btlr. are dimorphic also. 


D. G. Sevasroputo, P.O. Box 95026, Mombasa, Kenya. 


VoLuME 30, NuMBER 2 151 


NOTES AND NEWS 


Recent Letters 


Dear Sir, 


A paper by Benton and Jennings (1975, Lepid. Soc. 29: 192-194) ... . gives 
a photographic figure of a pupal anomaly. It also mentions in the text examples 
of hysterotely by Cockayne (1927) and Haggett (1954) with the comment “None 
of these cases is similar to the R. neomexicana pupal anomaly.” 

The LITERATURE CITED, at the end of the paper, omits several quite important 
references, among them: 


Hawkins, C. N. 1933. Trans. Roy. Ent. Soc. Lond. 81: 223. 
. 1938. Proc. Roy. Ent. Soc. Lond. 13: 92. 

Evans, E. 1940. Entomologist 73: 134. 

Cockayne, E. A. 1942. Ibid. 75: 49. 

Classey, E. W. 1942. Ibid. 75: 151. 


In the last mentioned paper, figures are given of the anomalies of segmentation 
in the larva, pupa and adult of the same specimen. 

From the figure of the pupa, it will be seen that the segmentation is very closely 
similar to that of Rhyacionia neomexicana shown in Benton and Jennings’ paper. 
The main differences are that an extra segment is involved and the spiral in the case 
of Classey (1942) for Malacosoma neustria Lin. is sinister and in the other, dexter. 


E. W. Classey 


Dear Dr. Godfrey, 


The pupa described and figured in Benton & Jennings’ paper (1975, J. Lepid. 
Soc. 29: 192-194) appears, from the photograph, to be a fairly typical example 
of spiral segmentation, the spiral affecting the 2nd, 3rd and 4th abdominal somites. 
The authors, whilst referring to Cockayne’s paper on prothetely and _ hysterotely, 
quite different phenomena, rather strangely make no mention of Cockayne’s papers 
dealing specifically with spiral segmentation (1929, Trans. Ent. Soc. Lond. 77: 
177-184 and 1934, Ibid. 82: 165-172). In these papers Cockayne describes and 
figures spirals in a number of insects belonging to various Orders, and discusses 
the probable origin and cause of the phenomenon. He is definitely of the opinion 
that it arises during the development of the ovum. There is a later paper by 
Cockayne (1942, Entomologist 75: 49-54) in which he records further cases that 
had come to light since his 1934 paper, and in which he records his opinion that 
the condition has a genetic basis, the gene probably being recessive and with very 
poor penetration. 

Since these papers were published, I can trace very few accounts of spiral seg- 
mentation in the English language journals available to me. Classey (1942, 
Entomologist 75: 151-152) describes and figures an example in the lasiocampid 
Malacosoma neustria L., and there are three papers dealing with examples that I 
have bred, viz. a brood of Leucania irregularis Wlk. (Noctuidae) (1948, Entomologist 
81: 38-40), a brood of Euxanthe wakefieldi Ward (Nymphalidae) (1965, Entomolo- 
gist 98: 107) and a single Acherontia atropos L. (Sphingidae) (1966, Entomologist 
98: 107). 


D. G. Sevastopulo 


152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


OBITUARY 


MAURICE LOUIS BRISTOL (1890-1975 ) 


Maurice Louis Bristol, a charter member of The Lepidopterists’ Society, died at 
his home in Elgin, Illinois on 13 February 1975. He was born in Naperville, Illinois, 
23 March 1890, and moved to Elgin in 1906. He was employed by the Elgin 
National Watch Company and later by the David C. Cook Publishing Company 
until his retirement. He never married and is survived by two brothers, Harris 
and George, and two sisters, Mrs. Hattie Graves and Mrs. Lucy Miller. 

Bristol began to collect butterflies as a boy of eight. He was an all-around 
naturalist of the old school, whose interests in addition to Lepidoptera included 
Coleoptera, birds, fossils, nature photography and plants, especially native prairie 
species, which he grew in his back yard. When the Audubon Society of Kane 
County, of which he was also a charter member, reconstructed a small patch of 
native prairie north of Elgin, he donated many of the plants for it, and it was 
named in his honor. He actively encouraged young people in developing an interest 
in nature and was well known in Elgin for his activities and his devotion to children. 

Bristol’s collecting in the Elgin area yielded such Illinois rarities as Glaucopsyche 
lygdamus couperi Grote and Chlosyne harrisii (Scudder ), as well as most of the known 
Illinois specimens of Calephilis muticum McAlpine. He rediscovered the classic 
Wakelee, Michigan type locality of Euptychia mitchelli (French) in the late 1920's 
and collected there regularly for many years. His particular interests were Apantesis. 
Catocala, Schinia, Papaipema and Theclinae. He maintained an extensive corre- 
spondence and exchange with other lepidopterists. 

I remember encountering Maurice at Wakelee on my first visit there a number of 
years ago. Then nearly 70, he was collecting with an energy that I, less than half 
his age, envied. He retained that vigor and continued active field work until the 
end of his life; at the time of his unexpected passing he was busily planning for 
his next season. His collection has been acquired by the Illinois Natural History 
Survey. Unfortunately, some of it was found to have suffered dermestid damage, 
but the greater portion was intact and will be incorporated into the Survey's 
research collection. 

The assistance of Bristol’s friend and collecting companion Irwin Leeuw, and his 
sister, Mrs. Hattie Graves, in the preparation of this article and in arranging for 
the transfer of his collection to the Survey is gratefully acknowledged. 


Roperick R. Irwin, Illinois Natural History Survey, Urbana, Illinois 61801. 


EDITORIAL STAFF OF THE JOURNAL 
GrorcE L. Goprrey, Editor 


Illinois Natural History Survey, Natural Resources Building 
Urbana, Illinois 61801 U.S.A. 


WituiaM H. ALLEN, Associate Editor JAMEs G. STERNBURG, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study 
of Lepidoptera. Contributors should prepare manuscripts according to the following 
instructions. 

Text: Manuscripts should be submitted in duplicate, and must be typewritten, 
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11 
inch paper. Titles should be explicit and descriptive of the article’s content, including 
the family name of the subject, but must be kept as short as possible. The first men- 
tion of a plant or animal in the text should include the full scientific name, with 
authors of zoological names. Insect measurements should be given in metric units; 
times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). 
Underline only where italics are intended. References to footnotes should be num- 
bered consecutively, and the footnotes typed on a separate sheet. 

Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically 
under the heading LirErature Crrep, in the following format: 


SHepparp, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, 
London. 209 p. 
196la. Some contributions to population genetics resulting from the 
study of the Lepidoptera. Adv. Genet. 10: 165-216. 


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

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

Tables: Tables should be numbered consecutively in Arabic numerals. Headings 
for tables should not be capitalized. Tabular material should be kept to a minimum 
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approximate desired position indicated in the text. Vertical rules should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
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the proofs. 

Page Charges: Authors with grant or institutional funds are requested to pay a 
charge of $27.50 per printed page (including tabular and black-and-white illustrative 
material) for articles up to 20 pages in length. This charge may be waived in the 
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be requested to pay this charge for material in excess of 20 printed pages. 

Address all correspondence relating to the Journal to the editor. Material not 
intended for permanent record, such as current events and notices, should be sent to 
the editor of the News: Ron Leuschner, 1900 John Street, Manhattan Beach, 
California 90266. 


ALLEN PRESS, INC. PRINTED LAWRENCE, KANSAS 
USA 


CONTENTS 


A Srupy OF THE PENINSULAR FLornmDA POPULATIONS OF THE MONARCH 
BUTTERFLY (DANAUS P. PLEXIPPUS; DANAWDAE). F. A. 
Urquhart and N. R. Urquhart ee 

FucitivE Coton IN THE MALes oF CERTAIN PieripAE. Harry K. 
Clench oo 

LarvaL More VARIATION IN CHLOSYNE LACINIA (NYMPHALIDAE). 
Raymond W. Neck _..- 

PORTABLE OutTpoor CAGcEs FOR MATING FEMALE GIANT SILKWORM 
Morus (SaTurNUDAE). Thomas A. Miller and William J. 
Cooper 0 

LARVAL FOODPLANTS, SPATIAL AND TEMPORAL DISTRIBUTION FOR FIVE 
SKIPPERS (HESPERIDAE) FROM Texas. Roy O. Kendall and 


MA.’ Rickard esc 2 i Oe nC ae Me! / 
COLLECTING COCOONS OF CALLOSAMIA SECURIFERA (SATURNIDAE). 
Richard 8S, Peigler | 
Winc Cotor VARIATION IN CALLOSAMIA (SATURNUDAE). Richard S. 
Peigher 0 8 
THE GENUS COPABLEPHARON IN TEXAS, WITH DESCRIPTION OF THREE 
New Species (Noctruwae). André Blanchard ______ 


NATHALIS IOLE ( PIERIDAE) IN THE SOUTHEASTERN UNITED STATES AND 
THE Banamas. Harry K. Clench 0. 
OBSERVATIONS OF EUPACKARDIA CALLETA IN SOUTHERN TEXAS (SAT- 
URNIDAE). Themas A. Miller. 
EXMERGENCE OF HYALOPHORA CECROPIA (SATURNIIDAE) BLOCKED BY 
SEEDS IN THE Cocoon VALve. G. P. Waldbauer and J. G. 
Sternburg 


DESCRIPTIONS OF THE IMMATURE STAGES AND BIOLOGY OF VITULA 
LUGUBRELLA (RaAGONOT) (PyYRALIDAE: PuyciTinAE). H. H. 
Neunzig 


GENERAL NOTES 


A gyandromorphic Lymantria dispar (Lymantriidae). Joseph Muller 
Mass hibernation site for Nymphalis vau-album (Nymphalidae). Noble 
8, Proctor to nO Ae OO oi a a 
Summer monarch (Danaus plexippus) in southern Texas (Danaidae). 
Raymond W. Neck io ee 
Libytheana bachmannii (Nymphalidae) in Connecticut. Noble S. Proctor 
A rahe for marking butterflies in eapture-recapture studies. Henry S. 
PRU a le A Ce Oe ea 0h nh rrr 
Papilio troilus (Papilionidae) in pursuit of fish crow. Noble S. Proctor 
a of butterflies as prey for ambush bugs (Heteroptera). John 
UR a NN i 
Papilio xuthus (Papilionidae) in Hawaii. Paul R. Ehrlich 
Charaxinae (Nymphalidae): Old World versus New. D. G. Sevastopulo 
Book Review 


73 
88 


91 


95 


105 
111 
114 
116 
121 


127 


131 


133 


Si A tg Ra fa 
Se ee ee 


SSS Ss 


ieee aS SSS 


& 


vw Oe eee ee 


Volume 30 1976 Number 3 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


¢. hodges 


22 September 1976 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


S. S. Nico.ay, President Cartos R. BEUTELSPACHER, 

MrmiamM RoruHscuitp, Ist Vice President Vice President 

THEODORE D. SARGENT, Vice President Joun M. Sniper, Treasurer 
Lee D. Miter, Secretary 


Members at large: 


J. T. BREWER D. F. HArpwIck R. A. ARNOLD 
K. S. Brown J. B. ZiEGLER E. D. CasHatr 
K. W. Pxinip F. S. CHEw R. E. STANFORD 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and 
formally constituted in December, 1950, is “to promote the science of lepidopterology 
in all its branches, . . . . to issue a periodical and other publications on Lepidoptera, 
to facilitate the exchange of specimens and ideas by both the professional worker and 
the amateur in the field; to secure cooperation in all measures” directed towards 
these aims. 

Membership in the Society is open to all persons interested in the study of 
Lepidoptera. All members receive the Journal and the News of the Lepidopterists’ 
Society. Institutions may subscribe to the Journal but may not become members. 
Prospective members should send to the Treasurer full dues for the current year, 
together with their full name, address, and special lepidopterological interests. In 
alternate years a list of members of the Society is issued, with addresses and special 
interests. There are four numbers in each volume of the Journal, scheduled for 
February, May, August and November, and six numbers of the News each year. 


Active members—annual dues $13.00 
Student members—annual dues $10.00 
Sustaining members—annual dues $20.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $18.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
John M. Snider, 3520 Mulldae Ave., San Pedro, Calif. 90732 U.S.A. 


Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA 


by Cyrit F. pos Passos 


Price: Society members, $5.00 U.S.; non-members, $7.50 U.S. Paper covers, revisions 
of the Melitaeinae and Lycaenidae supplied separately. 


Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni- 
versity of Louisville, Louisville, KY 40208, U.S.A. 


The Lepidopterists’ Society is a non-profit, scientific organization. The known 
office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second 
class postage paid at Lawrence, Kansas, U.S.A. 66044. 


JOURNAL OF 


Tue LEPIDOPTERISTS’ SOCIETY 


Volume 30 1976 Number 3 


THE OVERWINTERING SITE OF THE EASTERN POPULATION 
OF THE MONARCH BUTTERFLY (DANAUS P. PLEXIPPUS; 
DANAIDAE) IN SOUTHERN MEXICO 


F. A. URQUHART AND N. R. URQUHART 
Scarborough College, University of Toronto, West Hill, Ontario, Canada MIC 1A4 


Twenty-five years have passed since we started an intensive investi- 
gation of the movements of the monarch butterfly (Danaus p. plexippus 
(L.)) in North America, although experiments concerned with methods 
of identifying specimens of a moving population were conducted from 
1937-1939. The question to be answered was: “Where is the final 
destination of the migrating monarchs?” 

Over the years we have had the assistance of thousands of volunteers 
living in various parts of North America. Over 300,000 monarchs were 
alar tagged (Urquhart, 1960, 1966). In addition, through numerous 
field expeditions on which we traveled more than 100,000 mi. during 
the 25-year period, we investigated the presence of monarchs (larvae 
and imagoes) and the host plants in most parts of southern Canada, 
the United States, Mexico, and Central America. Some data were 
obtained as a result of visits to 56 colleges and universities to examine 
insect and plant collections and discuss the project with our colleagues. 

In January 1975, the overwintering colony of the monarch was finally 
located. We were able to join an expedition in January 1976 to view 
for ourselves this most spectacular sight—the final southern destination 
of the eastern population of the migrating monarch butterflies. 


Description of Site 


The overwintering colony described in the present paper was located 
on the slope of a volcanic mountain situated in the northern part of 
the State of Michoacan, Mexico, at a height of slightly over 3,000 m 
(Fig. 1). The predominant tree, averaging 26 m in height, was the 


154 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


\ ae 


4 
ery ~— 


Fig. 1. A, area over which monarch butterflies of the eastern population are 
known to breed; B, area of greatest concentration around the Great Lakes; C, over- 
wintering site of the eastern population; D, location of migrants in southeastern 
Mexico and Guatemala (Urquhart & Urquhart 1976); E, aberrant migration com- 
posed of a small portion of the eastern population that reaches the base of the 
Florida peninsula and continues southward; F, possible route taken by the aberrant 
Florida migrants. Note: the breeding area of the western population lies within 
the mountain valleys with overwintering on the California coast. 


Oyamel (Oyamel mexicano (Dougl.)). The colony occupied an area of 
roughly 20 acres. The trunks and branches of the Oyamel trees were 
completely covered with monarchs (Fig. 2)—only small portions of the 
crowns of the trees were bare as a result of the action of the mountain 
winds dislodging the butterflies. The monarchs were so closely packed 
together on the trunks that it was impossible to see even a small portion 
of the underlying bark. The branches were not only completely covered, 


VoLuME 30, NUMBER 3 155 


Fig. 2. Clusters of monarch butterflies on three of over a thousand Oyamel 
trees in the overwintering site. The trunks of the trees are also completely covered. 
Because of higher temperatures and exposure to the sun through a forest clearing, 
the monarchs in this photograph have spread their wings. During the evening and 
throughout the night and early morning, the wings remain closed. Color photographs 
appear in an article in the National Geographic Magazine (August 1976). 


156 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


from the smallest terminal twig to the base, but the butterflies clung 
to each other to a depth, in some instances, of 10 cm. One such branch, 
ca. 2 m in length with a base 4 cm in diam., was so heavily laden with 
butterflies that it broke, spilling countless thousands of monarchs upon 
the ground beneath the tree. The forest floor throughout the area 
was sprinkled with monarchs that, having been dislodged from the 
trees, were unable to fly because of the low temperatures at this alti- 
tude. In some locations the ground was covered to an average depth 
of 10 cm. It was impossible to walk over the area without crushing 
hundreds of monarchs underfoot, so, to avoid such wanton destruction, 
the field party limited its tagging activities to one small area near a 
forest clearing. 

During the early part of our visit (January 7-21) the temperature 
on the mountain slope varied from a low of -1° C at night and in the 
early morning to a high of 13° C in the afternoon. During cloudy 
periods and periods of rain or snow, the daytime temperature remained 
at an average of 10° C. In the cool periods the monarchs remained 
inactive with wings closed. With the advent of higher temperatures, 
many of them, particularly in areas where the sun shone through the 
Oyamel branches, opened their wings, presenting a magnificent display 
of color. At such times thousands would take to wing, filling the air 
over the forest clearing. As the temperature dropped, the monarchs 
returned to the trees and remained inactive during the night and early 
morning. Occasionally there was a slight snowfall. Monarchs shaken 
from the Oyamel branches were found fluttering on the snow surface. 
With the advent of warmer temperatures, the light snow cover would 
melt and the monarchs would return—either by flight, in areas where 
the sun could reach them, or by crawling—to roosting sites—trunks and 
branches of the Oyamel trees, bushes, or upright dead branches of 
fallen trees. 

To estimate the possible number of monarchs roosting upon one small 
Oyamel branch, the following procedure was conducted. A small branch, 
1.5 m in length, weighed down by its burden of butterflies, was located 
2m from the ground surface. The distance from the ground to the 
tip of the branch was measured. The roosting monarchs were then 
removed, which allowed the branch, relieved of its burden, to return 
to its unweighted position. One end of a cord was tied to the branch, 
about % from the tip, and the other end to a bag. A number of stones 
were placed in the bag, which caused the tip of the branch to return 
to its monarch-laden position. On returning to our base, we were able 
to weigh the bag and its content of stones, which came to 3,800 gm. 


VoLuME 30, NUMBER 3 Tey 


Knowing the weight of one butterfly to be 0.61 gm, based on the 
weight of 50 specimens of both sexes, the estimated number of monarch 
butterflies on this small branch is calculated to be 6,230. Since over 
a thousand trees, with branches much larger than the experimental one, 
together with the trunks and the forest floor, were covered with butter- 
flies, some conception of the number of overwintering monarchs in the 
area can be imagined. An accurate estimate is, of course impossible, 
but the numbers would undoubtedly be in the millions. 

On our last trip to the site, January 24, the afternoon temperature 
had risen to 22° C. Thousands of monarchs had left the trees and 
were flying about, filling the air with fluttering wings, a sight long 
to be remembered. A number of matings were taking place—pairs flew 
from one tree to another, and clusters of males and females copulated 
on the ground. As we climbed the slope of the mountain, we witnessed 
thousands upon thousands of monarchs heading on a compass bearing 
of true north. We had witnessed the first spring movement back to 
the breeding areas of the United States and Canada. 

Two members of the field party returned to the site on February 9 
to find that an estimated 75% of the butterflies had left the colony. 
Trees that were previously covered with monarchs were bare. 


Predators 


A few stray cattle were seen feeding extensively upon the monarchs 
that were on the ground and the lower portions of the tree trunks. 
The natives informed us that the cattle commonly fed upon monarchs 
and grew fat on the diet. 

In addition to the predation by cattle, a number of species of birds 
fed upon the roosting monarchs. We procured one bird (later identified 
as Myiarchus tyrannulus, the Brown-crested Flycatcher) and found its 
crop and stomach filled with the bodies and a few wing scales of 
monarchs. The wings of the monarchs had been removed by the birds 
before ingesting the bodies. It seems reasonable to conclude that other 
birds did the same thing because the ground was covered with monarch 
wings that we saw falling from the trees and fluttering to the ground 
like so many dead leaves. 


Tagging Program and Results 


Three monarchs that bore our identifying alar tags were recovered 
at the overwintering sites. During the expedition of January 1975, 
one specimen tagged in Nevada, Missouri on 9 September 1974 and 
another tagged in Fredericksburg, Texas on 8 October 1974 were re- 


158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


captured. During the January 1976 expedition one specimen tagged 
at Chaska, Minnesota on 6 September 1975 was recaptured. Such 
recaptures indicate that the site here described is the overwintering 
area for the monarch butterflies of the eastern population of North 
America. 

During our field trips to the site, a total of 25,000 specimens were 
alar tagged. In view of the fact that the specimens at the site were 
in excellent condition with very little indication of wing fractures, we 
are confident that tagged monarchs from the overwintering colony will 
eventually be recaptured at various localities throughout the United 
States and Canada, thus indicating a return migration for at least a 
part of the eastern population. Such recaptures will be reported in 
subsequent publications. 


ACKNOWLEDGMENTS 


The study presented in this paper is part of a long-term investigation 
of the migrations and ecology of the monarch butterfly sponsored through 
grants from the National Research Council of Canada and the National 
Geographic Society of the United States, and donations from members 
of the Insect Migration Association. Permission to use the photograph 
of the overwintering monarchs on the Oyamel trees was granted by the 
National Geographic Magazine. The final discovery of the site was 
made possible by the active cooperation of thousands of volunteer field 


assistants living in various parts of Canada, the United States, and 
Mexico. 


LITERATURE CITED 


Urgumartr, F. A. 1960. The monarch butterfly. Univ. Toronto Press, Toronto. 
B61" p: 

1966. A study of the migrations of the Gulf Coast populations of the 
monarch butterfly (Danaus plexippus L.) in North America. Ann. Zool. 
Fennici 3: 82-87. 

Urgunart, F. A. & N. R. Urqgumarr. 1976. <A study of the Peninsular Florida 


populations of the monarch butterfly (Danaus p. plexippus; Danaidae). J. 
Lepid. Soc. 30: 73-87. 


VoLuME 30, NUMBER 3 ; 159 


NOES ON THE LIFE CYCLE AND NATURAL HISTORY OF 
BUTTERFLIES OF EL SALVADOR. VIII. ARCHAEOPREPONA 
ANTIMACHE GULINA, SIDERONE MARTHESIA, ZARETIS 
CALLIDRYAS AND CONSUL ELECTRA (NYMPHALIDAE) 


ALBERTO MUYSHONDT 
101 Avenida Norte #322, San Salvador, El Salvador 


This article completes a series dealing with what my sons and I have 
learned about the early stages, larval food plants, and adult behavior 
of the Charaxinae found in El Salvador. As far as we know, the re- 
maining local species of this group for which we have been unable to 
find the early stages, even though we have collected adults, are Anaea 
(Anaea) aidea Guerin-Méneville, A. (Memphis) nobilis Bates and A. 
(M.) glycerium Doubleday. Another species of Archaeoprepona occurs 
here, but we unfortunately have been unable to capture an adult to 
have it determined. All these species are very scarce locally; only one 
or two specimens of some are known in local collections (including 
Ours ). 


Life Cycles 


Archaeoprepona antimache gulina Frihstorfer 


This species is like A. demophon centralis Frihstorfer (Muyshondt, 1976) except 
for the following points: 

Larva (Figs. 2-6). Dark brown dorsally during Ist and 2nd instars; 3rd instar 
lighter brown with a whitish spiracular area on 7th and 9th abdominal segments; 
5th instar larva whitish dorsally from 3rd—9th abdominal segments, rest of body 
brown. During all instars the mesal thoracic prominence is larger than in respective 
instars of A. d. centralis. 

Prepupa (Fig. 7). Prepupa changes to yellow instead of green. 

Pupa (Figs. 8,9). Pupa slightly more slender laterally than pupa of A. d. centralis. 

Adult (Figs. 10, 11). Same shape as A. d. centralis but colored triangle on dorsal 
surface of forewing and band on dorsum of hindwing are definitely iridescent light 
blue instead of iridescent greenish-blue. Ventral surfaces of wings mostly light gray 
instead of light brown with a complicated array of darker zones and black lines, 
more abundant than in A. d. centralis. The wingspan and developmental time are 
about the same as A. d. centralis. 


Siderone marthesia Cramer 


In all respects, this species is like Zaretis itys Cramer (Muyshondt, 1973b) but 
for the following: 

Egg. Considerably larger, but same shape and color. 

Larva (Figs. 12-16). Corresponding instars about 114 times larger, lighter brown 
than Z. itys; caudal end rounded instead of bifid. Later instars with a white patch 
formed by short and thin spinulets subdorsally on mesothorax. Head horns very 
thick and knobby; head and its horns covered with short setae. 


160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-6. Archaeoprepona antimache gulina. 1, Egg, width about 2.5 mm; 2, 
second instar larva, about 13 mm long; 3, third instar larva, about 30 mm long; 
A, fourth instar, about 47 mm long; 5, head of fifth instar larva, note lateral pro- 
jections; 6, fifth instar larva, about 80 mm long. 


Pupa (Figs. 18-20). Bigger than but about the same shape as pupa of Z. itys. 
Green with dark markings along the spiracular zone and on the lateral edges of the 
wingcases. The latter have a marked indentation, lacking in Z. itys. 

Adults (Figs. 21, 22). Sexually dimorphic. Wing of male: costal margin convex, 
apex pointed, outer margin S-shaped, tornus rounded and inner margin of forewing 
straight. Costal margin of hindwing strongly rounded basally, outer angle and 
margin rounded, anal angle slightly projecting, inner margin S-shaped. Female: 
larger with costal margin of forewing less convex than in the male, apex without a 
marked point, outer margin slightly convex, forewing not S-shaped. Hindwing as 
in the male. Dorsal color pattern of the male: forewing dark brown, almost black, 
with a slanting red bar from the mid-costal margin almost to the tornus and a red 
triangle basally, leaving a black band in between. Male hindwing mostly black or 
very dark brown with red markings of variable size starting from mid-costal margin, 
markings sometimes reduced to two or three spots or forming a band running toward 
the anal angle. Apex of front wing in some individuals red-tinged and with a very 
small white spot near costal margin of the subapical area. Females have same color 
pattern, but lighter brown with two subapical white spots on forewing and red 
markings of hindwing greatly reduced. Ventral wing surfaces of both sexes very 


VoLUuME 30, NUMBER 3 161 


Figs. 7-11. Archaeoprepona antimache gulina. 7, Prepupa, note tails located on 
either side of twig; 8, pupa, dorsal view, about 42 mm long; 9, pupa, ventral view; 
10, male, dorsal view, wingspan about 80 mm; 11, male, ventral view. 


similar but darker in male: ground color dark brown with darker lines resembling 
leaf veins and lighter zones, the latter more evident on hindwing. Apex of forewing 
with some white sprinkling. The whole of ventral wing surfaces gives silvery sheen 
at certain angles. Thorax reddish above, yellowish below; abdomen brown. Antennae 


light brown; palpi and proboscis yellow. Wingspan of female about 7.0 cm, male 
5.0 cm. 


Zaretis callidryas R. Felder 


A second instar of Z. callidryas was collected and at first thought to be a difterent 
morph of the common Z. itys. The larva was reared on Casearia silvestris and C. 
nitida. During the later instars differences were noticed and photographed: 

Larva (Fig. 23). Abdomen lighter dorsally than Z. itys; head smoother, without 
the typical Z. itys spines and the horns incurved posteriorly. 

Pupa. Closely resembles the green morph of Z. itys. 

Adult. The obtained adult was a tailed female. Its shape similar to Z. itys except 
for tail on M-3 of hindwing and less acute anal angle. Dorsal color predominantly 


162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 12-14. Siderone marthesia. 12, Fourth instar larva, about 32 mm long; 13, 
same, dorsal view; 14, fifth instar larva, two days after molting, about 44 mm long. 

Fig. 15. Left, Zaretis itys, fifth instar larva; right, Siderone marthesia, fifth instar 
larva, about 62 mm long. Compare caudal end of Zaretis with Siderone. 

Figs. 16-17. Close-up of larval heads: 16, Siderone and 17, Zaretis. 


white with pearly sheen; apex of forewing faintly gray. Outer margin of hindwing 
with submarginal row of small faint brown spots. Ventrally wings yellowish-white 
with faint markings as in Z. itys. Wingspan about 5 cm. 


Consul electra Westwood 


In its early stages, behavior and selection of food plant, this species is similar to 
C. fabius (Cramer) (Muyshondt, 1974) except for the following noticeable dif- 
ferences: 

Larva (Figs. 26, 27). During 5th instar, head predominantly green with black 
near the horns and yellow frontal bands. In C. fabius the head is mostly black with 
yellow frontal bands. 

Pupa (Wigs. 28-30). Green and brown pupal morphs occur. Slightly wider lat- 
erally than in C. fabius; shows two dark spots dorsally on thorax. 

Adult (Figs. 31-38). Adults not showing marked sexual dimorphism, but are 
amazingly polymorphic. Apex of forewing more or less elongate and acuminate; M-3 


VoLUME 30, NUMBER 3 163 


1 22 


Figs. 18-22. Siderone marthesia. 18, Pupa, dorsal view, about 25 mm long; 19, 
pupa, lateral view; 20, pupa, ventral view; 21, male, dorsal view, wingspan about 
do mm; 22, male, ventral view. 


of hindwing more or less elongate and spatulate, anal angle sometimes forming second 
tail. Dorsal and ventral colors more or less dark with some variations in color 
pattern. Variations in shape and color exist between individuals reared simulta- 
neously, thus, not attributable to seasonal variation. Wingspan of female about 7.0 
cm, male 5.5 cm. 


Natural History 


Archaeoprepona a. gulina behaves during its early stages and as an 
adult very much like Prepona omphale octavia (Muyshondt, 1973a) 
and A. demophon centralis (Muyshondt, 1976). It looks very much 
like the latter except for variations in color. The larva of A. a. gulina 
also reacts in a menacing manner by lifting the forked tail which is 
opened and closed scissor-like when prodded. During the later instars 


164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 23-25. Zaretis callidryas. 23, Fifth instar larva, about 40 mm long, note 
incurved horns, compare with Z. itys, Fig. 15; 24, female (tailed morph), dorsal 
view, wingspan about 50 mm; 25, female, ventral view. 


it also has a reptilian aspect when viewed dorsally. This is due to a 
prominent dorsal projection which covers the head. The lateral aspects 
of the dorsal projection bear false eyes which enhance the effect. This 
prominence also is retractable. 

A. a. gulina larvae feed on a Laurinaceae tree determined by Lic. 
Flores, Universidad Nacional, as Nectandra sinuata, which is the same 
food plant used by the larvae of Anaea morvus boisduvali Comstock 
(Muyshondt, 1975). Actually it was while searching for A. m. boisduwali 
eggs that the first eggs of A. a. gulina were found. The eggs of the 
latter species are also heavily parasitized by chalcidid wasps, and the 
small larvae are preyed upon by spiders. A. a. gulina occupies the same 
habitat as A. demophon centralis. 

Siderone marthesia and Zaretis callidryas behave like Z. itys (Muy- 
shondt, 1973b) and share its food plants: Casearia sylvestris and C. 
nitida although Zaretis itys appears to prefer the latter and Siderone 
marthesia and Zaretis callidryas the former. Both S. marthesia and Z. 
callidryas have been collected at an altitude of 1,000 m. Z. itys has a 
wider range. The females of S. marthesia are very seldom seen. Most 
of our observations have been of adult males; they have definite hill- 
topping behavior between 1030-1600 hr. They usually perch on the 
top branches of trees, and it is quite a feat to net them. Luckily 
they are attracted by fermenting banana baits, but we have never found 
females at the baits. However, Descimon et al. (1973) report several 
captures of females at baits near Colima, Mexico. We are unable to 


VoLUME 30, NUMBER 3 165 


Figs. 26-34. Consul electra. 26, Fifth instar larva, about 37 mm long; 27, close- 
up of larval head; 28, pupa, dorsal view, about 20 mm long; 29, pupa, lateral view: 
30, pupa, ventral view. Emergence of adult: 31-34, sequenced from left to right. 


comment on the behavior of adult Zaretis callidryas because the only 
specimen we have seen is the one reared in our insectary. 

The behavior of the early stages and adult of Consul electra resemble 
those of Anaea (Consul) fabius Cramer (Muyshondt, 1974). The larvae 
of both use a number of species of Piperaceae as food plants. C. electra 
has been collected at higher altitudes than fabius (500-2,600 m). Both 


166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


37 38 


Figs. 35-38. Consul electra. 35, Female, dorsal view, wingspan about 70 mm; 
36, female, ventral view; 37, male (acute apex morph), dorsal view, wingspan 
about 55 mm; 38, male, ventral view. 


species can be found simultaneously feeding on the same plants where 
their ranges merge at an altitude of 500-1,200 m. The adults do not 
visit flowers, but feed on fermenting fruits and excrement on the ground 
and on sap from tree cavities. The males are pugnacious and perch 
on shrubs at a height of 2-4 m. With C. fabius, C. electra is one of 
the slowest flyers among the local Charaxinae. The larvae and pupae 
are often parasitized by tachinid flies and chalcidid wasps. 


DISCUSSION 


Miiller (1886) described the life cycles of various Archaeoprepona 
(under the generic name of Prepona), including A. amphimachus 
Fabricius on Laurinaceae (probably Nectandra spp.), A. catachlora 
Staudinger on Mollinedia brasiliensis Jul. (Monimaceae) and A. demo- 
phon extincta Staudinger on Dugetia lanceolata St. Hilaire (Anonaceae). 


VoLuME 30, NUMBER 3 167 


His descriptions conform closely with our own observations of A. demo- 
phon centralis (Muyshondt, 1976) and A. antimache gulina except for 
the eggs which he describes as having “very fine and crowded longi- 
tudinal ribs” which we have been unable to detect, even under 15x 
magnification. Miller also described the early stages of Siderone ide 
Hubner and S. strigosus Gmelin and claimed both species fed on Cas- 
earia sylvestris. However, Siderone strigosus, according to Comstock 
(1961), is a synonym of Anaea (Zaretis) itys. Furthermore, because 
Miller states that the early stages of “Siderone strigosus’ are “as ide,” 
it appears that his descriptions actually deal with two species of Zaretis. 

After studying species of both genera (Zaretis itys (Muyshondt, 
1973b), Z. callidryas, and Siderone marthesia), we are certain that 
there are notable differences between the early stages of the larvae of 
the two genera that would not escape the attention of an investigator. 
The only part in Miller’s description of “ide” that would fit a Siderone 
pupa is “wing edge and stigma brown, yet he does not mention a 
drastic indentation (as seen dorsally and ventrally) at the brown mark 
on the wingcase of the pupa he described. 

According to Comstock (1961), the only information about the early 
stages of A. (Siderone) marthesia is Miiller’s, however, we are certain 
that Miuller’s description was not of a species of Siderone. In Rydon 
(1971) there are some good painted illustrations of the larvae of S. 
marthesia and S. nemesis and their respective pupae copied from the 
original drawings of Miss M. E. Fontaine which were published by 
the Royal Entomological Society around 1910. However, Miss Fontaine 
apparently did not write any descriptions. 

We are unaware of any previous publications on the early stages of 
Zaretis callidryas. The adult is figured in Seitz (1924) as an Anaea 
and is not likened to its close relative, Zaretis itys. Even in Lewis (1973) 
the tailless form of Z. callidryas is presented as an Anaea. The same 
is true with Consul electra. Seitz (1924) and Lewis (1973) present it 
as Anaea electra and do not relate it to its close relative, Consul fabius, 
which they picture as Protogonius hippona Fabricius. 

We hope our descriptions and illustrations of the species we have 
studied will help the experts clarify and settle the apparent congeneric 
questions of the Charaxinae. 


ACKNOWLEDGMENTS 


We are greatly thankful to Drs. A. H. B. Rydon and H. Descimon for 
supplying us with a number of publications that helped document the 
present article; to Drs. James G. Sternburg and G. L. Godfrey for dedi- 


168 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


cating part of their valuable time to reading, criticizing, and suggesting 
improvements for the manuscript; to Viktor Hellebuyck for sharing his 
own observations of the species mentioned; and to Alberto, Jr., Marilynn 
and Pierre, members of the family, for their enthusiastic help. 


LITERATURE CITED 


Comstock, W. P. 1961. Butterflies of the American Tropics, the genus Anaea, 
(Lepidoptera, Nymphalidae). The American Museum of Natural History, N.Y. 
QIA p. 

peers H., J. M. pe Magscur & J.-R. SrorFeL. 1973. Contribution a létude 
des Nymphalides neotropicales. Description de trois nouveaux Prepona mexi- 
cains. Alexanor 8: 155-159. 

Hemminc, F. 1967. The generic names of the butterflies and their type-species. 
(Lepidoptera: Rhopalocera). Bull. British Mus. (Nat. Hist.), Entomology 
Suppl. 9. 509 p. 

Lewis, H. L. 1973. Butterflies of the World. Harrap Books, London; and Follett, 
Chicago. xvi + 312 p. 

MULLER, W. 1886. Siidamerikanische Nymphalidenraupen. Versuch eines natiir- 
lichen Systems der Nymphaliden. Zool. Jahrb. Zeitschr. Syst., Geogr., Biol. der 
Thiere 1: 417-678. 

MuysHonpr, A. 1973a. Notes on the life cycle and natural history of butterflies 
of El Salvador. I. Prepona omphale octavia (Nymphalidae). J. Lepid. Soc. 27: 
210-219. | 

1973b. Notes on the life cycle and natural history of butterflies of El 

Salvador. II. Anaea (Zaretis) itys (Nymphalidae). J. Lepid. Soc. 27: 294-302. 

. 1974. Notes on the life cycle and natural history of butterflies of El 

Salvador. III. Anaea (Consul) fabius (Nymphalidae). J. Lepid. Soc. 28: 

81-89. 

1975. Notes on the life cycle and natural history of butterflies of El 

Salvador. V. Anaea (Memphis) morvus boisduvali (Nymphalidae). J. Lepid. 

Soc. 29: 32-39. 

1976. Notes on the life cycle and natural history of butterflies of El 
Salvador. VII. Archaeoprepona demophon centralis (Nymphalidae). J. Lepid. 
Soc. 30: 23-32. 

Rypon, A. H. B. 1971. The systematics of the Charaxidae (Lepidoptera: Nymph- 
aloidea). Ent. Rec. J. Var. 83: 219-233, 283-287, 310-316, 336-341, 384-388. 

Seitz, A. 1924. Macrolepidoptera of the World. Vol. 5. Stuttgart. 


VoLuME 30, NUMBER 3 169 


PATTERNED PERCHING BEHAVIOR IN TWO CALLOPHRYS 
(MITOURA) (LYCAENIDAE) 


Kurt JOHNSON’ 


Department of Biology, City University of New York, Convent Avenue and 
West 138th Street, New York, New York 10031 


AND 
PeTeER M. Borco? 
Department of Marine Studies, University of Delaware, Newark, Delaware 19711 


Several species of Nearctic butterflies are well-known for the habitual 
attachment of their adults to the larval food plant, which serves as a 
substrate for a behavior popularly called “perching.” Notable among 
these are tree-feeding species such as the nymphalids Asterocampa celtis 
(Boisduval & LeConte) and A. clyton (Boisduval & LeConte) on hack- 
berry (Celtis spp., Urticaceae) and lycaenids of the taxon Callophrys 
(Mitoura), which feed on various Cupressaceae. Except for visits to 
food sources, adults of these species confine their flight activities to 
the immediate vicinity of trees, where they are often seen flying out 
from the branches and returning. Although these habits are often noted 
in popular studies and regional works, no published research has dealt 
with their details or nature. The term “perching” has been used as a 
general category in mate finding in butterflies (Scott, 1973), but such 
“perching’ probably merits distinction from the particular behavior 
displayed by the aforementioned tree feeders. 

The perching species Callophrys (Mitoura) siva (Edwards) and C. 
(M.) gryneus (Hibner) are exclusive Juniperus feeders, utilizing a 
broad spectrum of morphologically and chemically divergent Juniperus 
species that replace each other geographically over a wide area of the 
Nearctic region (Johnson, 1976). The present paper treats the tree- 
perching behavior of C. (M.) siva and C. (M.) gryneus as studied on 
two of their larval food plants, J. scopulorum Sarg. and J. virginiana L., 
respectively. The purpose is to outline the outstanding characteristics 
of the behavior, consider what environmental stimulus-response relation- 
ships are present, and comment on their significance. 


Perching Behavior and Environmental Stimuli 


Many aspects of insect behavior have been described in the literature 
and defined into types and categories. Most of these are not applicable 


1 Museum Research Associate, Museum of Natural History, University of Wisconsin, Stevens 
Point, Wisconsin 54481 and Research Associate, Allyn Museum of Entomology, Sarasota, Florida 
33580. 

2 Present address: 535 Park Avenue, Burlington, Wisconsin 53105. 


170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


to specific observations until thorough physiological data is available 
on a species. Thus, it is best to introduce perching behavior by re- 
viewing literature that concern characteristics of behavior which are 
observable in the field. 

Specific activities of insects result from the continuous “release” of 
fixed behavior patterns and pattern combinations, which are the insects’ 
response to evolutionarily defined stimuli within its niche. Stimuli may 
include extrinsic factors (e.g., gravity, light, temperature, humidity, 
air movements, sound, chemicals, food, mates and other biota) or in- 
trinsic ones (e.g., hormones, feeding rhythms, and mating rhythms). 
Stimuli are processed into particular released behavior through specific 
adaptations and interactions of sense receptors. A process may involve 
a single stimulus and a single receptor, screening of stimuli by a single 
receptor or balancing of stimuli by several receptors. Generally, many 
factors, each of which could release such fixed patterns of behavior, 
are working on many receptors (Jander, 1957). 

For the field investigator, one subject for study is the role of light. 
Two factors may be active, depending on the physiological adaptations 
of the insects: a relationship to the sun’s polarized light (sky light) 
and/or the position or intensity of the sun itself. 

Sky-patterns of polarized light are caused by the varying directions 
of oscillation of light as it is distributed from the sun. Insects may 
respond to polarized light directly (von Frisch, 1960), to its different 
patterns of reflection from the substrate (Kalmus, 1958; Smith & Baylor, 
1960) or to both (Jander & Waterman, 1960). Especially important 
to insects is the relationship in the sky of points of equal degrees of 
polarization. When connected by hypothetical lines, these points form 
mathematical planes, and it is to these planes that many insects have 
become sensorily adapted. In other words, orientation to sky points 
of certain degrees of polarization causes an insect to react along a 
certain plane in relation to the source of light and the environment. 
The most important of these planes for many insects is the plane formed 
by connecting the points of maximum polarization—70%. This plane 
runs perpendicular to the straight line from sun to earth, a line which 
is generally referred to as the electric or “e” vector of light. Reactions 
and orientations along this perpendicular plane have received much 
study, and four basic posturing positions have been shown to occur in 
insects: the parallel position (0° from “e” vector ), the vertical position 
(90° from the “e” vector) and the angular positions (+45° from the 
“e” vector) (Jander & Waterman, 1960). In polarization-adapted insects, 
the ratio of occurrence of these positions varies with species and factors 


VoLUME 30, NUMBER 3 we 


such as temperature (Willington et al., 1951; Henson, 1954) and excite- 
ment (Jander & Waterman, 1960). 

Reaction to the position and/or intensity of the light source itself 
involves phototaxis. Insects may be positively or negatively phototactic, 
and the degree of their response may vary with increase in brightness 
of the source and/or higher contrast between the source and environ- 
ment (Robinson & Robinson, 1950; Robinson, 1952), temperature or 
excitation (Mell, 1954; Marten, 1956), or the physiological tolerances 
of the sense organs (Verheijen, 1958). These reactions to polarized 
light or the sun itself can go on simultaneously in the environment. 

Gravity is another factor that often cooperates with functions per- 
formed by the visual organs (Schwartzkopff, 1965) and is usually sensed 
through proprioreception. It is best to consider the influence of gravity 
in separate data since it may involve co-actions of various receptors, 
elicit patterns of response independent of other observations, or even 
be observed as alternative positive and negative influences in other 
observed patterns (Markl & Lindauer, 1965). Statistical data that indi- 
cate the roles of all of these factors in Callophrys (Mitoura) behavior 
were discovered in the present study and are presented below. 


METHODS AND MATERIALS 


Field-condition studies are neither the only nor most ideal method for 
studying complex behaviors. The number of inferences that can be 
drawn concerning possible physiological relationships is limited (Gold- 
smith, 1965), especially since such inferences depend largely on distinct 
knowledge of the physiology of the insect’s sense organs (Hodgson, 
1965). However, appropriately designed experiments are useful for 
discovering good subject species, basic behavior patterns, and possible 
parameters for laboratory experiments (Mazokhin-Porshnyakov, 1969, 
pp. lar f, 134 f. and 213 f£.). In the present study, data samples 
totaling 471 randomly selected perch positions were gathered in 1973 
and 1974. Initially, to compare the behavior of C. (M.) gryneus and 
C. (M.) siva, two juniper habitats were selected where sample trees 
could fulfill the following requirements: nearly equal mean tree height; 
presence of extremes of height, but similar range; all trees adjacent; 
and a workable population of Mitowra. These sites were: off County 
Rd. 547, near Lakehurst Naval Air Station, Ocean Co., New Jersey, 
and Catholic Cemetery, Chadron, Dawes Co., Nebraska. Because of 
the desire to investigate the possible role of perching behavior in 
geographic distributions and food plant selection, an extremely northern 


172 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(apparently transplanted) C. (M.) gryneus colony was studied at New 
London, Merrimack Co., New Hampshire. Similarly, an effort was made 
to study the behavior in an arid area, where small juniper morphs occur, 
but this attempt (by a third co-worker) failed as a result of inclement 
weather. 3 

The trees numbered at these sites comprised a “base sample” from 
which trees to be used for treatments could be drawn by random 
numbers. Because of daily movements by the butterflies in the habitat, 
treatment trees were actually drawn from a “sub-sample” made up of 
the number of trees being utilized for perching before each treatment. 
When the “treatment tree” was selected, the “treatment insect’ was 
designated arbitrarily, and randomness was assured by measuring and 
recording data on five consecutive choices of perch following the initial 
flight from the tree. Motivation for flying was either natural movement 
or from nudging of the appropriate twig by the investigator. Early 
samples confirmed that such “aided” and “unaided” movements showed 
no significant mathematical differences in analysis. If the treatment 
insect changed trees, the change was noted and the sample continued. 
The time between tree changing allowed study of the types of trees 
chosen for perching. Accordingly, a “selected mean tree height” was 
calculated for each study area based on the number of times certain 
trees were chosen for perching. If an insect flew out of the area and 
the five-choice treatment could not be completed, the remaining perches 
were considered to be zero for all measurements and observations. 
Perch locations were designated by lifting a colored loop of string or 
wire to the perch site; tree and perch heights were measured with a 
graduated measuring rod. The measurement method was tested against 
known heights by the “t” test and showed no significant differences 
when used to the nearest centimeter up to 4 m. Above this height, 
measurements were rounded to the nearest decimeter. The latter method 
had to be used only in the supplementary samples mentioned below. 

For comparison, supplementary areas (medium-size trees at Tuckerton, 
New Jersey and tall trees at Blackbird State Park, Delaware) were 
sampled, some by the above method; other data were culled from earlier 
observations at Long Pine Recreation Area, Brown Co., Nebraska. These 
data were not used in the random sample analysis of measurements, 
but were enumerated. 

In this paper, any statistical value listed without an accompanying 
probability was not significant above p = 0.20, except in the few cases 
where extraordinarily high x? values were significant beyond question. 


VoLUME 30, NuMBER 3 iio 


MEAN (Lakehurst) 
MEAN (Chadron) 


HEIGHT (M) 
DO 


ee oe iS ie i? 1s PO. 20 
TIME OF DAY ( x100=Hr. ) 


Fig. 1. Heights of perch plotted against time of day at Lakehurst and Chadron. 


RESULTS 


Perch Location 


Perch location and sunlight. The role of sunlight showed general and 
specific consequences in the behavior. At Chadron and Lakehurst, of 
239 perches recorded, only 3 were not in direct sunlight (x? = 226.8), 
and at New London this ratio was 232/1 (x? = 229.0). These samples 
were always made in sunny or partly sunny weather, and although some 
activity by the insects occurs during overcast warm weather, it may 
involve stimulus-response mechanisms that are not comparable to those 
subject to this investigation. 


JourNAL OF THE LEPIDOPTERISTS SOCIETY 


MEAN = 218.9 


HEIGHT (M) 


PEINCnk 


@ 2 
& =other (n) 


9 VOT 12-3 14 15 6. 7 ee 
MS OF WAN ( x1OOShir. 


Fig. 2. Heights of perch plotted against time of day at New London. 


Heights of perch were plotted against time of day as in Figs. 1 and 2. 
The dots form an “hourglass-on-its-side” configuration characterized by 
greater dispersal of height of perch in morning and late afternoon- 
evening than at midday. This gradual shift in degree of dispersal was 
confirmed by correlation coefficients (cf. Fig. 3). These were also used 
to designate three “periods” apparent in the behavior: morning “ex- 
pansion” (0700-1245 hr), midday “compression” (1245-1445 hr) and 
late afternoon—evening “expansion” (1445-1900 hr) [all ST]. These three 
periods are significant because they demonstrate changes in the behavior 
in relation to the angle of the sun, which will be discussed further. 


VoLUME 30, NUMBER 3 ies 


poe] 
Ol 

=) 

“I © 
By) 
e 


Ov 
>) 


STANDARD DEVIATION 
SA © = G 
ee. = GS 


e SOMA Ze TS Pavol 5 
380 °@ TIME OF DAY ( x100=Hr.) 


DEVIATION FROM THE MEAN 


9 10 Tl 12 13 
TIME OF DAY ( x100=Hr.) 


Fig. 3. Correlation coefficients. (A) Deviation from the sample mean of each 
perch at New London (morning expansion period) correlated with time: r= —.143: 
Pas O= 05-5 = 145. Small) dot = 1; medium dot = 2: large dot =-3.- (B) 
Standard deviation of each treatment from the sample mean at Chadron (morning 
expansion period) correlated with time: r = —.897; p = .001; n = 11. Not shown: 
deviation from the sample mean of the mean perch height of each treatment at 
Chadron (evening expansion period ) correlated with time (scaled opposite direction) : 
io) = .0b; nm-= 20. 


176 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


FREQUENCY DISTRIBUTIONS 
N= sample size X= mean S= std. deviation 


60;-A SO 183 N=55 
X=284.5 
40 20 S=48.70 
20 
Mn=.95m Mx=3.56m Mn=1.90m Mx=3.81m 
100-— D N=232 


X=218.90 


80 = 
15 C N=52 S=26.50 
X=448. | 60 
10 S=106.90 


Mn=2.7m Mx=7.0m Mn=.63m Mx=2.90m 


Fig. 4. Frequency distributions of heights of perch: A, Chadron; B, Lakehurst; 
C, Long Pine; D, New London. Lakehurst, Chadron, and New London: 20 cate- 
gories, each 20 cm. Long Pine: 20 categories, each 35 cm. Maximum and minimum 
perch height are indicated. 


Perch location and height. Frequency distributions of heights of 
perches (Fig. 4) skewed right, which indicated preference for height. 
This indication was further confirmed by calculation of “selected mean 
tree height’ for each sample area, in which the height of each tree 
was multiplied by the number of times it was selected for perching 
and the mean was computed. These were always higher than the mean 
tree height of the sample. Furthermore, frequencies of heights of indi- 
vidual trees selected for perching, plotted against availability of these 
heights in the sample (Fig. 5), showed similar preference. Together, 
these data indicate that choice of perch as a whole forms a gradually 
shifting and somewhat symmetrical pattern in relation to time of day 
and is distinctly preferenced for heights. 

Perch location and tree surface. Percent of height of tree used per 
time of day was evaluated as in Figure 6. The entire figure (dots 
and hatches) shows the frequency of use of heights (in percent) for 


VoLuME 30, NuMBER 3 LG, 


60/;-A N=158 CHOICE OF HEIGHT IN 
- RELATION TO AVAILABILITY 
and THE SELECTED MEAN 
4 
fs ey 76 


SELECTED MEAN = 394.8m 


30;- B: N=55 , 

. 50 

10 4 Z, 
2 7 


SELECTED MEAN = 386. 8m SELECTED MEAN = 280. 9m 


Fig. 5. Frequencies of heights of individual trees selected for perching plotted 
against availability of these heights. Left scale (black frequencies): height category 
chosen for perching. Right scale (hatched frequencies): number of trees in the 
sample of each category. (A) Chadron, 20 categories, each 20 cm. (B) Lakehurst, 
20 categories, each 20 cm. (C) New London, 20 categories, each 17 cm. Trees 
in the base sample not chosen for perching are excluded. “Selected mean perch 
height” of each sample is indicated: the mean of—the height of each tree times 
the number of times chosen for perching. 


the entire day; the stippled areas show morning and late afternoon— 
evening expansion periods, and the hatches show the midday compres- 
sion period. The forming of two separate “humps” indicates that two 
interrelating stimulus factors are probably involved. Use of the tree 
in the compression period is uniformly undispersed (as also apparent 
on the perch height per time of day (Figs. 1, 2)) and occurs mostly 
high in the trees. This compression period occurs when the sun is high 
above the sample area and the plane of reaction to 70% polarized 
light is distributed horizontally along the tops and ascending angles of 
the trees. Height-use of the trees in the two expansion periods, how- 
ever, forms two humps (there is some variation in the two samples), 
with the second hump being lower on the trees. The expansion periods 
occur when the sun is low in the sky and the plane of reaction to 70% 
polarized light is distributed vertically over the sides and ascending 
angles of the trees. Actually, this plane is continually changing, rotating 
from the vertical obliquely to the horizontal and back as the day 
continues. Apparently this rotation dictates the configuration of the 
perch locations through the day. 


178 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Height 
| Spread 
2% Op 
{ tree 


Fa 20) oe OO 90 150n Gn meEmnnG 
FREQUENCY DISTRIBUTIONS 


Fig. 6. Evidence of two factors influencing perch height: percent height of tree 
utilized per time of day. See text for discussion. 


Behavioral differences between the species. It is especially interesting 
that the samples of C. (M.) siva in Nebraska and C. (M.) gryneus in 
New Jersey could be lumped together for this study. The study areas 
were chosen with this in mind. Early comparisons of “heights of perch’/ 
“heights of tree” for a similar time period showed no significant dif- 
ference in the behavior of the two species in the nearly duplicate 
habitats ((e — 1.9359 —=).01054 1 — 00) 

Summary. The trees at Chadron and Lakehurst were extremely 
variable in height and often located meters apart. At New London 
all the trees were nearly the same height (except for one a bit higher), 
and all were immediately aside one another. This difference may be 
reflected in the respective patterns of selection of perch. There was 
little distance for flight between the trees at New London, and thus, 
perhaps, occurred the higher frequency of changing trees, the marked 
tendency to choose a perch relatively near the height of the former, 
and the tendency to ascend higher when the single taller tree was 
reached. Despite these variations, the general information of the “hour- 
glass-on-its-side” configuration is most compatible, in both areas, with 


VoLuME 30, NUMBER 3 179 


the hypothesis that the shifting plane of reaction to polarized light, 
vertical to horizontal to vertical and the position of the sun itself form 
the basic influences on the behavior. 


Perch Posture 


Posturing positions. Insects specifically adapted to polarized light 
and/or phototactic stimuli generally display certain fixed postures when 
they are stationary. Gotz (1936) and von Buddenbrock (1917) demon- 
strated a “turning tendency’ that follows locomotion, in which the 
insect moves to assume its “stable” position of posture in relation to 
the sun. These two Callophrys (Mitoura) species displayed the four 
basic positions mentioned in the Discussion. Such positions have been 
demonstrated in a number of other insects, all of which are polarized- 
light adapted. Willington et al. (1951) and Henson (1954) showed 
the occurrence of these positions in particular frequencies in larvae of 
Lepidoptera. They were also demonstrated in Bidessus and Geotrupes 
(Coleoptera) by Birukow (1953) and Jander & Waterman (1960), 
respectively. Jacobs-Jessen (1959) showed that in Halictus (Hyme- 
noptera ) only two postures (0° and 90°) occurred, the two 45° postures 
evidently disappearing with excitation, the role of which was also con- 
firmed by Jander & Waterman (1960). Similarly, effects of temperature 
or excitation have been shown in the aforementioned studies of Lepi- 
doptera larvae. 

In Callophrys (Mitoura), the 45° posture was only noted early in 
the periods and then disappeared. The gross ratios of 0, 45, and 90° 
postures were: Chadron—19/3/48; Lakehurst—23/4/17; Chadron and 
Lakehurst—42/7/65; Lakehurst and two supplemental sites—30/4/23; 
and New London—71/5/148. The 0° number is divided between the 
anterior and posterior toward the sun for these respective localities as 
roloms-mo) lon 0/14 lo/27, 17/13, and 35/36. 

It is clear that a discrepancy occurs if the Chadron and Lakehurst 
calculations are lumped together as was shown reasonable with perch 
location. This may not be due to species difference but because all 
Lakehurst (and supplemental) populations were morning expansion- 
period samples. Dropping the 45° posture, the combined Lakehurst 
and Chadron ratios of 90 to 0° posture is ca. 3:2. For New London 
this ratio is ca. 2:1. Thus, the difference, considering Lakehurst, may 
indicate that additional data would show certain postures more prevalent 
at certain times of day. 

A number of x? tests were performed on 1 X 2, 1 X 3, and 1 Xx 4 
combinations of data from these sites as well as within and between 


180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


species. It is not useful to state each calculation here except to mention 
that all showed a significance of at least p = 9.05—p = 0.10 to the 
extreme categories—0 and 90°. Similarly, the dominant frequency of 
90° when compared with the sum of all others was x? = 23.14, p = 
0.001 at New London and x? = 9.78, p = 0.01-0.001 at Chadron. 

Special mention should be made that no locomotion was evident by 
these insects at the perch. Of the 471 perch choices in this study, 
only 6 insects showed locomotion at the perch. Thus, any “turning 
tendency” of movement necessary to assuming the stable posture must 
occur in flight before alighting. 

Repetition of perch posture. There was a definite tendency for pos- 
tures to repeat, at least for a brief period. This was investigated by 
creating an “index of repetition (IR)” and “index of nonrepetition 
(INR).” Each five-choice sample was evaluated as to the total combi- 
nations of repetitions or changes. Each change or repetition was given a 
value of % X the number of figures in the combination. (In a 90°, 90°, 
90°, 0°, 0° sample the repetition value would be 74) supe 
% and the change value would be %.) These were then multiplied 
by 2 and added in decimals. The results were as follows: Chadron (n = 
A7)—IR = 15.4, INR = 6.6 (2 = 3.52, p = 0105); Malkcehunstay cams) 
—IR = 14.4, INR = 10.0 (x? = 0.80); and New London (n = 226)— 
IR = 76.2, INR =\345 (x? = 15.62, p = 0.001). Wheremigeessmanrc 
graphic difference at New London, possibly because of the closeness 
of the trees in the sample. 

There was little evidence that the insects (as commonly observed in 
the popular literature) often return to the exact perch from which 
they flew. Of the 471 perches in the sample, only 11 indicated such 
behavior. 

Thermodynamics and perch posture. Since the physiological function 
of “sunning of the wings” in butterflies is well-known (Clench, 1966), 
the high frequency of the 90° angle position has an obvious thermo- 
dynamic value. For Mitoura, sunning is accomplished by closing the 
wings above the thorax and thus displaying one ventral primary and 
secondary to the sun. 

Perch position and gravity. The investigation of positive and negative 
geotaxis in these insects was hampered by the subjectivity of the ob- 
servations. At first, three positions were noted in samples—head obvi- 
ously upward, head obviously downward, and insect horizontal. The 
latter was too subjective and was abandoned, which resulted in arbi- 
trary designation of one of the first two. The results suggest that the 


VoLUME 30, NuMBER 3 181 


positioning of the head occurs at random and alternates independently 
of the angle of the posture: Chadron—(method abandoned); Lake- 
hurst and two supplemental—head up = 33, head down = 22 (x? = 
2205p —0:2020.10), IR = 16.6, INR = 4.2 (x? = 7.40, p = 0.01); and 
New London—head up = 102, head down = 113 (x? = 0.56), IR = 
foo Nhe 16.5) (yx? = 33.96, p — 0.001). 

Other observations. Marked lethargy was shown by the insects in 
early morning and late evening, with refusal to move even upon touching. 
Borgo, in Delaware and New Jersey, noted individuals very low in the 
trees (0647-0702 EST: 0.45-1.95 m) and some covered with dew. These 
all later activated for daytime perching. Similarly, Johnson, in Nebraska, 
noted refusal of the insects to fly at dusk (ca. 7 pm MST). Behavior 
that suggests oviposition was also observed by Johnson. The insects 
favored no particular part of the tree, occurring on upper branches 
and on twigs near the ground. Females clung to the twig, with head 
upward (angle posture variable), and arched the abdomen in successive 
movements toward the undersurface of the twig. No oviposition was 
ever observed. Distinct avoidance behavior was noted where a culti- 
vated spruce tree (Abies sp., Pinaceae) occurred in the center of the 
Chadron habitat. On two occasions butterflies flew to it but veered 
off to perch on a nearby juniper. Other butterflies found on the 
junipers included Cynthia cardui (Nymphalidae), Asterocampa celtis 
(Nymphalidae), Cercyonis pegala olympus (Satyridae), and Strymon 
melinus (Lycaenidae). Each of these flew out of junipers as Callophrys 
(Mitoura) were being studied. 


SUMMARY AND CONCLUSIONS 


The data in this study strongly suggest that the perching behavior 
of C. (M.) siva and C. (M.) gryneus is distinctly patterned and not 
simply a random usage of the tree. This has strong implications with 
regard to all Callophrys (Mitoura) species that feed on Cupressaceae, 
since their behavior is generally considered similar by lepidopterists. 
The statistical data on changes in the behavior with time of day and 
on perching postures support the assumption that the insects orient to 
interactions of polarized light and the position of the sun. The behavior 
is distinctly preferenced for height, and taller trees are most often 
selected for perching. 

These species are suggested to the physiologist for laboratory experi- 
ments to examine in greater detail the precise environmental relations 
of the behavior and the stimulus-response mechanisms of the insects. 


182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ACKNOWLEDGMENTS 


We would like to thank the staff of the Museum of Natural History, 
University of Wisconsin, Stevens Point, especially Dr. Charles A. Long, 
Director, for logistical support of this research. We would similarly 
like to thank Mr. Richard Priestaf (Goleta, California) for his efforts 
to duplicate samples in California. Debt is owed to the staff of the 
Entomology Department of the American Museum of Natural History 
for aid with literature, most especially Dr. Frederick H. Rindge, Curator 
of Lepidoptera, who reviewed this manuscript. Thanks are also due 
Ms. Renate Rosner (New York City, New York) for checking our 
German translations. 


LITERATURE CITED 


Birruxow, G. 1953. Menotaxis in polaristen Licht bei Geotrupes silvaticus Panz. 
Naturwissenschaften 40: 611. 

CiencH, H. K. 1966. Behavioral thermoregulation in butterflies. Ecology 47: 
1021-1034. 

GoupsmirH, T. H. 1965. The visual system in insects. In Rockstein, M. (Ed.), 
The physiology of Insecta. Vol. I. Academic Press, New York. 640 p. 

Gorz, B. 1936. Beitrage zur Analyse des Verhaltens von Schmetterlingsraupen 
beim Aufsuchen des Futters und des Verpuppungsplatzes. Z. Vergl. Physiol. 23: 
429-503. 

Henson, W. R. 1954. The light reactions of larvae of the spotless fall webworm, 
Hyphantria textor Harr. (Lepidoptera: Arctiidae). Can. Ent. 86: 529-542. 
Hopcson, E. S. 1965. Chaemoreception. In Rockstein, M. (Ed.), The physiology 

of Insecta. Vol. I. Academic Press, New York. 640 p. 

Jacoss-JEssen, U. F. 1959. Zur Orientierung der Hummeln und einiger anderer 
Hymenopteren. Z. Vergl. Physiol. 41: 597-641. 

JANDER, R. 1957. Die optische Richtungsorientierung der roten Waldameise 
(Formica rufa L.). Z. Vergl. Physiol. 40: 162-238. 

JANDER, R. & T. H. WatrerMAn. 1960. Sensory discrimination between polarized 
ee as light intensity patterns by arthropods. J. Cell. Comp. Physiol. 56: 

37-159. 

Jounson, K. 1976. Specificity, geographic distributions and foodplant diversity in 
four Callophrys (Mitoura), Lycaenidae. J. Lepid. Soc. 30: (in press ). 

Katmus, H. 1958. Responses of insects to polarized light in the presence of dark 
reflecting surfaces. Nature 182: 1526-1527. 

Marky, H. & M. Linpaver. 1965. Physiology of insect behavior. In Rockstein, 
ee Ed.), The physiology of Insecta. Vol. II. Academic Press, New York. 
FUr p. 

Marten, W. 1956. Beobachtungen beim Lichtfang. Ein Versuch zur Losing der 
Frage nach dem “Warum” des Anfluges der Insekten an kunstliches Licht. 
Ent, Z,. 1662 121-138), 

Vee ae Reizwirkung des kunstlischen Lechtes auf Lepidopteren. Ent. Z. 
a1. Cds 

MAZOKHIN-PorsHNyAkoy, G. A. 1969. Insect vision. Plenum, New York. 306 p. 

Ropinson, H. S. & P. J. Rosprnson. 1950. The observed behavior of Lepidoptera 
in flight in the vicinity of light sources. Ent. Gaz. 1: 3-20. 

Ropinson, H. S. 1952. On the behaviour of night flying insects in the neighbor- 
hood of a bright light source. Proc. Roy. Ent. Soc. London, Ser. A, 27: 12-21. 


VoLuME 30, NUMBER 3 183 


SCHWARTZKOPFF, J. 1965. Mechanoreception. In Rockstein, M. (Ed.) The physi- 
ology of Insecta. Vol. I. Academic Press, New York. 640 p. 

Scott, J. A. 1972 (1973). Mating in butterflies. J. Res. Lep. 11: 99-127. 

SmirH, F. E. & E. R. Baytor. 1960. Bees, daphnia, and polarized light. Ecology 
Al: 360-363. 

von BuppENBRocKk, W. 1917. Die Lichtkompassbewegungen bei den Insekten, 
insbesondere den Schmetterlingsraupen. Heidelberg. 181 p. 

von Frisco, K. 1960. Uber der Farbensiun der Insekton. P. 19-40, in Mechanisms 
of colour discrimination. London-Oxford-N.Y.-Paris. 

VERHEIJEN, F. J. 1958. The mechanisms of the trapping effect of artificial light 
sources upon animals. Arch. Neerland Zool. 107 p. 

Wiiuincron, W. G., C. R. SuLLIVAN & G. W. GREEN. 1951. Polarized light and 
body temperature level as orientation factors in the light reactions of some 
hymenopterous and lepidopterous larvae. Can. J. Zool. 29: 339-351. 


HILLTOPPING IN LEBANON 


Considering the extensive literature on hilltopping (Shields (1967, J. Res. Lepid. 
6: 69-178) gives nearly 200 references), it seems worthwhile to give a few notes 
from Lebanon on the topic. I do not recollect having seen previous references from 
the Levantine area. 

After collecting the localized Euchloe belemia Esper (10 km E of Saida, 21 
March 1972) for a few hours, I ascended a small rounded hillock with typical 
garrigue vegetation. For more than 15 min. one Papilio machaon syriacus Verity, 
two Vanessa cardui Linné and one Vanessa atalanta Linné were observed circling 
the top. One or more might briefly settle, but mostly all four were on the wing. 
There was little beating of the wings, most movement being a strong glide. No 
other butterflies were seen on the hilltop, and the species in question were not 
noted elsewhere in the vicinity. The weather was fine, with a breeze from the west. 

A large and fresh male of Iphiclides podalirius virgatus Butler was caught on a 
small summit surrounded by a precipitous drop of more than 200 m on three sides, 
the eastern side being somewhat gentler (Cedar Mountain, 1,900 m, 17 July 1974). 
The specimen hardly moved its wings while gliding on the updraft produced by a 
breeze from the west. It must have come from Bscherré village, some 300 m lower, 
where the closest breeding colony is located. 

In the above cases I was struck by the apparent “joie de vivre” of the ebullient 
circling and by the method of flight, which differed so much from that ordinarily 
seen. Two rare species, Papilio alexanor maccabaeus Staudinger and Elphinstonia 
charlonia penia Freyer, are nearly always found about the summits of stony out- 
crops. Their flight in such situations is quite normal, however, which may simply 
indicate the presence of their food plants—so far unknown in Lebanon. 

I have little doubt that hilltopping is more common in the Middle East than 
the lack of recorded evidence suggests. The places where these singular aerobatic 
displays are performed are otherwise so unattractive that no entomologist would 
pay them a visit. 


TorBEN B. LaRsEN, c/o IPPF, 18-20 Lower Regent St., London SW 1, England. 


184 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


OBSERVATIONS ON HOST PLANT RELATIONSHIPS AND 
LARVAL NUTRITION IN CALLOSAMIA (SATURNIIDAE)* 


RicHARD S. PEIGLER 


Department of Entomology and Economic Zoology, Clemson University, 
Clemson, South Carolina 29631 


The genus Callosamia Packard is comprised of three closely related 
species that occur in eastern North America. Callosamia promethea 
(Drury) ranges from Canada to Florida and west to the Great Plains 
and accepts a wide range of hosts. Callosamia angulifera (Walker ) 
occurs east of the Mississippi River, most commonly from Pennsylvania 
to Georgia, and feeds exclusively on tuliptree (Liriodendron tulipifera 
L.). Callosamia securifera (Maassen) is found only in swamps and 
pine woods in the coastal areas of the Southeast (Peigler, 1975) and 
feeds exclusively on sweetbay (Magnolia virginiana L.). 

The purpose of the present paper is to discuss briefly some of my 
observations regarding the hosts of these moths. Because the Saturni- 
idae are popularly reared, I have attempted to discover alternate foods 
for the two monophagous species so that they might be reared where 
the preferred foods are not available. Information is also presented 
on the acceptability of these host plants as a function of larval age. 


Host Plants of Callosamia promethea 


The wide variety of host plants of C. promethea accounts for the 
much wider geographical distribution of this species when compared 
with its congeners. This variety is also a factor in the ability of C. 
promethea to exist in many different types of habitat. Although the 
host range is comparatively wide, a preference for Lauraceae is evident, 
and no conifers or monocots are known hosts. Most of the C. promethea 
foods are aromatic plants such as sassafras (Sassafras albidum (Nutt.) 
Nees.), horse sugar (Symplocos tinctoria (L.) L’Her.), sweetgum 
(Liquidambar styraciflua L.), spicebush (Lindera benzoin (L.) Blume) 
and wild black cherry (Prunus serotina Ehrhart). Such plants may 
possess an important olfactory stimulus which initiates oviposition by 
females or feeding by newly eclosed larvae. 

Callosamia promethea on horse sugar in coastal South Carolina was 
thought to represent a host-specific population (Ferguson, 1972), such 
as the population on lilac (Syringa vulgaris L.) around Milwaukee, 


_ | Published by permission of the Director of the South Carolina Agricultural Experiment Station. 
echnical Contribution No. 1251. 


VoLUME 30, NUMBER 3 185 


Wisconsin. However, I have found cocoons in the former area on wild 
black cherry. I have also found cocoons on horse sugar and other more 
common hosts in Oconee Co., South Carolina. It is interesting to note 
the corresponding disjunct ranges of C. promethea and horse sugar 
(Radford et al., 1964) in South Carolina, where both occur in the 
mountains and near the coast but are absent for over 100 mi. between. 
Callosamia promethea is not recorded on horse sugar anywhere else, 
except where I have found it in Brunswick Co., North Carolina. 

Certain hosts given by Packard (1914) and cited by later authors 
need to be verified and may have been based on cocoons of larvae 
that did not spin on the actual food plant. These are barberry (Ber- 
beris), maple (Acer), azalea, birch (Betula) and arbor-vitae (Thuja), 
the latter two seeming especially questionable. I have tried to rear 
Pennsylvania C. promethea on azalea and Acer rubrum L., but the 
larvae died in first instar. 

An unpublished host of C. promethea is sweetbay. Dale Schweitzer 
(pers. comm.) found C. promethea cocoons on sweetbay in Longwood 
Gardens, Kennett Square, Pennsylvania, where the tree is not native, 
and in the pine barrens of Burlington and Atlantic counties, New 
Jersey. However, I have tried several times to rear C. promethea 
(Pennsylvania stock) on sweetbay; the larvae always grew slowly and 
died within 15 days, usually in the second instar. 


Host Plants of Callosamia angulifera and C. securifera 


A brood of C. angulifera ¢ xX C. securifera 2° was reared on tulip- 
tree and sweetbay, the two respective parent foods. The larvae on 
tuliptree matured faster and spun their cocoons almost two weeks 
before their siblings on sweetbay, although the size of adults in both 
groups was the same. The physiological advantage of tuliptree over 
sweetbay was also shown in broods of pure C. angulifera and C. secu- 
rifera. Newly hatched larvae did not survive on the host plant of the 
opposite species, although some C. securifera were once reared from 
ova to adults on tuliptree (Jones, 1909). Later instar larvae of C. 
angulifera completed their larval development on sweetbay in most 
cases. However, C. securifera that fed for the first two weeks on sweet- 
bay readily completed their larval life on tuliptree. For comparison, 
larvae from these same broods were reared exclusively on their pre- 
ferred hosts. 

In addition to the trials of C. angulifera and C. securifera on their 
two hosts, these species were tested on other species of Magnoliaceae, 
both American and Oriental. Newly hatched larvae of both species 


186 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


were placed on potted seedlings or rooted cuttings of the two American 
plants, Magnolia grandiflora L. and Illicium floridanum Ellis., and the 
Asian M. stellata (Sieb. & Zucc.), M. soulangeana (M. denudata Des- 
rouss. X M. liliflora Desrouss.), Illicium anisatum L. and Kadsura 
japonica (L.) Dun. All except the first of these plants have tender 
leaves. The larvae fed sparingly, wandered and, in all cases, died within 
a few days in the first instar. Unfortunately, none of the several 
American species of deciduous Magnolia were available. 

The results obtained with these substitute hosts imply the lack of a 
phagostimulant rather than some essential nutrient(s). It is possible 
that the larvae would survive on such plants if they would feed freely. 
No evidence of toxicity was observed. 

Further testing the following year gave an alternate host for C. 
securifera. Ten newly hatched larvae were put into a jar with leaves 
of wild black cherry, Magnolia grandiflora, sweetgum, buttonbush 
(Cephalanthus occidentalis L.) and more of the Magnoliaceae tried 
earlier. Again larvae fed sparingly and wandered, and all but two 
died within three days. Of the two that remained, one was a first 
instar which ate buttonbush and the other was a second instar which 
fed on M. grandiflora. Sweetgum also showed that feeding had occurred 
on it. Fresh leaves of buttonbush, M. grandiflora, and sweetgum were 
offered to these larvae, and they chose sweetgum. Soon after, they 
were sleeved outdcors on sweetgum. The smaller larva, which had 
eaten buttonbush, died in the third instar. The other larva produced 
a slightly undersized female moth the following month. If one attempts 
to rear C. securifera on sweetgum, most or all of the larvae would be 
expected to die early. I do not believe this tree ever serves as an 
alternate host in nature. 

Lepidopterists have found that C. angulifera will sometimes accept 
spicebush, sassafras, and wild black cherry (Eliot & Soule, 1902). How- 
ever, since these plants are inferior substitutes for tuliptree, larvae on 
them would probably not grow as rapidly or as large. I seriously doubt 
that these alternate hosts are ever utilized in nature. 


DISCUSSION 


The Magnoliaceae is a very old angiosperm family with fossils dating 
to the Cretaceous period. The widest ranging species of the family in 
North America is tuliptree, the food of C. angulifera. The nocturnalism 
of C. angulifera suggests that it is closest to the ancestral form of the 
genus. Tuliptree is the best choice for rearing any Callosamia hybrid 
and is presently the only “common denominator” food for all three 


VoLuME 30, NUMBER 3 187 


species. Therefore, it is probable that tuliptree is the original food 
for the genus. 


ACKNOWLEDGMENTS 


I am grateful to Drs. Raymond Noblet and G. R. Carner, entomology 
professors at Clemson University, for much helpful criticism on the 
research and the manuscript. 


LITERATURE CITED 


Exior, I. M. & C. G. Soutz. 1902. Caterpillars and their moths. The Century 
Company, New York. 302 p. 
Frercuson, D. C. 1972. Bombycoidea, Saturniidae (in part). In R. B. Dominick 
et al., The moths of America north of Mexico, fasc. 20.2B: 155-269, 22 pls. 
Jones, F. M. 1909. Additional notes on Callosamia carolina. Ent. News 20: 49- 
Bie 

PacxarD, A. S. 1914. Monograph of the bombycine moths of North America, 
part 3 (Ed., T. D. A. Cockerell). Mem. Natl. Acad. Sci. 12: 1-516. 

PricLeR, R. S. 1975. The geographical distribution of Callosamia securifera (Sa- 
turniidae). J. Lepid. Soc. 29: 188-191. 

RaprorpD, A. E., H. E. Antes & C. R. Bett. 1964. Manual of the vascular flora 
of the Carolinas. The University of North Carolina Press, Chapel Hill. 825 p. 


TIME VARIATIONS OF PUPAL STAGE OF EUPACKARDIA CALLETA 
(SATURNIIDAE ) 


In December 1971 a friend presented me with 16 pupae of Eupackardia calleta 
(Westwood) which he had found while trimming shrubbery in his yard. These 
pupae had undoubtedly come from the same egg hatch, probably in the early fall 
of 1971, because all were found on the same bush and several on the same limb. 
I was never able to find out from what plant he collected the pupae. All the 
pupae appeared to be alive and in good condition. 

Eight of the pupae were sent to a friend and eight I kept for myself. Of the 
eight sent away two males emerged in September 1972, also one pupa produced 
several parasitic flies during the same month. In March 1973 one female emerged 
and one pupa had died and dried up. I have no record as to what has happened 
to the other three remaining pupae. 

A record of the eight pupae I kept for myself is as follows. One male emerged 
in August 1972 and another male in September 1972. In March 1973 one female 
emerged and on 19 August 1974 one male emerged. It was at this time, August 
1974, that I noted that one of the four remaining pupae had died. On 1 Sep- 
tember 1975 one male emerged and on 25 September 1975 a large female emerged. 
Of the six adults which have emerged the development time from egg to adult 
ranged from approximately 1-4 years. The remaining pupa recently has lost some 
of its weight and probably has died. 

All of my pupae have been kept in the same environmental conditions, and 
all of the adults have been normal and healthy. 


Jack B. Prentiss, 4222 Hermosa, Corpus Christi, Texas 78411. 


188 JouRNAL OF THE LEPIDOPTERISTS’ SOCIETY 


A REVISION OF THE GENUS DUNAMA SCHAUS 
(NOTODONTIDAE) 


je, 1b, on 


Systematic Entomology Laboratory, IIBIII, Agr. Res. Serv., USDA 
C/o U.S. National Museum, Washington, D.C. 20560 


Several years ago some moths reared from larvae from Mexico found 
on importations of Chamaedorea, a genus of small palms, were sent 
to me for identification by Mr. D. R. Johnston, San Antonio, Texas. 
The adults proved to be a new species of the monobasic genus Dunama 
Schaus of the family Notodontidae. A search of the collections of the 
Neotropical notodontids in the U.S. National Museum and of the British 
Museum (Natural History) revealed three other species belonging to 
the genus. One of the three is another undescribed species from 
northern South America. Of the three described species, one had been 
described twice, but the synonymy had not been recognized. The 
synonyms were described in different genera, and each subsequently 
had been moved to a different genus. All the generic assignments 
and transfers, except of the type-species, were incorrect. Five species 
of the genus Dunama are now recognized. 

Until recently the only known food plant was a single species of 
Chamaedorea. This plant genus is composed of a large number of 
species, and some have rather restricted or limited distributions. It is 
possible, therefore, that other species of Dunama may be recognized 
when some of the other species of Chamaedorea are examined for 
larvae or when collecting at light for adults is accomplished in the 
vicinity of such plant species. In 1971, Dunama angulinea (Schaus) 
was reared from larvae found feeding on bananas in Panama by Mr. 
C. S. Stephens. 

Descriptions of the larvae are not included in this paper. They will 
be described in a separate paper by D. M. Weisman. 


Dunama Schaus, 1912 


Ann. and Mag. Nat. Hist., ser. 8, vol. 9, p. 52. 
Type-species: D. angulinea Schaus, monotypy and original designation. 


Diagnosis: Small to moderate-sized notodontid moths, length of forewing 10-22 
mm; male antennae bipectinate for %4 length, pectinations of fifth or sixth segment 
longest, decreasing to simple at apical fourth, female antennae simple, scale tuft of 
first antennal segment small in both sexes; palpi upcurved to near middle of frons, 
second segment long, third segment small not more than V4 length second segment, 
slightly decumbent; tongue well developed; ocelli absent. Thorax without prominent 
tufts, vestiture rather loose; abdomen without tufts. F orewing with slender accessory 


VoLUME 30, NUMBER 3 189 


cell; M: from bottom third of accessory cell; hindwing with Sc from about middle 
of cell, diverging from Rs, nearly straight; Rs and M; connate from upper angle of 
cell; M; and Cu; connate from lower angle. Male genitalia distinctive (Figs. 10-14), 
uncus variable, an ovoid lobe or bifid; socii well developed, sclerotized curved 
processes; valves with costal margin sclerotized, ventral margin mainly membranous, 
but sometimes toothed near end of sacculus; juxta scarcely developed, fused with 
bases of valves when present; aedeagus moderately long, apical half heavily sclero- 
tized and usually somewhat reduced in diameter toward apex, shaft usually with 
dorsolateral spines and processes or a dorsal plate near middle; eighth sternite with 
caudal margin produced into heavily sclerotized process, sometimes bifid. Female 
genitalia (Figs. 15-18), rather reduced; ovipositor lobes and ninth segment sclero- 
tized; some ostial sclerotizations; corpus bursae small, membranous, without signa. 

This genus has been placed near Disphragis Hubner, but it is doubtful that it 
belongs to the Heterocampini. I believe it to be a member of the tribe Nystaleini, 
subfamily Notodontinae. It does not agree completely with the definition of that 
tribe by Forbes (1948, p. 206), but that definition was based primarily on a few 
North American genera and species and undoubtedly will need to be modified 
when the much larger group of Neotropical elements are studied. The present 
classification of the Neotropical notodontids is very unsatisfactory. Forbes (1948, 
p. 203), stated: “The classification of the North Temperate fauna is fairly well 
understood, but the much larger and richer tropical fauna is in complete confusion.” 
The almost complete lack of knowledge of the immature stages of the Neotropical 
notodontids presents a serious complication to the development of a more satisfactory 
classification of the family. 


KEYS TO LHE SPECIES 


MACULATION AND COLORATION 


LL, Misgiee™ Abe WE fe OS OS ra ee 2 
PSUS a aes 6 
2. Vertex of head, thorax and basal part of forewing with conspicuous areas 
of pale straw yellow; forewing with distinctive, slightly oblique, black bar 
from basad of reniform spot to near base of inner margin —___ D. tuna (Schaus) 
—Head, thorax and base of forewing mostly dark brown; forewing with median 
part of antemedial band at most a dark triangular spot only slightly longer 
ier URN CIG] CR a nt SA tee hada nn ee ci ala Sanaa i Ree neice t ea ave 3 
3. Subterminal transverse shade of forewing absent; forewing with a faint 
longitudinal pale streak present through cell from base of wing nearly to 
SSTTDGTT ak, ON a ee D. ravistriata, n. sp. 
—Subterminal transverse shade of forewing present, especially in anterior half 
of wing; forewing with longitudinal pale streak absent, or present only 


Rho MO EMBECTULEO TIN ASO En tee ee ome rete etcetera eh eA 4 
4. Longitudinal pale streak of forewing weak, but present distad of reniform 

DOE ae ce es rep pee Ae D. angulinea Schaus 

=imougiceudinal pale streak of forewime absent 2.) 5 


5. Costal area of forewing distad of reniform spot distinctly darker than re- 
mainder of distal part of wing; median transverse band immediately distad 
of antemedial band paler at middle than near inner margin 
__ a) ee ee Peale ee ei ee ) Ghasicernirata, :( Doenin ) 
—Costal area of forewing distad of reniform spot if darker than remainder of 
distal part of wing only slightly so; median transverse band immediately 
distad of antemedial band not especially paler at middle _— D. mexicana, n. sp. 
6. Forewing with vague to distinct pale longitudinal streaks distad of reniform 
SOGNE | | pee ee EE res she A es DC eee ed OO gsi ae i 


190 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


_Forewing with pale longitudinal streaks absent distad of reniform spot —- 

sp og pee Sah than. Sgt teeal RS 0 & at a D. mexicana, n. sp. 
7. An oblique black bar present from reniform spot to near base of forewing 

ee Di es 2 er ere ee le ie SR Sa D. tuna (Schaus ) 

-Forewing with at most a dark triangular spot, slightly longer than wide, 

from near middle of antemedial band =2 2. EEE 8 
8. Median part of dorsum of thorax with an oval spot of pale scales —_____ 

pcg se ee en Ee ere os D. angulinea Schaus 

—Median part of dorsum of thorax uniformly dark D. ravistriata, n. sp. 


1, Males _.... ee ee 2 
—Females ..2c::-- ee 6 
2. Sclerotized process of eighth abdominal sternite bifurcate —___-___-____. 3 
—Sclerotized process of eighth abdominal sternite not bifurcate 4 
3. Process of eighth abdominal sternite with toothed lateral flanges at base of 
bifureation : 22.22.20 oes Ge oe, eee Se ee D. tuna (Schaus ) 
—Process of eighth abdominal sternite without lateral flanges at base of bi- 
FTCA OI ae ee Os D. angulinea Schaus 
4, Uncus bifid; ventral margin of valves nommal = 5 


—Uncus clavate, the lateral margins more heavily sclerotized than middle; 
ventral margin of valves heavily sclerotized and irregularly toothed near end 
Of SaACCUlUS: eo 2. ete Ne ee ae D. claricentrata (Dognin) 
5. Uncus Y-shaped, the lobes slender, slightly knobbed at apices; aedeagus with 
two large elongate, spinelike dorsolateral processes from middle of shaft 
EN PE As cr a Si NR ERE in OE i us DS mexicaiomueesp: 
—Uncus with two stout slightly divergent lobes; aedeagus with one small blunt 
spinelike process near distal 14 of shaft 0-2 ee D. ravistriata, n. sp. 
6. Posterior margin of ninth abdominal tergum irregularly toothed; lateral angles 
of eighth abdominal sternum produced into flat, sharp-pointed triangular 


DIOCESSES. he ee ee D. mexicana, n. sp. 
—Posterior margin of ninth abdominal tergum and lateral angles of eighth 
abdominal sternum not'so modified 922 ee i 
7. Posterior margin of eighth sternum V-shaped and slightly to moderately 
irregularly toothed (0s Gpae ANG OAS ee Ne a eee a D. ravistriata, n. sp. 
—Posterior margin of eighth sternum transverse or slightly convex and slightly 
emarginate in middle) margin not toothed __~.~ ae 8 
8. Sternum of ninth abdominal segment a broad sclerotized band nearly uni- 
form im length’ basad of ovipositor lobes). D. tuna (Schaus ) 
—Sternum of ninth abdominal segment membranous at middle third basad of 
ovipositor lobes) / 83 ceul mma ne CMe 0 Sn eee D. angulinea Schaus 


Dunama angulinea Schaus 
rate en Tes 10), IS 
Dunama angulinea Schaus, 1912, p. 52. Draudt, 1932, p. 981. Gaede, 1934, p. 263. 


Diagnosis: Size, forewing length, ¢ 10-13 mm, ? 12-15 mm. The pattern of 
maculation (Fig. 1) is similar to that of Dunama claricentrata (Dognin) and D. 
mexicana, n. sp. in the males. Females are not so contrastingly marked and _ re- 
semble that sex of D. mexicana and D. ravistriata, n. sp., but the disc of the thorax 
has a tuft of pale scales not found in D. mexicana and the basal area of the fore- 
wing does not have a longitudinal ray of pale scales as in D. ravistriata. Both sexes 
are somewhat variable in the pattern of maculation. The most reliable characters 
for species identification are shape of the eighth abdominal sternite of the male 


VoLUME 30, NuMBER 3 191 


Figs. 1-5. Dorsal view of adult males of Dunama species: 1, angulinea, holo- 
type male, Guapiles, Costa Rica; 2, tuna, holotype male, Colombia; 3, mexicana, 
holotype male, México; 4, ravistriata, paratype male, Teffé, Amazonas, Brazil; 5, 
claricentrata, holotype male, French Guiana. 


(Fig. 10) and the genital structures of the female (Fig. 15). The bifurcate process 
of the eighth abdominal sternite of the male lacks toothed, lateral flanges as in D. 
tuna (Schaus). The female genitalia are similar to those of D. tuna, but the ninth 
abdominal sternite is membranous at middle basad of ovipositor lobes, not sclero- 
tized as in D. tuna. 

Type: The male holotype is in the U.S. National Museum. It was collected at 
Guapiles, Costa Rica. 

Distribution: Twenty-nine examples, 14 ¢ and 15 @, have been studied. They 
are from the following localities; Mrxico: Teapa, Tabasco. GUATEMALA: Cayuga; 
Quirigua. Cosra Rica: Guapiles; Sixaola River. Panama: Changuinola; Paraiso, 
Gr-Z; 

Food plant: This species has recently been reared from bananas in Panama, but 
it does not seem likely that the banana plant is the normal host. If that were so, 


192 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


, 


Figs. 6-9. Dorsal view of adult females of Dunama species: 6, mexicana, para- 
type female, México; 7, angulinea, female, Changuinola, Panama; 8, ravistriata, 
paratype female, Para, Brazil; 9, tuna, female, paralectotype of sagittula, Rio Toche, 
Colombia. 


considering the attention given to insects on that crop, surely larvae would have 
been discovered long ago. 


Dunama tuna (Schaus), n. comb. 
Figs. 2, 9, 11, 18 


Heterocampa tuna Schaus, 1901, p. 304. 

Disphragis tuna (Schaus), Draudt, 1932, p. 972. Gaede, 1934, p. 261. 
Naduna sagittula Dognin, 1914, p. 22. |New synonymy. | 

Tachuda sagittula (Dognin), Draudt, 1932, p. 929. Gaede, 1934, p. 222. 


Diagnosis: Size, forewing length, ¢ 12-19 mm, 2 22 mm. This is the largest 
species of the genus. The presence of areas of pale straw yellow scaling on the 
head, thorax and basal halves of the forewings is characteristic of this species. A 
prominent black, slightly oblique bar is located between the base of the reniform 
spot and the base of the inner margin of the forewing in both sexes. The male 
genitalia are quite similar to those of D. angulinea, but the eighth abdominal 
sternite bears toothed, lateral flanges at the bases of the distal bifurcation, and 
the aedeagus is slightly less armed with sharp-pointed processes on the apical half 
of the shaft. The female genitalia are most like those of D. angulinea (see Figs. 
15 & 18), but D. tuna has a broad sclerotized platelike ninth abdominal sternum. 
The middle of the ninth abdominal sternum of D. angulinea is mostly membranous 
with only some small subtriangular sclerotizations. 

Types: The holotype male of D. tuna from Colombia and the two syntypes of 
Naduna sagittula Dognin are in the collections of the U.S. National Museum. The 
syntype of the latter labeled: “Naduna sagittula Dgn. Type3”; “Dognin Collec- 


VoLUME 30, NuMBER 3 193 


tion’; “Canon del Tolima, Colomb., Cent. Cord., 1,700 m, Coll. Fassl”; “4 genitalia 
on slide 2153, Mar. 1966, ELT.” has been selected, labeled and is now designated 
the lectotype of that nominal species. 

Distribution: Only five specimens of this species have been examined. Three 
are from localities in Colombia, the other two, both males, are from Sixaola River, 
Costa Rica, and Porto Bello, Panama. 

Food plant: Unknown. 


Dunama mexicana Todd, new species 
Hisss3u6. 13. 17 


Description: Head with proboscis well developed; labial palpi slightly upcurved 
to near middle of frons, third segment very small, slightly porrect, second segment 
about 5 times as long and 2 times as wide, vestiture of short, loose, reddish-brown 
scales and a few scattered gray scales; frons flat, vestiture loose, scales directed 
mesoventrad from each side; eyes large, hemispherical, slightly wider than frons in 
male, subequal to frons in female; antennae of male bipectinate, the pectinations 
slender, longest at middle of antenna, about 3 times as long as width of article, 
antennae of female simple. Vestiture of patagia, tegulae and thorax of elongate 
scales and hairlike setae; patagia with dark brown scales medially, straw yellow 
scales basally and distally; tegulae nearly uniformly brown; thorax variegated with 
mixture of reddish-brown and yellowish-brown scales. Abdomen dark brown dor- 
sally, yellowish brown ventrally; dorsal tufts absent. Pectus clothed with long, 
sparse, yellowish-brown, hairlike setae. Legs of male pale yellowish-brown except 
trochanter of forelegs dark reddish brown, a dark brown band or apical patch at 
distal third of tibiae of all legs, and tarsal segments mostly dark brown, each with 
narrow pale ring at apex. Legs of female usually more uniformly dark brown. 
Pattern of maculation of dorsal surfaces of wings as illustrated (Figs. 3 & 6); males 
grayish brown, transverse median band of forewing paler, spots basad of band and 
in reniform spot dark brown, nearly black; females less maculate and considerably 
darker brown; ventral surface of wings essentially immaculate in both sexes except 
some small yellowish-brown, oblique marks on apical half of costal margin of fore- 
wing. Length of forewing: male, 11-15 mm, female, 12-17 mm. 

Male genitalia distinctive (Fig. 13). Uncus bifid, slender, Y-shaped; aedeagus 
with two large, spinelike processes from dorsal surface at apical third; ventral 
margin of valva simple, membranous; posterior margin of eighth abdominal sternite 
heavily sclerotized and produced medially, upcurved, apex concave. Female geni- 
talia as illustrated (Fig. 17). Posterior margin of ninth abdominal tergum sclerotized 
and irregularly toothed; ninth abdominal sternite well developed, a deep U-shaped 
emargination at middle; lateral angles of eighth abdominal sternite flat, sharp- 
pointed, triangular processes. 

Types: Holotype (USNM type no. 64649) male, 6 ¢ and 2 2 paratypes, México, 
reared from “Chamaedorea elegans’; 3 6 and 2 @ paratypes, Tuxapan, V. C., 
México, reared from “Chamaedorea elegans,’ March 1959, in the U.S. National 
Museum. One ¢ and 1 Q paratype, same data as holotype, in the British Museum 
(Natural History ). 

Food plant: Larvae were reared from plants imported by florists as “Chama- 
edorea elegans.” There is some question as to the correctness of the identification 
of the plant, and the original plant material is no longer available for study. Ac- 
cordingly, the food plant should be listed as Chamaedorea sp. 

Discussion: The characters given in the key based on maculation and coloration 
probably will separate most examples from the similar species, D. angulinea and D. 
claricentrata, but the genitalia, male or female should be examined for positive 
identification. There is some question as to the actual area of origin of the plant 


JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


— 
(de) 
io 


Figs. 10-14. Males, genitalia and ninth abdominal sternites of Dunama species: 
10, angulinea, Quirigua, Guatemala; 11, tuna, Porto Bello, Panama; 12, claricentrata, 
holotype, French Guiana; 13, mexicana, paratype, México; 14, ravistriata, holotype, 
French Guiana. 


material in México. These plants have rather restricted distributions and the col- 
lectors of the plants tend to keep their source secret. After several inquiries in an 
attempt to determine a type locality, I was informed only that the plants came from 
northern Veracruz. Later specimens stated to be from Tuxapdn, Veracruz were 


received, but it is not possible to state with certainty that either statement is 
reliable. 


VoLuME 30, NuMBER 3 195 


I8 


Figs. 15-18. Females, genitalia of Dunama species: 15, angulinea, female, 
Paraiso, C. Z., Panama; 16, ravistriata, paratype female, Para, Brazil; 17, mexicana, 
paratype female, México; 18, tuna, female, paralectotype of sagittula, Rio Toche, 
Colombia. 


Dunama ravistriata Todd, new species 
Hien 49 14 1G 


Description: Very much like D. mexicana except: vestiture of head and thorax 
generally darker; frons yellowish-brown; patagia pale at base only; medial band of 
male forewing conspicuous only at base of cell; a pale longitudinal streak present 
in both sexes from base of forewing through reniform spot toward termen; legs of 
male darker than in D. mexicana. Length of forewing: male, 14-15 mm, female, 
14-17 mm. 

Male genitalia distinctive (Fig. 14). Uncus bifid, bifurcations stout, only slightly 
divergent; socii longer than uncus, stouter than in D. mexicana; aedeagus with only 


196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


one small, thornlike dorsal process at apical fifth; sclerotized caudal process of 
eighth abdominal sternite with knob-shaped apex. Female genitalia as illustrated 
(Fig. 16). Posterior margin of eighth abdominal sternite broadly V-shaped and 
slightly irregularly toothed. 

Types: Holotype male, French Guiana, Bar [collector], ¢ genitalia on slide 2120 
E. L. Todd, Jan. 1966 (BMNH Notod. 313); 1 ¢ and 4 2 paratypes, same locality 
and collector; 1 4 and 9 @ paratypes, Para, Brazil, A. M. Moss; and 1 @ paratype, 
Teffé, Amaz., Brazil, M. de Mathan, in the British Museum (Natural History). 
One male and 1 @ paratypes, Parad [Brazil], A. M. Moss, and 1 @ paratype, 
Goedebert, Maroni, French Guiana, Collection Le Moult, in the U.S. National 
Museum. 

Food plant: Unknown. If a palm, it must be other than the chamaedoreoid 
group as they do not occur in the area from which the moth is known to occur. 


Dunama claricentrata (Dognin), n. comb. 
Mes, 5, 1 


P Eunotela claricentrata Dognin, 1916, p. 13. 


Diagnosis: Size, @, length of forewing 13 mm. Pattern of maculation as illus- 
trated, similar to that of D. angulinea and D. mexicana. Some differences in macu- 
lation have been noted and utilized in the key to species. It must be pointed out 
that D. angulinea and D. mexicana are both somewhat variable in maculation and 
that D. claricentrata may easily prove as variable when specimens other than the 
holotype are available for study. The male genitalia (Fig. 12) are very distinctive 
and easily permit the identification of the species. The simple action of brushing 
the hair and scales from the venter of the distal segments of the abdominal sternum 
and the valves of the genitalia, if slightly produced, would enable one to view the 
characters of significance. The uncus is spatulate, but more elongated and more 
membranous medially and distally than that structure in D. angulinea and D. tuna; 
the ventral plate of the abdominal sternite is similar to that structure in D. mexicana 
and D. ravistriata, not bifurcate as in the other two species; the strongly toothed 
ventral margin of the valve and the slightly bilobed, flat dorsal process of the 
aedeagus are not found in any other Dunama species. 

Type: The holotype male from Nouveau Chantier, French Guiana is in the 
collection of the U.S. National Museum. 

Food plant: Unknown. 


LITERATURE CITED 


Docnin, P. 1914. Heétérocéres nouveaux de ’Amérique du Sud. Fasc. VII, 32 p. 
Rennes: Oberthiir. 

IRSMNG  Wartalen Veeco, IUL, yA joy, 

Draupr, M. 1931-1934. Notodontidae. In Seitz, Die Gross-Schmetterlinge der 
Erde, 6: 901-1070, pls. 143-159. Stuttgart: Kernen. [p. 901-904, 1931; p. 
905-1016, 1932; p. 1017-1048, 1933; p. 1049-1070, 1934. |] 

Forbes, W. T. M. 1948. Lepidoptera of New York and neighboring states, Pt. 2. 
Cornell Univ. Agric. Exper. Sta. Memoir 274, 263 p., 255 figs. 

Garpe, M. 1934. Notodontidae. In Strand, Lepidopterorum Catalogus, Pt. 59: 
301 p. Berlin: Junk. 

ScuAus, W. 1901. A revision of the American Notodontidae. Trans. Ent. Soc. 
London, 1901, p. 257-343, pls, =. 

1912. New species of heterocera from Costa Rica—XII. Ann. & Mag. 

Nat. Hist,, ser:s8, vol: 9, p. 84257, 


VoLUME 30, NUMBER 3 197 


THE BUTTERFLIES OF MISSISSIPPI—SUPPLEMENT NO. 2! 


BrYANT MATHER? AND KATHARINE MATHER 
213 Mt. Salus Drive, Clinton, Mississippi 39056 


The first list of Mississippi butterflies (Weed, 1894) included 53 
species; the second (Hutchins, 1933) included 73; the third (Mather 
& Mather, 1958) included 122, 45 of which were not included in either 
of the previous lists. Mather & Mather (1959) removed two names 
from the 1958 list because the material upon which their inclusion had 
been based was found to have been incorrectly determined and added 
two names based on new collecting data. Since 1959 we have prepared 
and given informal limited circulation to a number of revised manu- 
script lists, but in each case there have been questions of nomenclature 
and relationships awaiting resolution through publication by others. 
It now seems appropriate to issue as a second supplement, a list of 
the names of the 143 species now known to have been found in 
Mississippi, arranged in current sequence and designated by current 
names. 

The 120 names included in the 1958 list still regarded as valid are 
given in their current form and position in the list without further 
comment. The two added in 1959 were Nastra neamathla (Skinner & 
Williams) (42) and Satyrium kingi (Klots & Clench) (86). The 21 
additional names, indicated by “*” in the list, are included on the 
basis of evidence of their occurrence as summarized below. Wallen- 
grenia egeremet (Scudder) (31) and Lethe anthedon (Clark) (136) 
were known prior to 1958 to be present in the Mississippi fauna; they 
are now regarded as species distinct from Wallengrenia otho (Smith) 
(30) and Lethe portlandia (Fabricius) (135) respectively, rather than 
forms of these. The remaining 19 additions represent new data on the 
Mississippi fauna. Work by Rick Kergosien added six: Amblyscirtes 
reversa Jones (10), Erynnis funeralis (Scudder & Burgess) (48), Urbanus 
d. dorantes (Stoll) (59), Brephidium isophthalma pseudofea (Morrison ) 
(100), Anartia jatrophae guantanamo Munroe (113), and Hypolimnas 
misippus (Linnaeus) (121). Work by Charles T. Bryson added five: 
Poanes hobomok (Harris) (21), Problema byssus (Edwards) (25), 
Leptotes marina (Reakirt) (101), Ewphydryas phaeton ozarkae Masters 
(127), and Lethe a. appalachia Chermock (138). We added four: 


1 Contribution No. 329, Bureau of Entomology, Division of Plant Industry, Florida Department 
of Agriculture and Consumer Services, Gainesville, Florida 32602. 

2 Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, Florida 
Department of Agriculture and Consumer Services, Gainesville. 


198 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Euphyes dukesi Lindsey (19), Poanes viator zizaniae Shapiro (24), 
Atrytone d. delaware (Edwards) (27), and Calycopis isobeon (Butler 
& Druce) (89). Work by MacDonald Fulton added Harkenclenus titus 
mopsus (Hiibner) (84) and Eumaeus atala florida Rober (88). Work 
by Roy Strickland added Megathymus y. yuccae (Boisduval & LeConte) 
(1), and work by Charles Daniel added Speyeria c. cybele (Fabricius ) 
(129). 

In the 1958 list there was only one name for which no preserved 
specimen could then be located: Phoebis philea (Johansson) (76); 
there is still no known Mississippi specimen of this species. Two of 
the added species fall into this category; their addition being based 
only on sight records. The basis for their inclusion is as follows: 

Eumaeus atala florida (88). Professor MacDonald Fulton, Department 
of Bacteriology, Mississippi State College for Women, Columbus, Mis- 
sissippi wrote to me in October 1969 as follows: “In 1966 I saw and 
observed for at least 10 minutes here on the campus, Eumaeus atala 
(I assume the species—the genus is easy). It was a perfect specimen. 
No net.” Dr. Fulton discussed this observation further with us and 
with Charles Bryson. There appears to be no doubt as to what he 
observed. 

Hypolimnas misippus (121). In July and August 1970, Rick Ker- 
gosien and a number of his colleagues in Bay St. Louis, Hancock Co., 
Mississippi took a series of specimens of several species of generally 
more tropical facies than customary for Mississippi. These included: 
Anteos maerula lacordairei (25 July 70) and Anartia jatrophae guan- 
tanamo (30 July 70 and substantial numbers later). On 6 August 1970 
at Bay St. Louis, he and two others observed a female Hypolimnas 
misippus. It lit, was swung at, was missed, and slowly flew away while 
being chased for 2-3 min. It went over a roof top and was not seen 
again. All three observers quickly went to Kergosien’s residence to 
examine a female H. misippus in his collection. All three were in 
complete agreement that this was what they saw. The foregoing is 


summarized from a letter dated 6 August 1970 from Rick Kergosien 
to us. 


Checklist of Mississippi Butterflies 


* 1. Megathymus yuccae yuccae 7. Amblyscirtes samoset (Scudder ) 
(Boisduval & LeConte ) 8. A. aesculapius (Fabricius ) 
2. Panoquina panoquin (Scudder ) 9. A. carolina (Skinner ) 
3. P. ocola (Edwards ) * 10. A. reversa Jones 
!. Calpodes ethlius (Stoll) ll. A. vialis (Edwards ) 
5. Oligoria maculata (Edwards ) 12. A. belli Freeman 
6. Lerodea eufala (Edwards ) 13. A. alternata (Grote & Robinson ) 


VoLuUME 30, NUMBER 3 


. Atrytonopsis hianna hianna 


( Scudder ) 


. A. loammi (Whitney ) 
. Euphyes arpa (Boisduval & 


LeConte ) 


. E. palatka (Edwards ) 

. E. dion alabamae (Lindsey ) 

. E. dukesi (Lindsey ) 

. E. vestris metacomet ( Harris ) 
. Poanes hobomok ( Harris ) 


P. zabulon ( Boisduval & LeConte ) 
P. yehl (Skinner ) 

P. viator zizaniae Shapiro 
Problema byssus (Edwards ) 


. Atrytone arogos (Boisduval & 


LeConte ) 


. A. delaware delaware (Edwards ) 
. Atalopedes campestris ( Boisduval ) 
. Pompeius verna sequoyah 


(Freeman ) 
Wallengrenia otho (Smith) 
W. egeremet (Scudder ) 


. Polites themistocles (Latreille ) 


P. origenes origenes ( Fabricius ) 
P. vibex vibex (Geyer) 
Hesperia metea Scudder 

H. attalus seminole (Scudder) 
Hylephila phyleus phyleus 
(Drury ) 

Copaeodes minima (Edwards ) 


. Ancyloxypha numitor (Fabricius ) 


Lerema accius ( Smith ) 

Nastra lherminier ( Latreille ) 

N. neamathla (Skinner & 
Williams ) 

Pholisora catullus (Fabricius ) 
Pyrgus communis communis 
(Grote ) 

P. oileus (Linnaeus ) 

Erynnis brizo brizo (Boisduval & 
LeConte ) 


. E. zarucco (Lucas ) 

. E. funeralis (Scudder & Burgess ) 
. E. martialis (Scudder ) 

. E. horatius (Scudder & Burgess ) 
. E. juvenalis juvenalis (Fabricius ) 


Staphylus hayhurstii (Edwards ) 
Thorybes bathyllus (Smith) 

T. pylades (Scudder) 

T. confusis Bell 


. Achalarus lyciades (Geyer ) 
. Autochton cellus (Boisduval & 


LeConte ) 
Urbanus proteus proteus 
( Linnaeus ) 


nOGe 
OO: 


199 


U. dorantes dorantes (Stoll) 
Epargyreus clarus clarus 
(Cramer ) 

Battus philenor philenor 

( Linnaeus ) 

P. polyxenes asterius Stoll 

P. cresphontes cresphontes 
Cramer 

P. glaucus glaucus Linnaeus 
P. troilus troilus Linnaeus 

P. palamedes palamedes Drury 
Graphium marcellus (Cramer ) 
Pieris protodice protodice 
Boisduval & LeConte 

P. rapae ( Linnaeus ) 


. Ascia monuste phileta (Fabricius ) 


Colias eurytheme eurytheme 
Boisduval 

C. philodice philodice Godart 
C. (Zerene) cesonia (Stoll) 


. Anteos maerula lacordairei 


(Boisduval ) 
Phoebis sennae eubule (Linnaeus ) 
P. philea (Johansson ) 


. Eurema daira daira (Godart ) 


E. mexicana ( Boisduval ) 

E. lisa lisa Boisduval & LeConte 
E. nicippe (Cramer ) 

Nathalis iole Boisduval 


. Anthocharis midea midea 


( Hubner ) 

Calephelis virginiensis Gray 
Harkenclenus titus mopsus 

( Hubner ) 

Satyrium liparops strigosa 

( Harris ) 

S. kingi (Klots & Clench) 

S. calanus falacer (Godart ) 
Eumaeus atala florida Rober 
Calycopis isobeon (Butler & 
Druce ) 

C. cecrops (Fabricius ) 
Callophrys henrici turneri Clench 
C. augustinus croesioides Scudder 
C. niphon niphon ( Hubner ) 

C. gryneus gryneus (Hubner ) 
Atlides halesus halesus (Cramer ) 
Euristrymon ontario ontario 
(Edwards ) 

Panthiades m-album m-album 
(Boisduval & LeConte ) 

Strymon melinus melinus Hiibner 
Lycaena thoe Guérin-Méneville 
Brephidium isophthalma 
pseudofea ( Morrison ) 


200 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


*101. Leptotes marina ( Reakirt ) *121. Hypolimnas misippus (Linnaeus ) 
102. Hemiargus ceraunus antibubastus 122. Chlosyne nycteis nycteis 
Hiuibner ( Doubleday ) 
103. H. isola alce (Edwards) 123. C. gorgone gorgone ( Hubner) 
104. Everes comyntas comyntas 124. Phyciodes texana seminole 
( Godart ) (Skinner ) 
105. Celastrina argiolus pseudargiolus 125. P. tharos tharos (Drury ) 
(Boisduval & LeConte ) 126. P. phaon (Edwards ) 
106. Feniseca tarquinius tarquinius *127. Euphydryas phaeton ozarkae 
( Fabricius ) Masters 
107. Libytheana bachmanii bachmanii 128. Speyeria diana (Cramer ) 
( Kirtland ) *129. S. cybele cybele (Fabricius ) 
108. Anaea andria andria Scudder 130. Euptoieta claudia (Cramer ) 
109. Asterocampa celtis celtis 131. Heliconius charitonius tuckeri 
( Boisduval & LeConte ) Comstock & Brown 
110. A. clyton clyton (Boisduval & 132. Agraulis vanillae nigrior Michener 
LeConte ) 133. Danaus plexippus plexippus 
111. Limenitis arthemis astyanax ( Linnaeus ) 
( Fabricius ) 134. D. gilippus berenice (Cramer) 
112. L. archippus watsoni (dos Passos) 135. Lethe portlandia missarkae 
*113. Anartia jatrophae guantanamo Heitzman & dos Passos 
Munroe *136. L. anthedon (Clark) 
114. Vanessa atalanta rubria 137. L. creola (Skinner ) 
( Fruhstorfer ) *138. L. appalachia appalachia 
115. Cynthia virginiensis (Drury ) Chermock 
116. C. cardui (Linnaeus ) 139. Cyllopsis gemma gemma 
117. Precis coenia (Hubner ) ( Hubner ) 
118. Nymphalis antiopa antiopa 140. Euptychia areolata areolata 
( Linnaeus ) (Smith ) 
119. Polygonia interrogationis 141. E. hermes sosybia (Fabricius ) 
( Fabricius ) 142. E. cymela cymela (Cramer) 
120. P. comma ( Harris ) 143. Cercyonis pegala abbotti Brown 


Note added in proof: Species No. 144, Erynnis baptisiae (Forbes), was added to 
the list of those known from Mississippi as a result of one male having been taken 
on 19 March 1976 by Mike Rickard at the Big Biloxi Recreation Area, Harrison Co., 
on blackberry blossoms along the railroad track. The determination was confirmed by 
Dr. John M. Burns who previously (1964) regarded it as “rather surprising” that it 
had not then yet been found in Mississippi. 


LITERATURE CITED 


Burns, J. M. Evolution in skipper butterflies of the genus Erynnis. Univ. of Calif. 
Publ. Ent., vol. 37, 214 p. 

Hurcuins, R. E. 1933. Annotated list of Mississippi Rhopalocera. Can. Ent. 65: 
210-213. 

Maruer, B. and K. Marner. 1958. The butterflies of Mississippi. Tulane Stud. 
Zool. 6: 63-109. 

AND . 1959. The butterflies of Mississippi—Supplement No. 1. J. 
Lepid. Soc. 13: 71-72. 

Weep, H. E. 1894. A preliminary list of the butterflies of north-eastern Mississippi. 
Psyche 7: 129-131. 


VoLuME 30, NUMBER 3 201 


RHOPALOCERA IN THE N. B. SANSON COLLECTION 


CHARLES D. Birp 


Department of Biology, University of Calgary, 
Calgary, Alberta, Canada T2N 1N4 


Norman Bethune Sanson served as Curator at the Banff National 
Park, Alberta, Museum from 1896-1931, during which time he gathered 
together much insect, plant, and other material. This collection includes 
585 labelled Alberta specimens of skippers and butterflies, representing 
71 different taxa. These collections are reported here, along with some 
biographical notes on Sanson, as little has been published on either 
the Rhopalocera of the Park (Bean, 1890-1893) or on Sanson himself. 

N. B. Sanson was born on 1 November 1861, in Toronto, Ontario. 
He came west in 1885 as a member of the Queen’s Own Regiment and 
participated in the fierce battles against the forces of Louis Riel at 
North Battleford and Prince Albert, Saskatchewan. In 1892 he travelled 
to Banff and became an accountant at the Sanatorium Hotel and at a 
general store. In 1896 he was hired by the Canadian Government and 
appointed as a Meteorological Officer and as Curator of the Museum. 
His work in the first position involved the keeping of detailed weather 
records, especially in an observatory erected in 1902-1903 on the top 
of Sulphur Mountain at an elevation of 8,030 ft., some 3,500 ft. above 
the town of Banff. It has been estimated that he climbed to this 
observatory some 815 times, usually every two weeks, to man the 
weather instruments. Many of his collections were made on these hikes. 
His work as Curator involved the gathering of extensive collections 
and data on the local fauna and flora. The earliest Rhopalocera col- 
lection made by him is dated 10 May 1897, the latest 12 June 1929, 
but most collections were made in the period from 1906-1912. His 
collection includes a small number of specimens collected by J. Macoun, 
T. E. Bean, and J. Fletcher, and it is assumed that he was personally 
acquainted with these individuals. He was also acquainted with F. H. 
Wolley Dod as some of his collections are cited for Banff in the first 
list of the species of the province which Wolley Dod published in 1901. 
Wolley Dod and Arthur Gibson helped identify his material according 
to some determination labels. Sanson died at the age of 88 on 30 May 
1949 and was buried in the Banff Cemetery. 


Annotated List 


The following list includes all of the Alberta specimens, mostly from 
Banff National Park, in the N. B. Sanson collection. The geographical 


202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


location of the collections is indicated along with the flight period. 
The scientific names employed, unless otherwise indicated, are those 
of the latest list of North American taxa by dos Passos (1964) and 
of subsequent revisions to it by the same author (1969, 1970). The 
identifications of all of the specimens have been either verified or 
revised by the author. 


HESPERIUDAE 


Polites coras (Cramer)—Coras Skipper. Banff. July 27-31. 

Hesperia manitoba (Scudder )—Manitoba Skipper. Banff, Mystic Lake, Sulphur 
Mountain, Tunnel Mountain. June 18—-August 24. The name of this taxon, listed 
as H. comma manitoba by dos Passos (1964), was altered to the above by MacNeill 
(1964). 

Carterocephalos palaemon mandan (Edwards )—Arctic Skipper. Banff, Mt. Rundle, 
Spray River Valley, Sulphur Mountain, Tunnel Mountain. June 5—July 27. 

Pyrgus centaurae loki Evans—Grizzled Skipper. Banff. June 12. 

Pyrgus ruralis (Boisduval)—Ruralis Checkered Skipper. Banff, trail to Lake 
Minnewanka, Simpson Pass, trail to Stony Squaw Mountain, Tunnel Mountain. 
May 8-July 13. 

Erynnis icelus (Scudder & Burgess)—Dreamy Dusky Wing. Banff. June 25. 

Erynnis persius fredericki H. A. Freeman—Persius Dusky Wing. Banff, Sulphur 
Mountain, Upper Anthracite Road. May 4-July 14. 


PAPILIONIDAE 


Parnassius phoebus smintheus Doubleday—Parnassian. Aylmer Pass, Cascade 
Mountain, Simpson Pass, Stony Squaw Mountain, Sulphur mountain. August 6— 
October (day not mentioned ). 

Papilio glaucus canadensis Rothschild & Jordan—Tiger Swallowtail. Banff. May 
23-July 5. 


PIERIDAE 


Neophasia menapia menapia (Felder & Felder)—Pine White. Sulphur Mountain. 
August 10-September 6. 

Pieris sisymbrii flavitincta J. A. Comstock—California White. Tunnel Mountain. 
June 7. 

Pieris protodice occidentalis Reakirt-Western Checkered White. Banff, Sulphur 
Mountain. April 25-September 20. Considered a separate species, P. occidentalis, 
by some recent workers. 

Pieris napi oleracea Harris-Mustard White. Sundance Canyon Road, Tunnel 
Mountain, Upper Anthracite Road. May 23—August 8. 

Colias meadii elis Strecker-Elis Sulphur. Banff, Cascade Mountain, Ptarmigan 
Valley, Sulphur Mountain. July 21-September 4. 

Colias philodice eriphyle Edwards—Alfalfa Butterfly. Banff; F. H. Wolley Dod’s 
Ranch, SW of Calgary. June (day not mentioned)—October 4. Ultraviolet photo- 
graphic studies by Ferris (1972) and others have shown C. philodice and C. eury- 
theme to be different species. The taxon eriphyle belongs to the philodice complex. 

Colias interior interior Scudder—Pink-edged Sulphur. Ptarmigan Valley. June 21. 

Colias alexandra christina Edwards—Christina Sulphur. Banff, Lake Minnewanka, 
Sulphur Mountain, Sundance Canyon Road. July 18—-August 20. Four (57%) of 
seven females were albinistic. The subspecific nature of this species in Alberta has 
been discussed by Ferris (1973). 


VoLuME 30, NUMBER 3 203 


Colias pelidne minisni Bean—Pelidne Sulphur. Sundance Canyon Road. July 18. 

Colias nastes streckeri Grum-Grschimailo—Nastes Sulphur. Cascade Mountain. 
August 25. 

Euchloe creusa (Doubleday )—Creusa Marble. Banff, 40 Mile Creek Campground, 
Sulphur Mountain, Tunnel Mountain, Upper Anthracite Road. May 21-Septem- 
ber 25. 

Euchloe ausonides ausonides Lucas—Marbled White. Banff. May 17-July 1. 


LYCAENIDAE 


Callophrys polios obscurus Ferris and Fisher-Hoary Elfin. Upper Anthracite 
Road. May 28—June 9. This subspecies was recently described by Ferris & Fisher 
(1973)).. 

Callophrys augustinus iroides (Boisduval)—Brown Elfin. Banff, Sundance Canyon 
Road, Upper Anthracite Road. May 5—-June 9. The relationship of this subspecies 
and of ssp. augustinus (Westwood) in Alberta are discussed in dos Passos (1943). 

Callophrys eryphon eryphon (Boisduval)—Western Pine Elfin. Banff, Spray River 
Valley. April 24-June 24. 

Lycaena mariposa mariposa Reakirt—Mariposa Copper. Banff, trail to Lake Minne- 
wanka, Mystic Lake, Stony Squaw Mountain. July 25-September 25. 

Lycaena dorcas dorcas Kirby—Dorcas Copper. Banff. July 19-25. 

Lycaena phlaeas arethusa (Wolley Dod)—Arethusa Copper. Vermilion Range. 
Date not mentioned. 

Lycaena snowi (Edwards)—Snow’s Copper. Upper Kananaskis Pass, Vermilion 
Range. August 3. Alberta material may belong to ssp. henryae (Cadbury) de- 
scribed from Caribou Pass, B.C. 

Lycaeides argyrognomon scudderii (Edwards )—Scudder’s Blue. Banff, Lake Min- 
newanka, Stoney Creek to Cascade Valley, Sulphur Mountain, Sundance Canyon. 
July 10-August 11. 

Plebejus saepiolus amica (Edwards )—Saepiolus Blue. Banff, Laggan (J. Fletcher), 
Tunnel Mountain. June 19-August 3. 

Plebejus acmon lutzi dos Passos-Acmon Blue. Banff. August 7. Goodpasture 
(1973) has shown that lutzi is the only subspecies of Plebejus acmon in Alberta. 

Plebejus aquilo megalo McDunnough—Mountain Arctic Blue. Banff, Ptarmigan 
Valley, Simpson Pass, Sulphur Mountain. June 21—-September 1. 

Plebejus aquilo rustica (Edwards )—Plains Arctic Blue. F. H. Wolley Dod’s Ranch, 
SW of Calgary. June 19. 

Everes amyntula albrighti Clench—Western Tailed Blue. Banff, Sulphur Mountain, 
Tunnel Mountain, Upper Anthracite Road, Whiteman’s Cabin. June 9—August 2. 

Glaucopsyche lygdamus couperi Grote-Silvery Blue. Banff, Sulphur Mountain, 
Tunnel Mountain, Whiteman’s Cabin. June 11-July 29. 

Celastrina argiolus lucia (Kirby)—Spring Azure. Banff, base of Stony Squaw 
Mountain. May 21-June 24. 


NYMPHALIDAE 


Limenitis arthemis rubrofasciata (Barnes & McDunnough)—White Admiral. Anth- 
racite, Banff. July 12-30. 

Vanessa atalanta rubria (Fruhstorfer)—Red Admiral. Stony Squaw Mountain, 
Sulphur Mountain. June 24—26. 

Cynthia cardui (Linnaeus )—Painted Lady. Banff, Stony Squaw Mountain. June 
26—July 1. Specimens of this occasional migrant to Alberta were collected only in 
1911 and 1914. 

Nymphalis vau-album j-album (Boisduval & Le Conte )—Compton’s Tortoise-shell. 
Banff. August 18—September 30. 


204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Nymphalis californica californica ( Boisduval )—California Tortoise-shell. Banff, Sul- 
phur Mountain. One specimen of this occasional migrant was collected on June 10, 
1912, while 11 were taken on September 12, 1911. 

Nymphalis milberti (Godart )—Milbert’s Tortoise-shell. Banff, Cascade Mountain, 
Sulphur Mountain. July 30-August 29. 

Nymphalis antiopa antiopa (Linnaeus)—Mourning Cloak. Banff. August 24— 
September 26. 

Polygonia satyrus satyrus (Edwards)—Satyr Angle-wing. Banff, base of Stony 
Squaw Mountain. May 12-27. 

Polygonia faunus rusticus (Edwards )—Green Comma. Banff, Sulphur Mountain. 
April 23-30, September 2-23. 

Polygonia zephyrus (Edwards )—Zephyrus Angle-wing. Banff, Sulphur Mountain. 
May 7, August 9-September 16. 

Phyciodes tharos pulchella (Boisduval)—Pearl Crescent. Banff. July 15-31. 

Phyciodes campestris camillus Edwards-Meadow Crescent. Banff, Lake Minne- 
wanka, Sundance Canyon Road. June 17—August 6. 

Euphydryas anicia anicia (Doubleday )—Anicia Checkerspot. Banff, Lake Minne- 
wanka, Ptarmigan Valley, Stony Squaw Mountain, Sulphur Mountain. May 29- 
July 30. The type locality for anicia is Banff, Alberta. 

Boloria selene atrocostalis (Huard )—Silver-bordered Fritillary. Banff. June 17. 

Boloria toddi jenistai Stallings & Turner—Meadow Fritillary. Calgary. June 8. 

Boloria frigga saga (Staudinger )—Frigga Fritillary. Banff, Spray River Valley. 
May 21—June 30. 

Boloria freija freija (Thunberg)—Freija Fritillary. Banff, Rundle Mountain trail. 
April 28—-June 12. 

Boloria astarte astarte (Doubleday )—Astarte Fritillary. Sulphur Mountain. July 20. 

Boloria titania grandis (Barnes & McDunnough)—Purple Lesser Fritillary. Banff, 
Simpson Pass, trail to Stony Squaw Mountain, Sulphur Mountain. June 24— 
August 22. 

Boloria eunomia dawseni (Barnes & McDunnough)-—Bog Fritillary. Banff, Cas- 
cade Mountain, Ptarmigan Valley, Simpson Pass. June 3—August 26. 

Speyeria atlantis beani (Barnes & Benjamin)—Bean’s Fritillary. Banff, Sulphur 
Mountain. June 25—-August 27. 

Speyeria atlantis helena dos Passos & Grey—Northwestern Silverspot. Banff, Sul- 
phur Mountain. July 27—August 2. The distinction between the two subspecies of 
atlantis is often tenuous. The three specimens labelled ssp. helena were lighter 
underneath than those called ssp. beani. 

Speyeria hydaspe sakuntala (Skinner )—Hydaspe Fritillary. Banff, Sulphur Moun- 
tain. July 23-August 22. 

Speyeria mormonia eurynome (Edwards )—Mormon Fritillary. Banff, Upper Kan- 
anaskis Pass. July 1l—August 18. 


SATYRIDAE 


Coenonympha tullia inornata Edwards—Ringlet. Banff. June 24. Though referred 
to as C. inornata in dos Passos (1964), Brown (1955) has shown that inornata 
should be included within C. tullia. 

Cercyonis oetus charon (Edwards)—Small Meadow Brown. Banff. July 18— 
August 8. Emmel (1969) refers Alberta material to the ssp. charon and regards 
it as distinct from ssp. oetus. The two were regarded as synonymous by dos 
Passos (1964). 

Oeneis uhleri varuna (Edwards)—Varuna Arctic. Kananaskis (J. Macoun, no 
date); I’. H. Wolley Dod’s Ranch, SW of Calgary. June (no day mentioned). 

Oeneis chryxus (Doubleday )—Chryxus Arctic. Banff, Lake Minnewanka, Ptar- 
migan Valley, Spray River Valley, Sulphur Mountain, Tunnel Mountain. May 23- 


VoLuME 30, NUMBER 3 205 


July 25. Banff area material is close to ssp. caryi described from Smith Landing 
in extreme northeastern Alberta. 

Oeneis jutta chermocki Wyatt—Jutta Arctic. Banff, Lake Louise (J. Fletcher), 
Ptarmigan Valley, Spray River Valley. June 7—August 2. This subspecies was 
described by Wyatt (1965) after the appearance of dos Passos’ (1964) Synonymic 
List. Masters (1969) regards it as “a weak but valid subspecies somewhat inter- 
mediate between Oeneis jutta ridingiana . . . and Oeneis jutta reducta.” 

Oeneis melissa beanii Elwes—Bean’s Arctic. Sulphur Mountain. June 28-July 29. 

Oeneis polixenes brucei (Edwards)—Bruce’s Arctic. Ptarmigan Valley, Sulphur 
Mountain. July 23-29. 

Erebia disa mancinus Doubleday—Mancinus Alpine. Banff, Spray River Valley, 
Sulphur Mountain. June 18—July 12. 

Erebia discoidalis macdunnoughi dos Passos—Red-disked Alpine. Banff, May 2-— 
June 20. 

Erebia epipsodea epipsodea Butler-Mountain Common Alpine. Banff, Ptarmigan 
Valley, Sulphur Mountain. June 5—August 11. 

Erebia epipsodea freemani Ehrlich—Plains Common Alpine. F.-H. Wolley Dod’s 
Ranch, SW of Calgary, June (day not mentioned). Dos Passos (1964) regarded 
this taxon as ssp. sineocellata but as it was described on the basis of aberrant 
material (Ehrlich, 1955) “without ocelli,” sineocellata should be regarded as a 
“form” name only. 


ACKNOWLEDGMENT 


Biographical information was graciously provided by Miss Aileen 
Harmon. 


LITERATURE CITED 


BEAN, T. E. 1890-1893. Butterflies of Laggan, N.W.T.; account of certain 
species inhabiting the Rocky Mountains in Latitude 51°25’. Can. Ent. 22: 
O99 126-132 25: 145-149) 155-156. 

Brown, F. M. 1955. Studies of Nearctic Coenonympha tullia (Rhopalocera, 
Satyridae). Coenonympha tullia inornata Edwards. Bull. Amer. Mus. Nat. Hist. 
105: 359-410. 

pos Passos, C. F. 1943. Some new subspecies of Incisalia from North America 
(Lepidoptera, Lycaenidae). Amer. Mus. Novitates, No. 1230. 5 p. 

pos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. 
Soc. Memoir 1. 145 p. 

pos Passos, C. F. 1969. A revised synonymic list of the Nearctic Melitaeinae 
with taxonomic notes (Nymphalidae). J. Lepid. Soc. 23: 115-125. 

pos Passos, C. F. 1970. A revised synonymic catalogue with taxonomic notes on 
some Nearctic Lycaenidae. J. Lepid. Soc. 24: 26-38. 

EuruicH, P. R. 1955. The distribution and subspeciation of Erebia epipsodea 
Butler (Lepidoptera: Satyridae). Univ. Kansas Sci. 37: 175-194. 

EMMEL, T. C. 1969. Taxonomy, distribution and biology of the genus Cercyonis 
(Satyridae). I. Characteristics of the genus. J. Lepid. Soc. 23: 165-175. 
Ferris, C. D. 1972. Notes on certain species of Colias (Lepidoptera: Pieridae ) 
found in Wyoming and associated regions. Bull. Allyn Museum No. 5. 
Ferris, C. D. 1973. A revision of the Colias alexandra complex (Pieridae) aided 
by ultraviolet reflectance photography with designation of a new subspecies. 

femliepid: Soc. 27 2257=73: 

Ferris, C. D. & M. S. FisHer. 1973. Callophrys (Incisalia) polios (Lycaenidae ): 
distribution in North America and description of a new subspecies. J. Lepid. 


Soest: LIQ=118. 


206 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


GooppasturE, C. 1973. Biology and systematics of the Plebejus (Icaricia) acmon 
group (Lepidoptera: Lycaenidae). I. Review of the group. J. Kansas Ent. Soc. 
46: 468-485. 

Masters, J. H. 1969. An unusual nomenclatural problem regarding Oeneis jutta, 
(Lepidoptera: Satyridae). Bull. Assoc. Minnesota Ent. 3(2): 23-24. 

MacNeiz, C. D. 1964. The skippers of the genus Hesperia in western North 
America. Univ. Calif. Publ. Entomol. Vol. 35. 

Wottey Dop, F. H. 1901. Preliminary list of the macro-lepidoptera of Alberta, 
N.W.T. Can. Ent. 33: 40-42, 157-172. 

Wyatt, C. W. 1965. Zwei neue Formen von holarktischen Tagfaltern. Zeitschrift 
Wiener Ent. Geselschaft 50: 69-71. 


EVIDENCE OF BREEDING MIGRANT POPULATIONS OF 
LEPTOTES CASSIUS (LYCAENIDAE) IN KANSAS 


Three newly emergent specimens of Leptotes cassius Cramer were captured 
during a field study conducted on 15 June 1975 within the city limits of Law- 
rence, Kansas. The range of this species is normally southern Florida and southern 
Texas, and these records represent a rarity in northeastern Kansas (Lawrence is 
situated in Douglas Co., ca. 35 miles west of Kansas City). 

All three specimens were female; two were released with the intention of propa- 
gating the species locally, and one was mounted in lamination for a permanent 
record. Other records of L. cassius occurring in Kansas are as follows: (1) 24 
July 1935, Douglas Co., one specimen; (2) 4 July 1935, Scott Co., one specimen 
(Field 1938, Studies in Kansas Insects, p. 163-164). 

The presence of three females might indicate a local breeding (migrant) popu- 
lation, although Dr. J. C. Downey (pers. comm.) suggests: (1) pupae of L. 
cassius may have been imported with Plumbago transplants from Florida and (2) 
the Jack of northern populations even in northern Florida indicates that one must 
be cautious when concluding that an indigenous population exists in northeastern 
Kansas. A search of the capture area failed to yield any Plumbago, neither in 
adjacent lots nor in any greenhouse in Lawrence. However, Phaseolus is common 
throughout the neighborhood and is listed as an alternate larval food plant (Klots 
1951, A Field Guide to the Butterflies, p. 157-158). 

It is extremely unlikely that these three female specimens of L. cassius represent 
windblown migrants at such an early date in Kansas. Subsequent field collection 
data in this area are necessary and will be continued over the next two years to 
establish the probable migrant breeding residency of this butterfly. 


MatrHew M. Douctias, Department of Entomology, University of Kansas, Law- 
rence, Kansas 66044. 


VoLuME 30, NuMBER 3 207 


NOTES ON THE BIOLOGY AND IMMATURE STAGES OF THE 
WHITE PEACOCK BUTTERFLY, ANARTIA JATROPHAE 
GUANTANAMO (NYMPHALIDAE ) 


GrEoRGE W. Rawson 
10405 Amherst Avenue, Silver Spring, Maryland 20902 


Observations on the white peacock butterfly, Anartia jatrophae guan- 
tanamo Munroe, are recorded here to fill gaps in the documentation of 
this species’ life history. A small colony near New Smyrna Beach, 
Florida was studied in June 1970. 

Anartia jatrophae guantanamo was described from Guantanamo, Cuba 
(Munroe, 1942). Distributed locally in Florida, it is sometimes common 
in the east and west coastal areas. It has been reported as far north 
as Tampa on the west coast, and on the east coast it occurs in de- 
creasing numbers not much farther north than Daytona. Strays have 
been taken by collectors as far north as Savannah, Georgia and southern 
New England. The range in the United States extends as far south 
as the Everglades and Keys. 

Local distribution of the butterfly in Florida is confined to low 
ground along shallow ditches and the borders of ponds and streams 
where its larval food plant, Bacopa Monniera (L.) Wettst. (Scrophu- 
lariaceae) grows. Because the food plant, commonly called hedge or 
water hysop, depends on adequate moisture for survival, relative scarcity 
of the butterfly during drought may result from adverse effects of dry 
weather on the plant as well as on the butterfly itself. Usually a gravid 
female oviposits a single ovum on the ventral surface of a water hysop 
leaf and flies to a plant some distance away to deposit another single 
ovum. This scattered distribution of eggs may provide a higher rate 
of survival. 


Descriptions 


Ovum (Figs. 1A, B). Color pale pearly yellow, turns darker usually on second 
or third day. Shape similar to squat barrel; ten (+) evenly spaced longitudinal 
keeled ribs arise from flattened base and expand upward in conformity to wider 
portion of ovum, then narrow toward top where each rib ends abruptly at boundary 
of flattened micropile. 

Dimensions: Height 0.60 mm, width 0.50 mm. 

First instar (Figs. 1C, D). On emerging from ovum, larva has large black head 
capsule, about one-third larger than first thoracic segment; disproportion becomes 
less evident as larva grows. 

Color of body pale greenish-yellow, at first; after completely consuming its egg 
shell and feeding on succulent green leaves of water hysop, color changes to darker 
green, then brown just prior to ecdysis. 


208 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


re—_Ii = 25mm 


Fig. 1. Anartia jatrophae guantanamo Munroe. (A) ovum, lateral view; (B) 
ovum, dorsal view; (C) larva, first instar, lateral view; (D) larva, first instar, dorsal 
view; (E) larva, second instar, lateral view; (F) larva, second instar, dorsal view; 
(G) larva, third instar, dorsal view. 


Throughout first instar, shiny black head contrasts with lighter body; entire larva 
covered by fine blackish hair-like spines arising from dark pyramid-like bases; spines 
bend over the back in a gentle curve toward the head. 

Dimensions: Length of newly-emerged larva, 1.5 mm (+); full-grown, 3.0 mm 
G2eel 

Duration of first stadium: 4 days (+). 

Second instar (Figs. 1E, F). Head, mandibles, and forelegs shiny black; head 
somewhat rounded, its two branched spines appreciably longer and thicker than in 
first instar, as are branched spines on body. Cervical shield on first thoracic seg- 
ment has chain of four, dark, wartlike nodules. Anal segment bears two dorsolateral 
branched spines surrounded by short hair-like bristles. 

Dimensions: Length of newly emerged larva, 3.0 mm (+); full-grown, 8-10 mm. 

Duration of second stadium: 4 days. 

Third instar (Figs. 1G, 2A, B, C). It is difficult to determine the change from 
second to third instar unless, in ecdysis, the cast skin is noticed. Careful observation 
is needed here because the skin soon is consumed by the larva after ecdysis. 


VoLuME 30, NUMBER 3 209 


Fig. 2. Anartia jatrophae guantanamo Munroe. (A) larva, third instar, facial 
mask; (B) larva, third instar, one of dorsal branched spines on head; (C) larva, 
third instar, one of branched spines on body; (D) pupa, from left to right, dorsal, 
lateral, ventral views. 


210 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Immediately after ecdysis, head, branched spines, legs, and prolegs conspicuous 
dull yellow; within 30 (+) minutes, the same structures turn dark brown to black. 

Basic body color dark brown to black, but ventral surface lighter dull yellowish- 
brown; chain of very small silver spots runs closely parallel to anterior and posterior 
margins of each body segment on dorsal and ventral surfaces; papillae or bases of 
spines and prolegs dull orange. Anal segment equipped with curved hooklets. From 
mesothorax to next-to-last abdominal segment, five parallel rows of spines run 
lengthwise along body; median dorsal row is flanked left and right by two dorso- 
lateral rows of somewhat larger spines. From head, prothorax, and anal segment 
arises a pair of dorsolateral spines, the cephalic ones terminating in clubs. 

Dimensions: Length of newly emerged larva, 10.0-12.0 mm; full-grown, 27.0- 
30.0 mm. Length of full-grown third instar larvae indicates sex, as larger individuals 
develop into female imagoes. 

Duration of third stadium: 4—5 days. 

Pupa (Fig. 2D). Surface smooth, unornamented, light green but darkens in color 
nearing time of eclusion; dorsal surface has 2—4 light spots on each segment divided 
by black central line extending from head to tail; scattered white spots on wing pads 
and on ventral surface. Posterior end of body terminates in dark brown cremaster. 

Dimensions: average length, 17.5-—20.0 mm. 

Duration of pupal stadium: 7-10 days. 

Pupa hangs head down suspended by cremaster embedded in pad of silk attached 
to some support. Eclosion of imago takes place in about 15 min., and 20 min. or 
more pass before its wings can sustain flight. 


ACKNOWLEDGMENTS 


I wish to express my thanks and appreciation for assistance rendered 
to me by Drs. H. A. Denmark and Howard V. Weems, Jr., Florida Depart- 
ment of Agriculture, Gainesville; and also, to Dr. Lee D. Miller, Allyn 
Museum for preparing photographs of Anartia 7. guantanamo genitalia. 
[ am particularly grateful to my colleague Mr. Marc Roth, Entomology 
Department, National Museum of Natural History for his assistance in 
correcting and improving my manuscript. 


LITERATURE CITED 


Munroe, E.G. 1942. The Caribbean Races of Anartia jatrophae Johansson (Lepi- 
doptera: Nymphalidae). Amer. Mus. Nov. no. 1179. 4 p. 


VoLuME 30, NUMBER 3 PE 


A NEW SPECIES OF THE GENUS BERTELIA B. & McD. 
(PYRALIDAE ) 


ANDRE BLANCHARD 
P.O. Box 20304, Houston, Texas 77025 


An examination of specimens of Bertelia grisella B. & McD. in the 
National Museum, including the cotype, shows that I had misidentified 
my series of specimens of the same genus, taken in the Presidio and 
Culberson cos. of Texas, recorded in my 1970 article. These specimens 
really belong to a new species, the description of which follows. 


Bertelia dupla A. Blanchard, new species 


The description of the habitus of Bertelia grisella, as published by Barnes & 
McDunnough (1913) and Heinrich (1956) applies extremely well to the new species. 
The specimens of B. grisella in the National Collection (ex Barnes Collection) are 
somewhat paler but this is probably due, at least in part, to fading. The maculaticn 
of the new species is shown in Figures 1 and 2. 

Male genitalia (Figs. 3-6): Uncus triangulate. Valves simple. Vinculum broadly 
rounded. Apical process of gnathos developed as a long, tapered hook narrowly 
and deeply notched at apex. Transtilla with strong sclerotization limited to two 
long processes embracing the aedeagus by their bases, extending dorsad of the 
aedeagus beyond the base of the gnathos hook, weakly united a trifle distad of 
their middle, where they are narrowest, by a short, narrow bridge, enlarged and 
flattened basad of this bridge, enlarged and tricuspid at their apices. Juxta U-shaped 
with long flattened lateral arms, widest near their middles, pointed at their apices. 
Aedeagus with a row of minute spines on each of two lateral, symmetrical edges. 
Penis with a few sclerotized wrinklings, otherwise unarmed. Eighth abdominal 
segment with a pair of ventrolateral hair tufts. 

Female genitalia (Figs. 8-10): Similar to those of Bertelia grisella. 

Wing expanse: 23-27 mm, average 25.5 mm. 

Holotype: Male, Shafter, Presidio Co., Texas, 19 Oct. 1973, deposited in the 
National Museum of Natural History (No. 73530). 

Paratypes: Shafter, Texas, 18 Oct. 1968, 3 4, 2 2; 15 Oct. 1969, 8 ¢, 289; 
16 Oct. 1973, 1 6, 4 2; 19 Oct. 1973, 19; Guadalupe Mts. Nat. Park, Bear 
Canyon, 2 Oct. 1969, 1 ¢, 3 @. All types collected by A. & M. E. Blanchard. 


B. grisella is the only other species in the genus Bertelia. It is to be 
expected that, when fresh, unfaded specimens of B. grisella are available, 
it will be necessary to dissect the males to distinguish them from B. 
dupla. The transtilla of B. grisella, shown in Figure 7, is abundantly 
different: the laterodorsal processes are much shorter and do not reach 
the base of the gnathos hook, and the shape of the enlarged apices 
differs considerably. 


aT? JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


\ 


NG 


RG 


S 


WS 


HK 
\ \\ 


Figs. 1-2. Bertelia dupla: 1, male holotype; 2, female paratype. 

Figs. 3-6. Bertelia dupla male genitalia, slide A. B. 3595: 3, genitalia, aedeagus 
omitted; 4, tufts of eighth abdominal segment; 5, aedeagus; 6, enlarged genitalia 
showing. transtilla. 

Fig. 7. Bertelia grisella, cotype male genitalia, aedeagus omitted, enlarged to 
show transtilla, slide USNM 52494 by A. B. 

Figs. 8-10. Bertelia dupla, female genitalia, slide A. B. 3594: 8, genitalia in- 
cluding seventh abdominal segment; 9, enlarged posterior part; 10, signum of bursa. 


VoLUME 30, NUMBER 3 IES 


ACKNOWLEDGMENTS 


My thanks are due to Mr. Roger Reisch, ranger in the Guadalupe 
Mts. National Park for helping me to set my traps in the hard to reach 
Bear Canyon, to Mr. Philip F. Van Cleave for permission to collect 
there, to the National Museum for the loan of seven specimens and 
to Dr. Douglas C. Ferguson for arranging the loan. 


LITERATURE CITED 


Barnes, W. & J. H. McDunnoucHu. 1913. Bertelia grisella. Contrib. Nat. Hist. 
Mepis Ne Ay 2(3): Oo. 

BLANCHARD, A. 1970. Observations on some Phycitinae (Pyralidae) of Texas. J. 
Lepid. Soc. 24: 254. 

Hernricu, C. 1956. American moths of the subfamily Phycitinae. U.S. Natl. 
Muss-pull, 207: 37. 


A POPULATION OF THE STRIPED HAIRSTREAK, SATYRIUM LIPAROPS 
LIPAROPS (LYCAENIDAE), IN WEST-CENTRAL FLORIDA 


As a resident species, Satyrium liparops liparops Boisduval & LeConte, has been 
previously reported only from the north Florida border and panhandle areas (Kimball 
1965, Vol. I, Div. Plant Industry, Gainesville, 363 p.). However, on 15 May 1973, 
a freshly emerged female S. /. liparops was captured at Chassahowitzka, Citrus Co., 
Florida along the border of a hydric forest at the headwaters of the Chassahowitzka 
River. Two other adults were observed but not collected in the same location on that 
date. They were present in an ecotone area of young and mature hammock trees 
dominated by basswood (Tilia floridana), southern magnolia (Magnolia grandiflora), 
water ash (Fraxinus caroliniana), sweet bay (Magnolia virginiana), water oak 
(Quercus nigra) and bald cypress (Taxodium distichum). 

The area was revisited in early June 1975, and two more S. I. liparops were col- 
lected and several others observed. These specimens were more wom than ithe 
female collected in May 1973. All adults observed or collected at this Florida west- 
coast locality are typical S. l. liparops having the conspicuous orange-brown patches 
on the upper sides of the wing rather than the subspecies, S. /. strigosa, which occurs 
over wide areas of Georgia. 

A careful examination of vegetation in the Chassahowitzka area produced two 
early instar larvae of S. liparops (identified by rearing) in mid-June 1975. They 
were found on tree blueberry (Vaccinium sp.) in the same area where the adult 
hairstreaks were previously encountered. In Georgia (Harris 1972, Univ. Okla. Press, 
Norman, 326 p.), S. liparops produces only one brood annually with adults flying 
from May-July. This is compatible with my Florida data. 

The presence of a population of S. liparops halfway down the west coast of penin- 
sular Florida suggests that the striped hairstreak may be present over a much wider 
area of the southern Gulf and Atlantic Coastal Plains than previously reported. 


Larry N. Brown, Department of Biology, University of South Florida, Tampa, 
Florida 33620. 


214 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


THE STATUS OF SATYRIUM BOREALE (LYCAENIDAE) 


LAWRENCE F. GALL 


Peabody Museum of Natural History, Yale University, 
New Haven, Connecticut 06520° 


The hairstreak butterfly Satyrium boreale (Latontaine) was described 
by its author as phenotypically similar to Satyrium falacer (Godart). 
In samples from the Peabody Museum and the collections of D. F. 
Schweitzer and the author, intergradations appear between the wing 
patterns of these two species. This observation, along with the noted 
plasticity in wing pattern of S. falacer and the related Satyrium carya- 
evorum (McDunnough), prompted a detailed investigation of the status 
of S. boreale. I propose that the characters of S. boreale fall within 
the pattern of variation of S. falacer. 


EXPERIMENTAL PROCEDURES 


I have chosen to designate phenotypic characters consistent with those 
described for S. boreale by Lafontaine as “type B” and those described 
for S. falacer as “type F.” From ca. 120 available male Satyriwm, a 
colleague selected a random sample of 96 specimens, unsorted with 
respect to their phenotypic characteristics. He then removed the ab- 
domens, placing them individually into glass vials, with the specimens 
and abdomens cross-referenced by number. I then attempted a genitalic 
determination for each abdomen, without knowing from which specimen 
it came. Next, I sorted the pinned specimens by type B and type F 
wing pattern without reference to the genitalic determinations. In both 
methods of determination, the characters distinguishing S. boreale from 
S. falacer (Lafontaine, 1970) were followed closely. The Satyrium 
examined were from the Peabody Museum and the author's collection. 
Of the 96 Satyriwm, 2 were genitalically determined as caryaevorum 
and set aside. Locality data for the sample specimens are as follows: 
Connecticut, 69; Virginia, 6; New York, 3; Massachusetts, 3; Wisconsin, 
2; Illinois, 2; Kansas, Pennsylvania, Colorado, and Missouri, each 1 
specimen. Five other specimens had no data labels. 


RESULTS AND Discussion 


In the S. falacer-boreale series, 65 specimens were identified by wing 
pattern as type F and 29 as type B. From genitalic determinations, 
43 were identified as type F and 51 as type B. 


Mgt address: Department of Biological Sciences, Stanford University, Stanford, California 
94305. 


VoLUME 30, NUMBER 3 215 


Taste 1. Test of correlation between wing pattern and saccus characters in male 
Satyrium.* 


Type B Wing Pattern Type F Wing Pattern 
(n= 29) (nr==; 6) 
Expected if Expected if 
Saccus uncorrelated Observed uncorrelated Observed 
ivperb (Cn = 51) 15.7 ley oo 34 
ivpers (m= 43) 13.3 1 29 31 
* Chi-square for three df = 0.344; p = .050; association is random (methods from Guenther, 


1965). 


These procedures permit a test of the hypothesis that S. boreale 
represents a wing-pattern variant of S. falacer and, therefore, that the 
two are genitalically indistinguishable. If this hypothesis is correct, 
one would expect to find the same ratio of genitalic types in each 
wing pattern class as in the total sample. The ratio for the total sample 
was 51 type B:43 type F, or 0.543:0.457. Table 1 compares the wing- 
pattern determinations with the genitalic determinations for the same 
specimens, listing expected and actual genitalic type totals for each wing 
type. A Chi-square test of the data from Table 1 demonstrates that 
the hypothesis may be accepted at the 95% confidence level. 

It must be stressed that Lafontaine gave no biological criteria for 
considering S. boreale to be a distinct species, and its status is based 
entirely on phenotypic characters of dead specimens. Therefore, testing 
the validity of S. boreale depends on utilizing Lafontaine’s explicit 
characters in a detailed examination of dead specimens. 

The genitalia of these Satyrium deserve particular attention. The 
sole genitalic character cited in Lafontaine’s redescription of S. boreale 
is the shape of the saccus. In S. boreale the saccus “narrows evenly 
throughout its length,” whereas in S. falacer it is “strongly constricted 
subbasally.” Figures of male S. falacer genitalia in the literature show 
considerable variation (Forbes, 1960; Klots & Clench, 1952; Lafontaine, 
1970). The genitalia of a male of S. falacer as figured by Forbes are 
intermediate between those figured by Lafontaine for S. boreale and 
S. falacer, and the saccus of S. falacer shown by Klots & Clench is 
much broader than in either of the former figures. Because of the 
variability of the diagrams cited above, short notes were kept on each 
dissection, e.g., “constriction consistent w/ Forbes.” 

In the genitalia examined, those with the saccus intermediate between 
Lafontaine’s S. boreale and Forbes’ S. falacer are more numerous (37 
of 94) than either distinct type F (28 of 94), or distinct type B (29 


16 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-5. Males of Satyrium “boreale”’: 1, Southford Falls State Park, New 
Haven Co., Conn., 2 July 1975, L. F. Gall & D. F. Schweitzer; 2, same data; 
3, West Rock, New Haven Co., Conn., 4 July 1975, L. F. Gall; 4, Southford Falls 
State Park, New Haven Co., Conn., 4 July 1975, L. F. Gall; 5, West Rock, New 
Haven Co., Conn., 19 July 1958, W. B. Watt. 


Figs. 6-10. Genitalia of same specimens in Figs. 1-5. 


Figs. 11-15. Males of Satyrium falacer: 11, Southford Falls State Park, New 
Haven Co., Conn., 4 July 1975, L. F. Gall; 12, Southford Falls State Park, New 
Haven Co., Conn., 2 July 1975, L. F. Gall & D. F. Schweitzer; 13, Dousman, 


VoLuME 30, NuMBER 3 DAG 


of 94) genitalia. These genitalic intergradations were observed for both 
groups sorted by wing pattern. Representative series of the intergrada- 
tions appear along with the specimens from which the genitalia came 
in Figures 1-20. Also, 13 specimens show saccus characters in strong 
contradiction to their respective wing patterns: 8 have the type B 
saccus and type F wings, sensu Lafontaine, and 5 have the reciprocal. 

Four other observations warrant attention. First, in Lafontaine’s 
redescription of boreale, he stated that the posterior (6th) spot of the 
subterminal lines on the primaries shows “a trace of white only, never 
with any trace of black.” I have looked at those specimens which I 
termed type B wing pattern under a dissecting microscope and have 
always noted at least three, often as many as nine or ten, dark scales 
in this area. In a specimen in the Peabody Museum determined by 
Lafontaine as S. boreale (taken 3 July 1965, Ottawa, Ontario, leg. J. D. 
Lafontaine), I noted four dark scales present; this specimen proved 
to have a type F saccus. Secondly, a captive female of type B wing 
pattern (taken at UV light, 3 July 1975, Southford, Connecticut, leg. 
D. F. Schweitzer) laid 8 eggs on shagbark hickory, Carya ovata ( Mill.) 
K. Koch, a food plant of S. falacer in Connecticut. The female was 
kept alive for 15 days in a flight cage containing red oak, scrub oak, 
white oak, shagbark hickory, butternut, beech, and an ash. Third, I 
found no difference in the ostium bursae or signum between four 
females with type F wing pattern and four females with type B wings 
(Illinois, 3; Connecticut, 2; Massachusetts, 2; and Maryland, 1). Lastly, 
W. J. Holland (1930) shows in Plate 54, Fig. 21, a specimen of S. falacer 
(= Thecla calanus) with ideal type B wing characters. 


SUMMARY 


The data show that no correlation exists between the wing pattern 
and saccus characters on which S. boreale was based. No evidence is 
presently available to support the claim that S. boreale is a distinct 
species. The S. boreale characters appear to fall within the normal 
pattern of variation of S. falacer. The data also show the extreme 
phenotypic variability of S. falacer. It is possible that the S. boreale 
characters predominate in the north or north-central part of the range 


< 


Waukesha Co., Wisc., 8 July 1950, N. Euting; 14, Greenwich, Fairfield Co., Conn., 
22 June 1941, leg. Starrett; 15, Greenwich, Fairfield Co., Conn., 15 July 1940, leg. 
Starrett. 


Figs. 16-20. Genitalia of same specimens in Figs. 11-15. 


218 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


of S. falacer, in which case the name might be preserved for the end 
of a clinal trend. 


ACKNOWLEDGMENTS 


I wish to thank Dr. Charles L. Remington for his help in preparing 
this paper. I also wish to thank Dale F. Schweitzer for the use of his 
specimens and his helpful discussions on Satyriwm. Lastly, thanks to 
David G. Furth for his technical assistance and sometimes pointed, 
but lighthearted, encouragement. 


LITERATURE CITED 


Forses, W. T. M. 1960. Lepidoptera of New York and neighboring states; 
Agaristidae through Nymphalidae, including butterflies. Cornell Univ. Agric. 
Exp. Sta. Memoir 371. 188 p. 

GuENTHER, W. C. 1965. Concepts of statistical inference. McGraw-Hill, New 
York, 393) p: 

HoLianp, W. J. 1930. The butterfly book, revised edition. Garden City, New 
York. 424 p. 

Kuors, A. B. & H. K. Crencn. 1952. A new species of Strymon Huebner from 
Georgia (Lepidoptera: Lycaenidae). Amer. Mus. Novitates, No. 1600. 

LAFONTAINE, J. D. 1970. <A redescription of Strymon borealis Lafontaine (Lycae- 
nidae). J. Lepid. Soc. 24: 83-86. 


OIKETICUS TOUMEYI: A BAGWORM MOTH NEW TO THE 
TEXAS FAUNA (PSYCHIDAE) 


In the first compilation of bagworm moths (Psychidae) of Texas, Jones & Park 
(1928, Tex. Agr. Exp. Sta. Bull. 382: 36 p.) listed 13 species that were then 
known to occur in the state with three additional species to be anticipated. One 
of these anticipated species was the mesquite bagworm, Oiketicus touwmeyi Jones. 
At that time the species was known only to be “widely distributed in southeastem 
Arizona.” Thirty-six years later, O. towumeyi has not yet been reported from Texas 
(Davis 1964, Bull. U.S.N.M. 244: 233 p.), although it was known from Las 
Cruces, New Mexico, some 40 km from the Texas-New Mexico line. 

On 24 October 1970, a single bag containing a live larva was collected on 
ocotillo (Fouqueriaceae: Fouqueria splendens Engelm.) in Presidio, Presidio Co., 
Texas on the Rio Grande River. The bag measured 62 mm in length and 12 mm 
at greatest width. The bag was only slightly ornamented with ocotillo thorns and 
looked quite similar to the bag illustrated by Davis (op. cit., Fig. 98), which had 
been reared on mesquite. The present report is also a new host plant record for 
O. toumeyi (Davis lists nine other food plants). 

The collection of O. toumeyi from Presidio indicates that this species most prob- 
ably occurs over a widespread area of the southwestern United States and northern 
Mexico, albeit sporadically. 

I wish to thank D. Otte, who collected the specimen. 


RAyMOND W. Neck, Texas Parks and Wildlife Department, John H. Reagan 
Building, Austin, Texas 78701. 


VoLuME 30, NUMBER 3 219 


THE GIANT BLASTOBASID MOTHS OF YUCCA 
(GELECHIOIDEA ) 


Jerry A. POWELL 
University of California, Berkeley, California 94720 


Members of the family Blastobasidae are mostly small and uniformly 
drab moths which are characterized by bands of stout spines on the 
abdominal tergites. The forewings are often gray or brownish with 
one or more transverse bands of whitish. Species of subfamily Pigritinae 
are usually less than 12 mm in wingspread and have small or minute 
labial palpi, and often their forewings are yellowish or brownish, uni- 
colorous or marked by a straight, transverse band. By contrast Blasto- 
basinae are mostly larger, exceeding 12 mm in wingspread, with gray 
forewings, frequently showing a chevron-shaped pale marking. Their 
labial palpi are of the typical gelechioid form, elongate and strongly 
upcurved, reaching the crown or higher. 

Taxonomic relationships within the family are only preliminarily 
known because no comprehensive assessment of genitalic characters 
has been attempted. The state of knowledge for the North American 
fauna is as poorly documented as that of any sizeable group of Micro- 
lepidoptera. The genera are based on wing venation and secondary 
male characters (Dietz, 1900, 1910; Forbes, 1923) and need to be 
readjusted by use of genital features. Preliminary steps in this direction 
were taken by McDunnough (1961), who based his decisions in pro- 
posing two new genera on too little material. About 100 species are 
described, but probably at least twice that many are represented in 
collections. 

In 1974 I examined all types of Nearctic Blastobasidae at the National 
Museum of Natural History and Museum of Comparative Zoology 
(Harvard) (about 70% of the names) in order to develop a framework 
for biological studies of western species. Dissections of virtually none 
of the types have been made, and a number of Chambers or Dietz 
types are lost or are without abdomens, which will complicate matters 
for future students, as will Dietz’s species concepts which were typo- 
logical and imponderably fine-lined or heterogeneous from one time to 
another. In general it appears that considerable synonymy will be 
necessary among species of eastern North America, especially in the 
Pigritinae, while the western fauna remains largely undescribed, and 
the few species that have been named probably are valid. 

Larvae of Blastobasinae are characterized by a well-developed sub- 


220 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


mental pit; otherwise they resemble those of the scavenger group of 
Oecophoridae. Species for which habits have been studied feed in a 
wide variety of situations, mostly associated with detritus, often in 
nests of external feeding caterpillars or in galleries of insect borers. 
Rearing records include flower or seed heads of sumac (Forbes, 1923), 
Labiatae, Leguminosae, and Dipterocarpaceae (Fletcher, 1920, 1933); 
in acorns usually but not always following weevil borings (Craighead, 
1950); a variety of conifer cones and foliage in association with various 
moth larvae (Fletcher, 1920; Lyons, 1957; Keen, 1958; Powell unpubl. 
data); in fallen fruit of Ficus (Fletcher, 1920), coffee (Busck & Oliviera, 
1925), palm (Common, 1970) and apple mummies (Forbes, 1923); 
and in association with coccid colonies (Riley, 1887; Essig, 1916; Costa 
Lima, 1945; Fletcher, 1920; Forbes, 1931). However, in at least one 
example a species thought to be a predator was later proved to be a 
general feeder (Basinger, 1924). One species is said to be a gall maker 
(Costa Lima, 1945 after Brethes, 1917), a relationship that needs 
confirmation. 

The general conclusion that one reaches is that larvae of most species 
are opportunists, with colonies basically established as scavengers, but 
individuals feeding in unaffected plant tissue when convenient or even 
acting in a predator role when a susceptible prey such as scale insects, 
beetle larvae (Lyons, 1957) or moth pupae (Keen, 1958) are en- 
countered. 

Apparently the biology of no member of the Pigritinae has been 
recorded. I reared Pigritia arizonella Dietz from root crowns of Chrys- 
othamnus (Compositae) which were tunnelled by larvae of Eucos- 
mini and Cochylidae (Hot Creek, Mono Co., Calif., VII-11-68, J. Powell 
no. 68G22). Although the feeding habits of P. arizonella were not 
observed, the record suggests that species of this subfamily also are 
scavengers, but feed on the ground or in subterranean situations. Thus 
their larvae have been overlooked because North American insect bi- 
ologists have tended to ignore this ecological horizon. 

In some situations species of Holcocera are general scavengers which 
exploit various habitats. For example, at Dune Lakes, San Luis Obispo 
County, California, I reared one species from bacterial cankers on 
Salix, abandoned nests of Lepidoptera on Baccharis and occupied ones 
on Eriogonum, and from sporophores of Polyporaceae used by ciid 
beetles. In general, however, data are too preliminary to permit con- 
clusions about habitat specificity because the taxonomy is incomplete 
and because there are few rearing records relative to the number of spe- 
cies for which adults are known. 


VoLUME 30, NUMBER 3 221 


Holcocera gigantella was described from Colorado by Chambers 
(1876), who found the adults perched and mating on leaves of yucca. 
The biology of this species appears to be exceptional in that a specific 
habitat is used and prior insect colonies may not be a necessity. Widely 
disjunct populations showing varying degrees of differentiation have 
been discovered in association with Agavaceae. Larvae feed in Yucca 
inflorescences after the pods are developing and in flowers of Agave. 

Some information on the biology of H. gigantella has been given 
elsewhere (Powell & Mackie, 1966). During a study of the moths 
associated with Yucca whipplei larvae were found in large numbers in 
old pods and in green fruit prior to seed maturation. At one seaside 
locality in southern California there appeared to be continuous gen- 
erations. Pupae that produced adults within a few days were collected 
in February, March, July, and October. 

A second species, described below, occurs sympatrically with H. 
gigantella in southern Arizona in association with Yucca schottii. Al- 
though the new species lives in the same habitat and also is one of the 
largest Nearctic blastobasids, the two may not be phylogenetically 
closely related. Markedly different morphological characteristics, espe- 
cially in female genitalia, are exhibited by the two. The present clari- 
fication of the status of these blastobasids provides background infor- 
mation for the Y. schottii study as well as a name for the second species. 


Holcocera gigantella (Chambers ) 


Blastobasis gigantella Chambers, 1876, Can. Ent. 8: 219. 

Holcocera gigantella; Dyar, 1903, Bull. U.S. Natl. Mus. 52: 529; Dietz, 1910, Trans. 
Amer. Ent. Soc. 36: 29 (taxonomy); Powell & Mackie, 1966, U. Calif. Publ. Ent. 
42: 41 (biology). 


This is the largest Nearctic blastobasid. The forewings are gray with variable, 
longitudinal blackish markings. 

Male. Length of forewing 11.1-14.7 mm. Head: Labial palpus moderately 
elongate, strongly upcurved, exceeding crown; II segment length 1.1—1.3 times eye 
diameter; III = 0.65-0.70 as long as II; smooth scaled, whitish with scattered dark 
gray scales. Antennal scape elongate, length 0.9 eye diameter, broadened distally: 
2nd segment short, broad, notched dorsally; 3rd segment claw-shaped, greatly 
broadened basally with a deep, rounded dorsal notch distally (Fig. 2); setae of 
shaft elongate, length greater than diameter of 4th and succeeding segments. 
Scaling of front and crown appressed, whitish to grayish. Thorax: Dorsal scaling 
white to gray; underside whitish, legs grayish exteriorly, hind tibia with enlarged, 
white dorsal fringe. Forewing: Narrow, length 4.5—4.9 times width. Ground color 
whitish to gray, usually with obscure to well-defined longitudinal, blackish lines 
along the veins, at least on distal half. In the typical form the lines are well- 
defined from base to apex, distinctly contrasting with the whitish ground and 
forming a short bar in middle of cell above fold and a transverse line or paired 
spot at end of cell; more often the markings obscured on basal half, with lines 
and spots of distal half strongly contrasted on a pale to dark gray ground; rarely 
with additional infuscation broadening the lines into spots that tend to form 


222 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Nae 


ate 
Whee / 
Me Lf HA / 


Figs. 1-3. Holcocera gigantella (Chambers): 1, setal patterns on thoracic seg- 
ments I-II, abdominal segments 1, 2, 6-10; 2, outline of antennal segments 1-3, 
eye, dorsolateral aspect; 3, setation of second abdominal tergite. Fig. 4. H. paradoxa 
Powell, setation of second abdominal tergite (posterior margin at top). 


VoLuME 30, NUMBER 3 ZAG 


transverse bands at middle and in distal third of wing. Fringe pale gray, without 
extensions of the longitudinal markings. Underside dark gray, in pale forms with 
suggestion of whitish lines between veins in terminal area. Hindwing: Costal 
margin moderately excavate on distal third; width at end of cell about equal to 
forewing. Veins M; + Cu; stalked a varying distance, M: free, connate at base 
of M; + Cu: or from their stem. Base of anal margin and 3A with elongate, 
whitish ochreous hair tufts. Upperside scaling whitish and roughened on_ basal 
third, becoming pale gray and smooth distally; fringe paler, whitish. Underside 
similar, darker. Abdomen: Tergite 1-2 with a dense row of spines posteriorly 
and scattered smaller spines over whole surface (Fig. 3), 3-7 with compact, narrow 
posterior spine row only. Scaling shining whitish gray, obscuring the spine rows 
in fresh specimens. Genitalia as in Fig. 5 (drawn from lectotype, JAP prep. no. 
4008, 9 preparations examined); transverse band of gnathos broad, with posterior 
notch; valva with a basal, triangular-shaped flap; costal process and free portion 
of sacculus relatively elongate, ca. 0.3 the total length of valva; sclerotized band 
of aedeagus forming a narrow ring at base. 

Female. Length of forewing 11.7-14.7 mm. Essentially as described for male, 
differing as follows: labial palpus more elongate, II segment length 1.4-1.5 times 
eye diameter, III = 0.80-0.85 as long as II. Second and 3rd antennal segments 
unmodified; antennal ciliation short, less than 0.5 segment diameter. Forewing 
pattern variable, tending to be darker than male in any given population; fore- 
wing broader, length 4.3-4.6 times width. Genitalia as in Fig. 7 (drawn from 
paralectotype, JAP prep. no. 4009, 6 preparations examined); base of ductus bursae 
unmodified, without spiculae or scobination, gradually enlarged and abruptly turned 
before entering corpus bursae; signum broad, with a deep median crease, smooth 
surfaced. 

Lectotype. Male, by present designation; Colorado, in MCZ; bearing labels: 
“Chambers Color.,” “Type 1551,” “Blastobasis gigantella Col.” (the last in Cham- 
bers’ handwriting); and Genitalia 4008, JAP ’75. There are two females and a 
specimen lacking metathorax and abdomen with the same data, in MCZ, and a 
specimen in poor condition with the Chambers determination label in NMNH. 
In the original description Chambers stated that he had met with the species only 
once, on the road to Monument Park, about 3 mi. N of Colorado Springs. 

The type series is of the phenotype with whitish forewings marked by strongly 
contrasting dark longitudinal lines. Most specimens from certain California localities 
(San Luis Obispo Co., Santa Barbara Co., San Bernardino Mts., Mojave Desert ) 
are similar, usually with more blackish markings. The material from San Diego 
County and Baja California is generally darker, most individuals having a uniform 
dark gray ground marked by diffuse longitudinal lines. A few southern California 
specimens and those from southern Arizona are intermediate, having a pale gray 
ground and with the dark markings more well-defined than in most San Diego 
examples and more extensive than in the typical form, tending to form transverse 
spots. The single specimen from Wyoming is similar to the types, which are the 
largest specimens. Those from California are moderately large, ranging up to 
14.4 mm in forewing length; the short series from Arizona average smaller, ranging 
to 12.7 mm in the male and 13.4 mm in the female. Two males from Baja 
California reared from Agave orcuttiana flowers are the smallest (FW _ 11.1-11.7 
mm), possibly a function of the food (flowers vs. pods) or rearing conditions, 
but they do not differ structurally. 

Larva. The following diagnosis of the larval characteristics is based on a com- 
posite series of 20 last instar individuals from Cardiff, San Diego Co., California, 
collected in June 1963, and March and October 1967 (J. Powell nos. 63F13, 67C36 
and 67K80). 

Last instar larva: Length including head 13.0-23.8 mm (distended in KAAD 
preservative). Head: Width of head capsule (HC) 3.2-3.8 mm; deep amber to 


994 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


orange brown dorsally with faint darker mottling; adfrontal plates narrow and 
irregularly bordered exteriorly. Setal arrangements as in H. chalcofrontella (Clemens ) 
(MacKay, 1972). Submental pit well-defined, the submentum sclerotized laterally 
and posteriorly to the pit. Body: Thoracic shield and rings below SD; seta of 
Th3 to Ab7 dark amber, darker than the pinacula and anal shield, which vary 
in color independently, from yellowish to brown. Integument variably mottled with 
pale to dark purplish, forming on abdomen 3 broad, longitudinal bands, 2 in SD 
area and a weaker one in L enclosing spiracles; the intervening D and SD bands 
(latter enclosing SD, setae and their sclerotized crescents) usually unpigmented, 
but white in intensely colored individuals. Setal arrangements as in Fig. 1. 
Abdominal spiracles slightly oblong, tilted dorso-anteriorly, about 2 the size of 
most setal bases. Abdominal crotchets in a circle, 38-46 (usually 38-42), partially 
biordinal in an inconsistent pattern. Anal crotchets 27-35, partially biordinal to 
nearly triordinal. Anal fork absent. 

Larvae collected in October were smaller (HC: 3.2-3.35 mm; length 13.0-16.7 
mm) and were more deeply pigmented. Samples taken in March and June con- 
sisted of larger individuals (HC: 3.4-3.8 mm; length 16.0-23.8 mm) and con- 
tained no specimens with white longitudinal markings between the purplish bands. 

In larval characters H. gigantella does not differ appreciably from the much 
smaller H. chalcofrontella as characterized by MacKay (1972) except in pigmenta- 
tion and by the oval, relatively larger spiracles in H. gigantella. 

Material examined. Arizona: Bog Springs, Madera Cyn., Santa Rita Mts., 
VII-10/26-64 (adults in flowers and reared from fruit of Yucca schottii; D. R. 
Davis); VII-30-73 (1 @ at light; Coville, Szerlip and Powell); 2 mi. SW Portal, 
Cochise Co., VII-25-72 (1 2 in flowers of Y. schottii; Powell). Ca tirornta: 
short series, unknown locality in Los Angeles Co., VI-VII-1888 (reared from Y. 
brevifolia; Coquillett and Riley). Large series from various localities in association 
with Y. whipplei, as follows: Cardiff, San Diego Co. (reared from old and green 
fruit); 7 mi. E Morro Bay, S. L. O. Co., VI-23-65 (at light, Buckett); Mt. Lowe, 
VI-6-24 (Piazza) and 2 mi. NW Devil’s Punchbowl, L. A. Co., V—1-68 (at light; 
Opler, Powell); Forest Home (reared; Dammers) and Mill Creek, San Bernardino 
Co., (reared; Keifer); 3 mi. N Refugio Beach, Santa Barbara Co., VI-21/28-65 
(at light; Buckett and Powell). Conorapno: type series, cited above. Wyominc: 
6 mi. NW Newcastle, Weston Co., VII-13-65 (1 @ at light; Hodges). Mexico, 
Baya CairorniA Norte: 2 mi. E El Rosario, IJJ-26-73 (reared from flowers 
of Agave orcuttiana; Doyen and Powell); 5 mi. E El Rosario, III-18-72 (1 @ at 
light; Doyen and Powell). 

In addition there is a series of specimens from Hidalgo, Mexico, which differ 
by having the forewing nearly immaculate white (Ruins of Tula, 7500’, VII-31-63, 
reared from yucca pods; D. R. Davis). Despite the differing phenotype and widely 
disjunct distribution, I did not find structural features to distinguish this popula- 
tion from H. gigantella (two genitalia preparations of each sex examined). The 
forewing in most individuals is nearly entirely pale grayish white, with only the 
two dark dots at the end of the cell which are characteristic of the genus. One 
female is slightly darker grayish, showing faint longitudinal grayish lines. 

Two larvae from the intervening geographical region were examined. These 
were collected from fruit of Yucca filifera on the road to Salinas, 79 km NW of 
San Luis Potosi, 18 June 1963, by L. E. Caltagirone. They are distinguishable 
from H. gigantella as characterized above from California only in having the in- 
tegumental purplish pigmentation restricted to more well-defined SD and L bands. 


Holeocera paradoxa Powell, new species 


ae moderately large blastobasid having pale forewings with blackish longitudinal 
ee and two black dots at the end of the cell, and short antennal ciliation in 
the male. 


VoLuME 30, NuMBER 3 225 


Male. Length of forewing 9.7 mm. Head: Labial palpus elongate, strongly 
curved, exceeding crown; II segment length 1.4 times eye diameter; II] = 0.9 as 
long as II; smooth scaled, whitish with scattered dark scales. Antenna with scape 
elongate, subequal to eye diameter, broadened with elongate pecten on anterior 
margin, 2nd segment short, less than 0.25 length of Ist; 3rd claw-shaped, broadened 
basally with a deep notch on dorsal side distally; short ciliate, the setae less than 
0.5 diameter of 4th segment. Scaling of front and crown whitish tan. Thorax: 
Dorsal scaling whitish tan with scattered brownish. Venter whitish tan; legs 
speckled with dark gray exteriorly; hind tibia with enlarged dorsal fringe. Fore- 
wing: Narrow, length 4.4 times width. Scaling whitish tan, dark gray between 
the veins, forming moderately distinct longitudinal lines, especially on costal half 
and beyond cell; a whitish blotch at end of cell enclosing two dark dots at upper 
and lower corners of cell. Fringe pale grayish. Underside dark gray to margins; 
fringe pale. Hindwing: Slightly narrower than forewing at end of cell; vein M: 
separate, nearly adjacent to connate base of M; and Cu. Dorsal scaling shining 
whitish gray. Fringe broad, whitish ochreous. Underside gray, fringe as above. 
Abdomen: Scale coloration not recorded. Dorsal spine bands narrow, compact, 
well-defined (about 3-4 spines wide) on segments 2—7 (Fig. 4); basal segment 
without spines. Genitalia as in Fig. 6 (drawn from holotype, JAP prep. no. 
3874, one preparation examined); transverse connection of gnathos narrow without 
posterior notch; valva with costal process and free portion of sacculus relatively 
short, ca. 0.25 valva length; sclerotized median fold relatively elongate, 0.5 valva 
length; aedeagus with sclerotized band broad at base. 

Female. Length of forewing 10.6 mm. Essentially as described for male, dif- 
fering as follows: 2nd and 3rd segments of antenna not modified, antennal ciliation 
short. Forewing pattern more diffuse, the dark lines and spots less distinct. Hind- 
wing with veins M; + Cu, stalked, the stem about 0.25 the length of Cui; M2 
connate with M; + Cu: stem. Abdomen with dorsal spine rows on segments 2-7. 
Genitalia as in Fig. 8 (drawn from allotype, JAP prep. no. 3847, one preparation 
examined ); sterigma weakly sclerotized, scobinate; ductus bursae narrow through- 
out, rugose but not spiculate; signum a broad, concave plate, densely spurred 
interiorly. 

Holotype. Male, Arizona, Madera Canyon, Santa Rita Mountains, 6 June 1968, 
reared from remnants of 1967 pods Yucca schottii, emgd. 24 September 1968 (J. 
Powell no. 68F47). Allotype female, same locality, 3 October 1968, reared from 
1968 pods Y. schottii, emgd. 3 June 1969 (J. Powell no. 68K15). Holotype placed 
at the California Academy of Sciences on indefinite loan from the Essig Museum of 
Entomology, University of California, Berkeley, which retains the allotype. 

The peculiar wing venation in the hindwing of the male is outside the range 
of that shown in a long series of males of H. gigantella, which although somewhat 
variable, always have M; + Cu; at least short-stalked. The female venation in 
H. paradoxa is typical of Holcocera and Holcocerina as defined by McDunnough 
(1961), but the female genitalia in these two species possess a mixture of features 
of the two genera. The status of Holcocerina will have to be reassessed by com- 
prehensive study of North American species. The new species is most similar to 
H. gigantella among Nearctic blastobasids for which salient features have been 
described. H. paradoxa differs by its smaller size, more elongate labial palpus, 
and short antennal ciliation in the male, and compact spine row on abdominal 
segment 2. In genitalia, paradoxa is distinguished by its narrow gnathos band and 
relatively short free parts on the valva and by the distinctive features in female 
genitalia: sterigma form, ductus form and surface texture, and the spurred signum. 

Preserved larvae believed to represent H. paradoxa were examined from the type 
locality (Oct. 1968, Sept. 1969, Aug. 1970), Pena Blanca Lake area, Santa Cruz 
Co., Ariz. (Sept. 1969) and Paradise Road, Chiricahua Mts., Ariz. (Aug. 1971). 
All were collected as eggs or larvae from green pods of Yucca schottii. These larvae 


bo 
bo 
ep) 


JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 5-6. Holcocera, male genitalia, ventral aspect, valvae spread, aedeagus (a) 


removed and shown in lateral aspect: 
H. paradoxa Powell, holotype. 


5, H. gigantella (Chambers), lectotype; 6, 


VoLUME 30, NUMBER 3 227 


4 


= => 
Se 
Z ean 
oe 


7, H. gigantella (Cham- 


o*"S 


Figs. 7-8. Holcocera, female genitalia, ventral aspect: 
bers), paralectotype; 8, H. paradoxa Powell, allotype. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


bo 
bo 
CO 


do not differ structurally from those of H. gigantella, as characterized above from 
California. However, Arizona specimens differ by lacking most of or all of the 
integumental pigmentation. The color and extent of sclerotization of the prothoracic 
shield and kappa group pinaculum is similar in the two geographic samples, but 
the extensive purplish markings that form longitudinal bands in H. gigantella are 
lacking or represented by only faint traces in H. paradoxa. The dark sclerotized 
crescents (SD;) of the metathorax and abdominal segments 1-7 and the conspicu- 
ously brown pinacula in H. gigantella are unpigmented in H. paradoxa. The integu- 
mental pigment is darker in the penultimate and antepenultimate stadia in H. 
gigantella, These stages lack integumental markings in the Arizona populations, so 
instar differences between collections is not a factor in the two forms. 

After adults were reared from small samples of pod material in 1968, larger 
collections were made in September 1969, but all larvae died, probably due to 
overheating during field transit. The larval habits of H. paradoxa are briefly de- 
scribed elsewhere (Powell, 1976). 


ACKNOWLEDGMENTS 


Field work during this study was supported in part by National 
Science Foundation Grant GB-6813X. J. T. Doyen (University of Cali- 
fornia, Berkeley), P. A. Opler (Office of Endangered Species, Wash- 
ington, D.C.), and R. A. Mackie (Public Health Service, Guam) 
assisted with field collections. The illustrations were executed by 
Celeste Green. 

Cooperation by J. F. Lawrence, Museum of Comparative Zoology, 
Harvard and R. W. Hodges, National Museum of Natural History, 
Washington, D.C. enabled study of specimens in those institutions. 


LITERATURE CITED 


Basincer, A. J. 1924. A supposedly beneficial insect discovered to be a citrus 
joese, |i. Jaeom, laine, ive C7 ae9), 

Buscx, A. & M. L. Otrviera Firuo. 1925. Da Auximobasis coffeaella Busck. 
Mariposa dos fructos de cafe abandonados sua determina cao e biologia. Sec. 
Agric., Commercio Obras Publicas, S. Paulo. 19 p. 

CHAmBers, V. I. 1876. Winema. ‘Can.’ Ent, 8; 2117-220) 

Common, I. F. B. 1970. Lepidoptera. In, Insects of Australia. Melbourne U. 
Press. xiii + 1029 p. 

Costa Lima, A. 1945. Insetos do Brasil. 5th Tomo. Lepidopteros, Ist parte. 
Esc. Nac. Agron., Serie Didatica, 7. 379 p. 

CraAIGHEAD, F. C. 1950. Insect enemies of eastern forests. U.S.D.A., Misc. Publ. 
6p: (67 9"p, 

Dietz, W. G. 1900. On Pigritia Clem. Trans. Amer. Ent. Soc. 27: 100-120. 

. 1910. Revision of the Blastobasidae of North America. Trans. Amer. 
Emt. Soc, i667. 1-72; 

Kssic, E. O. 1916. A coccid-feeding moth. Holcocera iceryaeella (Riley). J. 
Econ. Ent., 9: 369-370. 

MLercner, T. B. 1920. Life histories of Indian insects. Microlepidoptera. Mem. 
Dept. Agric. India, Ent. Ser. 6. 217 p. 

——. 1933. Life histories of Indian Microlepidoptera (Second Series). Cos- 
mopterygidae to Neopseustidae. Imperial Counc. Agric. Res., Sci. Monogr. 4. 
§5 p. 


VoLuME 30, NUMBER 3 229 


Forses, W. T. M. 1923. Lepidoptera of New York and neighboring states. 
@omell U. Agric. Exp. Sta. Mem. 68. 729 p. 

1931. Supplementary report on the Heterocera or moths of Puerto Rico. 
J. Dept. Agric. Puerto Rico. 4: 339-394. 

Keen, F. P. 1958. Cone and seed insects of western forest trees. U.S.D.A., Tech. 
Bull. 1169. 168 p. 

Lyons, L. A. 1957. Insects affecting seed production in red pine III. Can. Ent. 
89: 150-164. 

MacKay, M. R. 1972. Larval sketches of some Microlepidoptera, chiefly North 
American. Mem. Ent. Soc. Can. 88. 83 p. 

McDunnoucu, J. 1961. A study of the Blastobasinae of Nova Scotia, with par- 
ticular reference to genitalia characters (Microlepidoptera, Blastobasidae ). 
Amer. Mus. Novit., No. 2045. 20 p. 

PoweE.L.L, J. A. 1976. Biological interrelationships of moths and Yucca _ schottii 
(Lepidoptera: Incurvariidae, Cochylidae, Blastobasidae). U. Calif. Publ. 
Entomol., in press. 

PowE.., J. A. & R. A. Mackie. 1966. Biological interrelationships of moths and 
Yucca whippiei (Lepidoptera: Gelechiidae, Blastobasidae, Prodoxidae). U. 
Calif. Publ. Entomol., 42: 1-46. 

Ritey, C. V. 1887. Report of the Commission of Agriculture, 1886: 485 (not 


seen ). 


NOTES AND NEWS 


Recent Letter 
Dear Dr. Godfrey, 

With reference to Mr. Manley’s note on “The ‘greasy’ wing gene of Utetheisa 
ornatrix ( Arctiidae)” (1975, J. Lepid. Soc. 29: 77), I do not think that any special 
significance should be attached to the 1:1 ratio in females ex various collections, 
except that it shows that the aberration is confined to the female, but whether 
sex-linked or sex-controlled can only be ascertained by breeding. It is most unusual 
for the proportion of an aberration to the type in museum collections to correspond 
with the proportion in nature; there is an inevitable bias in favour of the aberration. 
Also no inference as regards dominance can be drawn from such figures, as many 
morphs, genetically dominant, are far rarer than the recessive allelomorph in 
nature. A good example is f. salaami Suff. of Papilio dardanus Brown, which is far 
rarer than f. hippocoon F., to which it is dominant. 

Dr. Sargent’s experience with Papaipema duovata (1975, J. Lepid. Soc. 29: 9) 
appears to confirm the comment made by J. W. Tutt, the famous English en- 
tomologist, early in the century “that no species is rare if you know where to 
look for it.” 

An entomologist, working with light, both mercury vapour and incandescent, 
would be quite justified in concluding that the sphingid Nephele peneus Cr. did 
not occur in Mombasa. Its congeners argentifera Wlk., bipartita Btlr., funebris F. 
and comma Hpffr., all visit light freely, whilst aequivalens WI1k., oenopion Hbn. 
and rosae Btlr. occur more rarely, but during 20 years collecting I have known 
a single peneus to visit light. Yet it is quite common and an examination of its 
foodplant in the proper season will always provide large numbers of ova and larvae 
at all stages of growth. 

D. G. SEVASTOPULO 


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GENERAL NOTES 


A SURVEY OF THE SPHINGIDAE OF SANIBEL ISLAND, FLORIDA 


Sanibel Island, which lies ca. 3 mi. off the coast of Florida at Fort Myers, is 
one of a series of islands that form a chain reaching from north of Charlotte 
Harbor to slightly south of the mouth of the Caloosahatchee River. The island, 
which is roughly 12 mi. long by 3 mi. wide at its widest point, is currently 
suffering considerable habitat destruction from commercial development. However, 
large areas of relatively unspoiled land still remain. A survey was conducted to 
obtain data on the Sphingidae of Sanibel Island, including the relative abundance 
and feeding behavior of each species. The recent rapid commercial development 
of much of the island makes these data, obtained before this latest and largest 
wave of habitat destruction, especially important. 


METHODS 


Two methods of data collection were utilized in making this survey. The pri- 
mary method involved observing and collecting specimens as they fed at flowers 
(mainly sea periwinkle, Vinca rosea), and collecting at lights was used as a sec- 
ondary method. 

Three major sites were used in the primary method. Two of these sites were 
large open areas, largely covered with V. rosea. Both of these sites were within 
300 yd. of the Gulf of Mexico but were surrounded by large trees so that ocean 
breezes, which would influence feeding behavior, were negligible. The third site 
was farther inland. This site was partially covered with V. rosea, but numerous 
bushes and small trees were scattered over it as well. The sites were generally 
checked at dusk and dawn (many Sphingidae are crepuscular) and were often 
checked continuously for several hours beginning at dusk. Most species were easily 
disturbed by artificial light. Therefore, the author collected in the dark, using 
movement of the flowers to locate specimens. 

Specimens collected at lights were mainly obtained by collecting in the parking 
lot of a shopping center located roughly 1 mi. inland from the Gulf of Mexico. 
This parking lot was brilliantly illuminated by a series of mercury vapor lights 
on tall posts. The author also obtained specimens from lights located at many 
other points on the island, these lights being patrolled periodically by car. 

This survey was conducted by the author during the entire months of August 
of 1961 through 1966; June 20 through August 28, 1967; the latter halves of 
August of 1968 and 1969; the first half of August, 1971; and one week each in 
December, 1960 and December, 1966. Most of the specimens collected during 
this survey are in the author's private collection. 


RESULTS AND Discussion 


One of the most interesting results of this survey concerns feeding behavior of 
the different species. Each species has a characteristic pattern of flight while 
feeding. Some species (e.g., Madoryx pseudothyreus (Grote) ) stay under leaves 
and between stems whenever possible, whereas other species (e.g., Pachylia ficus 
(Linnaeus) ) avoid such situations and remain in the open, and still others (e.g., 
Manduca brontes (Drury) ) seemingly have no preference in this regard. Further- 
more, some species skip from one group of flowers to another as a normal part of 
feeding (e.g., Enyo lugubris (Linnaeus) ), whereas other species feed on nearly all 
the flowers in one area before moving to another (e.g., Erinnyis obscura (Fabricius ) ). 
Another interesting behavioral difference involves movement of flowers while feed- 
ing. ach species moves a flower in a characteristic manner as it hovers over the 
flower to feed, interspecific differences in this regard probably being due to dif- 


VoLUME 30, NUMBER 3 Did 


ferences in proboscis length and body size. The differences in flight patterns and 
flower movements are consistent enough to allow generally accurate specific identi- 
fication based on these factors alone, although considerable practice is needed to 
master this art. 

A list of species collected and observed by the author during this survey follows. 
Names used are those found in Hodges (1971, Sphingoidea. In R. B. Dominick, 
et al., The moths of America north of Mexico, fasc. 21). Times are Eastern Standard 
Time. Comments on such things as behavior and abundance are included for each 
species. The author believes that this list is essentially a complete one for the 
month of August. However, it is, of course, quite possible that other species are 
present at different times of year. 

1. Agrius cingulatus (Fabricius) is common both at lights and at feeding sites, 
especially during the latter part of August. In general, this species feeds only well 
after dark. 

2. Cocytius antaeus (Drury) is rare during August. The author collected only 
three specimens during the survey, all at lights. Two additional specimens were 
observed flying just before dusk. Large quantities of bright yellow pollen were found 
in the proboscises of two of the collected specimens, but the author never observed 
this species feeding. 

3. Manduca sexta (Linnaeus) is very common at lights and common at feeding 
sites. 

4, Manduca quinquemaculata (Haworth) is abundant at lights and common at 
feeding sites. It is not unusual to have 10 or 12 specimens in view simultaneously 
at lights. 

5. Manduca rustica (Fabricius) is common at lights and moderately common at 
feeding sites. It generally feeds well after dark, in August often between 2200 and 
2300 hr. 

6. Manduca brontes (Drury) is common at feeding sites and somewhat less com- 
mon at lights. It generally begins feeding shortly after dusk. 

7. Pseudosphinx tetrio (Linnaeus) is rare. The author collected only one speci- 
men during the survey. This specimen was feeding on V. rosea at ca. 2230 hr. 

8. Erinnyis alope (Drury) is uncommon but is found at feeding sites and at lights 
with about equal frequency. Some individuals are extremely difficult to approach 
while feeding, whereas others at the same sites and under apparently similar condi- 
tions are relatively easy to approach. 

9. Erinnyis ello (Linnaeus) is common at lights and very common to abundant 
at feeding sites. It usually starts to feed just as true darkness begins. 

10. Erinnyis obscura (Fabricius) is one of the three most abundant sphingid 
species on Sanibel Island. It is abundant at lights and incredibly abundant at feed- 
ing sites. The author once had 29 specimens of this species in view simultaneously, 
and groups of 12 to 15 specimens are common. Erinnyis obscura begins feeding 
well before dark, usually as the sun sets, and is one of the first species to begin 
feeding each night. 

11. Phryxus caicus (Cramer) is variable in August, sometimes locally common 
and at other times quite uncommon. This species is unusual in that it barely moves 
the flower as it feeds. 

12. Pachylia ficus (Linnaeus) is uncommon to rare. It begins feeding before 
sunset and often will fly erratically for distances of several hundred feet, ending the 
flight at a flower a foot or so from the one it fed on just prior to the flight. The 
species has two modes of flight. When flying at normal speed the abdomen extends 
out behind the thorax as it normally would, but, when slowing down or flying slowly, 
the abdomen is bent toward the ground and the stroke of the wings changes, the 
hindwings being used to reduce speed by adding drag. This change in manner of 
flight is extremely obvious during the erratic “feeding flights” described above. 

13. Madoryx pseudothyreus (Grote) is generally uncommon. When feeding, it 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


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hovers under leaves or between stems whenever possible and, because of this habit, 
may easily be overlooked by the collector. : 

14. Aellopos tantalus (Linnaeus) is uncommon and diurnal, feeding on several 
species of flowers. 

15. Enyo lugubris (Linnaeus) is one of the three most common sphingid species 
on the island. It begins feeding well before dusk and is not uncommonly found 
feeding during the day also. As this species flies, it makes a loud whirring noise 
that sounds similar to the noise produced by a diving nighthawk, Chordeiles minor 
minor ( Forster ). 

16. Hemaris thysbe (Fabricius) is diurnal and uncommon. Since its feeding habits 
are well-known, they will not be treated here. 

17. Hemaris diffinis (Boisduval) is diurnal and reasonably common. Its well- 
known feeding habits will not be treated here. 

18. Eumorpha achemon (Drury) is rare. The author collected only two specimens 
during the survey, one feeding on V. rosea and one at lights. 

19. Eumorpha vitis (Linnaeus) is fairly uncommon. Specimens feeding on V. 
rosea tend to remain close to the ground and may remain within a small area for 
several minutes at a time. Feeding begins after nightfall. 

20. Eumorpha fasciata (Sulzer) is variable, although generally somewhat uncom- 
mon. It is more commonly found at feeding sites than at lights and begins feeding 
at dark. 

21. Cautethia grotei Henry Edwards is not too uncommon. It begins feeding at 
dusk and feeds intermittently, well into the night. 

22. Xylophanes tersa (Linnaeus) is one of the three most common sphingids on 
Sanibel. It is abundant at both lights and feeding sites. Feeding begins just as the 
sun sets. ; 

23. Hyles lineata (Fabricius) is variable, although, in general, it is moderately 
common at feeding sites and common at lights. It is often found feeding diurnally, 
although nocturnal and crepuscular feeding are more common. 


Hints on Collecting at Feeding Sites 


Collection of feeding sphingids at night requires techniques quite different from 
those generally employed to collect other Lepidoptera. Therefore, it may be of 
interest to briefly review the basic techniques involved. 

Equipment. Dark clothes are helpful, since light colored moving objects tend to 
disturb feeding sphingids. The author uses a lightweight net with a 4-ft. handle. 
Light colored netting should not be used. A series of killing bottles should be tied 
to a belt in such a way that a bottle can be opened rapidly with one hand. Large 
corks used in place of screw tops are helpful in this regard. If artificial light is used, 
it should be extremely weak, since most lights disturb feeding behavior in many 
species. Unfortunately, most insect repellents repel sphingids; therefore, mosquito 
repellents can only be used sparingly. If mosquitoes are bad, a headnet offers 
some protection but reduces vision. 

Techniques. The key to successful collecting is to move slowly, stalk, and use 
a short, rapid stroke to capture specimens. Squatting close to the ground consider- 
ably improves the collector's chances of seeing both moving flowers and hovering 
moths. Identify all moths visible before stalking a specimen, and decide which speci- 
men is most desired. Take care not to disturb other moths because one alarmed 
moth will often dart about and alarm the others. Hold the net low but keep it 
ready for use. Be sure to hold the net bag so that it also does not alarm specimens. 
The author holds the netting against the net handle with the first finger of the right 
hand and does not release it until beginning the capture stroke, which should be 
short and swift (2 ft. or less) for best results. If killing bottles are worn on the 
left side, specimens can be transferred to them rapidly from the net. This transfer 
can largely be done by touch, so that the collector can be bottling one specimen as 


VoLuME 30, NUMBER 3 233 


he plans his next stalk. Knowledge of the behavior of each species is important in 
planning captures; some species will come to you if you position yourself properly 
in the feeding site. If a moth changes its flight pattern and becomes restless, ex- 
perience is your best guide. Some species will return after such behavior; others 
will not. 

Times to Collect. Each species feeds only at certain times during the night. 
On Sanibel, one group of species feeds at dusk, a second shortly after dark, and a 
third group ca. 2% hr after dark. Practically no species feed between 0100 and 
0330 hrs on Sanibel. The dawn feeding schedule is basically the reverse of the dusk 
schedule, except that some species feed at dusk only. 

Selection of a Collecting Site. Obviously, local conditions greatly influence choice 
of sites. If possible, the site should have large numbers of flowers accessible to the 
collector. Some species prefer large open areas, and others prefer sites with tall 
vegetation. One good way to increase the number of species collected is to use 
several different types of sites and to check each site at various times and under 
different weather conditions. 


C. Hucu Brown, Department of Biology, Valdosta State College, Valdosta, Georgia 
31601. 


FIRST RECORDS OF BOLORIA FRIGGA (NYMPHALIDAE) IN WISCONSIN 


During the last week of May 1975, Boloria frigga (Thunberg) (Fig. 1) was dis- 
covered in a number of northern Wisconsin localities. On 24 May two fresh males 
were collected at the edge of a wet, open bog in Conover Township, Vilas Co., by 
George Balogh of Milwaukee. Other specimens were seen farther out in the bog 
but were not collected. On 25 May we (LAF, RMK) discovered a colony of B. 
frigga in a bog located in Upham Township, Langlade Co. Thirteen freshly emerged 
males and two females were collected. The next weekend, one of us (LAF) in- 
vestigated other northern Wisconsin bogs in search of this species. Another colony 
was found in Lincoln Township, Vilas Co., on 31 May. Four males and one female 
were taken, and several other specimens were seen. Some of the females observed 


Figs. 1-2. 1, Boloria frigga (Thunberg) males, Vilas Co., Wisconsin, 31 May 
1975 (above), and Langlade Co., Wisconsin, 25 May 1975 (below); 2, bog 
habitat of B. frigga, Vilas Co., Wisconsin. 


934 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


were somewhat worn. Also on 31 May, two worn males and a fresh female were 
collected in Bradley Township, Lincoln Co. These specimens, taken by Fay 
Karpuleon of Eau Claire, were found in the open portion of a large acid bog. These 
records indicate that B. frigga probably exists throughout the northern quarter of 
Wisconsin wherever there are suitable bogs. 

These dates of collection appear to be more advanced than normal, probably 
due to unseasonably high temperatures during the third week of May. An unusually 
early Oeneis jutta (Hiibner) was also taken on 25 May at the Langlade Co. locality. 
The earliest date that O. jutta was collected in neighboring Marathon Co. was 31 
May. 

Bolo frigga appears to be restricted to open, sedgy sphagnum bogs (Fig. 2). 
The bogs at all localities tended to be quite wet. At the localities visited by the 
authors, the bogs supported scattered Larix laricina (Tamarack), clumps of Betula 
pumila (Dwarf Birch) and Salix pedicellaris (Bog Willow). Also present were 
Kalmia_ polifolia (Swamp Laurel), Andromeda glaucophylla (Bog Rosemary), 
Chamaedaphne caliculata (Leather Leaf) and an occasional Sarrcenia purpurea 
( Pitcher-plant ). 

The specimens of Boloria frigga are presently retained by their respective collectors. 


Lesutrz A. Ferce, Rt. 5, Town Line Road, Wausau, Wisconsin 54401. 
Rocer M. Kueun, 5042 N. 61st Street, Milwaukee, Wisconsin 53218. 


ADDITIONAL NEW BUTTERFLY RECORDS FROM FLORIDA 


Florida continues to be a source of new records of Lepidoptera in spite of 
burgeoning land development and destruction of the natural habitat. However, 
some species may benefit from this as certain ornamental plants become more 
widespread. We report herein one new U.S.A. record (Pieridae), one new Florida 
record (Lycaenidae), and additional records of some recently reported species. 


Pieridae 


Phoebis orbis (Poey). A single fresh male of this species (previously unrecorded 
from the U.S.A.) was taken on 25 April 1973 at about 1200 EST, on Big Pine 
Key, Monroe Co., Fla. It was captured visiting flowers with other Phoebis species, 
on Sands Rd. across from a pine forest. No others were seen in the area. 

P. orbis is illustrated by Lewis (1973, Butterflies of the World. Chicago. 312 p.). 
However, the specimen taken has a milky white ground color rather than the yellow 
shown by Lewis, and the basal patch on the upper forewing is orange rather than 
dark yellow. 

P. orbis has previously been reported from Cuba and Hispaniola (Scott 1971, J. 
Res. Lepid. 9: 249-256), and the Florida specimen reported in the present paper 
is evidently a stray. 

Anteos maerula maerula (Fabricius). Five specimens were taken on 31 August 
1973 on Big Pine Key, Monroe Co., Fla. These were caught during intermittent 
rain showers while visiting flowers along the road. 


Nymphalidae 


Anartia lytrea chrysopelea (Hiibner). Six fresh specimens of this recently re- 
ported species (Anderson 1974, J. Lepid. Soc., 28: 354-359) were taken on 24 
and 25 April 1973, on Big Pine Key, Monroe Co., Fla. Two specimens were 


7 


deposited in the Carnegie Museum and two in the M. Howard Collection. 


VoLuME 30, NUMBER 3 25D 


Lycaenidae 


Tmolus azia (Hewitson). The first specimen of Tmolus azia taken in Florida 
was a worn female on 28 June 1974 in Fairchild Gardens, Dade Co. This butterfly 
was found on an ornamental acacia. A second specimen, also female, was taken 
on 27 April 1975 at the same location on Montezuma speciosissima Moc. & Sesse 
( Malvaceae ). 

A third record of T. azia was contributed by Mr. Charles Covell, who captured 
a single fresh specimen on 12 May 1975 near Homestead, Dade Co., Fla. 

T. azia is found in South and Central America extending into the extreme 
southern portions of Texas and Arizona (Erlich & Erlich 1961, How to Know the 
Butterflies. Dubuque, Iowa. 200 p.), but there are no records of this species 
from the Antilles (Scott 1971, J. Res. Lepid. 9: 249-256). T. azia may, there- 
fore, represent a recent introduction to Florida, perhaps associated with exotic 
ornamental plants. 

Electrostrymon angelia (Hewitson). This species, also reported by Anderson 
(1974, J. Lepid. Soc., 28: 354-359), is taken commonly at Fairchild Gardens, 
Dade Co., Fla., where it is associated with Derris sp. (Leguminosae), a group of 
trees and shrubs native to India. However, immatures have not been found on 
this plant. E. angelia was also taken on 8 May 1975, in a sawgrass marsh located 
near the junction of U.S. Highway 41 and State Road 27 in Dade Co., Fla. 


ACKNOWLEDGMENTS 


We wish to thank Dr. Lee Miller of the Allyn Museum of Entomology for help 
with determination of Tmolus azia, Phoebis orbis, and Anartia lytrea chrysopelea 
and Dr. Harry Clench of the Carnegie Museum of Natural History for help with 
determination of Anartia lytrea chrysopelea. 


Rospert BENNETT, 1228 Miles Avenue, Kalamazoo, Michigan 49001. 
Epwarp C. Knupson, 7211 Cecil Street, Apt. 3, Houston, Texas 77025. 


NOCTURNAL ACTIVITY OF A MONARCH BUTTERFLY (DANAIDAE) 


Rhopalocera are generally diurnal in habits, even becoming inactive during total 
solar eclipses (Moucha 1964, J. Lepid. Soc. 18: 109-110). Scattered reports of 
Rhopalocera at artificial light, including light traps, have indicated some nocturnal 
activity of these insects (Kendall & Glick 1972, J. Res. Lepid. 10: 273-283 and 
references therein). Some reports have included pairs in copula (Heitzman 1969, 
J. Lepid. Soc. 23: 105-106; Chambers 1962, J. Lepid. Soc. 16: 200). However, 
activity directed toward artificial light may not indicate normal nocturnal activity. 
Some species collected at artificial light are those normally active at dusk or low 
light level habitats [Opsiphanes by Welling (1963, J. Lepid. Soc. 17: 37-38) and 
Melanitis by Donahue (1962, J. Lepid. Soc. 16: 131-135)]. Diurnal species 
collected at light traps may well include only specimens that have somehow be- 
come disturbed (Kendall & Glick, op. cit.). 

On 29 October 1971 at the Brackenridge Field Laboratory of the University of 
Texas at Austin, I observed an adult male monarch, Danaus plexippus plexippus 
L., feeding at inflorescences of shrubby boneset, Eupatorium havanense H.B.K. 
(Compositae), at 2130 CDT. The time of observation was 2 hr 44 min after 
sunset. Moonlight was apparent but not bright (between first quarter and full), 
with a clear sky (0% cloud cover). The temperature at recording stations 50 
and 125 m away was 21.1° C (70° F) with 84% RH. No artificial light was 
present at the site. The white coloration of the inflorescences is significant, since 
white blossoms are more visible at night than flowers of other colors. Flowers 


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pollinated by nocturnal insects are generally white (Faegri & van der Pijl 1971, 
The Principles of Pollination Ecology, 2nd ed., Pergamon Press). Monarchs have 
been reported previously at artificial light in Texas, Missouri and Mexico (Kendall 
& Glick, op. cit.; Heitzman 1965, J. Lepid. Soc. 19: 179-180). Lack of previous 
reports of nocturnal activity for monarchs in a natural setting indicates that such 
activity is not normal (Urquhart 1960, The Monarch Butterfly, U. Toronto Press). 

Several environmental factors may have resulted in the behavior cited in the 
present note. Drought conditions from late 1970 to mid-1971, followed by heavy 
rains in early August 1971 resulted in massive numbers of butterflies in late 
August and September. Although plant growth, including blossom production, was 
greatly enhanced, little rain occurred from mid-August through September. Flower 
production was retarded. Feeding pressure from local butterflies as well as the 
immigrant monarchs resulted in a nectar shortage. The daylight hours of 29 
October 1971 were overcast with fog persisting until late morning and cloudiness 
(80%) as late as mid-afternoon. Only 32% (3.6 hr) of possible sunshine was 
recorded that day. Conditions for nectar foraging were definitely inferior. As a 
result, at least one individual fed at flowers at night. The time (pre- or post- 
sunset) of arrival of the butterfly at the inflorescence is unknown. 


RayMonp W. Neck, Texas Parks and Wildlife Department, John H. Reagan 
Building, Austin, Texas 78701. 


CRAB SPIDER PREYS ON NEOPHASIA MENAPIA (PIERIDAE ) 


On 22 August 1974, an immature crab spider, Misumenops sp. (Araneae: Thomi- 
sidae), was observed feeding on an adult male pine butterfly, Neophasia menapia 
(Felder & Felder) (Pieridae). The spider and butterfly prey were on a flower of 
goldenrod, Solidago rigida L., in ponderosa pine forest at Sheridan Lake (TIS, 
R5E, sec. 13), el. 1,400 m, Black Hills National Forest, Pennington Co., South 
Dakota. Examination of other goldenrods failed to yield additional spiders with 
butterfly prey, although another N. menapia cadaver was found. 

Evenden (1926, J. Agr. Res. 33: 339-344) described the life history of N. 
menapia. Additional information on habits, parasites, and predators of this pierid 
are given by Orr (1954, USDA, For. Seryv., Intermountain For. and Range Exp. 
Stn., Misc. Publ. No. 1, 12 p.), Cole (1956, USDA, For. Serv., Intermountain 
For. and Range Exp. Stn., Res. Note No. 29, 8 p.; 1971, USDA, For. Serv., For. 
Pest Leafl. 66, 3 p.), and Bousfield & Dewey (1972, USDA, For. Serv., Northern 
Region Insect and Disease Rept. No. I-72-12, 9 p.). Natural enemies include 
various hymenopterous and dipterous parasitoids that attack the larval and pupal 
stages, and pentatomids and snakeflies which prey on the eggs. We found no 
previous records of spiders preying on any of the life stages of the pine butterfly. 

The spider feeding on N. menapia was captured alive, but an attempt to rear 
it to maturity failed. Since species determinations of spiders are based chiefly on 
characters of the genitalia, which are not fully developed until maturity, the specific 
identity of the spider is unknown although we suspect that it is M. asperatus 
(Hentz), a common inhabitant of goldenrod. A related misumenid crab spider, 
Misumenoides formosipes (Walckenaer), was also found on goldenrod, but without 
prey. This latter species is readily distinguished from Misumenops by the presence 
of a white clypeal carina. Records of South Dakota crab spiders are given by 
Buckman (1966, Proc. S. D. Acad. Sci. 45: 118-123) and include both Misumenops 
asperatus and Misumenoides aleatorius (Hentz) (= formosipes (Walckenaer) ). 

_ Crab spiders of the subfamily Misumeninae are ambushers and are commonly 
found on flowering plants, such as goldenrod, where they lie in wait for visiting 
insects. Some species blend with the background plant color and are capable of 


VoLuME 30, NUMBER 3 ZL 


changing from white to yellow or vice versa. Gertsch (1939, Bull. Amer. Mus. 
Nat. Hist. 76: 277-442) reports that the misumenids have a powerful venom 
and are capable of quickly subduing insects, including bumblebees, moths, and 
butterflies, that are much larger than the spiders. 

An interesting spider-plant-butterfly relationship is indicated by this collection. 
Gertsch (1939, Bull. Amer. Mus. Nat. Hist. 76: 277-442) maintains that the 
habitat of a spider determines the kind of prey that becomes available to it. 
Flower-inhabiting spiders feed on insects attracted to flowers for nectar, pollen, 
or other food sources. Although larvae of N. menapia are destructive defoliators 
of pine, Orr (1954, USDA, For. Serv., Intermountain For. and Range Exp. Stn., 
Misc. Publ. No. 1, 12 p.) reports that the adults feed only on flowers. This habit 
renders them vulnerable to predation by flower-inhabiting predators, such as crab 
spiders. 

Spider and butterfly are deposited in the collection of the American Museum 
of Natural History, New York. 


DanrEL T. JENNINGS, U.S. Department of Agriculture, Forest Service, Rocky 
Mountain Forest and Range Experiment Station, 240 W. Prospect, Fort Collins, 
Colorado 80521. 

MicuaeL E. Touiver, Department of Entomology, Morrill Hall, University of 
Illinois, Urbana, Illinois 61801. 


BUTTERFLIES ASSOCIATED WITH AN ARMY ANT SWARM RAID 
IN HONDURAS 


The swarm raids of the army ant Eciton burchelli (Westwood) (Formicidae: 
Dorylinae) are a striking feature of tropical forests throughout Central and South 
America. Associated with these raiding swarms of Eciton are various animals that 
exploit the swarm raid for the purposes of feeding or reproduction. For example, 
ant birds (Formicariidae) forage at the leading edge of such swarms and feed 
on insects flushed from the vegetation by the army ant juggernaut. Flying above 
the swarm raid are various species of tachinid flies (Diptera: Tachinidae) and 
other insects, e.g., staphylinid beetles (Coleoptera: Staphylinidae), the life cycles 
of which may be regularly intermeshed with those of the ants (Akre & Ret- 
tenmeyer 1966, J. Kansas Entomol. Soc. 39: 745-782). Some of these flies, 
for instance, are known to hover above the swarm, then dart down and quickly 
lay an egg on a prey item being carried back to the army ant bivouac. The egg 
then develops among the doryline brood (Schneirla 1971, Army Ants, Freeman & 
Co., San Francisco ). 

During the early afternoon of 24 May 1972, I observed a large swarm raid of 
E. burchelli in tropical broadleaf forest located on the west shore of Lago Yojoa, 
Santa Barbara Province, Honduras. Flying low over the leading edge of the swarm 
were six butterflies: two male Graphium philolaus (Boisduval) (Papilionidae: 
Papilioninae), two female Mechanitis isthmia isthmia Bates, and two female 
Mechanitis polymnia doryssus Bates (Nymphalidae: Ithomiinae). These three 
species were the most common of the several butterfly species in the area, but 
only the six individuals listed above were flying in the vicinity of the swarm 
during the observation period. Flying in general ca. 2 ft. above the ground and 
occasionally dipping down to ground level (but without alighting), the butterflies 
stayed above the leading edge of the swarm as it moved steadily southward some 
20 ft. during the 2 hr that I was able to watch it. The behavior of all three 
species during this time was similar, although G. philolaus had a more soaring 
and wide-ranging flight than the two ithomiines and, as a result, seemed to be 
tracking the movements of the ant swarm less closely. Ant birds foraged at the 


238 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


head of the swarm, but none of the three or four birds present were ever seen 
to take or attempt to take any of the six butterflies. 

Eciton burchelli has a distinctive odor that can be recognized by a sensitive 
human nose as an army ant odor (Carl W. Rettenmeyer, pers. comm.) and has 
been described in the old literature (Rettenmeyer, op. cit.) as similar to the odor 
of human feces. Although Dr. Rettenmeyer thinks this description incorrect, the 
odor is at least unpleasant. This distinctive odor is probably enhanced by the 
large swarm size and may thereby attract certain animals (Rettenmeyer 1961, 
Univ. Kansas Sci. Bull. 42: 993-1066). Indeed, the tachinid flies and other insects 
mentioned above may even respond to the odor as an olfactory signal that initiates 
oviposition behavior. Perhaps the odor contains elements similar to those of the 
androconial tufts of Mechanitis males, which might explain why only female 
Mechanitis were following the swarm. Longstaff (1912, Butterfly-hunting in 
Many Lands, Longmans & Co., London) recorded that some clearwing ithomiines 
in Venezuela had scents of a “disagreeable character, recalling stables or pig-sties,” 
that he believed to be associated with the hindwing androconial brushes (found 
only in males). However, this does not explain why the male Graphium followed 
the swarm, since in Graphium, like the ithomiines, it is the males that have the 
scent scales. Possibly the Graphium males were attracted by a component of the 
ant odor that elicited food searching behavior. 

During 14 months of field research in the tropical rain forests of Eastern 
Ecuador in 1973-1974, I observed dozens of swarm raids by several colonies of 
E. burchelli in areas rich in ithomiines (but not in Graphium) yet never saw any 
butterflies that seemed in any way attracted to or associated with the army ant 
swarms. Dr. Rettenmeyer informs me that in his many years of field work on 
army ants he has never observed any butterflies which appeared to be associated 
with swarm raids. Thus, it appears that my observation may be unique and, 
therefore, interesting only as a curiosity, at least until such time as the chemical 
components of the pheromones of E. burchelli are better known. 

Specimens of the Honduran army ant population are in the collection of C. W. 
Rettenmeyer. All six of the observed butterflies are in the personal collection of 
the author. 


ACKNOWLEDGMENTS 


These observations were made while I was a student in the Organization for 
Tropical Studies course 72-2. I thank: C. W. Rettenmeyer for identifying the 
army ant and for discussion and T. C. Emmel for reviewing the manuscript. 


Boyce A. DrumMmonp III, Department of Zoology, University of Florida, Gaines- 
ville, Florida 32611. 


VoLuME 30, NUMBER 3 239 


OBITUARY 


ROBERT GRANT WIND (1912-1975) 


Fig. 1. Robert and Clo Wind. Photograph taken in 1936. 


Robert Grant Wind was born in San Francisco, California, 17 June 1912, the 
first son of Walter W. Wind and Helen Ables Wind. His father was owner of 
the Berkeley Plumbing Company, which has been continued as a family business 
by his brother Howard. Bob, as he was known to his associates, was raised in 
the community of Berkeley. His family occupied a large house on Santa Clara 
Street. The basement, attic, and Bob’s bedroom provided the storage space needed 
for his large butterfly and insect collection with its many glass-topped drawers, 
used Simmons mattress boxes (providing storage for row on row of filed butterflies 
in envelopes), and 5 gallon metal storage containers (probably containing exotic 
specimens ). At times his bedroom was filled with recently emerged saturniid moths. 

Following his graduation from Berkeley High School, he attended the University 
of California, Berkeley from 1930-1933. In the summer of 1933 he was a Nature 
Counselor at the Boy Scouts of America Camp Wolfeboro in the Sierra Nevada of 
Stanislaus County. It was there that Davies and his lifelong colleague William 
A. Hammer first met Bob. He gave them information on the desirable butterflies 
of the area and provided their first cyanide bottles for collecting. 

In 1933 Bob began the business—Pacific Coast Biological Company—for sales 
of all types of biological supplies. He also issued extensive lists of world butterflies, 
moths, and beetles that were offered for sale. He had the use of a large red 
touring car, and this provided transportation for his entomological friends to field 
collect or to attend meetings of the Pacific Coast Entomological Society in San 
Francisco. These included R. M. and G. E. Bohart, R. L. Usinger, J. W. MacSwain, 


240 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Art and Edgar Smith, and others. Some also served as his agents collecting 
specimens to fill orders, or handling orders for his company (Art and Edgar Smith 
serving the San Joaquin Valley area). 

In 1937 Bob married Clo Mifflin of Piedmont. In the following year he sold 
his business in order to finance and organize the “Papuan-Australian Expedition.” 
It had been his lifetime ambition to explore these areas for their entomological 
treasures, and, so, in 1938 Bob and Clo left for Sydney, at that time a four week 
sea voyage. From Sydney they travelled by narrow-gauge railway to Redlynch 
in North Queensland, in the Cairns area, renting a house from August-December 
1938. This permitted the collecting and rearing of the Cairns Birdwing (Ornithop- 
tera priamus euphorion (Gray)) and other butterflies, and it was a very pleasant 
period in their journey. In January 1939, collections were made on Thursday 
and the Prince of Wales Islands in the Torres Straits. A chartered fishing boat 
then brought them to Merauke on the mainland of New Guinea where they 
collected during the months of February and March. They proceeded to the Aru 
Islands in April where Clo stayed with an Australian family while Bob went on 
to Fakfak to collect. There he became ill with dengue and he returned to Aru. 
Bob and Clo then proceeded to the northern part of New Guinea to the village 
of Babo, in a low marshy area with several rivers, where there was a Dutch oil 
field. They were permitted the use of houseboats for housing. They journeyed 
up the Wasian River to Camp 5 at the village of Wasian, and also up the Aimau 
River by outrigger to Soedoe Point, staying in native huts. Clo became ill with 
dengue at the latter locality. On her recovery they proceeded to the lowland area 
of Inanwatan. Bob had contemplated a trip to Bougainville in the Solomon Islands, 
but due to the late arrival of funds from a supporting institution the trip was 
cancelled. Meanwhile he had contacted a European collector in Kieta who supplied 
extensive materials from that area. Leaving New Guinea they travelled west to 
Ceram, staying for some time at Amboina. From Amboina they proceeded north 
to Batjan, which they considered a beautiful area. There they enjoyed the gen- 
erous hospitality of the Dutch inhabitants. Collecting during December was 
excellent, providing an abundance of specimens of Ornithoptera priamus croesus 
(Wallace). The islands of Obi, Ternate and Halmahera were visited. At the 
latter Bob became ill with yellow jaundice and an oxcart had to be hired to return 
him to the boat for Ternate. The Winds decided to visit the Minahasa Peninsula, 
staying at Tondano in the mountainous northern Celebes so that Bob could regain 
his health in a more favorable climate. This area is only 1° north of the equator. 
At nearby Lake Dono (Lake of Man) remarkable collections were made including 
a very large lycaenid. They left the Dutch East Indies for Manila in 1940 under 
blackout conditions, as the Second World War had started in Europe. Some 
collections were made about Manila prior to their return to California. 

The Winds returned to Berkeley and Bob undertook lecture tours showing movies 
of their Papuan-Australian Expedition. Their son Robert M. was born in 1943. 
The health of both Bob and Clo had been so impaired by their travels that they 
required hospitalization at the Del Valle Sanatorium in Livermore in 1942 for 
tuberculosis. Bob conducted a successful philatelic business from the sanatorium. 
Davies recalls a visit when Bob was sitting in bed surrounded by boxes of stamps 
and catalogs relative to this mail-order business. During his convalescence he 
resided in Livermore. He purchased a house there, continuing the philatelic as 
well as the entomological sales. Davies also recalls the visits made to make butterfly 
purchases from the extensive Wind expedition materials. At this time Bob’s health, 
even though he had lost the use of one lung, improved sufficiently that he could 
again start field work. 

Davies and Bob collected frequently in the Los Mochos Canyon area, Mitchell’s 
Canyon in Contra Costa County, twice at Strawberry Lake at Pinecrest where 
the new Melitaea leanira daviesi Wind was collected, and at Sonora Pass. Sub- 


VoLuME 30, NUMBER 3 241 


sequently in the early 1950’s they made trips to collect at Partington Canyon in 
Monterey County. It was in August, 1948 that Arnaud, while camping at Pine- 
crest and collecting in terrain at the far end of Strawberry Lake, first met Bob, 
who was collecting butterflies. 

It was in the period of 1945-1947 that Bob actively published, as the bibliography 
shows, on the Indo-Australian Lycaenidae (authored with Harry K. Clench), as 
well as on North American Satyridae and Nymphalidae. Eighteen species and 
subspecies were described in 5 papers. 

From 1950-1955 the Winds operated the Butterfly Tree Park Museum and 
Gift Shop in Pacific Grove. This was associated with a motel and a grove of 
Monterey Pines frequented by the monarch in its winter stay. ‘here were ex- 
tensive displays of tropical butterflies, beetles, and other “Oh My” insects. A 
12 page pamphlet, “Wandering Wings, The Story of Pacific Grove’s World-Famous 
Buttertly Trees” was written and published by Bob at this time. 

Starting in 1952 and in following years Bob was interested in magic and was 
an active member of the Monterey Bay Sahareen (Sorcerers) Club. In 1953 the 
Winds established their own business—the Funny Abalone—on Fisherman’s Wharf 
in Monterey. Here they sold shells, gifts and butterfly novelties. This successful 
venture led to the Winds becoming wholesale distributors of natural history ma- 
terials, particularly shells. To have room for all the stocks required a large facility, 
and this led to their renting a large warehouse on Cannery Row in Monterey 
and the establishment of Bob Wind’s Butterfly Shop, with floor space of over 
20,000 sq. ft. Extensive stocks of insect specimens were handled from all world 
areas. In 1955 the Winds opened another gift shop—the Trade Winds—also on 
the wharf at Monterey. 

Robert Wind became well-known in his field of business, and this led to awards 
and articles in newspapers and to TV coverage. He was asked to participate on 
Art Baker’s nationally televised show “You Asked For It” on 5 January 1958, with 
a portion of the program showing the operation of his Butterfly Shop on Cannery 
Row. Articles also appeared by John Keefauver on “Butterfly collector settles 
down with a shop in Monterey” in the Monterey Peninsula Herald, 15 November 
1957, and by Lonnie Wilson on “Butterflies on Cannery Row” in the Sunday 
Parade section of the Oakland Tribune on 2 March 1958. The 12 August 1958 
Monterey Peninsula Herald reported that Bob Wind was awarded a gold certificate 
by the Craft, Model, and Hobby Industry Magazine for “outstanding product 
developments aiding the growth and welfare of the hobby industry.” This cer- 
tificate was for his butterfly collecting and mounting kits. 

In May 1959, in a partnership, Bob opened the restaurant “The Outrigger” 
located at 700 Cannery Row on the ocean end of the Old Monterey Cannery 
overhanging the waters of Monterey Bay. The front end of the Cannery was 
occupied by his wholesale and retail Butterfly Shop, while the restaurant was 
capable of seating 600 persons (with a 400 person banquet room) with dining 
on 2 levels overlooking the bay. In October 1960, the Winds opened the much 
larger Trade Winds Gift Shop in the Carmel Plaza area. The financing of the 
restaurant business necessitated the gradual sale of the gift shops. In 1965 a 
business change was required and Bob decided to renew his activities from his 
own collecting of butterflies and insects from areas in Mexico. He established 
himself first at Ajijic in Jalisco on Lake Chapala, and later in the state of Chiapas. 
In 1975 he was moving the site of his collecting from the higher altitudes of 
Chiapas to new areas of Guatemala. He arrived in Antigua to recuperate his 
health and to begin collecting there. There were volcanic eruptions in the area 
at the time, and with his weakened health complicated by respiratory problems, 
he died on July 4th at Antigua. He was buried there. 

Bob will be missed by his many professional and amateur entomological friends 
throughout the world. He is survived by his widow Clo Wind (now Mrs. Morrie 


949 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


+ 


J. Carroll), his son Robert Mifflin Wind, a brother Howard Wind and two sisters— 
Mrs. Ray Hetman and Mrs. Marge Harville. We would like to thank Mrs. Carroll 
for providing data and verifying other information included in this article. 

Specimens that Bob collected personally or handled in his dealership are dis- 
tributed in many major institutional collections and in countless numbers of private 
collections throughout the world. For materials deposited in the California Academy 
of Sciences collections, we have the record that between 16 April 1941 and 21 
June 1960, in nine transactions (exchanges, gifts, and purchases) the Academy 
acquired about 9600 insect specimens from localities in Madagascar, New Guinea, 
Australia, Celebes, and Peru. 

His collection of primarily western North American butterflies assembled in the 
1930’s and 40’s, through arrangements made by Davies with Mrs. Clo Carroll and 
with a donation from Mr. Donald Patterson, was transferred to the Department of 
Entomology on 1 February 1976. It numbers 3841 pinned specimens. Among 
the many species represented are the type series of Megisto rubricata smithorum 
Wind (holotype, allotype and 5 paratypes; now assigned CAS Ent. type no. 12575), 
Minois meadii melania Wind (holotype, allotype and 5 paratypes; now assigned 
CAS Ent. type no. 12576), and Neonympha henshawi texana Wind (holotype and 
3 paratypes; now assigned CAS Ent. type no. 12574), 6 paratypes of Coenonympha 
inornata nipisiquit McDunnough, 1 paratype of Incisalia niphon clarki Freeman, 
2 paratypes of Megathymus evansi Freeman (now in Agathymus), 5 paratypes of 
Melitaea leanira daviesi Wind (now in Thessalia), 2 paratypes of Plebeius scudderi 
empetri Freeman (now Lycaeides argyrognomon empetri), 2 paratypes of Speyeria 
cybele pugetensis Chermock & Frechin, 1 specimen of the aberration Nymphalis 
antiopa hygiaea (Heydenreich), 14 specimens of the extinct Glaucopsyche xerces 
(Boisduval) and its forms from San Francisco, and 8 specimens of the rare or 
possibly extinct Speyeria adiaste atossa (Edwards) from the Tehachapi Mountains. 
Each specimen of this collection is receiving a label indicating that it formed 
part of the Robert G. Wind collection and was accessioned by the California 
Academy of Sciences in 1976. 


Some Taxa Named After R. G. Wind 


windi Clench, Incisalia doudoroffi, 1943, Can. Ent. 75: 173. New _ subspecies, 
holotype female from Placer County, California and three paratypes from 
Gold Lake, Plumas County; Mt. Elwell; and “Sier. Nev. Cal.” Now considered 
a subspecies of Incisalia fotis (Strecker). 

windi Gunder, Euphydryas anicia, 1932, Can. Ent. 64: 283. New race, holotype 
male, allotype female, and four paratypes collected at Timber Island, Teton 
County, Wyoming. Now considered a subspecies of Euphydryas anicia 
( Doubleday ). 

windi Gunder, Plebeius maricopa, 1933, Can. Ent. 65: 173. New transitional form, 
holotype male from Berkeley, California. Now considered an aberration of 
Plebejus pardalis pardalis (Behr). 


New Taxa—Lepidoptera—Described by R. G. Wind 


The collection data are given only for the holotypes. 


Lycaenidae 


arfakiana Wind & Clench, Callictita cyara, new subspecies; 1947, Psyche 54: 60-61. 
Holotype, male, “Mt. Siwi, Arfak, Dutch New Guinea, 800 meters, May 4, 1928 
(Dr. E. Mayr).” 

ariadne Wind & Clench, Philiris, new species; 1947, Bull. Brooklyn Ent. Soc. 42: 
7-8. Holotype, male, “Wau, Morobe District, New Guinea, May 6, 1932 (H. 


Stevens ).”’ 


VoLUME 30, NUMBER 3 243 


azula Wind & Clench, Philiris, new species; 1947, Bull. Brooklyn Ent. Soc. 42: 8— 
9. Holotype, male, “Wau, Morobe District, New Guinea, Oct. 15, 1932 (H. 
Stevens ).” 

bicolorata Wind & Clench, Philiris fulgens, new subspecies; 1947, Bull. Brooklyn 
Ent. Soc. 42: 9-10. Holotype, male, “Dobo, Aru Islands, June 3, 1939 (R. G. 
Wind ).” 

birou Wind & Clench, Philiris intensa, new subspecies; 1947, Bull. Brooklyn Ent. 
Soc. 42: 10-11. Holotype, male, “Wau, Morobe District, New Guinea, Aug. 
8, 1932 (H. Stevens ).” 

deliciosa Wind & Clench, Thaumaina uranothauma, new subspecies; 1945, Pan- 
Pacific Ent. 21: 14-16. Holotype, male, “Wau, Morobe District, New Guinea, 
January 30, 1933 (H. Stevens ).” 

evinculis Wind & Clench, Philiris innotatus, new subspecies; 1947, Bull. Brooklyn 
Ent. Soc. 42: 11-12. Holotype, male, “Redlynch, North Queensland, Australia, 
August 14, 1938 (R. G. Wind).” 

kunupiensis Wind & Clench, Candalides meeki, new subspecies; 1947, Bull. Brooklyn 
Ent. Soc. 42: 3-4. Holotype, male, “Mt. Kunupi, Menoo Valley, Weyland Mts., 
Dutch New Guinea, 6000 ft., Nov._Dec. 1920 (C., F., and J. Pratt), ex coll. 
E. I. Huntington, Acc. 34,909.” 

mayri Wind & Clench, Philiris, new species; 1947, Bull. Brooklyn Ent. Soc. 42: 
14-15. Holotype, male, “Mt. Siwi, Arfak Mts., Dutch New Guinea, 800 m., 
April-June 1928 (Dr. E. Mayr), Acc. 31075.” 

misimensis Wind & Clench, Philiris, new species; 1947, Bull. Brooklyn Ent. Soc. 42: 
15-16. Holotype, male, “Mt. Misim, Morobe District, New Guinea, 5—6000 feet 
(H. Stevens)” [date of collection omitted]. 

morobea Wind & Clench, Candalides grandissima, new subspecies; 1947, Bull. Brook- 
lyn Ent. Soc. 42: 4-6. Holotype, male, “Wau, Morobe District, New Guinea, 
April 18, 1932 (H. Stevens ).” 

papuanus Wind & Clench, Philiris diana, new subspecies; 1947, Bull. Brooklyn Ent. 
Soc. 42: 6. Holotype, male, “Wau, Morobe District, New Guinea, June 2, 1932 
(H. Stevens ).” 

putih Wind & Clench, Philiris moira, new subspecies; 1947, Bull. Brooklyn Ent. Soc. 
42: 12-13. Holotype, male, “Pt. Moresby, British New Guinea, April 26, 1939 
(R. G. Wind).” 

stevensi Wind & Clench, Candalides erinus, new subspecies; 1947, Bull. Brooklyn 
Ent. Soc. 42: 1-2. Holotype, male, “Wau, Morobe District, New Guinea, 
April 4, 1932 (H. Stevens).” 


Nymphalidae 


daviesi Wind, Melitaea leanira, new subspecies; 1947, Pan-Pacific Ent. 23: 171. 
Holotype, male, “Strawberry Lake, Tuolumne County, California, el. 5500 ft., 
June 26, 1945.” 


Satyridae 
melania Wind, Minois meadii, new subspecies; 1946, Pan-Pacific Ent. 22: 25-26. 
Holotype, male, “Marfa Alpine, Texas, July 17, 1941” (Arthur & Edgar Smith). 
smithorum Wind, Megisto rubricata, new subspecies; 1946, Pan-Pacific Ent. 22: 26. 
Holotype, male, “Marfa Alpine, Texas, July 17, 1941” (Arthur & Edgar Smith). 


texana Wind, Neonympha henshawi, new subspecies; 1946, Pan-Pacific Ent. 22: 27. 
Holotype, male, “Marfa Alpine, Texas, July 17, 1941” (Arthur & Edgar Smith). 


Bibliography of Papers by R. G. Wind with New Taxa 


Winp, R. G. & H. K. Cirencu. 1945. Notes on the genus Thauwmaina (Lepidop- 
tera: Lycaenidae). Pan-Pacific Ent. 21: 14-16. (One new subspecies. ) 


244 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Winp, R. G. 1946. Some new species of North American Satyridae (Lepidoptera ). 
Pan-Pacific Ent. 22: 25-27. (Three new subspecies. ) 

Winp, R. G. 1947. A new subspecies of Melitaea (Lepidoptera). Pan-Pacific 
Ent. 23: 171. (One new subspecies. ) 

Winp, R. G. & H. K. Crencn. 1947. New Indo-Australian Lycaenidae (Lepi- 
doptera). Bull. Brooklyn Ent. Soc. 42: 1-16. (Four new species; eight new 
subspecies. ) 

Winn, R. G. & H. K. Crencn. 1947. The genus Callictita (Lepidoptera, Lycae- 
nidae). Psyche 54: 57-61. (One new subspecies. ) 


Paut H. Arnavup, Jk. AND Tuomas W. Davies, Department of Entomology, 
California Academy of Sciences, Golden Gate Park, San Francisco, California 94118. 


BOOK REVIEW 


History or Enromo.ocy, Editors: RF. Smith, T. E. Mittler and C. N. Smith. 
1973, Annual Reviews, Palo Alto, Calif. 517 p., 42 figs. Price $10.00 (U-S.). 


Lepidopterists should enjoy browsing through this multi-chaptered (20), multi- 
authored (25) tome, because it covers such a wide span of interesting entomo- 
logical observations and xsesearch. Many of the topics (systematics, paleoentomology, 
anatomy and morphology, physiology, behavior, etc.) are applicable as background 
thinking for and appreciation of problems related to moths and _ butterflies. 

A further point of interest, the agreed intent of the editorial committee (Preface ) 
was that emphasis be placed on the personalities of those who have contributed to 
entomology. Such was ably accomplished in most chapters (many “greats” figured, 
personal viewpoints and traits noted—even current members of The Lepidopterists’ 
Society, e.g., C. P. Alexander, cited). 

Specifically for the Lepidoptera, Lindroth (section 6) devotes several pages to 
systematists; the soul (psyche) was named from the moth, “phalaene” (p. 38); 
butterflies were light-trapped in ancient times (p. 52); court trials of destructive 
caterpillars took place in the 15th century (p. 81); and so forth. Silkworms are 
discussed in several chapters—that by Yokayama is most instructive for students 
of lepidopteran biology. 

Overall, this History of Entomology is a fine book for all interested in the 
development and developers of the scientific study of insects. 


WittiAM B. Nutrinc, Department of Zoology, University of Massachusetts, 
Amherst, Massachusetts 01002. 


EDITORIAL STAFF OF THE JOURNAL 
GrorcE L. Goprrey, Editor 


Illinois Natural History Survey, Natural Resources Building 
Urbana, Illinois 61801 U.S.A. 


Wii1i1AM H. ALLEN, Associate Editor JAMeEs G. STERNBURG, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study 
of Lepidoptera. Contributors should prepare manuscripts according to the following 
instructions. 

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SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, 
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196la. Some contributions to population genetics resulting from the 
study of the Lepidoptera. Adv. Genet. 10: 165-216. 


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U.S. A 


CONTENTS 


THE OVERWINTERING SITE OF THE EASTERN POPULATION OF THE 
MonarcH BuTTERFLY (DANAUS P. PLEXIPPUS; DANAIAE) IN 
SOUTHERN Mexico. F. A. Urquhart and N. R. Urquhart —_- 

NOTES ON THE LirE Cycle AND Natural History or BUTTERFLIES 
OF Ex Satvapor. VIII. ARCHAEOPREPONA ANTIMACHE GULINA, 
SIDERONE MARTHESIA, ZARETIS CALLIDRYAS AND CONSUL 
ELECTRA (NYMPHALIDAE). Alberto Muyshondt ___ 

PATTERNED PERCHING IN Two CALLoPHRYS (Mirour4) (LYCAENI- 
DAE). Kurt Johnson and Peter M. Borgo 

OBSERVATIONS ON Host PLANT RELATIONSHIPS AND LARVAL NUTRI- 
TION IN CALLOSAMIA (SATURNUDAE). Richard S. Piegler __ 

A REvIsION OF THE GENUS Dunama ScHaus (NoropontmaeE). E. L. 
Todd 82000 i 

THE BuTrerFLics OF MississippI—SUPPLEMENT No. 2. Bryant 
Mather and Katharine Mather _. ae 

RHOPALOCERA IN THE N. B. SANSON CoLLEcTION. Charles D. Bird — 

NOTES ON THE BIOLOGY AND IMMATURE STAGES OF THE WHITE 
PEACOCK BUTTERFLY, ANARTIA JATROPHAE GUANTANAMO 
(NYMPHALIDAE). George W. Rawson ___-------- 

A New SPpEcIEs OF THE GENUS BERTELIA B. & McD. (Pybsen eee 
André Blanchard 00 3 

THE STATUS OF SATYRIUM BOREALE (LYCAENIDAE). Lawrence F. 
Gall iyi oe 

THE GIANT Biastopasiy Motus oF Yucca (GELECHIOWEA). Jerry 

A. Powell 0 S80 NO ON 


GENERAL NOTES 


Hilltopping in Lebanon. Torben B. Larsen 
Time variations of pupal stage of Eupackardia calleta (Satumiidae). 
Jack: B. Prentiss i 
Evidence of breeding migrant populations of Leptotes cassius (Lycaenidae) 
in Kansas: Matthew: M. Douglas 0. 

A population of the striped hairstreak, Satyrium liparops liparops ( Lycaeni- 
dae), in west-central Florida. Larry N. Brown 
Oiketicus toumeyi: A bagworm moth new to the Texas fauna (Psychidae). 
Raymond W. Neck 22.000 

A survey of the Sphingidae of Sanibel Island, Florida. C. Hugh Brown _ 
First records of Boloria frigga (Nymphalidae) in Wisconsin. Leslie A. 
Ferge and Roger: M, Kuehn ee 
Additional new butterfly records from Florida. Robert Bennett and Edward 
Ci Knudson ie i ts is ear 
Nocturnal activity of a monarch butterfly (Danaidae). Raymond W. Neck 
Crab spider preys on Neophasia menapia (Pieridae). Daniel T. Jennings 
and ‘Michael E.: .Toliver: 3. a 
Butterflies associated with an army ant swarm raid in Honduras. Boyce 
A, Drummond TEL ee 
NOTES AND (NEWS) 


OBITUARY IO EET MEIN. Th CR a ae | 


153 


201 


207 


Volume 30 1976 Number 4 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


\ 


( dl @. hodges 


3 December 1976 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL e 


x 
S. S. Nico.ay, President Cartos R. BEUTELSPACHER, 
Miriam RorHscuHitD, Ist Vice President Vice President 


THEODORE D. SarcENT, Vice President Joun M. Sniper, Treasurer 
Jutian P. DonanueE, Secretary 


Members at large: 


J. T. Brewer D. F. Harpwick BR. A. ARNOLD 
K. S. BROWN J. B. ZrecLer E. D. CasHatTr : 
K. W. Pxuivip F. S. CHEw R. E. STANFORD 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and 
formally constituted in December, 1950, is “to promote the science of lepidopterology 
in all its branches, . . . . to issue a periodical and other publications on Lepidoptera, 
to facilitate the chee of specimens and ideas by both the professional worker an 
the amateur in the field; to secure cooperation in all measures” directed towards 
these aims. 

Membership in the Society is open to all persons interested in the study of 
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Society. Institutions may subscribe to the Journal but may not become members. 
Prospective members should send to the Treasurer full dues for the current year, 
together with their full name, address, and special lepidopterological interests. In 
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February, May, August and November, and six numbers of the News each year. 


Active members—annual dues $13.00 
Student members—annual dues $10.00 
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Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
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Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA ~~ 


by Cyr F. pos Passos 


Price: Society members, $5.00 U.S.; non-members, $7.50 U.S. Paper covers, revisions 
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Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni- 
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JOURNAL OF 


Tue LEpPIDOPTERISTS’ SOCIETY 


Volume 30 1976 Number 4 


PRESIDENTIAL ADDRESS 1976—WHAT INSECTS 
CAN WE IDENTIFY? 


Ronatp W. HopcEs 
Systematic Entomology Laboratory, IIBIII, Agr. Res. Serv., USDA? 


I work for an organization, the Systematic Entomology Laboratory 
of the U.S. Department of Agriculture, which is vitally concerned with 
recognition and differentiation of insect species. Although the numbers 
fluctuate from year to year, the 28 scientists in the laboratory identify 
approximately 250-300 thousand specimens each year. I realize that 
many of you must wonder as you hear me give these figures why is it 
that when you send specimens to be identified we do not respond im- 
mediately to a request for identification. Much of the very large number 
of specimens with which we deal comes from agricultural sources such 
as the plant quarantine stations of Animal and Plant Health Inspection 
Service, the Agricultural Research Service, the Forest Service, state 
agencies, international ports of entry, and museums. Our major resources 
for making these identifications are the National Collection of Insects 
(approximately 24 million specimens), the combined libraries of the 
Smithsonian Institution, the Library of Congress, and the National Agri- 
cultural Library, and the numerous files of host plants, catalogues, cards, 
and separates built up by the scientists working with the collection over 
the last 90 years. 

When we talk or think about numbers of insects, we usually refer 
to the large number of undescribed species. The general estimate is 
that about 1 million names have been proposed for insects to date and 
that there may be 1-10 million species to be described. These figures 
are impressive by any standard and are often cited as one of the major 
problems in making identifications. But, what about the names in the 


1 Mail address: c/o U.S. National Museum, Washington, D.C. 20560. 


246 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TaBLeE 1. Percent of specimens associable with described species by museum 
workers for a series of faunas. 


N. America Palearctic Neotropical Exotic 
Described Described Described Described 
% Species % Species % Species % Species 
Diptera 
Culicidae 95 250 50 3500 
Mycetophilidae 5 612 8 
Cecidomyiidae 5 1200 il 
Ceratopogonidae TS 400 25 4000 
Tipulidae 95 1500 80 14000 
Syrphidae 40 1000 25 6000 
Muscidae 90 622 40 
Sarcophagidae 95 Oli 60 
Calliphoridae 95 78 65 
Tachinidae 82 1281 28 2864 
Coleoptera 
Curculionidae 75 2600 50-60 0=5 0-5 
Scarabaeidae 90 1400 90 500 50 2000 5-10 
Coccinelidae 99 400 95 200 50 600 25 
Bruchidae 100 100 90 40-50 900 
Buprestidae 80 660 2 11400 
Colydiidae 95 110 25 JS 
Dermestidae 95 130 AO 877 
Histeridae 50 360 5 3500 
Heteroptera 15 60 50 25-50 
Homoptera ~ 
Aphidae 0) 1500 45-50 2000 
Aleyrodidae 90 500 
Coccoidea 80 1500 80+ 1500 20 1000 40-50 2000+ 
Cicadellidae (¢ 3) 80 3500 12 20000 
Hymenoptera 
Symphyta Tus) 1000 60 2500 25 1000 40 2500 
Formicidae 
workers 80 650 30 2000 15 2700 10 3000 
29 15 5 il 1 
é 4 10 5 il il 
Ichneumonidae 50 2850 15 15000 
Chalcidoidea 50-60 50-60 5-15 
Braconidae 65 2000 10 10000 
Isoptera 
soldiers 90 45 15 1000 15 1500 
alates 90 15 15 
workers 5 5 ] 
Lepidoptera 
Lycaenidae 95 300 98+ 70 80-85 
Noctuoidea 50 3100 30 50000 
Geometridae 90 1200 + 50 10-15 10000 


Gelechiidae 80 750 10-15 4000 


VoLUME 30, NUMBER 4 247 


literature? What do they mean as far as an identification is concerned? 
When you peruse the McDunnough check list or parts of the Lepidop- 
terorum Catalogus, what do the names mean to you or to anyone? The 
answer is that they represent various states of knowledge. In some 
rare instances the names can be associated with biological entities in 
contemporary terms. A higher percentage can be associated with mor- 
photypes and identified as such. A yet larger percentage of the names 
represent nearly nothing to an individual trying to make identifications. 

When a field worker in ecology or biological control, or someone 
making an environmental impact statement, wants to have specimens 
determined, he normally sends them to one place, usually where sys- 
tematists are willing or obligated to make determinations. In the United 
States the Systematic Entomology Laboratory is a major source of 
determinations on a broad level. In Canada scientists of the Biosys- 
tematics Research Institute make determinations. And, there are several 
regional identification centers such as the California Department of 
Agriculture, the Florida Department of Agriculture, the Illinois Natural 
History Survey, the New York State Science Survey, as well as individual 
systematists at numerous universities. 

Because I have access to information about the National Collection 
and workers associated with it, I asked the following question of our 
scientists: On the basis of the collection and the literature available to 
you, what percent of the names in the literature can you associate with 
specimens for a series of faunas with a fair degree of certainty? The 
answers were couched with various degrees of uncertainty and were 
not for a consistent series of zoogeographic areas. For North America 
north of Mexico I have listed data for each family group. For areas 
other than this I have data for different associations of areas. Some of 
the responses are as follows (Table 1): 

The numbers become monotonous, but they serve to emphasize the 
point that we can identify relatively well the described North American 
fauna and very poorly the fauna from other parts of the world. Also, 
these figures are for adults. When the comparison is made for taxa for 
which larvae are known, the contrast is striking (Table 2). 

What do these figures mean in practical terms? Currently, we cannot 
identify with certainty a relatively large percentage of the described 
world fauna. 

Without question there is a need to know what these insects are. 
When an insect is intercepted at a port of entry, the question arises, 
“Is it of economic importance or potentially of economic importance?” 
If it is, certain measures will be taken. If it is not, and the only way 


948 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TaBLE 2. Percent comparison of larval and adult specimens associable with 
described species. 


N. America Palearctic Neotropical Exotic 
Described Described Described Described 
% Species % Species % Species % Species 

Symphyta 

adults 75 1000 60 2500 25 1000 ZA) 2500 

immatures 10 10 1 1 
Formicidae 

workers 80 650 30 2000 15 2700 10 3000 

immatures il 0 0 0 


one can say that it is not is to know what it is, then the commodity can 
enter the country directly. 

A generic or family level determination is not adequate for sampling 
work. Much of the time and money spent on numerous surveys has 
been and continues to be wasted for lack of specific determinations. 
Meaningful comparisons for most purposes can be made only at the 
specific level. The main reason for stopping short of this level is lack 
of available expertise and/or cost of determinations. Use of parasitic 
insects as biological control agents requires specific determinations. The 
pendulum is swinging back insofar as needing to know exactly what 
an insect is for control programs in agriculture. No longer can all 
insecticides be used indiscriminately for pests. Each target insect must 
be identified on the label, and the insecticide must be used where 
needed—not ubiquitously. The Systematic Entomology Laboratory was 
started when there was a need to know insects for agricultural purposes. 
With the advent of DDT and successors, many entomologists thought that 
all the crop problems caused by insects would be solved by their use. 
We know better now. Control or suppression—not eradication—is a 
major goal in agricultural research today. 

Unfortunately, for economic and social reasons, it apparently is not 
justifiable to spend a research career on basic taxonomic work in most 
educational institutions. I strongly argue this concept. As long as there 
is a need to know what animals are, then we must continue the process 
of making known what has been described, refining our means of recog- 
nizing species, genera, and higher categories, and integrating the un- 
described species into a system. 

Within the Lepidoptera the families are in varying stages of knowl- 
edge. The butterflies are probably the best known with the papilionids 


VOLUME 30, NUMBER 4 249 


at the top of the list. For the large superfamilies, Noctuoidea, Geometroi- 
dea, Pyraloidea, Tortricoidea, Gelechioidea, Yponomeutoidea, and Tine- 
oidea, many problems are extant in determining what a name represents 
or with which genus a new species should be associated. In many in- 
stances specimens in collections have been identified by comparison with 
colored illustrations, by direct visual comparison of specimens with the 
holotype, by comparison of specimens with other determined specimens 
(often at the British Museum (Natural History) ), by comparison with 
written descriptions, and rarely by direct comparison with the holotype 
of several character systems. Most of the major papers written before 
1940 have been done without examination of type-specimens, including 
some that appear useful such as Heinrich’s revision of the North Ameri- 
can Olethreutidae, or without study of the male and female genitalia. 
With much of the literature nearly worthless except to validate scientific 
names and with many specimens in the collection questionably deter- 
mined, I contend that the base of our science is very weak. 

Someone must accumulate a large amount of material for each group 
in need of revision, draw together the available names (sometimes names 
are “hiding” in other families or superfamilies), study the type-specimen 
for each name, and associate each name with one or more specimens in 
the accumulated material. For a group as large as the Noctuidae with 
more than 5,300 generic names and 60,000 specific names the initial 
stages require an immense amount of time and dedication. Variation 
among specimens must be assessed. To my knowledge species vary in 
nearly all characters, and for this reason the male or female genitalia 
sometimes are no more final for specific determination than the shading 
of the color pattern, wing length, or other characters. Also, reliance on 
single characters for specific or generic distinction undoubtedly produces 
untenable classifications. Many of the species and particularly many 
of the genera are more widely distributed than our predecessors recog- 
nized, and often names proposed for specimens from other zoogeographic 
regions will prove to be senior synonyms of names proposed for specimens 
from North America. Conversely, many names have been applied too 
broadly in the past. These factors indicate that the studies should be 
done on as broad a base as possible, particularly at the generic level. 
Also, working with large numbers of species and genera gives the student 
a better perspective for his treatment of all categories. 

Special problems that Lepidoptera give to workers are their relatively 
large size, obvious color pattern, and scale covering. These have enabled 
many to work without recourse to study of other characters. Or, for 
wing venation, specimens have not been properly prepared for study. 


950 JOURNAL OF THE LEPOPTERISTS SOCIETY 


Many wings have been studied by temporary clearing of a part of the 
wing with volatile solvents rather than removing the wing, clearing, 
staining, and mounting it on a slide so that all veins could be studied 
properly. Many workers didn’t use microscopes. Edward Meyrick, who 
described more than 15,000 species of Lepidoptera, refused to use a 
microscope until his later years, and he refused to acknowledge that 
genital characters were worthwhile. He based much of his classification 
on the venation as seen through a hand lens. Meyrick died in 1938. 
If Lepidoptera were smaller, then workers would have been forced to 
study them at greater magnification initially and perhaps done a better 
job of comparative work. 

Lepidoptera, in general are very poorly collected. Although there 
are series of butterflies from several localities, this is not the case for 
the moths. There are many instances in which only the holotype is 
known or the extant specimens are less than 5. Before we can understand 
the species and their relationships, we must have much better repre- 
sentation of each species from numerous localities throughout its geo- 
graphic range. This part of the cycle is going to be very difficult to 
fill because one of the major sources of material is amateur collectors who 
find it very unrewarding to collect specimens and not be able to identify 
them. At the moment systematists cannot provide names for many 
species, or the number of systematists relative to those who would like 
to have names for specimens is so small that were they to do nothing 
but name specimens they would have no time for revisionary study. 

As we progress in our knowledge of the Lepidoptera, it will become 
necessary for those who want or need names for specimens to submit 
them properly prepared, with wings spread, in as good condition as 
possible, and with the genitalia prepared for study. It is not reasonable 
to expect the systematist to spend 1-2 hours preparing a specimen, so 
that he can begin to make a determination unless the correspondent 
wants to spend $30-40 for an identified specimen. 

One can ask, “Where do we go from here?” At the present rate of 
study the answer is “not very far.” With the current small number of 
systematists doing revisionary work, the likelihood that the world fauna 
will be known in a comprehensive manner is very low. The described, 
world fauna of Lepidoptera is more than 140,000 species, and if projec- 
tions are correct, the total may be as high as 280,000-1,400,000 species 
for the world. The demands on systematists’ time are such that to be 
able to revise 60-100 species within a year often is not possible. Also, 
the first needs are for general studies covering the higher categories 
through the generic level for the world. If done for smaller zoogeographic 


VOLUME 30, NUMBER 4 yAS 


areas, the revisions must take into account the genera of the world. 
From this point others can revise genera or groups of genera for the 
world or smaller areas. We also need general manuals for use by a broad 
spectrum of persons from the amateur to specialist. However, under 
current administrative requisites for job evaluation large projects are 
not favored. Many, short publications are preferred over few substantia] 
ones. Unless these attitudes change, I do not see how we can accomplish 
the work that needs doing. At the same time I feel that a broad audience 
should be made aware of the poor foundation of systematic entomology. 


952 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


CONCERNING THE NAME ANTHOCARIS COLORADENSIS 
HY. EDWARDS WITH DESIGNATION OF A NEW SUBSPECIES 
(PIERIDAE) 


Kurt JOHNSON? ? 


Department of Biology, City University of New York City College, Convent Avenue 
and 138th Street, New York, New York 10031 


Rocky Mountain populations of Euchloe ausonides Lucas found east of 
the continental divide have long been given the subspecies designation 
coloradensis. During a study of speciation in butterflies of the Black 
Hills of South Dakota and coniferous forest relicts on the western Great 
Plains, the original description and type specimens of coloradensis were 
found to lend considerable ambiguity to that name. Edwards’ (1881) 
original description of coloradensis emphasized dorsal characters of the 
wing which are generally useless when large series of Rocky Mountain 
material are considered. Likewise, the two syntype specimens labelled 
“Colorado” by Edwards and now in the American Museum of Natural 
History type collection were the only Rocky Mountain representatives in 
Edwards hands at the time of the description. Further, one of them 
represents a minority morph when the larger view of Rocky Mountain 
E. ausonides is considered. 

Thus, in describing a new subspecies from conifer forest relicts on the 
western Great Plains, I am first rediagnosing the name coloradensis, 
based on eastern Rocky Mountain populations in Colorado and Wyoming 
(Opler, 1968). 


Euchloe ausonides coloradensis (Hy. Edwards) 


Anthocaris coloradensis Hy. Edwards, 1881, p. 50. 

Anthocharis (sic) ausonides: Cary, 1901, p. 310. 

Anthocharis (sic) ausonides var. coloradensis: Beutenmiiller, 1892, p. 168; 1898, 
p. 241. 

Synchloe ausonides coloradensis: Dyar, “1902” 1903, p. 7. 

Euchloe ausonides coloradensis: Barnes & McDunnough, 1917, p. 3. Barnes & 
Benjamin, 1926, p. 7. Klots, 1930, p. 154. Leussler, 1938, p. 76. Defoliart, 1956, 
p. 98. Brown, Eff, and Rotger, 1957, p. 181. Puckering & Post, 1960, p. 8. 
dos Passos, 1964, p. 49. Opler, 1968, p. 69. Shields, Emmel, & Breedlove, 1969 
(1970), p. 31. Ferris, 1971, p. 15. Johnson, 1972 (1973), p. 28. 

Euchloe belia var. belioides race montana Verity, “1905-1911,” Dp) ooo 

Euchloe ausonides montana: Barnes & McDunnough, 1917, p. 3 (placed as synonym 
of coloradensis ). 


‘Museum Research Associate, Museum of Natural History, University of Wisconsin, Stevens 
Point, 54481, 


* Research Associate, Allyn Museum of Entomology, Sarasota, Florida, 33580. 


VOLUME 30, NUMBER 4 Ze 


Diagnosis. Distinguishable from E. a. ausonides Lucas and the following sub- 
species by a combination of traits: Under surface, hindwings: (1) green patch 
between veins RS and M: usually isolated on both wings, sometimes only on one; 
located more caudodistally toward marginal green patch between veins M: and Mb, 
not usually joined to more basad markings as on new subspecies. (2) White patches 
along inner angle generally invaded by white ground color, not recognizable as six 
alternating large and small smoothly edged patches as on new subspecies. (3) Green 
marbling less invaded by white ground color than on ausonides, more than on new 
subspecies. Under surface, forewings: (4) black crescent, distal end discal cell, at 
largest touching both veins Re and Ms. Upper surface, forewings: (5) apical area, 
black patterning darker than ausonides and new subspecies. 

Male. Upper surface of the wings: white; forewings, dark apical-subapical 
markings: discal cell, distal end, black crescent. Hindwings darker in areas where 
“marbling” occurs on under surface. 

Under surface of the wings: white; forewings, olivaceous apical-subapical mark- 
ings; discal cell, distal end, black crescent. Hindwings white with green “marbling” 
incised by white ground color; distal green patch between veins M: and Mz usually 
isolated, white surrounding it. 

Length of forewing: 20 mm (male type). 

Female. Wing characters identical with male. 

Length of forewing: 21 mm (female type). 

Male genitalia (Fig. 2). Tegumen flattened dorsally; uncus long, gradually 
tapered, often exceeding posterior end of valvae; valvae, apex directly slanted an- 
teriorly toward dorsal articulation, anterior margin slightly concave, aedeagus, phal- 
labase, noticeable “two-step” structure. 

Female genitalia (Fig. 2F). Not useful for infraspecific diagnosis. 

Early stages. Not specifically studied in relation to nominate E. ausonides, see 
Opler, “1974” (1975). 

Food plant. In Colorado, Arabis, Descurainia (Cruciferae) (Shields, Emmel, & 
Breedlove “1969” (1970) ); Arabis, Sisymbrium, Erysimum (Cruciferae) (Remington 
(1952)).)- ; 

Types. In the American Museum of Natural History collection (AMNH): type, 
male, “Colorado”; type, female, “Colorado.” In his description of coloradensis, 
Edwards implies that he possessed only two specimens at the time of his description 
but knew of, or had seen, others. The collection of the American Museum of Natural 
History contains two specimens labelled by Hy. Edwards as male type and female 
type. Because the female type does not accurately represent the coloradensis morph 
I hereby designate the male type as the lectotype. 

Distribution. By present diagnosis and material examined: Rocky Mountains 
of Colorado and Wyoming east of the continental divide. Varying somewhat west- 
ward in Colorado and blending into a. ausonides in western Wyoming (Ferris, 1971; 
Opler, 1968). 

Flight period. Early June—September (in Colorado (Brown, Eff, & Rotger, 1957) ). 

Remarks. 126 specimens were submitted to character analysis (see Discussion). 
20 male and 10 female genitalia were examined. 


Euchloe ausonides palaeoreios Johnson new subspecies 
(Figs. 1 & 2A, D) 


Anthocharis ausonides: Cary, 1901, p. 310. 
Euchloe ausonides ausonides: Puckering & Post, 1960, p. 8. 
Euchloe ausonides coloradensis: Leussler, 1938, p. 76. Johnson, 1972 (1973), p. 28. 


Diagnosis. This subspecies can be differentiated from E. a. coloradensis by a 
combination of traits: Under surface, hindwings: (1) green patch between veins 
RS and Miz usually heavily joined with median green patch between veins Mi: and 


954 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. E. a. palaeoreios, n. subsp. Top upper left: upper surface of the wings, 
holotype, male. Top upper right: upper surface of the wings, allotype, female. 
Bottom, center (variation in under surface of the wings): top left, under surface 
of the wings, paratype male, Black Hills near Sundance, Wyoming; top right, same, 
upper surface of the wings; bottom left, under surface of the wings, allotype, female; 
bottom right, under surface of the wings, holotype, male. 


M: (along cell) on one, or more often both wings and located more cephalobasad 
on the wing. (2) White patches along inner angle most often appear as six smoothly 
edged patches alternating large and small. (3) Edges of green marbling quite entire 
and noticeably smooth. Under surface, forewings: (4) black crescent, apical end, 
discal cell, nearly always broadly edging veins Re and M; with black, sometimes ex- 
tending to vein CU;. Upper surface, forewings: (5) apical area black patterning 
lighter and less extensive. 

Male. Upper surface of the wings: white; forewings, dark apical-subapical 
markings; discal cell, distal end, expansive black crescent. Hindwings darker in areas 
where “marbling” occurs on undersurface. 

Under surface of the wings: white; forewings, olivaceous apical-subapical mark- 
ings; discal cell, distal end, expansive black crescent. Hindwings white with green 
marbling little invaded by white ground color, distal green patch between veins Mi 
and M2 usually joined cephalobasad with green patch between M: and Mb. 

Length of forewing: 20 mm (holotype); 15 mm-—22 mm (x = 19 mm), paratype 
males. 

Female. Wing characters identical with male. 

Length of forewing: 18 mm (allotype); 21 mm-23 mm (x = 22 mm), paratype 
females. 


VOLUME 30, NUMBER 4 PADS, 


Fig. 2. (A) Genitalia, E. a. palaeoreios, n. subsp., holotype (AMNH), lateral 
view, aedeagus removed. (B) Male genitalia, E. a. coloradensis, Hy. Edwards 
“Colorado” (AMNH, K-27), lateral view, aedeagus removed. (C) Male genitalia, 
E. a. ausonides (Boisduval), Oakland, California (AMNH, K-3), lateral view, aede- 
agus removed. (D) Valva, lateral view, E. a. palaeoreios, Sundance, Wyoming 
(AMNH, KJ #171b). (E) Same, E. a. coloradensis, Laramie, Wyoming (AMNH, 
KJ #23). (F) Female genitalia, E. a. ausonides. 


Male genitalia (Fig. 2A, D). Tegumen somewhat humped, rounded dorsally; 
uncus, apex more toothed than tapered; valvae, anterior margin deeply concave, 
apex broad, not immediately slanted toward dorsal articulation; aedeagus, phallabase, 
roughly tapered. 

Female genitalia. As typical of the species (Fig. 2F). Not useful for infraspecific 
diagnosis. 

Early stages and foodplant. Not specifically known. 

Types. Holotype, male, Spearfish Canyon, near Spearfish, Lawrence Co., South 
Dakota, 26 June 1939 (AMNH). The genitalia are in vial K. Johnson #4. Holotype 
and genitalia are in the collection of the American Museum of Natural History. 
Allotype, female, Spearfish Canyon, nr. Spearfish, Lawrence Co., South Dakota, 
26 June 1939 (AMNH). Genitalia are in vial K. Johnson #6. Deposited as above. 
Paratypes (all Lawrence Co., South Dakota): American Museum of Natural History, 
males: 3 specimens, near Lead, 24 June 1939; 1 specimen, near Lead, 22 June 1939 
(all A. C. Frederick); 2 specimens, Spearfish Canyon, 26 June 1939 (collector 
unknown); 1 specimen, Spearfish Canyon, 1 July (year unknown) (collector un- 
known); 1 specimen, Ice Box Canyon, 28 June 1939 (A. C. Frederick). Females: 
1 specimen, Custer State Park, 1 July 1962 (F. H. Rindge). Los Angeles County 
Museum, male: 1 specimen, near Lead, 24 June 1939 (A. C. Frederick); female: 
1 specimen, near Lead, 24 June 1939 (A. C. Frederick). Allyn Museum of Entomol- 


256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 3. Distributional and post-Pleistocene relationships of Euchloe ausonides 
subspp. Left: Distributions of E. a. ausonides (circles), E. a. coloradensis (squares ), 
and E. a. mayii (half-open circles) in continental United States and southern Canada. 
Eastward, in United States, pepper-dotted configuration shows probable maximum 
extent of post-Pleistocene conifer forest on the present-day western Great Plains: 
darkly blackened areas—upland climax conifer forest, lightly dotted areas—pine- 
juniper open woodland. Right: Inset of area on western Great Plains, showing west- 
ern North Dakota, South Dakota, and Nebraska, plus Colorado, Wyoming, and Mon- 
tana. Blackened areas show present montane conifer forests surviving on the Great 
Plains; black lines show present distribution of scarp woodlands in same region. 
Known localities of E. a. palaeoreios are indicated by white dots. Left of figure 
adapted from Opler (1968) by permission. 


ogy, male: 1 specimen, Terry Peak, 24 June 1939 (A. C. Frederick); female: 1 
specimen, Terry Peak, 24 June 1939 (A. C. Frederick). 

Distribution. The Black Hills of South Dakota and Wyoming, the Pine Ridge 
of Nebraska (not recently recorded), the Little Missouri River escarpments in North 
Dakota, and possibly the Lone Pine Hills in South Dakota and Montana, and the 
Killdeer Mountains in North Dakota. One plains population is known, represented 
by two specimens from Port Roch (sic) [Port Roche], Saskatchewan, deposited in 
the AMNH. 

Flight period. Dates on specimens range from 5 June-8 July. 

Remarks. 26 specimens, 7 male and 7 female genitalia were studied from the 
Black Hills; 6 specimens, 3 male and 3 female genitalia were studied from Saskatche- 
wan; 28, 3, and 3, respectively from Manitoba; 27, 3, and 3 from Alberta; and 5, 1, 
and 1 from Ontario (all AMNH). Six specimens, 2 male and 2 female genitalia 
were studied from Nebraska (Canyon Region north of Harrison, Sioux County), ob- 
tained from Ohio State University. I have included in the distribution of this entity 
only those areas where wing characters and genitalia are very near the description, 
not areas where the phenotype diverges or possibly intergrades with other taxa or 
unnamed populations. 


Etymology. The name is from the Greek words palaeos (old) and oreios (of the 


VOLUME 30, NUMBER 4 257 


mountains ), meaning “of the old mountains.” It refers to the relict montane areas 
from which this stock evidently originates (Johnson, 1976). 


DISCUSSION 


Following the retreat of the Wisconsin glacier, vast climax pine forests 
connected the present-day Pine Ridge and Wildcat Hills of Nebraska, 
Black Hills of South Dakota, and escarpments of western North Dakota. 
They also extended eastward along the present Niobrara River and into 
east-central Nebraska (Johnson, 1976). 

These conifer areas had formerly been mixed with prevailing boreal 
forest at the beginning of the glacial retreat (11-12,600 years ago) but 
became predominant about 10,000 years ago as the boreal forests were 
destroyed. Climax pine forests then became centered on the western up- 
lands of these states and connected westward to the Rocky Mountains 
through moist pine-juniper woodland. Thus, an eastward region of the 
Rocky Mountain environment was present at that time. However, a 
gradual trend toward aridity eventually cut off these eastern forests from 
their western allies and eventually from each other, leaving all as sub- 
climax pine woodlands except the climax-forested Black Hills. This 
trend was gradual at first, a slow drying and decimation of the moist 
savannahs lasting perhaps 5-7000 years. However, an arid cycle (sup- 
plemented by fire) beginning about 2000 years ago rapidly isolated the 
present relicts themselves, which have continued to decline. 

Because of this, the subspecies palaeoreios just described is distinct 
from Rocky Mountain E. a. coloradensis and, similarly, does not bear the 
relationship to Canadian prairie populations that might be inferred from 
present-day geography. Thus, the new name palaeoreios helps define 
both E. a. coloradensis and the divergent phenotype from the Ridings 
Mountains of Manitoba represented by the name E. a. mayii (Chermock 
& Chermock). 

It has not been my intention to introduce confusion into the identity 
of Canadian prairie populations by naming palaeoreios. I believe the 
geographic origin of palaeoreios is distinct from both E. a. coloradensis 
and Canadian prairie populations which probably have a more northern 
origin. Thus, I have included Canadian material in its distribution only 
where the genitalia are almost identical to topotypical dissections. The 
former climax forests, spoken of above, had widespread pine-juniper 
savannah extensions, of which the central Nebraska “arm” has been the 
only one studied by paleobotanists (Johnson, 1976). These former ex- 
tensions are now plains, but the distinct affinity of their remaining mon- 
tane butterflies to those in the scattered relict scarp woodlands is clear. 

The frequencies of six traits, in 132 specimens from six populations, 


258 JouURNAL OF THE LEPIOPTERISTS SOCIETY 


100% 


oO 


FREQUENCY 
| eee 
[@) 
in © 
ro) ~ 


oe) 


iE Iie IEICIE IV V VI 


ABC ABC ABC. AS Ce eee ABC 


Fig. 4. Frequency distributions of six traits in two major wing characters in 
Euchloe a. coloradensis and E. a. palaeoreios. Six populations are analyzed: (1) 
Rocky Mountain National Park, Colorado (n = 39); (II) Gunnison County, Colorado 
(n= 14); (III) Custer County, Colorado (n = 20); (IV) Edwards’ Type Series 
(n=10); (V) Laramie, Wyoming (n= 23); (VI) Black Hills, South Dakota, 
Wyoming (n = 26). Categories: (bottom) Relationship of patch between RS and M, 
with that median between M, and M,—(A) These patches joined on both wings, 
(B) one wing, (C) neither wing. (top) Relationship of patch between RS and M, with 
that marginal between M, and M,—(A) These patches joined on both wings, (B) 
one wing, (C) neither wing. 


are compared in Fig. 4. In the new entity, the marked relationship of 
(1) joining of the patch between RS and M, with that median between 
M, and Mb», and (2) its cephalobasad location away from any joining 
with the marginal patch between M, and Mb is clear. These frequencies 
also show how the Edwards female type did not accurately represent 
the E. a. coloradensis morph. This problem has been solved by the 
designation of a lectotype. The exact number and subsequent location 
of specimens Edwards knew of at the time of the description are unclear. 
However, a total of 10 specimens in the American Museum of Natural 
History Collection bear his handwriting and the label “Colorado.” These 
have been included in the above analysis to place his apparent idea of 
the name in a more realistic perspective. This, and the rediagnosis, 
should make the name coloradensis much more meaningful for future 
workers; the new name palaeoreios serves to separate coloradensis from 
the distinct scarp woodland and prairie population which has a distinct 
eastern origin. 
ACKNOWLEDGMENTS 


[ especially want to thank Dr. Frederick H. Rindge (Curator of Lepi- 
doptera the American Museum of Natural History) for his helpful com- 


VoLUME 30, NUMBER 4 259 


ments on this research, and Dr. Paul A. Opler (United States Department 
of Interior) and Mr. William D. Field (Curator in Entomology, United 
States National Museum of Natural History, Smithsonian Institution ) 
for reading this manuscript. Thanks are also due Dr. C. A. Triplehorn 
(Ohio State University) for the loan of Nebraska specimens. 


LITERATURE CITED 


Barnes, W. & F. H. Benyamin. 1926. Check list of the diurnal Lepidoptera of 
boreal America. Bull. So. Calif. Acad. Sci. 25(1): 3-27. 

Barnes, W. & J. H. McDuNnNoucH. 1917. Check list of the Lepidoptera of boreal 
America. Herald Press, Decatur, Illinois. vii + 392 p. 

BEUTENMULLER, W. 1892. List of types of Lepidoptera in the Edwards collection 
of insects. Bull. Amer. Mus. Nat. Hist. X( XIII): 235-248. 

. 1898. Revision of the species of Euchloe inhabiting America, north of 
Mexico. Bull. Amer. Mus. Nat. Hist. X( XIII): 235-248. 

Brown, F. M., D. Err, & B. Rorcer. 1957. Colorado butterflies. Denver Museum 
of Natural History, Denver. 368 p. 

Cary, M. 1901. Notes on the butterflies of Sioux County, Nebaska. Can. Ent. 
33: 305-311. 

DerouiartT, G. 1956. An annotated list of southeastern Wyoming Rhopalocera. 
Lepid. News 10: 91-101. 

pos Passos, C. 1964. A synonymic list of the Nearctic Rhopalocera. Memoirs 
Bepids Soc., N. 1, 145 p. 

Dyar, H. G. “1902” (1903). A list of North American Lepidoptera and key to 
the literature of this order of insects. Washington, U. S. Gov. Printing Office. 
T2ap. 

Epwarps, H. 1881. On some apparently new forms of diurnal Lepidoptera. Papilio 
1(4): 50-55. ; 

Ferris, C. D. 1971. An annotated checklist of the Rhopalocera (butterflies) of 
Wyoming. Agric. Exp. Stat., Univ. of Wyoming; Sci. Monogr. 23, 75 p. 

Jounson, K. 1972 (1973). The butterflies of Nebraska. J. Res. Lepid. 11: 1-64. 

—. 1976. Post-Pleistocene environments and montane butterfly relicts on 

the western Great Plains. J. Res. Lepid. (in press). 

Kiots, A. B. 1930. Diurnal Lepidoptera from Wyoming and Colorado. Bull. 
Brooklyn Ent. Soc. 25: 147-170. 

LeussterR, R. A. 1938. An annotated list of the butterflies of Nebraska with the 
description of a new species (Lepid.: Rhopalocera). Ent. News 49: 76-80. 
OpteR, P. A. 1968. Studies on Nearctic Euchloe. Part 5. Distribution. J. Res. 

Lepid. 7: 65-86. 

1974. Studies on Nearctic Euchloe. Part 7. Comparative life histories, 
hosts, and the morphology of immature stages. J. Res. Lepid. 13: 1-20. 

PuckerINnG, D. L. & R. L. Posr. 1960. Butterflies of North Dakota. Dept. Agric. 
Ent., North Dakota Agric. College Publ. No. 1, 32 p. 

REMINGTON, C. L. 1952. The biology of Nearctic Lepidoptera. I. Foodplants and 
life-histories of Colorado Papilionoidea. Psyche 59: 61-70. 

SHrELDs, O., J. F. Emmet, & D. E. Breeptove. 1969 (1970). Butterfly larval 
foodplant records and a procedure for reporting foodplants. J. Res. Lepid. 
8: 21-36. 

Verity, R. “1905-1911.” Rhopalocera Palaearctica, Papilionidae et Pieridae. (by 
author), Florence, Italy. 359 p. 


260 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


PHOTOPERIODIC REGULATION OF SEASONAL POLYPHENISM 
IN PHYCIODES THAROS (NYMPHALIDAE) 


CHARLES G. OLIVER! 
R. D. 1, Box 78, Scottdale, Pennsylvania 15683 


The useful but not widely known term “polyphenism” was introduced 
by Mayr (1963) and refers to any sort of intrapopulation phenotypic 
variation that is not directly genetic. It thus includes all environmentally 
regulated variation that is shown in the phenotype. Seasonal polyphenism 
in Lepidoptera is regulated by environmental factors, and, as I have 
discussed in a previous paper (Oliver, 1970), may be manifested either 
as a discontinuous shift in phenotypic appearance or as a continuous 
shift in the range of phenotypic variation along a gradient. In either case 
the environment triggers an initial genetic response in the organism 
which canalizes development toward a “seasonal form.” 

Seasonal polyphenism in multivoltine populations of Phyciodes tharos 
Drury belongs to the discontinuous type. The wings of summer adults 
(form “morpheus” Fabricius, Figs. 1, 2) tend to be less intensely colored 
than those of spring and fall adults (form “marcia” Edwards, Figs. 3, 4). 
This tendency is more pronounced on the ventral sides of the wings and 
is most extreme in late winter individuals from the southern part of the 
range (e.g., Cedar Key, Florida). Table 1 gives a comparison of the 
phenotypic appearances of the two forms. A great range in color varia- 
tion is shown in any sample at any location at any time of year. The 
adaptive significance of a basically similar phenomenon in Colias ( Pieri- 
dae) has been discussed by Watt (1968). 

The chief factors responsible for the regulation of seasonal poly- 
phenism in Lepidoptera are photoperiod (Pease, 1962; Shapiro, 1968) 
or temperature (McLeod, 1968) alone or a synergism between the two 
(Ae, 1957). Klots (1951) implies that it is exposure to cold in the pupal 
stage that induces the “marcia” form, but no experimental evidence is 
presented. The experiments described here were designed to test the 
response of P. tharos to different photoperiod and temperature regimes 
during the larval and pupal stages. 


PROCEDURE AND MATERIALS 
Laboratory breeding stock was derived from three wild-inseminated 
females. One female was collected 4 mi. E Cedar Key, Levy Co., Florida, 


es ig Groh Assistant Professor, Department of Biology, West Virginia University, Morgantown, 
es irginia, 


VoLuME 30, NUMBER 4 261 


TaBLE 1. Comparison of phenotypic appearance of most contrasting specimens 
of seasonal forms “marcia” and “morpheus” of Phyciodes tharos. 


“marcia” “morpheus” 
Dorsal 

1. Submarginal spots clear, Spots obsolescent, dark gray 
contrastingly pale. if present. 

2. Marginal and submarginal black Black lines fused into a band. 
lines light and separated. 

3. Male forewing postmedian black Dark markings obsolescent, 
markings well expressed. making an cpen light area. 


Ventral (Hindwing ) 


Males 
1. Brown markings heavy, smudgy. Brown markings thin, crisp. 
2. Silver crescent spot Crescent covered by submarginal 
bright, blurry. dark patch. 
Females 
1. Brown markings heavy, Markings forming a thin, crisp 
very smudgy, filling areas brown reticulation. 
between light spot bands. 
2. Basal, median, submarginal light Crescent spot only silver-white 
bands suffused with silver-white. present, light bands straw yellow. 


29 March 1969, and the others 4 mi. N Emporia, Sussex Co., Virginia, 3 
May 1969. Populations from these areas appear similar in phenotypic 
appearance and are indistinguishable in the laboratory. Potted Aster 
ericoides L. served as the oviposition site and larval foodplant. The 
newly-hatched larvae in each brood were divided into a long day, short 
night group (15hL, 9hD) and a short day, long night group (10hL, 
14hD). Both groups were given a 25°C day and a 22°C night. When 
the short day group entered the third instar, it was divided into two 
lots, one of which was kept on the same temperature regime, and the 
other of which was given a 25°C day and a 5°C night. These regimes 
were maintained through the pupal stage and until adult emergence. 
Cultures were maintained in climate-controlled growth chambers with 
a temperature fluctuation of less than 2°C during each temperature 
period. Artificial lighting was provided with fluorescent tubes of the 
“sunlight” type. Chilling to 5°C was carried out in an unlighted domestic 
refrigerator. 


RESULTS 


The adults from the long day, short night group (N = 21) were of 
the “morpheus” form (Figs. 1, 2), whereas those from the short day, 
long night group (N = 46) were all of the “marcia” form (Figs. 3, 4). 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


i) 
ep) 
bo 


Figs. 1-4. Phyciodes tharos: 1 & 2, form “morpheus” reared on photoperiod re- 
gime of 15hL, S9hD; 3 & 4, “marcia” reared on 10hL, 14hD. 


No phenotypic differences were observed in the responses of the two 
population samples. Differences in temperature regime in the short 
day group had no apparent effect on adult phenotype. These labora- 
tory-reared broods of P. tharos showed great phenotypic homogeneity 
compared to wild-collected samples. 


DISCUSSION 


Seasonal polyphenism in Phyciodes tharos is controlled by photo- 
period. The fall and spring “marcia” form is induced by the compara- 
tively short days and long nights of late summer and early spring. Fall 
“marcia” have apparently developed without larval diapause, whereas 
spring “marcia” have undergone diapause. “Morpheus” is due to the long 
days and short nights of early and mid summer. 

Phenotypic variation in wild seasonal samples of P. tharos may be 
attributable to differences in larval photoperiod exposure as well as 
to individual genetic variation. The intensification of the “marcia” ap- 
pearance in southern U.S.A. samples (where the growing season ends 
and begins with shorter days and longer nights than in the northern 
U.S.A.) indicates that the effect is a graded one in nature. 


VOLUME 30, NUMBER 4 263 


There was no incidence of diapause among the P. tharos larvae on 
either photoperiod regime. Regulation of diapause in this species has 
not been investigated, but it may be influenced by larval temperature 
exposure during the first two instars. In any case diapause and poly- 
phenism seem to have no direct link in this species. 


SUMMARY 


Photoperiod regulates expression of the seasonal forms of Phyciodes 
tharos. A larval and pupal photoperiod regime similar to that of early 
summer produced adults all of the “morpheus” form, whereas a regime 
similar to that of late summer produced only “marcia.” Larval and 
pupal temperature exposure had no effect on adult phenotype. 


LITERATURE CITED 


Ag, S. A. 1957. Effects of photoperiod on Colias eurytheme (Lepidoptera, Pieri- 
dae). Lepid. News 11: 207-214. 

Kxorts, A. B. 1951. A field guide to the butterflies. Houghton Mifflin Co., Bos- 
ton, Mass. 349 p. 

McLeop, L. 1968. Controlled environment experiments with Precis octavia Cram. 
(Nymphalidae). J. Res. Lep. 7: 1-18. 

Mayr, E. 1963. Animal species and evolution. Harvard U. Press, Cambridge, 
Mass. 797 p. 

OniveR, C. G. 1970. The environmental regulation of seasonal dimorphism in 
Pieris napi oleracea (Pieridae). J. Lepid. Soc. 24: 77-81. 

Pease, R. W. 1962. Factors causing seasonal forms in Ascia monuste (Lepidop- 
tera). Science 137: 987-988. 

SHApmRO, A. M. 1968. Photoperiodic induction of vernal phenotype in Pieris 
protodice Boisduval and LeConte (Lepidoptera: Pieridae). Wasmann J. Biol. 
26: 137-149. 

Watt, W. B. 1968. Adapative significance of pigment polymorphisms in Colias 
butterflies. I. Variation of melanin pigment in relation to thermoregulation. 
Evolution 22: 437-458. 


264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


LARVAL FOODPLANTS AND LIFE HISTORY NOTES FOR 
EIGHT MOTHS FROM TEXAS AND MEXICO! 


Roy O. KENDALL? 
Route 4, Box 104-EB, San Antonio, Texas 78228 


So far as I can determine, nothing has been published on the life his- 
tory of the species in the present paper except for Madoryx oiclus 
(Cramer). Apparently this species uses the same larval foodplant in 
Mexico as it does in Venezuela (Lichy, 1944). Blanchard (1973) re- 
corded and illustrated 3 of these species (Syntomeida melanthus 
(Cramer), Rhescipha servia Cramer, and Scordylia atalanta Guenée) 
as rare or possibly new to Texas and the United States. Blanchard (1968) 
also described and illustrated Grotella margueritaria from Texas. Mun- 
roe (1972, 1973) described and illustrated Odontivalvia radialis from 
Texas. Guenée (1857) described the family Hedylidae and placed 
therein Venodes napiaria (Brazil), Phellinodes satellitiata (Brazil), and 
Hedyle heliconiaria (French Guiana) which he described ibidem. It is 
believed the latter species is here recorded from Mexico for the first time. 


Sphingidae 

This report confirms the infrequent occurrence of Sphinx lugens Walker within 
the United States and gives its local larval foodplant, Forestiera pubescens. Life 
history notes, including a possible larval foodplant, Tabebuia pentaphylla, are given 
also for the cocoon spinning Mexican Madoryx oiclus (Cramer). An adult 2, pupa, 
and 2 cocoons are illustrated. 

Sphinx lugens Walker 1856. Hodges (1971) indicated that the immature stages 
of this species were unknown, and that Strecker (1876:115) incorrectly cited Salvia 
as a host when he accepted eremitoides as a junior synonym of lugens. He states 
further: “Although I have seen no authentic specimens of lugens from the south- 
western United States, it is to be expected in southern Arizona and New Mexico or 
Texas. Some of the earlier literature citations to this species probably refer to other, 
closely related species.” 

On 8 August 71 in my lab garden at San Antonio, Texas I found 1 larva feeding 
on the foliage of a small (60 cm), cutover plant of Forestiera pubescens Nutt., 
OLEACEAE. This larva continued to eat until 12 August when it entered soil which 
had been provided, and in which it pupated unobserved; a ? emerged 28 August 
71. This remains the only example found, although I have examined the foodplant 
frequently for immatures since then. Other species of Forestiera, including F. pu- 
bescens, are found in Mexico, and perhaps may be acceptable to S. lugens through- 
out its range. 

Madoryx oiclus (Cramer) [1780]. Hodges (1971) in treating the genus Madoryx, 
stated: “The one known larva [imagine not given] is peculiar for a sphingid inas- 


' Contribution No. 341. Bureau of Entomology, Division of Plant Industry, Florida Department 
of ence ene Consumer Services, Gainesville 32602. 

“Research Associate, Florida State Collection of Arthronods, Divisio f Pl i 
Department of Agriculture and Consumer Services. , ; mami 


VOLUME 30, NUMBER 4 265 


& 


ee Me Fe cae 


Figs. 1-4. Madoryx oiclus: 1, 9, dorsal view (expanse 93 mm); 2, pupa, ventral 
view (length 57 mm); 3, cocoon spun on tree trunk (length 85 mm); 4, cocoon 
spun on whitewashed wall. 


much as it resembles that of a species of Catocala in the Noctuidae.... The pupa 
is dark and glossy, banded with pale orange at the base of some abdominal seg- 
ments, and the base of the tongue projects forward somewhat.” Lichy (1944) re- 
ported Tecoma pentaphylla (= Tabebuia pentaphylla (L.) Hemsl.), BriGNONIACEAE, 
as the larval foodplant in Venezuela. 


966 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


On 15 November 74 at Los Arcos Courts, Ciudad Mante, Tamaulipas, Mexico, I 
found 1 spun-up larva; it had selected a spot, ca. 60 cm above ground, on the white- 
washed outside wall of the court where we stayed. This larva pupated 16 November 
and a @ emerged 16 December 74 (Fig. 1). Also, on the same wall about 2 m away 
from the first, another cocoon was found near the eve, but this one had emerged. 
A third cocoon, containing a live pupa (Fig. 2) was found 5 December 74 about 
10 m from the first 2 and about 2 m above ground on a tree trunk, Tabebuia penta- 
phylla. This tree is planted extensively in Mexico as an ornamental and known 
locally as “Palo de rosa.” Still later, Mrs. Kendall found a fourth empty cocoon spun 
on the side of a concrete stepping block near the ground. In each instance, the 
cocoons were found on or within 2 m of T. pentaphylla, undoubtedly its larval 
foodplant. 

Most interesting was the larva’s ability to camouflage its cocoon to match the 
background on which it was spun. Note the darkened color and bits of tree bark 
in the silk of the cocoon spun on the tree (Fig. 3); compare this cocoon with Fig. 
4, the one spun on the outside wall, and note its lighter color and bits of whitewash 
in the silk. The darkened areas of the latter are caused by the empty pupal case 
inside. I had overlooked 3 of these cocoons several times earlier because the sun- 
light had not struck them at the proper angle. 


Ctenuchidae 


Syntomeida melanthus (Cramer) 1779. Near Ciudad Mante, Tamaulipas, Mexico 
single larvae were collected crawling on the ground as if in search of food on 11 
January 74, 26 January 74, and 12 February 74. These larvae proved to be para- 
sitized by dipterons; both host and parasites were preserved. Again on 21 November 
74, Mrs. Kendall found a cluster (ca. 40) of first instar larvae feeding on the foliage 
of Ipomoea populina House, CONVOLVULACEAE. This plant was fairly well defoliated, 
and later searching proved other plants to be in the same state of dormancy. Be- 
cause of the scarcity of food most larvae died of malnutrition. Seven larvae spun 
cocoons between 21 December 74 and 4 January 75, but only 3 pupated; adults 
emerged 16 January 75 ( 2 ), 17 January ( ¢ ), and 18 January ( @ ). 

In general appearance the larva of this species resembles that of an Halisidota. 
The last instar larva is clothed with short gray pile; there is a lateral row of black 
tufts; 3 mid-dorsal segments have paired white tufts lightly overlaid with gray hairs; 
the first 2 and last segment have paired long black hair pencils; head black. Larval 
hairs form the basis of the cocoon which is generally formed on the long axis of a 
twig. 


Noctuidae 


Grotella margueritaria Blanchard 1968. While on a joint field trip with André 
and May Elise Blanchard, Mrs. Kendall and I were fortunate in discovering 2 larval 
foodplants for this recently described species. On 17 September 71, ca. 3 km N 
of Study Butte, Brewster Co., Texas, we collected a few larvae resting on the stems 
of Anulocaulis leisolenus (Torr.) Standley, NycraGINACEAE. These larvae were lost 
because it was not realized at the time that they eat blossom buds, not foliage. At the 
type locality in Big Bend National Park, 12 more larvae were collected 21 Septem- 
ber 71, feeding on the blossom buds of Anulocaulis eriosolenus Standley. One para- 
sitized larva produced a dipteron 8 October 71. Seven other larvae burrowed in dirt 
on 21 and 22 September 71 where they pupated in earthen chambers. Later, when 
the dirt was screened, 2 pupae were exposed, and they later became parasitized; adult 
parasites were found 20 August 72. Other pupae, in their sealed earthen chambers, 
produced adults: 19 October 71 ( g ), 22 October (3), 24 October ( g ), 29' Octo- 
ber (2), 24 August 73 (2 ), and 6 September 73 (3). Two larvae and 1 deformed 
pupa were preserved. It was interesting that 2 remained in pupal diapause for al- 


VOLUME 30, NUMBER 4 267 


most 2 years. This would indicate that the species is well adapted to Chihuahuan 
Desert conditions. 

Because there is some question as to the proper systematic placement of Grotella, 
a completely illustrated and described life history for G. margueritaria will be pub- 
lished as soon as it can be reared from eggs of a known female. 

Rescipha servia Cramer 1782. On 13 November 71 at Santa Ana National Wild- 
life Refuge, Hidalgo Co., Texas, I collected 1 pupa in a leaf nest on Rivina humilis 
L.; a ¢ emerged later the same day. After finding this pupa I had thought R. humilis 
might be the larval foodplant. On 18 June 72, however, I found a larva at my lab 
in San Antonio, Bexar Co. feeding on the foliage of Diospyros texana Scheele, 
EBENACEAE. A few days later a second larva was collected feeding on the same plant. 
The first larva spun a cocoon in the leaves of the foodplant 1 July, and a ¢ emerged 
15 July 72. The second larva was preserved after passing through 7 instars and at 
the time it started spinning its cocoon. It is interesting to note that the larva of this 
species is catocala-like, both in appearance and habits; when not feeding it rests 
flat along the foodplant branches. 

Adults have been taken at my lab doorlight: 25 October 71 ( ¢), 17 November 
Maen eionune 7216.19), i july 72 (3S). 2 July-72- (23), 8 July 72 (4), 
10 July 72 (2), 7 August 72 (¢), 7 August 73 (@), 30 October 73 (6) and 1 
July 75 (2). Based on these dates at least 2 broods are indicated. 


Geometridae 


Scordylia atalanta Guenée 1857. On 28 January 75 near Ciudad Mante, Tamau- 
lipas, Mexico, about midafternoon I observed a @ ovipositing on the foliage of Ser- 
jania racemosa Schumacher, SAPINDACEAE. In the field lab this @ deposited 14 
more eggs the same day, but it became quiescent at sunset. On the following day, 
38 more eggs were deposited during daylight hours; that evening, the @ was killed 
and papered. Several eggs were preserved, and the remaining ones started hatching 
1 February 75, ca. 2030 hrs. The young larvae were offered foliage of Urvillea 
ulmacea H.B.K., SAPINDACEAE which they readily ate. On 14 February the larvae 
were offered swelling blossom and leaf buds of Ungnadia speciosa Endl., also Sap- 
INDACEAE, which they ate and on which they matured. Fifteen larvae pupated be- 
tween 21 and 28 February 75; adults emerged (746, 52) from 3-11 March 75. 
Twelve eggs, 12 larvae and 3 pupae were preserved. 

At the same time the gravid @ was collected, 2 larvae were found on a bit of 
the larval foodplant, S. racemosa, gathered at the site and placed with the captive 
female. These 2 larvae pupated 14 February 75, and 2 @ emerged 26 and 27 Feb- 
MITA Mos 


Hedylidae 


Hedyle heliconiaria Guenée 1857. Recently I had the good fortune to rear this 
most interesting species (Figs. 5, 6). Although a few of the diurnal adults were 
collected earlier, I never associated them with the larvae (Figs. 7-11) collected 
later. In fact, I thought the larvae might represent a satyr species, but when the 
first larva pupated I was sure it was a pierid. The first adult emerged as a complete 
surprise. The pupa (Figs. 12-15) is secured by girdle and cremaster, not unlike 
a pierid. Several egg shells, presumably of this species, were found deposited singly 
on top of leaves. Larvae rest on top of the leaves, oriented along the mid-vein, their 
color and configuration providing excellent camouflage, at least to the human eye. 
Larval feeding consists of eating, at random, a series of small holes in the leaf on 
which it rests. 

On 6 February 74 at Rancho Pico de Oro, near the Rio Sabinis, Tamaulipas, Mexico, 
Mrs. Kendall and I collected 13 larvae feeding on the foliage of Buettneria aculiata 


968 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Migs. 5-6. Hedyle heliconiaria: 5, 2, ex larva 11 March 1974, Ranch Pico de Oro, 
Tamps., Mexico (expanse 32 mm); 6, wing venation, same specimen. 


VOLUME 30, NUMBER 4 269 


Figs. 7-11. Hedyle heliconiaria: 7, last instar larva, dorsolateral view; 8, ventro- 
lateral view; 9, anal end and process; 10, larval head, laterodorsal view; 11, larval 
head, lateroventral view. 


Jacq., STERCULIACEAE. Six of these larvae pupated between 9 and 20 February 74; 
adults emerged 19 February ( ¢ ), 25 February (2), 2 March ( ¢), and 11 March 
1974 (@). Seven larvae and 2 pupae, most of which were parasitized, were pre- 
served. Again on 10 November 74, near Ciudad Mante, Tamaulipas, about 12 more 
(Ist and 3rd instar) larvae were collected on B. aculiata; 2 of these were preserved 
the same day. On 17 November it was discovered that all remaining larvae except 
2 had been eaten by 3 predatory fly larvae, predators on the foliage which had gone 
unnoticed at the time of collecting. The 2 remaining larvae pupated 26 November 
74; 1 was preserved and the other proved to be parasitized. 


Pyralidae 


Odontivalvia radialis (Munroe) 1972. On 15 September 71 at Dagger Flat, Big 
Bend National Park, Brewster Co., Texas, while collecting Thessalia chinatiensis 
(Tinkham ) (Nymphalidae) larvae on Leucophyllum minus Gray, SCROPHULARIACEAE, 
I found 3 micro larvae in silken tunnels covered with frass and attached to the 
branches of this plant. These 3 larvae were taken to the lab in San Antonio where 
little activity was observed in the rearing container (glass jar with screened lid). 
The small amount of foodplant brought to the lab soon dried, and because no ac- 
tivity was observed it was not until 8 June 73 that I decided to clean the jar. At 
this time (some 21 mos. later) I found 1 larva had pupated, 1 was dead, and the 
third was still in diapause. The pupa and diapausing larva were placed on a moist 
sponge. The following day a ¢ emerged. On 15 June 73 the larva had not pupated, 


70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 12-15. Hedyle heliconiaria: 12, pupa, frontal view; 13, dorsal view; 14, 
lateral view; 15, ventral view. 


but upon my returning from a field trip 6 July, it had; another ¢ emerged 7 July 
73. This is another species well adapted to Chihuahuan Desert conditions. 


ACKNOWLEDGMENTS 


I wish to thank the United States Department of the Interior, Santa 
Ana National Wildlife Refuge, especially Carrell Ryan and Wayne A. 
Shifflett, and Big Bend National Park, especially William M. Rabenstein 
for their sincere interest and cooperation in my continuing study of 
Texas Lepidoptera. I am also indebted to the United States Department 
of Agriculture, especially Jack E. Lipes for providing the necessary im- 
portation permits for Mexican material. To E. L. Todd and D. C. Fer- 
guson, Systematic Laboratory, United States Department of Agriculture, 
at the National Museum of Natural History, and André Blanchard of 
Houston, Texas, many thanks for comparing some of my specimens with 
those in the National collection. I am especially indebted to André for 
providing the morphological photographs and drawing used in this 
article. To William H. Sieker, for identifying one of these species, and 
to Carlos R. Beutelspacher for certain reference citations I am indeed 
grateful. To Sr. & Sra. Fernando Reyes Bugarin, Gte., Los Arcos Courts, 
for giving Mrs. Kendall and me full access to their beautiful gardens, and 
for permitting us to remove foliage from their prized plants to feed cater- 


VoLUME 30, NUMBER 4 271 


pillars, and to our equally dear friends, Sr. & Sra. Carlos Gonzales, we 
are most grateful for their warm hospitality and permission to do re- 
search at Rancho Pico de Oro. 


LITERATURE CITED 


BLANCHARD, A. 1968. New moths from Texas (Noctuidae, Tortricidae). J. Lepid. 
Soe. 22: 133-145. 

1973. Record and illustration of some interesting moths flying in Texas 
(Sphingidae, Ctenchidae, Noctuidae, Notodontidae, Geometridae, Pyralidae, 
Cossidae). J. Lepid. Soc. 27: 103-109. 

CorrELL, D. S. & M. C. Jounston. 1970. Manual of the vascular plants of Texas. 
Texas Research Foundation, Renner, Texas. 1881 p. 

GuENEE, M. A. 1857. Histoire naturelle des insectes. Spécies général des lépidop- 
terés. 10: 522. 

Hopces, R. W. in Dominick, R. B., et al. 1971. The moths of America north of 
Mexico, Fasc. 21, Sphingoidea. Classey, London. xii + 158 p., 14 color plates. 

Licuy, R. 1944. Documents pour servir a l’étude des Sphingidae du Venezuela 
(Lepid., Heter.) (6e. note). Sur un cas d’adaptation a un nouveau regime 
alimentaire chez Madoryx oiclus Cr. Etude Biologique partielle. Bol. Ent. Venez. 
3: 195-202, 2 figs. 

Munro, E. in Dominick, R. B., et al. 1972, 1973. The moths of America north 
of Mexico, Fasc. 13.1B, Pyraloidea (1972); Fasc. 13.1C, Pyraloidea (1973). 

1973. A new genus for Noctueliopsis radialis (Lepidoptera: Pyralidae: 
Odontiinae). Can. Ent. 105: 1361-1362. 

STANDLEY, P. C. 1920-1926. Trees and shrubs of Mexico. Contributions, U.S. 
Nat. Herbarium 23: 1-1721. Gov't. Printing Office, Washington, D.C. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


bo 
~] 
bo 


A KEY TO THE LAST INSTAR LARVAE 
OF WEST COAST SATURNIIDAE 


PauLt M. TuskEs 


Department of Environmental Toxicology, 
University of California, Davis, California 95616 


This following key will facilitate the identification of 16 species of 
saturniid larvae found in California, Oregon, and Washington. The 
identification of either live or preserved larvae should be possible since 
morphological characters have been stressed. Observation of many of 
the characters requires a 10 power hand lens. Secondary emphasis has 
been placed on color, since those colors derived from plant compounds, 
such as green and yellow, fade quickly to white or cream in preservatives. 
Pigments which are black or brown, however, remain rather constant. 
The first character in each couplet is assumed to be the most important. 

In addition to host plant and distributional information for each spe- 
cies, a reference to a paper containing a larval description is included. 
Other references which would aid in the identification of saturniid lar- 
vae include Peterson (1962) and Ferguson (1971, 1972). Ferguson in- 
cludes keys for the identification of many eastern species, with the 
Citheroniinae receiving the most complete treatment. Peterson’s work 
provides some information on preservation techniques, as well as a few 
illustrations of the more common eastern species. 

All material examined during the construction of this key was from 
California populations. The author would appreciate receiving any 
larva which does not key out correctly. It is unfortunate that at this 
time a complete key to the western species could not be published, but 
preserved larvae of a number of species from Arizona and New Mexico 
are lacking. 


KEY TO THE LAST INSTAR LARVAE OF WEST COAST SATURNIIDAE 


I. Dorsal thoracic scoli unarmed, consisting of 2 pairs of enlarged, caudally 
recurved spikelike projections on both meso- and metathorax (Fig. 1a); 
spikelike median dorsal scolus on abdominal segment VIII; ground color 
pad soomel (Orhdavergovaningeve jae ae a Sphingicampa hubbardi (Dyar) 

—Dorsal thoracic scoli branched or armed with spines (Figs. 1b, c, d, e, £); 
median dorsal scolus on abdominal segment VIII present or absent; ground 


color variable (ou 2 
2. Dorsal scoli not lorealiak but rounded, cylindrical, or bulblike with spines 
(Figs. 1b, c, d); ground color green or yellow-orange (Saturniinae) _..__. 3 


—Dorsal scoli branched (Figs. le, f); ground color black, gray, brown, or 
yellow (Hemileucinae ) 


VoLUME 30, NUMBER 4 2713 


10. 


Mee 


13; 


14. 


Dorsal thoracic scoli bulblike or cylindrical and 114-3 times longer than 
wide; spines on dorsal scoli equal to or less than width of scoli ( Figs. 1b, c); 
ventral scoli of thoracic segments without spines or with 2 spines shorter 
Hianlengthe of scoli CHijalophora) 2.228 oS he) ee 4 
—Dorsal thoracic scoli rounded, length approximately equal to width; spines 

on dorsal scoli 244-8 times longer than width of scoli (Fig. 1d); ventral 
scoli of thoracic segments with 2 or more spines 2-3 times longer than 
fener ot scoli (Antheraea & Saturnia), = 2 ee 5 
Dorsal meso- and metathoracic scoli cylindrical with complete black band 

at midpoint (Fig. lc); caudal scoli length usually less than width 
NE SAS DD I te eet thre Hyalophora euryalus (Boisduval ) 
—Dorsal meso- and metathoracic scoli bulblike; band at midpoint incomplete 

on two or more scoli (Fig. 1b); caudal scoli length greater than width ___ 
oases se Ue Le Sn Te Oe EE H. gloveri (Strecker ) 
Abdominal segment VIII with median dorsal scolus; ventral surface green 

voce eect gr OO MEER aes Antheraea polyphemus (Cramer ) 
—Abdominal segment VIII not bearing median dorsal scolus; ventral surface 


oEtOMaarkabrowmn (SUUMNIG)) 2a. 2k steer Ri bah OS 6 

Two or fewer dark setae on dorsal portion of any proleg; setae not extend- 

Cm TORIC CHOY OLE for eked Ne LR I cea ci eee Sn ee ac 7 
—Four or more dark setae on dorsal portion of any proleg; setae may extend 

beyond tip of proleg (Fig. 2a) Saturnia mendocino Behrens 


Lateral bands on abdominal segments extending ventrally from lateral scoli 
to sublateral scoli and touching the posterior edge of spiracle; some setae 


on prolegs extending from chalazae (Fig. 2b) __. Saturnia albofasciata (Johnson ) 
—Lateral bands absent; setae on prolegs extending from pinaculae rather than 
ohn eMmCLIP WC) resi ns bs Sows ski, S. walterorum Hogue & Johnson 


Dorsal and lateral scoli with short barrel-like branches, 2 or less times 
longer than wide (Fig. le); lateral ocelli in heavily sclerotized semicircular 
2G. te eee Coloradia pandora lindseyi Barnes & Benjamin 
—Dorsal scoli in rosettes, lateral scoli with branches 3 or more times longer 
than wide; sclerotized area surounding ocelli similar to rest of head capsule 


UB leCzEAT0) 10.) aes Sn Ee ees ARMED Re hs a TN 2 el Ae 9 
Secondary setae not arising from circular white or yellow spots 10 
—Secondary setae arising from circular white or yellow spots giving skin a 
Tene teumEADpCATANCe us. NDE ee eS ewe dS 1 
Clypeus with 6-10 setae; ventral abdominal surface black to brown; prolegs 
[aleve ee Sl a fo a lt ae eae Hemileuca nuttalli (Strecker ) 
—Clypeus with 4 setae; ventral abdominal surface white, red, or black; pro- 
“em teal Opp lollevcl = GN a2 Os NES PE ea eet eae ee eee Cet Bee iL 


Ventral intersegmental area white or cream colored; prolegs black; body 
with 3 lateral white bands; median dorsal line incomplete; microscopic 
cream colored dots ventral to spiracle Hemileuca hera (Harris ) 
—Ventral intersegmental area red to light brown; prolegs red; 1-3 complete 
lateral bands (may be absent in coastal populations); median dorsal line 
absent; no microscopic dots ventral to spiracle _____. H. eglanterina ( Boisduval ) 
Clypeus with 4 setae; white band covering 30% or less of clypeus —-.----_-___- 13 
—Clypeus with 6-8 setae; white band covering 40% or more of clypeus 14 
Branches of sublateral and ventral scoli white; dorsal scoli of rosette type 
only on abdominal segments I-VI __. Hemileuca electra Wright 
—Branches of sublateral and ventral scoli with basal %—% dark, tip dark 
brown; dorsal scoli with branching setae extending from rosettes on all 
OTM MA ls SE CMMETIES) eee eee al eA er ite, eel nme a) co H. burnsi Watson 
Dorsal rosette setae white-yellow with black tip; ground color yellow 

amr a ert 00 tien TUT ANE, i rtp Sel TN ee bn aver ee Hemileuca nevadensis Stretch 


974 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


—Dorsal rosette setae dark brown at base, distal 144-14 of setae dark, mid- 
portion white; ground color gray to brown Ee 15 

15. Secondary setae on ventral intersegmental area hyline brown; prolegs red; 
proximal %-% of rosette setae brown __ . Hemileuca neumoegeni (Edwards ) 

—Secondary setae on ventral intersegmental area brown; prolegs brown; 
proximal % of rosette setae black H. juno Packard 


Distributional and Host Plant Information 


Sphingicampa hubbardi. Distribution: California (new state record) 1 6, 1 Q, 
Bonanza King Mine, Providence Mts., San Bernardino Co., Cal., VIII-20-37; same 
locality, 1 ¢, VIII-23-37; 1 ¢, Wheaton Springs, at general store, Mescal Range, 
San Bernardino Co., Cal., elev. 4000’, VIII-30-41, C. Henne collector. All 4 speci- 
mens are in the collection of Mr. Henne. The specimens from the Providence Mts. 
were taken by the watchman, name unknown. Habitat: high desert. Host: pos- 
sibly Acacia greggii. Larval description: Comstock (1947). 

Hyalophora euryalus. Distribution: general. Habitat: chaparral, oak woodland, 
and pine forest. Hosts: Ceanothus, Salix, Rhus, Ribes, Schinus, Arctostaphylos, 
Arbutus, Prunus, and many minor hosts including Quercus. Larval description: 
Packard (1914). 

Hyalophora gloveri. Distribution: California, east slope of Sierra Nevada, from 
Inyo Co. to Alpine Co. and probably farther north. No available records for Wash- 
ington or Oregon, but it should occur in the extreme eastern portion of each state. 
Habitat: Great Basin and pine forest. Hosts: Salix, Purshia tridentata, Prunus vir- 
giniana, and Rosa. This species frequently hybridizes with H. euryalus where they 
are sympatric in California. Larval description: Packard (1914). 

Antheraea polyphemus. Distribution: general. Habitat: oak woodland, riparian, 
and residential. Hosts: Quercus, Salix, and Betula. Larval description: Packard 
(1914). 

Saturnia albofasciata. Distribution: California only, known from Lake and El 
Dorado counties south ta Los Angeles and San Bernardino counties. Habitat: mixed 
chaparral from 1300-7000’. Host: Ceanothus cuneatus and Cercocarpus betuloides. 
Larval description: Hogue et al. (1965). 

Saturnia mendocino. Distribution: California, Shasta Co. south to Santa Cruz 
Co., also western slopes of the Sierra Nevada south to at least Mariposa Co. Oregon, 
one record (Ashland, Jackson Co., IV-14-1946, Coll. Martin). Habitat: chaparral, 
pine, or mixed oak woodland. Hosts: Arctostaphylos and Arbutus. Larval descrip- 
tion: Comstock (1960). 

Saturnia walterorum. Distribution: California only, San Luis Obispo Co. south 
to San Diego Co, Habitat: chaparral to pine forest, sea level to 6000’. Hosts: 
Rhus laurina, R. integrifolia, and Arctostaphylos. Larval description: Sala & Hogue 
(1958). 

Coloradia pandora lindseyi. Distribution: southern Oregon and California; spotty 
over much of its range. Habitat: pine forest. Hosts: Pinus ponderosa and P. jef- 
freyi. Larval description: Patterson (1929). 

Hemileuca hera. Distribution: California, eastern slopes of Sierra Nevada. Ore- 
gon and Washington, eastern half of each state, especially the Columbia River basin. 
Populations are frequently very localized and scattered. Host: Artemisia tridentata. 
Larval description: McFarland (1964). 

Hemileuca eglanterina. Distribution: general. Habitat: varied, riparian in dry 
areas, to moist oak-pine forests. Hosts: Salix, Ceanothus, Purshia, Prunus, and 
Symphoricarpos. Two subspecies occur in California and Oregon: H. eglanterina 
shastaensis (Grote) is known from Shasta, Plumas, Lassen and Siskiyou counties 
in California, also Klamath and Jackson counties in Oregon. H. eglanterina annulata 


VOLUME 30, NUMBER 4 Dis 


Figs. la—f. Dorsal metathoracic scoli of: la, Sphingicampa hubbardi; 1b, Hyalo- 
phora gloveri; 1c, Hyalophora euryalus; 1d, Saturnia mendocino; le, Coloradia pan- 
dora; 1f, Hemileuca sp. 

Figs. 2a—2c. Prolegs of abdominal segments III and IV: 2a, Saturnia mendocino; 
2b, S. albofasciata. 2c, S. walterorum. 


Ferguson is known from the east slope of the Sierra Nevada Mts. from Inyo to 
Alpine Co. Larval description: Ferguson (1971). 

Hemileuca nuttalli. Distribution: eastern Oregon, Washington, and California, 
similar to that of H. hera. Habitat: sagebrush areas. Hosts: Symphoricarpos and 
Purshia. Larval description: McFarland (1974). 

Hemileuca electra. Distribution: California only, Los Angeles, Riverside, San 
Bernardino, and San Diego counties. This species may occur slightly farther north 
along the coast. Habitat: chaparral. Host: Eriogonum fasciculatum. H. electra 
clio Barnes & McDunnough occurs in desert areas of the above four counties. Larval 
description: Comstock & Dammers (1939). 

Hemileuca nevadensis. Distribution: spotty over much of California and Oregon. 
One sight record from southern Washington. Habitat: usually riparian. Hosts: 
Salix and possibly Populus. Larval description: Comstock & Dammers (1939). 


276 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Hemileuca burnsi. Distribution: California, Los Angeles Co. east, and north 
along the eastern slope of the Sierra Nevada. Habitat: high desert. Hosts: Tetra- 
dymia, occasionally Dalea, or Prunus fasciculata. Larval description: Comstock & 
Dammers (1937). 

Hemileuca juno. Distribution: California, Imperial Co. near Yuma. This species 
may occur in portions of Riverside and San Bernardino counties. Habitat: desert. 
Hosts: Prosopis and possibly Cercidium. Larval description: Comstock & Dammers 

1939 ). 
ee neumoegeni. Distribution: California, Providence Mts., San Bernardino 
Co. Habitat: High desert-chaparral. Hosts: Rhus trilobata and Prunus fasciculata. 
Undescribed. 


ACKNOWLEDGMENTS 


I would like to acknowledge the assistance of Mr. Christopher Henne 
of Pearblossom, California and Mr. Michael Van Buskirk of Tucson, Ari- 
zona, both of whom provided me with field notes and preserved larvae. 
I would also like to thank Julian Donahue of the Los Angeles County 
Museum of Natural History for providing preserved larvae. 


LITERATURE CITED 


Comstock, J. A. 1947. Notes on the early stages of Adelocephala heiligbrodti 
f. hubbardi Dyar. Bull. So. Calif. Acad. Sci. 46: 72-77. 

. 1960. Life history notes on a saturniid and two lasiocampid moths from 
California. Bull. So. Calif. Acad. Sci. 59: 170-181. 

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

. 1939. Studies in the metamorphoses of six California moths. Bull. So. 
Calif. Acad. Sci. 37: 105-128. 

Frercuson, D. C. 1971. The moths of America north of Mexico. Fascicle 20.2a, 
Bombycoidea (in part). Classey, London, p. 1-154. 

1972. The moths of America north of Mexico. Fascicle 20.2b, Bomby- 
coidea (in part). Classey, London. p. 155-275. 

Hocuge, C. L., F. P. Sata, N. McFarvanp, & C. HENNE. 1965. Systematics and 
life history of Saturnia albofasciata in California. J. Res. Lepid. 4: 173-184. 

McFaruanp, N. 1974. Notes on three species of Hemileuca from eastern Oregon 
and California. J. Lepid. Soc. 28: 136-141. 

PackaArD, A. S. 1914. Monograph of the bombycine moths of North America, 
part 3. Mem. Natl. Acad. Sci., 12: p. i-ix, 1-276, 503-516, pls. 1-113. 

PATTERSON, J. E. 1929. The Pandora moth, a periodic pest of western pine forests. 
United States Dept. Agric. Tech. Bull. 137, 19 p. 

PeTERSON, A. 1962. Larvae of insects. Part 1, Lepidoptera and Hymenoptera. 
A. Peterson Pub. Columbus, Ohio. 315 p. 

Sata, F. P. & C. L. Hocur. 1958. Description of the early stages and male imago 
and notes on the life history of Saturnia walterorum. Lepid. News 12: 17-25. 


VoLUME 30, NUMBER 4 DA 


LOW COST VACUUM FREEZE-DRYING 


FRANK R. HEDGES 
11852 Hempstead Highway, Lot J-9, Houston, Texas 77092 


The major cost of most vacuum freeze-drying systems is the vacuum 
pump. By replacing this expensive pump with a water aspirator, the 
cost of vacuum freeze-drying can be reduced by 90% or more. This will 
provide a system at a price the average lepidopterist can afford. 

An excellent treatment of vacuum freeze-drying was done by Dominick 
(1972). He includes many theoretical and practical ideas which help 
in understanding this process. The following paper will deal with my 
modifications of the vacuum freeze-drying process, and complications 
which arise from these modifications. 


Theory 


Although a fine treatment of the theoretical aspects of vacuum freeze- 
drying was given by Dominick (1972), a review of the theory as it ap- 
plies to this modified system is presented here. 

At a fixed temperature and in a closed container, the vapor pressure 
caused by the evaporation of water (or ice) will be constant, regardless 
of the amount of vacuum initially applied (Dalton’s Law of Partial 
Pressures). When a desiccating agent is placed in this container (but 
not in direct contact with the water or ice), the desiccant absorbs some 
of the water vapor. This causes more water (or ice) to evaporate in an 
attempt to maintain the partial pressure which is characteristic for that 
temperature. This means that the major driving force of dehydration 
is independent of the initial vacuum in the container. However, even 
though the driving force is independent of the initial vacuum, a con- 
tainer which is initially evacuated will evaporate a piece of ice faster 
than one with no vacuum. As Dominick explains it, “the more molecules 
of other gases present (air), the more difficult it is for the molecules 
evaporating from the water (or ice) to dissipate; then a state of quasi- 
saturation will be reached near the water (or ice) surface and further 
evaporation will cease.” Actually, the evaporation will only slow down 
rather than cease. This means that vacuum freeze-drying can take place 
at any level of vacuum; however, the lower the pressure the faster the 
process. Table 1 compares the times necessary to dehydrate an average 
butterfly larva at different vacuum levels. 

A water aspirator will attain a vacuum of around 15 mm Hg. My basic 
idea was to use this as a source of vacuum, in spite of the longer drying 


978 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TaBLE 1. Comparative times for dehydrating average butterfly larva. 


.001 mm Hg 15 mm Hg 760 mm Hg" 


2 days 14 days 100 days 
1Data at 760 mm Hg was obtained by Flaschka & Floyd (1969). 


times, in order to reduce the cost of a vacuum source from over $200 to 


less than $15. 


Equipment 


At this point a physical description of an aspirator vacuum freeze- 
drying system is in order. The basic components can be seen in Fig. 1. 
In order to aid in the selection of suitable equipment, each of the four 
components of the system (aspirator, trap, valve, and desiccator jar) 
will be discussed. 

The aspirator can be one of a number of different models. One of the 
main differences in aspirators is the vacuum they will produce with a 
given water pressure. Unfortunately, this data is hard to get from many 
suppliers. For my work I examined four different models, and chose an 
aspirator made of glass, by Kimax. It has a free air flow of 6.5 liters/min. 
with 40 lbs. of water pressure and is rated to pull down a 16 oz. jar to 
16 mm Hg. in a maximum time of 1% minutes. This glass aspirator re- 
quires a hose clamp, a hose, and an adapter for a kitchen faucet. These 
can be purchased from most hardware stores. Other aspirators are easier 
to connect since they come with threads on the aspirator and can be at- 
tached directly to the water faucet. 

The next component is the trap. This insures that if something goes 
wrong (such as a sudden change in the water pressure), water will not 
back up into the desiccator jar. A vacuum trap is excellent, but a home- 
made trap is almost as good and is very inexpensive. A good home-made 
trap consists of a thick-walled jar with a two-hole rubber stopper in- 
serted into it. Two pieces of glass or copper tubing are tightly inserted 
into the holes. One of the pieces of tubing should almost touch the bot- 
tom of the jar, and should be connected to the hose going to the aspirator. 
The other piece of tubing should only be inserted far enough to barely 
clear the rubber stopper, and should be connected to the hose going to 
the desiccator jar. In this way, water backing up will have to nearly fill 
the trap before it can back up into the desiccator, allowing time to shut 
off the valve to the desiccator jar. Do not reverse the order of the hose 
connections to the trap, or its purpose will be defeated. 

The third component is the valve. This item is hard to find at a low 


VOLUME 30, NUMBER 4 279 


HOSE 


FAUCET HOSE 


——— 


|__} HOSE 


1/4” PIPE F 


| JFITTING Ss a 
iB mI 
TUBING me VALVE 
RUBBER 7 
Logg cg t PSTOPPER SSS 
RUBBER 


ASPIRATOR TRAP DESICCATOR JAR 


Fig. 1. Diagram of an aspirator vacuum freeze-drying system. 


cost. The simplest choice is to buy a stopcock, but current costs for stop- 
cocks are around $8-$15 (U.S.) each. Polyethylene one-way valves are 
inexpensive and in theory should work, but I have had numerous ex- 
periments ruined because they failed to operate properly. Several other 
types were tried with some success, but my best results have been ob- 
tained with equipment that can be bought at many hardware stores. 
Most good hardware stores carry various valves to fit 4” pipe. Obtain 
one with two female connections. Also purchase a three inch piece of 
%”’ pipe. Cut this pipe in half, and attach half to each end of the valve. 
Bore a %4” hole through the rubber stopper you are going to use as a 
desiccator lid. As seen in Fig. 1, the pipe (which is attached to the valve ) 
is inserted through the hole in the stopper. The valve, pipe, and stopper 
cost about $2.50. 

The last component is the desiccating jar. This jar should be thick- 
walled and fitted with a one-hole rubber stopper. I use the same type 
of jar as in the trap. These are thick-walled square 8 oz. bottles with a 
wide mouth, have a cap size of 43 mm, and take a #7 rubber stopper. 
One word of caution: before using any jar with a vacuum, test it by 
wrapping some towels around it and connecting it to the vacuum source. 
Even jars of the same brand should be checked individually to make sure 


280 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Efficiency of desiccants. 


Residual Water per Liter 


Material of Air, 30.5°C (in mg) 
CaCl:-H2O IL) 
Ba(ClO,)»2, anhydrous 0.82 
NaOH, sticks 0.80 
CaCl, anhydrous 0.36 
Mg(Cl10,)2:3H:O 0.03 
Silica gel 0.03 
KOH, sticks 0.014 
AIO; 0.005 
CaSQ,, anhydrous 0.005 
CaO 0.003 
Mg(ClO,)2, anhydrous 0.002 
BaO 0.0007 
P.O; lower 


they don’t implode. Tape can be wrapped around the jar as a further 
safety measure. 

A vacuum hose or a large diameter thick walled hose should be used 
in connecting the trap to both the aspirator and the desiccator jar. A 
washing machine hose is excellent for connecting the aspirator to the 
faucet. 

One additional part of the system needs to be discussed. This is the 
desiccant. I use CaSO, which is impregnated with an indicator. The 
desiccant is blue when it is active, and turns red when it can no longer 
absorb water. This desiccant can be reactivated by placing it in a shal- 
low dish in an oven set for 450°F. One brand name for this desiccant 
is Drierite (with indicator). Other desiccants can be used, but CaSQx, 
as seen from Table 2 (Bower, 1934), is one of the most efficient. The 
last point in this discussion of desiccants is to be sure not to overload 
the system. The 8 oz. jar should be no more than one third full of desic- 
cant. This should be enough for any large butterfly larva or a couple 
of small ones. With very large larva such as those of Sphingidae or 
Saturniidae, be sure to start with freshly activated desiccant. A larger 
jar might be required on some of the largest larva. Remember that at 
best the CaSO, can absorb only about % of its weight in water. 

One piece of equipment not needed in this vacuum freeze-drying sys- 
tem is a large freezer. Several 8 oz. jars can be stored in a home refrigera- 
tor freezer and still leave plenty of room for frozen foods. 


VOLUME 30, NUMBER 4 281 


Procedure 


The basic vacuum freeze-drying process consists of the following steps: 
(1) freezing the larva in the desiccating jar; (2) pulling a vacuum; (3) 
keeping this jar frozen until the specimens are dehydrated; (4) thaw- 
ing and releasing the vacuum. Each of these will be discussed in turn. 

The first step, freezing the larva, seems fairly simple, but there are 
a few cautions. Be sure that the jar is tightly sealed. Also, allow plenty 
of time for the larva to freeze. I usually wait until the next day before 
pulling a vacuum. 

The second step, pulling the vacuum, involves several complications. 
The first major problem is to determine the amount of vacuum you are 
creating. The same aspirator will produce widely differing vacuum with 
different water pressures. One method of checking the vacuum is to 
connect the aspirator to a simple manometer. Since this involves an 
added expense (unless you happen to have one laying around the house), 
I use a less accurate but simpler method. I connect an empty 8 oz. 
desiccator jar, equipped with a valve, to the aspirator. The water is 
turned on for 40 seconds, and the valve is closed. Next, the hose is dis- 
connected, the jar and valve are submerged in a pan of water, and the 
valve is now opened. Water will then rush into the jar. A good aspirator 
operating with the correct water pressure should create a vacuum suf- 
ficient to fill an 8 oz. jar to within “4 oz. of being completely full. This 
is about 98% full if one is working with another size jar. 

Because of changes in water pressure one should check the vacuum 
just prior to pulling down the desiccator jar. If the faucet is equipped 
with an aerator, this should be removed to prevent it from reducing the 
water pressure. Be sure to note the position of the cold water faucet 
handle when checking the vacuum, so that it can be returned to the same 
position when the desiccator jar is connected. It is best to work late at 
night when the water pressure is the highest. A most important aspect 
in evacuating the desiccator jar is speed. Have everything set up and 
tested before removing the jar from the freezer. Only 40 seconds are 
needed to evacuate the jar, and the total time out of the freezer should 
not exceed one minute. 

The third step, keeping the jar in the freezer until the specimens are 
dehydrated, deserves a few comments. The length of time required is 
dependent upon numerous factors. The main variables are the tempera- 
ture of the freezer, degree of vacuum, size of larve, and type of desiccant. 
In the system I have described, I have found one week insufficient but 
two weeks adequate for most types of butterfly larva. One last word of 


282 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TasBLE 3. Comparative costs of freeze-drying systems (1974 U.S.A. prices). 


Item Conventional System Aspirator System 

Pump $200.00 $12.00 

Trap none 0.50 

2 Valves none 5.00 

2, Desiccators 50.00 1.00 

Freezer 130.00 none 

Hose & Connections 7.00 12.00 

Total $387.00 (minimum ) $30.50 (maximum ) 


caution: be sure the jar remains frozen—do not pack warm foods around 
it. 

The last step is to remove the jar from the freezer after sufficient time 
has elapsed for dehydration. Before letting it thaw, test to be sure the 
desiccator jar still has a vacuum. I recommend moistening a finger tip 
and then with the finger over the valve, open and close the valve. If upon 
removing the finger, a distinct pop is heard, one can be sure the vacuum 
is still being held. If for some reason there is no vacuum present, one 
can return the jar to the freezer and repull the vacuum later. If the vac- 
uum has to be repulled, replace the stopper and valve with dry ones 
just prior to repulling the vacuum. I have found this procedure to be 
satisfactory as long as the larvae are not allowed to thaw before they 
are completely dehydrated. Upon removing the jar from the freezer, 
do not release the vacuum for 24 hours. Experiments I have done in- 
dicate that the larvae should be sufficiently dehydrated at this point to 
prevent discoloration or spoilage, however the wait of 24 hours allows 
the dehydration to come to a completion. When releasing the vacuum, 
proceed slowly to avoid collapsing the specimens. 


Additional Comments and Results 


Since the main reason for using an aspirator vacuum freeze-drying 
system rather than a conventional system is cost, a comparison of costs 
is given in Table 3. The figures given are based on 1974 prices. Another 
benefit of this system is its portability. It weighs less than one pound 
and can be used anywhere there is running water and a refrigerator 
freezer. 

I have successfully used this system on a wide variety of butterfly 
and moth larvae. A few suggestions can be made for avoiding poor re- 
sults. Larvae that are preparing to moult or that are parasitized often 
do not turn out well. If the frozen larva becomes covered by desiccant, 


VOLUME 30, NUMBER 4 283 


such as by tilting the jar during the process of pulling the vacuum, the 
grains of desiccant can dent it. Leaves of larval foodplants can curl] 
during dehydration, therefore when possible avoid putting leaves in 
the desiccator. Finally a little fading of some colors (especially greens) 
occurs during vacuum freeze-drying. 


ACKNOWLEDGMENTS 


A special thanks goes to the late Dr. Richard Dominick for his many 
helpful suggestions. Two other friends, Mike Rickard and Jim Estep, 
deserve thanks for their help in the writing and proof reading of this 
article. 


LITERATURE CITED 


Bower, J. H. 1934. Bur. Standards J. Research 12: 241. 

Dominick, R. B. 1972. Practical freeze-drying and vacuum dehydration of cater- 
pillars. J. Lepid. Soc. 26: 69-79. 

FiascuKxa, H. A. & J. FLoyp. 1969. A simplified method of freeze-drying cater- 
pillars. J. Lepid. Soc. 23: 43-48. 


984 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TWO NEW SPECIES OF PHYCITINE MOTHS WITH 
DESCRIPTION OF A NEW GENUS (PYRALIDAE) 


ANDRE BLANCHARD 
P. O. Box 20304, Houston, Texas 77025 


Pimodes A. Blanchard, new genus 


Tongue well developed. Antenna finely pubescent; on male with a shallow sinus 
at base of shaft, fringed on both sides with long scales hiding a blunt, black, shiny 
process on each of four or five segments. Labial palpus porrect, extending at least 
twice length of head beyond it; second segment closely scaled, more than twice as 
long as third, longitudinally grooved in male to receive maxillary palpus. Maxillary 
palpus of male in form of a large aigrette, of female minute, squamous. Scaling of 
vertex and front forms with labial palpi a streamlined snout. Forewing smooth; cell 
about two-thirds length of wing; Cuz from before lower outer angle of cell; Cu: from 
before angle but close to it; Mz and M: from angle, approximate for about one-fourth 
their length; discocellular vein extremely weak; M: from a little below upper outer 
angle of cell, straight; R; from upper angle, stalked with R:,. for about one-half 
length of Rs; Re from cell much closer to stalk of R:., and Rs than to Ri, remain- 
ing approximate to this stalk for about one-half its length; Ri from cell; male without 
costal fold. Hindwing cell slightly less than half length of wing; discocellular vein 
weak, deeply concave, greatly extended at lower outer angle of cell; Cuz from before 
lower outer angle; Cu: from angle, much shorter than Cus, connate with the stalk 
of M:; and Mz which are anastomosed for about two-thirds their length; M: and Rs 
anastomosed for some distance beyond upper outer angle of cell; Rs and Sc approxi- 
mate for over half the free part of Rs. Eighth abdominal segment of male with 
paired, ventrolateral, compound tufts (Fig. 6). 

Male genitalia (Figs. 2—5): Uncus hoodlike, its terminal margin notched. 
Apical process of gnathos a stout hook. Transtilla absent. Valve (Fig. 5) elongate, 
tapering to bluntly rounded apex; distal fourth of valve virtually reduced to strongly 
sclerotized costa; sacculus a little over half length of valve, broadest at two-thirds 
its length from its base; a clasper, rooted in basal third of valve, angled and pointed 
at apex, runs parallel to costa, is about as wide and half as long as costa. Inner sur- 
faces of sacculus and clasper studded with numerous spinelike hairs. Vinculum much 
longer than its greatest width, with strongly sclerotized margins. Aedeagus straight, 
stout, with round, ventral extension at its distal rim; vesica (Fig. 4) armed with one 
strong cornutus and a bunch of about four smaller cornuti held together by a com- 
mon sclerotized base. Juxta subquadrate, broadest at base, weakly sclerotized. 

Female genitalia (Fig. 7): Bursa almost three times as long as seventh abdominal 
segment, over three times as long as its average width, slightly bulging ventrally in 
its middle, membranous except for a sclerotized, scobinate-granulate patch on its 
left side, cephalad from junction of ductus bursae; this signum is part of the bursa 
membrane anteriorly, but becomes detached from it and protrudes inside the bursa 
posteriorly; ductus seminalis from apex of a lobe at caudal end of bursa, left of 
ductus bursae; ductus bursae about one-fifth as long as bursa, membranous except 
ventrally at genital opening. 


This genus is closest to Pima Hulst, but there are significant differences: 
the third segment of the labial palpus is much shorter than the second 


in Pimodes, about equal in Pima; the male maxillary palpus is a large 
aigrette in Pimodes, minute and scaled in Pima; My and Ms of the fore- 


VoLUME 30, NUMBER 4 285 


wing are approximate in Pimodes, separate in Pima; the valve has no 
clasper in Pima and its vesica is armed with two subequal cornuti; the 
ductus bursae is short and membranous in Pimodes, long, ribbonlike, 
sclerotized and granulate in Pima. The preceding description, of the 
male and female genitalia of this new genus, is in fact that of the genitalia 
of the new species described below. It is understood that, if and when 
more species are discovered in this genus, a choice will need to be made 
in order to eliminate those characters which will prove te be specific 
rather than generic. 


Pimodes insularis A. Blanchard, new species 
(Figs. 1-7) 


Labial palpi closely and thickly clothed with long, whitish tipped, dark gray 
scales, very slightly paler beneath; apex of head, antennae at base, collar, thorax 
and tegulae concolorous with palpi above; abdomen ochreous yellow above, yellowish 
gray beneath; legs mostly concolorous with labial palpi beneath; middle and hind 
tibiae with long loose scales dorsally. Forewing above with median lines obsolete; 
basally concolorous with thorax, becoming progressively a little paler distally to 
still paler fringe; some reddish scales are scattered on the lower half, forming two 
poorly defined patches at about one-third distance to anal angle; a long, white, fusi- 
form fascie, more or less heavily sprinkled with reddish scales, extends from base 
to apex, thinning out at both ends, being widest at middle of costa. Hindwing above, 
translucent grayish white, somewhat darker along costa and on outer margin near 
apex; a fine dark brownish gray terminal line; fringe white distally and along a fine 
line at base, darker between. Forewing beneath dark brownish gray, paler basally. 
Hindwing beneath as above, except fringe entirely whitish. 

Wing expanse: 13 specimens, males and females measure 17.5-19 mm; the 
holotype measures 21 mm. 

Male genitalia: As described for the genus. 

Female genitalia: As described for the genus. 

Holotype: Male, Padre Island National Seashore, Kleberg Co., Texas, 29 September 
1975, genitalia on slide A. B. 3636, deposited in National Museum of Natural His- 
tory, Type No. 73652, A. & M. E. Blanchard collectors. 

Paratypes: Same locality, 7 July 1975, 1 3, 3 99; 29 September 1975, 3 ¢ 4, 
3 2 2; 2 October 1975, 1 6,1 2; 22 June 1976,2 3 46,1 9; 24 June 1976, 2 ¢ 64, 
1 9; 19 July 1976, 2 646, 16 99; A.& M. E. Blanchard collectors. 


Macrorrhinia signifera A. Blanchard, new species 
(Figs. 8-11) 


I have only five specimens of this new species before me and their maculation 
is quite variable, hence the description must needs take these variations into con- 
sideration. 

Palpi, head, collar, thorax and tegulae clothed with pale ochreous gray scales, 
mixed in variable proportion with darker scales, of same hue in three specimens, 
definitely reddish in one female and blackish in holotype. Forewing irregularly 
mottled with scales of same two colors. No recognizable am. band, but where 
M. aureofasciella Ragonot has an orange am. band preceded by a black line, two 
specimens of new species, including holotype, shows a faint trace of same pattern. 
Discal dots and pm. line mostly obsolete, except on holotype whose straight, pale 
pm. line is followed by an array of blackish spots. Terminal line of blackish inter- 


986 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Ps 


Figs. 1-7. Pimodes insularis: 1, holotype; 2, male genitalia, most firmly planted 
outer scales retained; 3, male genitalia, aedeagus removed, outer scales removed 
using forceps; 4, aedeagus, vesica inflated; 5, inner view of left valve: 6, paired 
ventrolateral, compound tufts of male eighth abdominal segment; 7, female genitalia. 


VOLUME 30, NUMBER 4 287 


.- | 


Figs. 8-ll. Macrorrhinia signifera: 8, holotype; 9, male genitalia; 10, pair of 
short ventrolateral hair tufts and lateral pair of eversible lobes with long hair tufts; 
11, female genitalia. 


venular dots. Fringe concolorous. Hindwing translucent, much paler ochreous than 
forewing, darker at apex and along upper half of outer margin; fringe concolorous. 
Forewing beneath ochreous gray, darker along costa and outer margin; terminal 
intervenular dots as above, fringe concolorous. Hindwing beneath as above. 

Wing expanse: 18.5-20.0 mm. 

Male genitalia (Fig. 9): Differ from those of M. aureofasciella (Heinrich 1956, 
p. 190, fig. 437) in that the cucullus of the valve is much less broadly expanded 
and the three distal lobes of the juxta are much narrower. 


988 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Female genitalia (Fig. 11): Differ from those of M. aureofasciella (Heinrich 
1956, p. 190, fig. 924) by the presence of a strongly sclerotized signum consisting 
of a linear array of minute claws on either side of junction of bursa with ductus 
bursae. 

Holotype: Male (Fig. 8), Town Bluff, Tyler Co., Texas, 7 August 1975, deposited 
in National Museum of Natural History, Type No. 73651, A. & M. E. Blanchard 
collectors. 

Paratypes: Santa Ana National Wildlife Refuge, Hidalgo Co., Texas, 18 November 
1966, 1 9; Town Bluff, Tyler Co., Texas, 7 August 1975, 2 ¢6¢, 2 99, Conroe, 
Montgomery Co., Texas, 9 September 1975, 1 9, A. & M. E. Blanchard collectors. 


Only two species of the genus Macrorrhinia have yet been described: 
M. aureofasciella Ragonot (Ragonot 1901, p. 190) and M. placidella 
(Zeller) (Zeller 1848, p. 874). The preceding comparative description 
leaves no doubt that the new species is different from aureofasciella. 
Dr. D. C. Ferguson made for me a sketch of the habitus of M. placidella 
figured in Ragonot (1901, p. 190); comparing this sketch with my speci- 
mens of the new species leaves no doubt that they are different. 


LITERATURE CITED 


Hernricu, C. 1956. American moths of the subfamily Phycitinae. U.S. Natl. 
Mus. Bull. 207, 581 p. 

Raconort, E. L. 1901. Monographie des Phycitinae et des Galleriinae. In N. M. 
Romanoff, Mémoires sur les Lépidoptéres, Vol. 8, xli + 602 p., pls. 24-57 [MS. 
completed by Sir G. F. Hampson]. 

ZELLER, P. C. 1948. Isis von Oken. 


VoLUME 30, NUMBER 4 289 


THE BIOLOGICAL STATUS OF NEARCTIC TAXA IN THE 
PIERIS PROTODICE-OCCIDENTALIS GROUP (PIERIDAE) 


ARTHUR M. SHAPIRO 
Department of Zoology, University of California, Davis, California 95616 


The taxonomy of Pieris protodice Boisduval & LeConte and its rela- 
tives in western North America has long been confused, with the rela- 
tionship of that taxon to P. occidentalis Reakirt being a matter of partic- 
ular contention. Both current “field guides” (Klots, 1951; Ehrlich & 
Ehrlich, 1961) treat them as conspecific, as does the North American 
check-list (dos Passos, 1965), while recent faunistic papers by authors 
in western North America (e.g., Garth & Tilden, 1963) have regarded 
them as separate species. Chang (1963) provided morphological evi- 
dence in support of the latter position and mapped the distribution of 
P. occidentalis. His analysis was followed by Howe (1975). The situa- 
tion is complicated by the occurrence of named seasonal and/or alti- 
tudinal phenotypes in both, as well as one valid subspecies. The present 
paper summarizes the result of a series of field and laboratory studies 
in which the biological relationships among these entities have been 
clarified; these studies are cited individually in the text. It is not intended 
as a formal taxonomic revision; such a revision should, if undertaken, 
be on a world-wide basis. All of the taxa are abundantly illustrated in 
the papers cited. 

The only previous American treatment of this group was by Abbott 
(1957) and Abbott, Dillon, & Shrode (1960). This incompetent study, 
which “synonymizes” the very different species Pieris beckerii Edwards 
and P. sisymbrii Boisduval with P. protodice and P. occidentalis with 
total disregard for their biology and extensive sympatry, makes no useful 
contribution to the taxonomy of the group. Those names in the group 
which were authored by William H. Edwards have been very thoroughly 
treated taxonomically by Brown (1973). McHenry (1962) has prepared 
a bibliography of the original descriptions of all taxa placed in Pieris in 
North America. 

The biological entities recognized in this paper are: 

I. Pieris protodice Boisduval & LeConte 
f. vern./aut. vernalis W. H. Edwards 
nasturtii “Boisduval MS.” (W. H. Edwards ) 
II. Pieris occidentalis Reakirt 


f. vern./aut./alt. calyce W. H. Edwards 
IIA. ssp. nelsoni W. H. Edwards 


290 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


I. Pieris protodice Boisduval & LeConte (Fig. 1). 


1829. Hist. Gen. Icon. Lepid. Chen. Amer. Sept. 1(5): 45-46; pl. 17, figs. 1-3. 
Type locality New York and Connecticut. Both sexes described and figured. 

The type locality makes this name biologically unambiguous, as there 
is only one member of this species-group in the eastern United States. 
Pieris protodice is distributed over most of the U.S. generally below 
2000 m but reaching 3000 m in New Mexico and Arizona. It is absent 
from the Pacific Northwest, north of the Central Valley of California, 
and from the northeastern states north of southwestern and southeastern 
Pennsylvania except along the immediate coast, north rarely to Massa- 
chusetts. It occurs in southern Ontario, at least sporadically. Its northern 
and upslope borders are extremely unstable; in cold-winter areas it is 
generally dependent on immigration and although it may breed, it over- 
winters only exceptionally. It is reported southward to southern Baja 
California, and on the mainland to Guatemala (Hovanitz, 1962), but 
is rare or absent in subtropical Florida although it has been collected 
on Cuba. It is striking that Boisduval recorded this species, along with 
Colias eurytheme (“edusa’), so far north so early. We will probably 
never know whether they occurred naturally or had already been in- 
troduced from further south. 

Pieris protodice is always associated with sunny, warm, and dry en- 
vironments. It is a “weedy,” “colonizing,” or “fugitive” species commonly 
found in highly disturbed, early-successional habitats, especially on 
sandy soils—often by roadsides, along railroad rights-of-way, or in urban 
vacant lots. In the west it often occurs in the dry washes and around 
corrals. In the northeast it is frequent on beaches. In most of its range 
its preferred hosts are the annual or winter-annual cruciferous weeds 
Lepidium virginicum L. and L. densiflorum Schrad. In California at . 
low elevations it breeds extensively on Brassica geniculata (Desf.) Ball 
where no summer Lepidium spp. grow, and on the southwestern and 
Great Basin deserts on native Lepidium spp. and Physaria spp. and on 
Sisymbrium altissimum L. It rarely breeds on Brassica nigra (L.) Koch 
(a misdetermination by Hovanitz, 1962) but is capable of accepting a 
great variety of crucifers in captivity. 

Pieris protodice is not known to be a permanent resident anywhere 
where the breeding season is too short for multiple broods. It has four 
to six broods in central California and about four at New York and 
Philadelphia. It is always much commoner and more widespread in late 
summer and autumn than earlier in the season. Winter is spent as a 
pupa in diapause under photoperiodic control. 

Stocks of P. protodice from California, Arizona, Colorado, Texas, New 


VOLUME 30, NUMBER 4 291 


So Ae 


SE SES 


ee 


ee 


Fig. 1. Pieris protodice from New York (left-hand pair) and California (right- 
hand pair), summer phenotypes, dorsal and ventral surfaces. The New York male 
is phenotypically “intermediate,” similar to the description of male nasturtii (see 
text). 


292 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Mexico, Pennsylvania, New Jersey, and New York have been crossed in 
various combinations with no significant loss of fertility or viability in 
continuous culture; wide geographic crosses may, however, disrupt the 
diapause response (Shapiro, unpublished ). 

Aspects of the biology of P. protodice in California are discussed in 
Shapiro, 1975a and of its reproductive biology in Shapiro, 1970. 

Pieris vernalis W. H. Edwards (Fig. 2). 


1864. Proc. Ent. Soc. Phila. 2(4): 501-502. Type locality Red Bank, N.J. (May). 
Both sexes described. 

This is the phenotype of P. protodice which is produced when the larva 
develops on a long-night regime, regardless of temperature, and may 
be induced in the progeny of any female of any brood by appropriate 
treatment (Shapiro, 1968). Its production is not dependent on the oc- 
currence of diapause, which may be inhibited under inducing photo- 
periods by high temperatures. Intergrades to the vernalis phenotype, and 
occasionally quite dark examples, are thus produced in autumn. This 
phenotype has not been reported for many localities where the summer 
broods of P. protodice occur; this is scarcely surprising given the fugitive 
nature of populations in this species. There are no differences in the 
expression of the phenotypes in Californian and northeastern stocks. 

No type of Pieris vernalis exists. Despite information given Brown 
(1973) by New Jersey collectors, Pieris protodice is by no means “nearly 
extinct’ in that state; in 1965 I collected over 500 in an afternoon in 
Camden. I have not, however, seen specimens of vernalis from Red Bank 
or elsewhere in Monmouth County, although I have several from nearby 
Staten Island. Because Staten Island is in New York state, I have re- 
frained from designating any of these as a neotype in the hope that a 
genuine Red Bank specimen may turn up. 

Pieris nasturtii W. H. Edwards. 

1864. Proc. Ent. Soc. Phila. 2(4): 501. Type locality San Francisco, California. 
Both sexes described. 

This was a Boisduval manuscript name, resurrected by Edwards. Its 
tangled history is given by Brown (1973). Apparently Boisduval ap- 
plied it to an animal in the Pieris napi group, but the specimens sent 
by Behr to Edwards as “nasturtii” were “an odd variety of protodice,” 
as Edwards later wrote. No type, nor any specimen which can definitely 
be linked to this name, is extant. 

The only member of this group found in or near San Francisco is 
P. protodice (except for the possible occurrence of P. occidentalis in 
the Santa Cruz mountains, see below). The descriptions strongly sug- 


VoLUME 30, NUMBER 4 293 


Fig. 2. Pieris protodice from New York (left) and California (right), vernalis 
phenotypes, dorsal and ventral surfaces. 


gest the late September-November phenotypes of Bay Area P. protodice, 
which are somewhat transitional to vernalis. The “coppery” color of the 
female is characteristic of old, faded-in-life specimens, as is the hyalinity. 
It thus appears that the name nasturtii W. H. Edwards refers to the 


994 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


autumn brood of protodice and is, thus, infrasubspecific and without 
taxonomic standing. Its revival would in no way benefit taxonomy or 
biology, especially since variation from summer to winter phenotypes 
is essentially continuous in autumn in continuously breeding populations. 
In my publications the male description of nasturtii applies to my “inter- 
mediate” phenotypic grade. 


II. Pieris occidentalis Reakirt (Fig. 3). 


1866. Proc. Ent. Soc. Phila. 6: 133-134. Type locality “Rocky Mountains, Colo- 
rado Territory, California.” Both sexes described. 

Pieris occidentalis occurs upslope and northward of P. protodice in 
western North America. The two species have been found sympatric 
at various locations from 1000-2500 m in the Rocky Mountains and Sierra 
Nevada, e.g., Donner Pass, California, where P. occidentalis is a perma- 
nent resident and P. protodice a breeding immigrant (Shapiro, 1975a). 
Pieris occidentalis ranges from Arctic Alaska and adjacent Canada south 
at increasing elevation to the southern Sierra Nevada of California and 
the Colorado Rockies (both above 2000 m), east to the Black Hills of 
South Dakota. It may extend into northem New Mexico. Its eastern 
limits in Canada are poorly understood. Its southern extent in the Cali- 
fornia Coast Ranges is unknown; it may reach Santa Cruz County, where 
it was apparently collected (by M. Doudoroff?) in 1930 (U. C. Berkeley 
collection). However, I have not seen any other Coast Range specimens 
from Mendocino County southward. 

As noted in the introduction, Chang (1963) has described morpho- 
logical differences between these two species. There are useful color 
and pattern differences; both sexes characteristically are more heavily 
and completely marked in P. occidentalis than in P. protodice; the wings 
of P. occidentalis appear thicker and more heavily scaled; the body is 
proportionally larger and usually hairier. The larva is more contrastingly 
colored and the pupa tends to be shorter and broader than in P. protodice. 
The chaetotaxy is univestigated. In areas of sympatry occasional inter- 
specific matings may occur, but only 2 of 339 specimens collected at 
Donner Pass in 1973 were phenotypically ambiguous (Shapiro, 1975a). 
Although conspecific pairs are readily formed in cages, interspecific ones 
are not. 

Pieris occidentalis is characteristically found at low densities in moun- 
tainous regions, where most captures are of “hilltopping” males. It 
normally breeds on montane crucifers such as Arabis and Streptanthus 
species and Thlaspi alpestre L., but becomes “weedy” and breeds at high 
density when presented with the opportunity, as in the railroad yard at 


VoLUME 30, NUMBER 4 295 


Fig. 3. Pieris occidentalis from the California Sierras, summer phenotypes, dorsal 
and ventral surfaces. 


Donner Pass where its host is Lepidium virginicum. It is double-brooded 
at moderate elevations southward and possibly partially triple-brooded 
at its lower elevational limit in both Colorado and California (see below). 
Winter is spent as a diapausing pupa. 

Pieris calyce W. H. Edwards (Fig. 4). 


1870. Trans. Amer. Ent. Soc. 3, signature 25: 189, no. 1. Type locality “Nevada,” 
restricted by F. M. Brown (1973) to vicinity Virginia City, Nev. Male described. 

Edwards (1876) speculated that calyce might be a spring form of Pieris 
occidentalis, and in this he was correct. Brown (1973) demonstrates that 
the type was probably collected by Henry Edwards at Virginia City 
(elev. 1921 m) in March 1868 or 1869. At Virginia City a March speci- 
men would be a very early example of the phenotype emerging from 
overwintering pupae in a bi- or trivoltine population of occidentalis. In 
such populations Shapiro (1973) has shown that calyce is the equivalent 
of vernalis, a seasonal, photoperiod-induced phenotype (although in 
occidentalis the control is less absolute). It can thus be obtained in the 
laboratory from the progeny of any female occidentalis. In its original 


296 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 4. Pieris occidentalis from the California Sierras, calyce phenotypes, dorsal 
and ventral surfaces. 


sense, then, although proposed as a species-group name, calyce clearly 
refers to a seasonal phenotype and is infrasubspecific. 

The name calyce has been used by some authors (e.g., Garth & Tilden, 
1963) in a subspecific sense to apply to the univoltine animal of this 
group which occurs at or above tree-line in the Rockies, Sierras and Great 
Basin ranges. This animal is phenotypically indistinguishable from speci- 
mens collected one or two months earlier 1000 m lower. When stock of 
univoltine “calyce” from Loveland Pass, Colorado (3600 m) was reared 
under continuous light at 25°C it did not diapause, but developed directly 
in less than a month and produced light, occidentalis “summer” pheno- 
types (Shapiro, 1975b). These animals were successfully crossed with 
P. occidentalis from Donner Pass, California, with no decrease in fertility 
or viability. An apparently spring-univoltine “calyce” stock from Hay- 
stack Mountain in the eastern foothills of the Colorado Rockies has also 
been studied (Shapiro, 1976b). 

The various populations of univoltine “calyce” are completely dis- 
junct from one another on mountaintops. Present evidence implies that 


VOLUME 30, NUMBER 4 297 


Fig. 5. Pieris occidentalis nelsoni, Fairbanks, Alaska, dorsal and ventral surfaces. 


each is independently derived locally from the multivoltine populations 
found downslope from it. Whether or not one is willing to accept “poly- 
topic’ subspecies, the use of calyce as a subspecific name is rendered 
inappropriate by the holotype data presented by Brown and should be 
discontinued. The name calyce is appropriately applicable to a pheno- 
type, not a population. 

Univoltine high-elevation populations have been found associated 
with Thlaspi alpestre and Smelowskia calycina (Desv.) Meyer in Colo- 
rado and Erysimum perenne (Wats. ex Cov.) Abrams at Sonora Peak, 
California. No definite host records are known to me. Erysimum is not 
a normal Pieris host (Chew, 1975), though P. rapae has been found on 
it at least once (Shapiro, 1975a). 


IIA. Pieris occidentalis nelsoni (Fig. 5). 
Pieris nelsoni W. H. Edwards. 

1883. Butt. North America 2(11): 71, pl. 15, Pieris I, figs. 6, 7. Type locality 
St. Michael’s, Alaska. Male described and figured. 

This entity has been “lost” since its original description, although many 
reports of P. occidentalis in Alaska have been made. In July 1974, I 
studied a population at Fairbanks, Alaska in which “nelsoni” is the most 
frequent male phenotype, and subsequently bred it in the laboratory, 
crossed it with Sierran occidentalis, and obtained genetic data which are 
being reported elsewhere (Shapiro, 1976a). The nelsoni phenotype is 


298 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


expressed with or without diapause. Ventrally, nelsoni is about as dark 
as calyce, but some specimens reared without diapause are as light as 
summer occidentalis (Shapiro, 1975c). 

St. Michael’s, now St. Michael, is on the south coast of Norton Sound 
in western Alaska. Now that nelsoni has also been found in interior 
Alaska, it appears that it is not an aberration but a valid and recognizable 
geographic subspecies, and I so treat it. It remains to be seen how far 
it extends into western Canada and the nature of its contact with nomi- 
nate occidentalis. Some nelsoni characters are recognizable in popu- 
lations of occidentalis as far south as Modoc Co., California (Shapiro, 
unpublished ). 

P. o. nelsoni has some resemblance to the vernalis phenotype of P. 
protodice, especially in the male. This is presumably the basis for its 
classification as a subspecies of protodice by Howe (1975). Although 
nelsoni has not been tested genetically with protodice, its conspecificity 
with that taxon can be ruled out on Chang’s characters, on the shape of 
the pupa, and on geographical grounds—nelsoni is distributed some 2000 
mi. from the nearest known protodice populations, and in a totally dif- 
ferent climatic and vegetational region. 

The only confirmed host of nelsoni is Lepidium densiflorum, a weed 
in the railroad yard at Fairbanks. As noted in Shapiro, 1975c and 1975d 
there is reason to suspect that this population may be facultatively bi- 
voltine. 


Relationships with Palaearctic Taxa 


Higgins & Riley (1970) treat P. occidentalis (and by implication P. o. 
“calyce” ) as subspecies of the Palaearctic Pieris callidice Hiibner. Brown 
(1973) follows this usage, under which there would be two Nearctic 
subspecies, occidentalis and nelsoni, as interpreted in the present paper. 
There are no genetic data bearing on the relationship. It is evident that 
there is great phenotypic similarity among these taxa, particularly be- 
tween callidice and nelsoni. All populations of callidice known to me 
have a yellow ventral hindwing, a character unknown in any North 
American population; in this respect they parallel many Palaearctic 
members of the Pieris napi complex. W. H. Edwards wrote Henry Ed- 
wards concerning Nelson’s specimen, March 15, 1882: “There is 1 male 
Pieris which I think is certainly Callidice. The upper side agrees per- 
fectly with a male Callidice I have from Europe. The underside is not 
so heavily green dusted on the nervures and branches. If this is Callidice, 
it is the first American example I ever saw.” The plausibility of the 
conspecificity hypothesis is increased by a report from K. M. Philip, 


VOLUME 30, NUMBER 4 299 


of the University of Alaska, that he has a male callidice from the River 
Omolon, Magadansk Oblast, NE Siberia—bridging the gap between 
the Alaskan populations and the nearest callidice recorded by Higgins 
and Riley, in Mongolia. It is very likely that further study will confirm 
the Higgins-Riley-Brown usage. 


ACKNOWLEDGMENTS 


I thank: K. M. Philip for data on Alaskan and Siberian material; R. & 
K. Stanford and M. Fisher for help in Colorado; and F. M. Brown for 
many kinds of aid, including the letter from W. H. Edwards, quoted 
above. This study has been supported by Grant D-804 from the Com- 
mittee on Research, U.C. Davis. 


LITERATURE CITED 


Axspott, W. P. 1957. Ecology and geographic variation in Pieris protodice Bois- 
duval and LeConte. M.S. thesis, Texas A. & M. University. 95 p. 
AxssoTt, W. P., L. S. Ditton, & R. R. SHRopDE. 1960. Geographic variation in 
Pieris protodice Boisduval and LeConte. Wasmann J. Biol. 18: 103-127. 
Brown, F. M. 1973. The types of the Pierid butterflies named by William Henry 
Edwards. Trans. Amer. Ent. Soc. 99: 29-118. 

CuHanc, V. C. 1963. Quantitative analysis of certain wing and genitalia characters 
of Pieris in western North America. J. Res. Lepid. 2: 97-125. 

Cuew, F. 1975. Coevolution of Pierid butterflies and their Cruciferous food 
plants. Oecologia 20: 117-127. 

pos Passos, C. F. 1965. A synonymic list of the Nearctic Rhopalocera. Mem. 
Lepid. Soc. 1: 1-145. 

Epwarps, W. H. 1876. Catalogue of the diurnal Lepidoptera of America north 
of Mexico. Trans. Amer. Ent. Soc. 6: 1-68. 

EnrRuicu, P. R. & A. H. Exrticu. 1961. How to know the butterflies. W.C. Brown 
Co., Dubuque, Iowa. 261 p. 

GarTH, J. S. & J. W. Tupen. 1963. Yosemite butterflies. J. Res. Lepid. 2: 1-96. 

Hiccrns, L. G. & N. D. Ritey. 1970. A field guide to the butterflies of Britain 
and Europe. Houghton Mifflin, Boston. 380 p. 

Hovanirz, W. 1962. The distribution of the species of the genus Pieris in North 
America. J. Res. Lepid. 1: 73-84. 

Howe, W. H. 1975. The butterflies of North America. Doubleday, Garden City, 


Nee YencO3o p: 
Knots, A. B. 1951. A field guide to the butterflies. Houghton Mifflin, Boston. 
349 p. 


McHenry, P. 1962. The generic, specific and lower category names of the Nearctic 
butterflies. I. The genus Pieris. J. Res. Lepid. 1: 63-72. 

SHAPIRO, A. M. 1968. Photoperiodic induction of vernal phenotype in Pieris 
protodice Boisduval and LeConte. Wasmann J. Biol. 26: 137-149. 

1970. The role of sexual behavior in density-related dispersal of Pierid 

butterflies. Amer. Nat. 104: 367-372. 

1973. Photoperiodic control of seasonal polyphenism in Pieris occidentalis 

Reakirt. Wasmann J. Biol. 31: 291-299. 

1975a. Ecological and behavioral aspects of coexistence in six Crucifer- 

feeding Pierid butterflies in the central Sierra Nevada. Amer. Midl. Nat. 93: 

424-433. 


300 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


. 1975b. Ecotypic variation in montane butterflies. Wasmann J. Biol. 32: 
267-280. 

1975c. Photoperiodic control] of development and phenotype in a sub- 
arctic population of Pieris occidentalis (Lepidoptera: Pieridae). Can. Ent. 107: 
Wip—119: 

1975d. Notes on the biology of a “weedy” butterfly, Pieris occidentalis 
(Lepidoptera, Pieridae), at Fairbanks, Alaska. Arctic and Alpine Res. 7: 273- 
278. 


1976a. The genetics of subspecific phenotype differences in Pieris occi- 
dentalis Reakirt and of variation in P. 0. nelsoni W. H. Edwards (Lepidoptera: 
Pieridae). J. Res. Lepid. 14: 61-83. 

1976b. Photoperiodic responses of phenologically aberrant populations 
of Pierid butterflies (Lepidoptera). Great Basin Naturalist 35: 310-316. 


VoLUME 30, NUMBER 4 301 


POPULATION STRUCTURE OF THE PRIMROSE MOTH, 
SCHINIA FLORIDA (NOCTUIDAE) 


STEVEN N. HANDEL! 


Section of Ecology and Systematics, Comstock Hall, 
Cornell University, Ithaca, New York 14853 


The primrose moth, Schinia florida (Guenée), uses the evening prim- 
rose, Oenothera biennis L. (Onagraceae), as both a larval food plant 
and a resting and oviposition site for adults (Hardwick, 1970). Larvae 
and adults are well-known mimics of the developing seed capsules and 
the flower petals (Kellicott, 1879; Hardwick, 1970). 

As many species of the Heliothidinae are associated with flowers 
(Hardwick, 1958), and may be of considerable importance to the re- 
productive biology of their hosts, I conducted a study of S. florida at 
an isolated population of O. biennis, including a capture-mark-recapture 
study during the summer of 1975. 

Study site. I conducted mark-release-recapture work at a population 
of O. biennis of approximately 2000 plants. This population was on a 
rocky island in the middle of Six Mile Creek, Tompkins Co., New York, 
320 m E from Van Natta’s Dam. The island was 112 m long and 16— 
28 m wide. It was formed during the extensive flooding of Six Mile 
Creek during June 1972. The island was covered with a diverse array 
of colonizing plants. I identified 16 woody and 73 herbaceous species 
present in July 1975. The dominant woody species included 32 staghorn 
sumac (Rhus typhina L.) and 27 black locust (Robinia pseudo-acacia 
L.) saplings and many aspen (Populus deltoides Marsh.) shoots. 

O. biennis was the most conspicuous and common herb. Many sweet 
clover (Melilotus alba Desr.) plants were also present. 

The ravine surrounding the island contains an unlogged beech—maple 
forest. No other populations of O. biennis were found in this section of 
the ravine from Van Natta’s Dam to the Ithaca City reservoir, except 
for one small cluster of 160 plants at the head of the dam. 

Population parameters. Daily, from 7 July—26 July 1975, I slowly paced 
across the island, examining each O. biennis plant for S. florida adults. 
These moths can easily be seen resting in or near the flowers. All moths 
found were caught and numbered with a marking pen, using the standard 
techniques (Ehrlich & Davidson, 1960). Moths first were found on 
the island 7 July and were last seen 25 July. This period closely matched 
the peak flowering time of O. biennis. During this period 53 moths (25 


1 Present address: Department of Biology, University of South Carolina, Columbia, South 
Carolina 29208. 


302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TaBLE 1. Population parameter estimates of Schinia florida at the Six Mile Creek 
study site. 


Date Alphat M? Ne Phit B> 
July 7 0.00 00.0 0.29 7: 
8 0.29 2.0 WAY) 0.14 2 
10 0.00 1.0 55 1.00 3 
IU 0.50 2.0 4.0 0.75 9.0 
1s 0.25 3.0 12.0 0.71 10.6 
iL) 0.22 4.3 UG) 3 1.00 10.8 
14 0.43 17.0 39.7 0.14 13 
15 0.43 3.0 Wa0 0.14 3.0 
16 0.25 1.0 4.0 0.25 3.0 
7, VAS 1.0 4.0 0.25 1.0 
18 0.50 1.0 2.0 0.00 = 


1 Proportion of marked animals. 

2 Total marked population. 

3 Total population. 

4 Probability of survival. 

5 Number of new animals joining the population. 
* Insufficient data to allow calculation. 


6, 28 2) were found and marked. Of these, 5 males (20.0%) and 8 
females (28.6%) were recaptured at least once. No moth was recaptured 
more than three days after its first capture; 11 moths were recaptured 
one or two days after marking. 

Table 1 summarizes data from this population, using statistics from 
the stochastic model of Jolly (1965). The size of the population (N) 
peaked at about 40 animals on 14 July, then rapidly fell. Although 2-3 
moths were found 19-21 July, no moths were found 22-24 July; 2 found 
on 25 July were the last seen that summer. These captures from the 
end of the flight season are too few to permit inclusion in the Table 1 
analysis. The short flight season of the adults complements a short de- 
velopment period from egg to pupa. Hardwick (1970) reported a 25 
day period from egg to pupa in his laboratory rearing. In my study, 
average survival rate was .58, which corresponds to an expected life- 
span of 3.26 days (Cook et al., 1967). Most recaptured animals looked 
extremely pale, and many had tattered wing edges. 

Additional observations. Among the moths captured in this study 
were 12 male-female pairs resting in the same or adjacent flowers. After 
marking and release, most moths quickly flew down to the lower leaves 
of herbs within a few meters of the release point. A few animals flew 
several meters down the island. 

On 20 July, I surveyed roadside O. biennis populations in Tompkins 
Co. I found 75 populations of 1-9 plants, of which only 4 (5.3%) con- 
tained S. florida individuals. Five of nine larger plant populations (10- 
59 individuals) contained S. florida, as did one cluster of 170 plants. 


VOLUME 30, NUMBER 4 303 


Much larger densities of this moth are occasionally seen in certain years 
and in certain areas (J. G. Franclemont and T. McCabe, pers. comm. ). 
I have also seen S. florida adults sequestered among Gaura biennis L. 
(Onagraceae ) flowers along Six Mile Creek. Gaura and Oenothera have 
very different floral morphologies and color, although they contain the 
same pigment, isosalipurposide, in their petals which produces ultraviolet 
patterns (Dement & Raven, 1973, 1974). 

Conclusions. S. florida has short lived adults, that rapidly leave, by 
death or migration, the plant population where they have eclosed. This 
behavior is not unexpected for insects which rely on early successional 
or ephemeral food plants (Brussard & Ehrlich, 1970). The close asso- 
ciation of S. florida with O. biennis flowers, and the moths’ possible 
movement from plant to plant and among populations may allow for 
pollen movement between these plants. Pollen is often found brushed 
on the moths’ thorax. Although O. biennis is a well-studied case of a self- 
pollinated plant with special cytogenetic features (Cleland, 1972), there 
are experimental results showing that crossing will occur at a regular 
low rate in test gardens of diverse O. biennis races (Hoff, 1962). Also, 
isozyme investigations confirm that small amounts of crossing do occur 
(Levin, 1975; Levy and Levin, 1975). A vehicle for such pollen move- 
ments is needed; S. florida is a likely possibility. 


ACKNOWLEDGMENTS 


I thank Diane F. Nielsen and Joan L. Handel for help in the field 
work, and A. T. Vawter for his helpful comments. 


LITERATURE CITED 


Brussarp, P. F. & P. R. Enruice. 1970. The population structure of Erebia 
epipsodea (Lepidoptera: Satyrinae). Ecology 51: 119-129. 

CLELAND, R. 1972. Ocnothera cytogenetics and evolution. Academic Press, New 
York. x +- 370 p. 

Cook, L. M., L. P. Brower, & H. J. Croze. 1967. The accuracy of a population 
estimation from multiple recapture data. J. Anim. Ecol. 36: 57-60. 

DEMENT, W. A. & P. H. Raven. 1973. Distribution of the chalcone, isosalipur- 
poside, in the Onagraceae. Phytochemistry 12: 807—808. 

DEMENT, W. A. & P. H. Raven. 1974. Pigments responsible for ultraviolet pat- 
terns in flowers of Oenothera (Onagraceae). Nature 252: 705-706. 

EnRuicuH, P. R. & S. E. Davipson. 1960. Techniques for capture-recapture studies 
of Lepidoptera populations. J. Lepid. Soc. 14: 227-229. 

Harpwick, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed 
species of Schinia (Lepidoptera: Noctuidae), with notes of the Heliothidinae. 
Can. Ent. Suppl. 6. 

1970. The life history of Schinia florida (Noctuidae). J. Lepid. Soe. 
24: 282-287. 

Horr, V. 1962. An analysis of outcrossing in certain complex-heterozygous 
euoenotheras. I. Frequency of outcrossing. Amer. J. Bot. 49: 715-724. 


304 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Jotty, G. 1965. Explicit estimates from capture-recapture data with both death 
and immigration-stochastic model. Biometrika 52: 225-247. 

Keiuicotr, D. S. 1879. An example of protective mimicry. N. Amer. Entom. 
1: 30-31. 

Levin, D. A. 1975. Genic heterozygosity and protein polymorphism among local 
populations of Oenothera biennis. Genetics 79: 477-491. 

Levy, M. & D. A. Levin. 1975. Genic heterozygosity and variation in permanent 


translocation heterozygotes of the Oenothera biennis complex. Genetics 79: 
493-512. 


VOLUME 30, NUMBER 4 305 


FEMALE ANAL HAIR TUFT IN NORDMANNIA MYRATLE 
(LYCAENIDAE): EGG-CAMOUFLAGING FUNCTION AND 
TAXONOMIC SIGNIFICANCE 


IcHtRo NAKAMURA 
41 Sunrise Boulevard, Williamsville, Buffalo, New York 14221 


Females of certain Palaearctic eumaeinid butterflies have a cluster 
of specialized scales, the so-called anal hair tuft, at the tip of their ab- 
domens. The best known example is the South European Nordmannia 
acaciae (Fabricius). The actual function of the structure apparently 
remains undocumented, even though similar structures in other genera 
(see below) are known to be used for camouflaging eggs with the scales. 
The purpose of this short note is to show that the female anal hair tuft 
of a closely allied species, Nordmannia myrtale (Klug), does function 
as an egg-camouflaging device. The taxonomic significance of the hair 
tuft will also be discussed briefly, as the genus Nordmannia Tutt tra- 
ditionally includes species with and without the structure in the female 
(Tutt, 1907; Higgins, 1975). N. myrtale is the type species of the genus. 

N. myrtale was originally described from “Syria” (today’s Lebanon) 
and its range extends to Turkey (A. Kocak, unpublished) and perhaps 
to Armenia (armena Rebel). The southern distribution limit is Mt. 
Hermon on the frontier (since 1967) of Lebanon, Syria, and Israel where 
it occurs above c. 1600 m on the southern slopes. The food plant found 
at 2000 m is Cerasus prostrata (Lab.) Ser. (Rosaceae), a typically Irano- 
Turanian subalpine dwarf shrub from the plant geographical point of 
view. In late June 1975, some freshly laid eggs were obtained from the 
plants. Several shells and dead eggs (some parasitized) remaining from 
the previous year were also found. As usual in this group, the eggs were 
located on wrinkled parts of the twigs and at branching points. Some 
were as close to the ground as a few centimeters; the plant itself is only 
20-30 cm high. 

Interestingly enough, the fresh pale brown eggs were invariably cov- 
ered with long scales (Fig. 1), although weathering left only a few 
damaged scales or none at all on the old shells and dead eggs. The 
morphology of the scales on the eggs corresponds exactly to those of 
the female abdominal tip. Such egg-camouflaging is presumably a 
defense mechanism against parasitic insects and predators, although its 
effectiveness could be questioned in certain cases (see below). It also 
seems likely that the female anal hair tuft has other functions as well; 
in Nordmannia the color of the hair tuft is either black or white de- 


306 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


asi 


Figs. 1-4. Egg of Nordmannia myrtale (Klug) (Mt. Hermon, 2000 m, 28. vi. 
1975; on Cerasus prostrata; I. N. leg.): 1, whole egg in situ, magnification 55 xX 
(egg diameter 0.72 mm); 2, the same, part, magnification 123 x; 3, cell structure, 
magnification 1320 x; 4, micropyle, magnification 1320 x. 


pending on the species and therefore it contrasts conspicuously with the 
rest of the abdomen, suggesting a role in courtship. 

Morphologically the egg (Figs. 1-4) resembles those of related spe- 
cies but differs in a number of structural aspects (for SEM photographs 
of an egg of spini Denis & Schiffermuller group, usually included in genus 
Strymonidia Tutt, see Nakamura, 1976). Since the egg provides ex- 


VOLUME 30, NUMBER 4 307 


tremely useful taxonomic characters, its fine structures are also included 
in the figures. The number of radiating micropylar cells varies between 
five and six. 

Certain other groups of butterflies are also known to camouflage eggs 
by an analogous method, usually but not always in association with a 
grossly visible anal hair tuft. They include the following Palaearctic, 
Oriental and African genera of Lycaenidae and Hesperiidae: Japonica 
Tutt (Theclinae, Theclini) in which the two Japanese species lacking 
a conspicuous hair tuft collect dust as well as scales to conceal the eggs, 
although rather poorly (see e.g., Shirozu & Hara, 1960-62); Chaetoprocta 
de Nicéville (Theclinae, Theclini) in which the female of the sole spe- 
cies of the genus carries a conspicuous hair-tuft and uses it effectively 
for camouflaging eggs (Wynter-Blyth, 1957); Daimio Murray (sens. 
str.) and Tagiades Hiibner (Pyrginae, Tagiades group) in which the 
eggs of known examples are covered by scales to such an extent that 
the shell may not be visible at all (see e.g., Shirozu and Hara, 1960-62). 
The species of the African genus Pseudaletis Druce (Theclinae, Aph- 
naeini) have a female anal hair tuft, but a camouflaging function does 
not seem to be on record (Stempffer, 1967). 

Thus, the degree of perfection in camouflaging eggs differs consid- 
erably among genera; the way it is achieved may also vary. According 
to Wynter-Blyth (1957), the scales stick to the egg automatically as the 
female Chaetoprocta odata (Hewitson) lifts the abdomen from the egg. 
To my knowledge, the females of Daimio and Tagiades, as well as 
Japonica, make deliberate efforts to conceal their eggs. I have not 
witnessed the oviposition behavior of N. myrtale, but the orientation 
and clustering of scales on the egg (Fig. 1) clearly mark several brush- 
ing strokes by the female abdomen. It is therefore possible that the 
egg-camouflaging behavior and the anal hair tuft have evolved partly 
independently of each other. The sporadic presence of the female anal 
hair tuft in widely separated genera indicates that it is a case of con- 
vergence as previously suggested (Eliot, 1973). Yet, it is of interest to 
note that the two Theclini genera mentioned above have been consid- 
ered to be rather primitive on structural grounds (Shirozu & Yamamoto, 
1956). 

Regardless of its evolutionary significance, the hair tuft as a taxonomic 
character seems to be constant within each of the genera and calls for 
a re-examination of the current view of the genus Nordmannia. The 
genus as originally introduced (Tutt, 1907) and as currently in use 
(Higgins, 1975) is based on characters of doubtful significance, and 
consequently includes heterogeneous elements. Therefore, it is worth 


308 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


pointing out that the species possessing female, anal hair tufts comprise, 
on other grounds as well, a homogeneous group distinct from those 
without such structure (ilicis Esper, esculi Hubner). 

Firstly, the specific status of many named forms in the myrtale-acaciae 
group is still obscure due largely to their extreme superficial uniformity 
and practically identical genitalia. On the other hand, ilicis and esculi 
were once considered as conspecific, reflecting their superficial resem- 
blance. Although the genitalia of both groups are internally quite uni- 
form, there are some notable differences between the groups as, for 
instance, in the phallus. Geographically, ilicis and esculi are typically 
Mediterranean in distribution, although the former has established it- 
self in Central Europe as have several other Mediterranean species. The 
myrtale-acaciae group is centered within the area conveniently called 
the Irano-Anatolian region of the Middle East. Again, the extension of 
acaciae into Southwest Europe along the northern shore of the Medi- 
terranean has a number of parallel examples among groups centered in 
the Middle East. This is evidently due to the climatic history of the 
Mediterranean region (e.g., Bonatti, 1966) and the persistence of sim- 
ilar ecological niches in the two regions today. Ecologically, the ilicis- 
esculi group is basically an inhabitant of warm, relatively mesic Medi- 
terranean maquis and forests, associated with evergreen, sclerophyllous 
oaks which are the main larval food plants of ilicis (those of esculi are 
apparently unknown). The fact that ilicis has been reported to feed on 
Prunus L. is hardly surprising since food plant specificity seems rela- 
tively plastic in the whole group of related genera. This is probably 
one of the factors that permitted northward expansion of ilicis in Europe. 
In contrast, the myrtale-acaciae group prefers, as a whole, more xeric 
habitats and is better adapted to the cold, being associated with certain 
vegetation types such as steppe forest characteristic of Irano-Anatolian 
region. The food plants are Prunus L., Crataegus L. (for acaciae in 
Yugoslav Macedonia; Nakamura, unpublished observation), and other 
arboreal Rosaceae. There is as yet no indication of oaks being utilized 
by this group. In short, the myrtale-acaciae group and the ilicis-esculi 
group are natural groups distinct from each other by criteria which may 
justify generic separation. It is a matter which should be decided in a 
revision of the entire, much larger group of related genera. 


ACKNOWLEDGMENTS 
[ thank A. Shmida, Department of Botany, Hebrew University of Jeru- 
salem, for botanical identifications and Tova Rivnay, Department of En- 
tomology, Faculty of Agriculture, Hebrew University Campus at Rehovot, 


VoLUME 30, NUMBER 4 309 


for assistance in taking the SEM pictures. The unpublished data pro- 
vided by Dr. A. Kogak, Ankara University, are much appreciated. 


LITERATURE CITED 


BonatTti, E. 1966. North Mediterranean climate during the last Wirm glaciation. 
Nature, London 209: 984—985. 

ExiotT, J. N. 1973. The higher classification of the Lycaenidae (Lepidoptera): A 
tentative arrangement. Bull. Brit. Mus. Nat. Hist. (Ent.) 28: 373-505. 

Hiccins, L. G. 1975. The classification of European butterflies. Collins, London. 
320 p. 

Nakamura, I. 1976. Descriptions of two new species of butterflies (Lepidoptera, 
Lycaenidae) from the South Sinai. J. Ent. (B) 44(1975): 283-295. 

SHirozu, T. & A. Hara. 1960-62. Early stages of Japanese butterflies in colour, 
vols. I & II. Hoikusha, Osaka. 285 p., 120 col. pls. (Text in Japanese). 

Sutrozu, T. & H. YaAMAMotTo. 1956. A generic revision and the phylogeny of the 
tribe Theclini (Lepidoptera; Lycaenidae). Sieboldia, Fukuoka 1: 329-421, pls. 
35-85. 

STEMPFFER, H. 1967. The genera of the African Lycaenidae (Lepidoptera: 
Rhopalocera). Bull. Brit. Mus. Nat. Hist. (Ent.) Suppl. 10. 

Tutt, J. W. 1907-08. A natural history of the British Lepidoptera, vol. 9. Swan 
Sonnenschein & Co., London. x + 495 p., 28 pls. 

Wynter-BiytH, M. A. 1957. Butterflies of the Indian Region. The Bombay 
Natural History Society, Bombay. xx + 523 p., 72 pls. 


NOTES AND NEWS 


On behalf of the Society, I thank Elaine R. Hodges for generously providing her 
drawing of Cosmopterix bacata Hodges that appears on the front cover of each issue 
of Volume 30 of the Journal. I am very grateful to the following persons who silently 
contributed by reviewing one or more manuscripts during the past year: A. M. 
Blanchard, J. K. Bouseman, F. M. Brown, K. S. Brown, Jr., H. K. Clench, D. R. Davis, 
J. P. Donahue, J. C. Downey, D. C. Ferguson, C. D. Ferris, R. S. Funk, H. N. 
Greenbaum, D. F. Hardwick, R. R. Irwin, R. O. Kendall, L. D. Miller, T. A. Miller, 
W. E. Miller, R. W. Neck, H. H. Neunzig, T. Pliske, J. C. E. Riotte, T. D. Sargent, 
D. K. Sell, A. M. Shapiro, F. W. Stehr, J. G. Sternburg, J. R. G. Turner, 
F. A. Urquhart, and A. M. Young. Assistance with the improvement of various sub- 
mitted illustrations was contributed by L. LeMere, Illinois Natural History Survey, 
and G. L. Venable, Smithsonian Institution. B. P. Sweeney and C. L. Kyse are thanked 
for their typing services. Finally, I acknowledge my associate editors, W. H. Allen 
and J. G. Sternburg, for supporting me in numerous and invaluable ways. 


G. L. GopFREY 
Editor 


310 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


GENERAL NOTES 


ECOLOGICAL NOTES ON CELASTRINA EBENINA (LYCAENIDAE) 


Among plebejine blues, Celastrina ebenina Clench is notable for having females 
that are partly lustrous pale blue above, whereas the males are dull blackish or 
grayish-brown: the reverse of the usual situation in species with dimorphic sexes. 
This taxon (for which an appropriate English common name might be “dusky azure’ ) 
was not recognized as a species until 1972. Its geographical range, however, ex- 
tends roughly from Indiana to Pennsylvania, south to North Carolina, biologically 
one of the better known regions in North America. Compared with its close rela- 
tive, Celastrina pseudargiolus, the new species is rare and local, known from only 
13 localities (including several new ones reported below). 

We attempted to obtain information on habitat and behavioral differences, if any, 
between the rare species and the common species. Where C. ebenina occurs, C. 
pseudargiolus is apparently always present. Wagner visited two C. ebenina localities 
in the spring of 1975. The first of these was an unreported one discovered by 
Showalter in Poverty Hollow, Montgomery Co., Virginia. Showalter took a single 
male there in flight on 23 April 1972, and the following year another single male 
was taken at the same spot on 15 April. Efforts to rediscover the butterfly in Poverty 
Hollow during 13-15 April 1975 failed, even though such other butterflies as C. 
pseudargiolus, Glaucopsyche lygdamus, Pyrgus centaurae, and an unidentified Cal- 
lophrys were flying. The precise spot where Showalter found his specimens was re- 
visited many times but without results. The habitat is a dry second-growth area 
where burning and logging took place a few years ago. Now it is grown up with 
young pines, brambles, and various acid-soil members of the heath family such as 
blueberries. It seems possible that the individuals found were casuals, and that 
the metropolis of the species in Poverty Hollow is elsewhere, perhaps in a more 
mature hardwood forest area nearby. 

On 3-5 May 1975, Wagner went to Clench’s locality near Zaleski, Vinton Co., 
Ohio (Clench 1972, Ann. Carnegie Mus. 44: 33-44) and, following his directions 
(in litt.), located the exact site. The first two days were cloudy, and only a few 
butterflies were observed. On the third day the weather cleared, and ca. 20 readily 
identifiable individuals of C. ebenina of both sexes were seen. Also found in the 
general vicinity were Pyrgus centaurae, Colias philodice, Vanessa (Cynthia) virgin- 
iensis, Papilio spp., Strymon melinus, Panthiades m-album, and Callophrys henvici. 
Skippers of the genus Erynnis (especially E. juvenalis) were abundant everywhere 
along roadsides. Celastrina pseudargiolus was common throughout the area, both in 
woodlands and along roadsides, where the males were clustered on moist earth. 

Celastrina ebenina was confined strictly to the tiny valley site described by Clench 
with only one exception. A solitary male was “mudding” with far more numerous 
C. pseudargiolus ca. % mile away along a dirt road. Recognition of females in the 
field is somewhat difficult, but the blue color has a grayish-green cast, and the flight 
pattern may be different (see below). A few specimens were taken, but most of 
the butterflies were only observed. 

Males are much less conspicuous on the wing than males of C. pseudargiolus be- 
cause they lack the blue reflectance. We found males of C. ebenina difficult to de- 
tect and follow in flight except in unusually good circumstances. Both sexes flew 
lower and faster than C. pseudargiolus. This was observed especially well in the 
open areas of the habitat. The males tended to have a direct, swift flight within 
one foot of the ground vegetation, with much “exploring.” Celastrina pseudargiolus 
generally had slower, rather fluttery, more up and down flight, and usually in the 
upper shrub layer. 

No actual instances of C. ebenina feeding on flowers were observed, but several 
males hovered around Jacob’s ladder (Polemonium reptans) flowers and stemless 


VOLUME 30, NUMBER 4 311 


blue violets (Viola cf. cucullata). One male landed on the petal of a Cranesbill 
flower (Geranium maculatum). 

Egg-lying was not observed, and thus no information on potential larval foods 
was obtained. Every plant species we saw both in and out of the small valley was 
common and widespread. It is possible that foodplants are various and that the 
limiting factor in local distribution involves something special about the habitat. 
Certainly the confinement of the Zaleski population to its little valley is extraordinary. 
We searched in many gulleys and woodlands in the region but found no additional 
populations, which indicates that other populations, if present, are very scattered. 

The Zaleski habitat is difficult to assess botanically because the north-facing slope 
has a very different flora from the one facing south. The north-facing slope has a 
maple-basswood association. Such rich woodland forbs as species of Viola, Dicentra, 
Orchis, and Trillium are prominent. The south-facing slope is much more exposed 
and dry, and it has an oak-hickory association. Fewer forbs occur at ground level, 
and there is a strong development of ericads. The center of the C. ebenina habitat 
is a cool, moist opening, just below a rocky waterfall. Here the dominant plant is 
spicebush, Lindera benzoin. If C. ebenina originates in the adjacent woods, we are 
inclined to associate the species with the north-facing slope. We actually saw a 
few specimens ca. 50’ above the bottom of the valley on that slope, and most of the 
specimens arriving below the waterfall came from that side. 

There is some question about how far west C. ebenina occurs. Until recently, the 
Zaleski site was the farthest west of thoroughly documented localities. Clench (1972) 
reports the species far to the northwest in Wabash Co., Indiana. In 1972, David K. 
Parshall (pers. comm.) encountered a small colony in the Fort Hills State Park in 
Highland Co., Ohio, southeast of Hillsboro. This is over 50 miles to the west of 
the Zaleski site and is the second record for Ohio. Parshall informed us that the 
butterfly is “intensely local” and seems to appear “a little later than C. pseudargiolus.” 
His locality, along with the two in Kentucky described below, represent the western 
extent of C. ebenina based upon actual specimens. 

We herewith report C. ebenina for the first time from Kentucky. On 21 April 
1974, Gerald B. Straley and Wagner visited a richly wooded area along Rt. 77 in 
Menifee Co. ca. % mile north of the Menifee-Powell line, where the following 
butterflies were flying: Amblyscirtes vialis, Erynnis juvenalis, E. icelus, Epigyreus 
clarus, Everes comyntas, and Pieris virginiensis. Among the blues collected by Straley, 
one was later identified as a female C. ebenina. 

In Breathitt Co. near Elkatawa, Showalter took a single female C. ebenina on the 
wing on 26 April 1975. In Menifee Co. near the junction of Rt. 77 and the Powell 
Co. line, close to where Straley found the species, two males were taken on mud on 
27 April 1975. 

From the evidence so far, the species appears to be closely associated with more 
or less mature, rich, deciduous forest. The general impression is that this scarce 
butterfly is most often encountered singly or in groups of a few individuals. Sizable 
populations are unusual, as is the case with such other lycaenids in the eastern United 
States as Erora laeta and Euristrymon ontario. Price (1974 J. Lepid. Soc. 28: 268) 
found C. ebenina several times in Buncombe Co., North Carolina, but observed only 
one or two individuals at a time. S. S. Nicolay (in litt.), however, saw the species 
in considerable numbers during the early 1950s in Pendleton Co., West Virginia. 

We should like to recommend that care be taken to preserve the natural popula- 
tions of C. ebenina. A better pursuit than merely making large collections of them 
would be to observe the insect and learn more of its habitat, larval food, and adult 
behavior, especially in comparison with the ever-present C. pseudargiolus, with which 
it was so long confused. 

We wish to acknowledge the help of Harry K. Clench, John Evans, Donald J. 


g12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Harvey, S. S. Nicolay, David K. Parshall, Gerald B. Straley, and Florence S. Wagner 
in making this study. 


W. H. Wacner, Jr., Department of Botany, The University of Michigan, Ann 
Arbor, Michigan 48104. 

Amos H. SHowALTER, Department of Entomology, University of Kentucky, Lex- 
ington, Kentucky 40506. 


A MIGRATION OF VANESSA CARDUI (NYMPHALIDAE ) 


Recently, in arranging a 25 year collection of Manitoba butterflies, a short series 
of eight specimens of Vanessa cardui (Linnaeus) was located in the writer's col- 
lection. Two are dated 4 June 1952; five are dated 5 June 1952 and one is dated 
7 June 1952. These are small specimens as compared to locally emerged autumn 
specimens. 

The above eight specimens (a ninth has just been found in the Sam Waller Mu- 
seum at The Pas from the same series and collection dates) were netted from among 
dozens that were flying at shoulder height from the east into the northwest. This 
is against the prevailing winds in this part of Canada. 

The migration started on the first of June 1952 and continued for the next seven 
days. The flight was a light one and the specimens are in remarkably good con- 
dition. The flight went through the eastern part of the town of four thousand people. 
One of two butterflies could be seen crossing a town block at any one time. They 
would also come in spurts of three or four and then there would be pauses when 
none would be seen. The flight was most pronounced at about 1600 hours. The 
butterflies then flew lower and into the setting sun. The flight ended at the end of 
the week as suddenly as it had started. 

Vanessa cardui cannot survive our severe winters this far north in Canada. The 
autumn generation here is produced from spring migrants. In some years not one 
V. cardui is seen all summer. Those that are locally produced in August are very 
large and brilliantly marked. 

The migration of 1962 was of continental dimensions and reached The Pas. It 
was locally abundant and coincided with the blooming of the dandelions in the first 
week of June. In this regard it resembled the 1952 local flight. 

There is, however, a vast difference between specimens of the two flights sep- 
arated by a 10-year period. In the 1952 flight the specimens were small, well marked 
and fresh looking. Specimens of the 1962 flight were large, much worn and tattered 
and were in greater numbers than the 1952 flight. In 1962 almost every dandelion 
had its butterfly. No directional flight was noted. They suddenly appeared in vast 
numbers in The Pas area and gradually became more and more tattered. The flight 
died out naturally here, not moving elsewhere. The local autumn flight is at best 
a small one in most years. There are not as many thistles around as there were in 
the 1950's. The use of herbicides along roadsides possibly accounts for this. A 
day’s collecting in the fall produces a few specimens of Vanessa cardui in the best 
of years. 

The 1952 June migration, strangely enough, did not produce a large autumn 
flight. This was expected and proved a disappointment when it failed to materialize. 

It is probable that the remarkable heat wave, breaking all previous records, on 
19 April 1952, influenced insect movements locally. At The Pas the heat persisted 
for three days, “bringing out” many species of noctuid moths in large numbers. 
Rare species appeared in numbers that have not been taken since. 

The temperature rose to 80°F... producing a sultry night. This is unheard of 
in these parts in April. It resembled an August night before a storm! These weather 
conditions may have initiated the Vanessa cardui migration some weeks later. 


Waxrer V. Krivoa, P.O. Box 864, The Pas, Manitoba, Canada. 


VOLUME 30, NUMBER 4 313 


BOOK REVIEW 


MACROLEPIDOPTERA OF FIJI AND RoTruMA: A TAXONOMIC AND BIOGEOGRAPHIC STUDY, 
by Gaden S. Robinson. 1975. E. W. Classey Ltd, Park Road, Faringdon, Oxon, 
Great Britain. vii + 362 p., 357 plate figures, 173 text figures, 15 maps. Price 
$25-95"°(U.S:). 

The base data for the biogeographic study are 400 species of macrolepidoptera of 
Fiji and Rotuma. These are treated in an abbreviated manner with the citation 
of the original description, whether type specimen was examined (usually indicated), 
description of male and female, diagnosis, world distribution, Fijian distribution, 
biology (if known) and remarks for each species. Keys for some of the larger 
genera are given. Each species is illustrated by a half-tone of the adult and some 
by diagnostic line drawings of the genitalia. Two genera, 72 species and 10 sub- 
species are newly described. Most of the more than 1 million specimens were 
collected by G. and H. S. Robinson between 1966 and 1972. Additionally, material 
accumulated by nearly all earlier collectors was studied. 

Robinson uses cluster analysis of the accumulated data to define groups of species 
with common distribution patterns for combinations of islands and then generalizes 
from them. To compare the faunas of many Pacific islands he subdivides the macro- 
lepidoptera into: a) generally large, strong flying moths, b) generally smaller, weaker 
flying moths than in group a, and c) butterflies. Major conclusions are: 1. Fiji 
has an unusually high percentage of endemic species, 46% (182 species) and is 
second only to the Hawaiian Islands among the Pacific islands in species endemicity. 
2. The largest number of Fijian endemic species are associated with the rain forest. 
3. Fiji has an island fauna that is derived mainly from the New Hebrides, Solomons 
and Papuasia. 4. Species common to Fiji and many Pacific islands are associated 
with secondary vegetation and are mainly those that can colonize “weedy” areas. 
5. One gateway to Polynesia was from the southern Solomon Islands via Rotuma 
and several other islands to Samoa during periods of maximum glaciation. 6. The 
butterfly fauna of Fiji is relatively impoverished. 7. Rotuma has a low percentage, 
8% (6 species), of endemic species. 

This book is a major contribution to our knowledge of the biogeography of the 
Pacific islands and to the knowledge of the macrolepidoptera of Fiji and Rotuma. 
It is well documented. Some minor points of criticism are: I seriously doubt that 
those species for which the type specimens were not examined are unquestionably 
correctly identified; I presume that an editorial decision caused Hiibner to be spelled 
“Hubner” and Guenée to be spelled “Guenee”; and for ease of reference, numbering 
each page throughout the work would have been helpful. 


RonaLtp W. Honces, Systematic Entomology Laboratory, IIBIIl, USDA, c/o U.S. 
National Museum, Washington, D. C. 20560. 


314 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


INDEX TO VOLUME 30 


(New names in boldface) 


aberrations, 151 
Agraulis vanillae, 59 
Amblyscirtes belli, 68 
Anartia jatrophae guantanamo, 207 
Andersen, W. A., 19 
Anthocaris coloradensis, 252 
Arbogast, R. T., 4, 61 
Archaeoprepona antimache gulina, 159 
demophon centralis, 23 
Argyreus hyperbius niugini, 12 
Arnaud, P. H., Jr., 244 
Battus philenor, 70 
behavior, 169, 183, 235, 237 
Bennett, R., 235 
Bertelia dupla, 211 
biologyss23, 50) 127loo sooo sean 
301, 310 
Bird, C. D., 201 
Blanchard, A., 1, 116, 211, 284 
Blastobasidae, 219 
Boloria frigga, 233 
book reviews, 72, 138, 143, 244, 313 
Borgo, P. M., 169 
Bristol, M. L., 152 
Brower, A. E., 33 
Browne. Ei 233 
Brown, L. N., 213 
Callophrys eryphon, 16 
fotis, 68 
gryneus, 169) 
polios, 68 
siva, 169 
Callosamia angulifera, 114, 184 
promethea, 114, 184 
securifera, 111, 114, 184 
capture-recapture, 145, 301 
Catocala ealiforniensis, 36 
erichi, 36 
johnsoniana, 34 
lincolnana, 34 
texarkana, 33 
Celastrina ebenina, 310 
Charaxinae, 150 
checklists, 38, 197, 230 
Chlosyne lacinia, 91 
Classey, E. W., 151 
Clench, H. K., 88, 121 
cocoons, 131 
collecting, 111, 145, 157, 232 
collections, 201 
coloration, 88, 114 
Colias alexandra, 68 
Consul electra, 159 
Cooper, W. J., 95 
Copablepharon albisericea, 116 
gillaspyi, 119 


grandis, 116 
serraticornis, 119 
Cosmopterix bacata, 309 
Ctenuchidae, 266 
Danaidae, 59, 73, 137, 153, 235 
Danaus plexippus plexippus, 59, 137, 153, 
235 
Davies, T. W., 244 
distribution, 5, 16, 18, 38, 50, 61, 62, 68, 
69, 105, 121, 145) a4e oe 2015 
206, 213, 218, 233) Deano 
Douglas, M. M., 206 
Drummond, B. A., 238 
Dunama angulinea, 190 
claricentrata, 196 
mexicana, 193 
ravistriata, 195 
tuna, 192 
egg-camouflaging, 305 
Ehrlich, P. R., 150 
Epiblema desertanum, 50 
discretivanum, 51 
scudderianum, 50 
Erebia callias, 68 
Euchloe ausinoides coloradensis, 252 
ausinoides palaeoreios, 253 
Eupackardia calleta, 127, 187 
Fales, J. H., 149 
Feeny, P. P., 71 
Ferge, L. A., 234 
Ferris, C; Di, 3866545 
Formicidae, 237 
Freeman, H. A., 62 
Cally hit 
genitalia, 117, 118, 194, 195, 212, 226, 
227, 255, 286) 28M 
Geometridae, 267 
Handel, S. N., 301 
Hapalia nigristriatalis, 11 
Hedges, F. R., 277 
Hedylidae, 267 
Henderson, R. A., 68 
Heppner, J. B., 18 
Hesperiidae, 5, 19, 42, 62, 68, 105, 202 
Heteroptera, 147 
hibernation, 126 
hilltopping, 183 
Hodges, E. R., 309 
Hodges, R. W., 245, 313 
Holocera gigantella, 221 
paradoxa, 224 
Horn, H. S., 146 
Hyalophora cecropia, 131 
hybrids, 4 
Irwin, R. R., 152 
jennings: Dp) Te 257, 
Johnson, K., 169, 252 


VOLUME 30, NUMBER 4 


Kendall, R. O., 105, 264 

Keel W. J., 16 

Kundson, E. C., 235 

Riive@a, W. V., 312 

Kuehn, R. M., 234 

Larsen, T. B., 183 

larval foodplants, 23, 50, 58, 70, 105, 
eto 184, 188, 207, 213, 219, 
2647712. 301 

Leptotes cassius, 206 

cassius theonus, 61 

Libytheana bachmannii, 145 

Libytheidae, 47 

life histories, 23, 159, 187, 207, 264 

Limenitis archippus, 4 

arthemis astyanax, 4 

Lycaenidae, 16, 46, 61, 68, 169, 203, 206, 
Promo 4 235, 249. 305, 310 

Lymantria dispar, 113 

Lymantriidae, 113 

Macrorrhinia signifera, 285 

Mather, B., 197 

Mather, K., 197 

Mathildana newmanella, 18 

mating cages, 95 

McGuire, W. W., 5 

Megathymidae, 40 

Melanis pixe, 69 

mugration, o9 73, 137, 153, 312 

Miller, L. D., 144 

Miller, T. A., 95, 127 

Miller, W. E., 50 

mortality, 58 

Muller, J., 113 

Munroe, E., 11 

Muyshondt, A., 23, 159 

Nakamura, I., 305 

Nathalis iole, 121 

Necks Re W., 70, 91, 137, 218, 236 

Neophasia menapia, 236 

INeunzie He He 133 

new genera, 284 

new species, 33, 34, 36, 116, 119, 198, 
hare Lh 294-9285 

new subspecies, 12, 19, 253 

Noctuidae, 33, 116, 266, 301 

Nordmannia myratle, 305 

Notes and News, 151, 229, 309 

Notodontidae, 188 

Nutting, W. B., 143, 244 

Nymphalidae, 4, 12, 23, 47, 91, 126, 145, 
150, 159, 203, 207, 233, 234, 235, 
237, 243, 260, 312 

Nymphalis vau-album, 126 

obituaries, 152, 239 

Oecophoridae, 18 

Oiketicus toumeyi, 218 

Olethreutidae, 50 

Oliver, C. G., 260 

oviposition, 70 


Papilio troilus, 58, 147 
xuthus, 149 
Papilionidae, 44, 58, 70, 72, 147, 202, 237 
Peigler, R. S., 111, 114, 184 
Phoebis sennae eubule, 59 
Phyciodes tharos, 260 
Pieridae, 45, 68, 88, 121, 202, 234, 235, 
236, 252, 289 
Pieris occidentalis, 289 
protodice, 289 
Pimodes, 284 
insularis, 285 
Poanes massasoit chermocki, 19 
polymorphism, 91, 114 
polyphenism, 260 
Powell, J. A., 219 
predation, 147, 157, 236 
Prentiss, J. B., 187 
presidential addresses, 1, 245 
Proctowe Ne See L260, Ab: TAz 
Psychidae, 218 
pupae, 187 
Pyralidae, 11) 133, 2115 269: 284 
Rawson, G. W., 207 
Rickard, M. A., 5, 105 
Riodinidae, 46, 69 
Samson G., 12 
Sanson, N. B., 201 
Saturniidae, 95, 111, 114, 127, 131, 184, 
LS 272, 
Satyridae, 48, 68, 204, 243 
Satyrium boreale, 214 
liparops liparops, 213 
Schinia florida, 301 
Scriber, J. M., 58, 71 
Sevastopulo, D. G., 150, 151, 229 
Shapiro, A. M., 289 
Showalter, A. H., 312 
Siderone marthesia, 159 
Simmons, R. S., 19 
Sphingidae, 230, 264 
Sternburg, J. G., 131 
systematics, 11, 12, 19, 33, 50, 116, 188, 
DNAS NG 252. 22. aoa AG CSOD 
territorality, 147 
Thomisidae, 236 
Modd. Ee 2 188 
Toliver, M. E., 237 
Tuskes, P. M., 272 
Udea angustalis, 11 
Urbanus proteus, 60 
Urquhart, F. A., 59, 73, 153 
Urquhart, N. R., 59, 73, 153 
vacuum freeze-drying, 277 
Vanessa cardui, 312 
Vitula lugubrella, 133 
Wagener, W. H., Jr., 3 
Waldbauer, G. P., 131 
Wind, R. G., 239 
Zaretis callidryas, 159 


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ALLEN PRESS, INC. he NaS LAWRENCE, KANSAS 
Us. & 


CONTENTS 


PRESIDENTIAL AppRESs 1976—Wuat Insects Can WE IpENTIFY? 
Ronald W. Hodges 0 0 eee 


CONCERNING THE NAME ANTHOCARIS COLORADENSIS Hy. EDWARDS 
WITH DESIGNATION OF A NEW SUBSPECIES (PIERIDAE). Kurt 
Johnson oye ee 


PHOTOPERIODIC REGULATION OF SEASONAL POLYPHENISM IN PHYCIO- 
DES THAROS (NyMpHALIDAE). Charles G. Oliver _ 


LarRvAL FOODPLANTS AND LirE History Notes ror EicHur Morus 
FROM TExAs AND Mexico. Roy O. Kendall 


A Key To THe Last INstar LARVAE OF WEST COAST SATURNIIDAE. 
Paul M. Tuskes 


Low Cost Vacuum FREEZE-pRyING. Frank R. Hedges 


Two New Species oF Puycirinr Morus wirH DESCRIPTION OF A 
New Genus (Pyratmar). André Blanchard 


Tue BIoLocicAL STATUS OF NEARCTIC TAXA IN THE PIERIS PROTO- 
DICE-OCCIDENTALIS Group (PieRmDAE). Arthur M. Shapiro __ 


POPULATION STRUCTURE OF THE PRIMROSE Motu, SCHINIA FLORIDA 
(Nocruwae). Steven N. Handel 


FEMALE ANAL Harr Turr In NORDMANNIA MYRATLE (LYCAENIDAE): » 


EGG-CAMOUFLAGING FUNCTION AND TAXONOMIC SIGNIFICANCE. 
Ichiro Nakamura 


Nores AND NEws 


GENERAL NOTES 


Ecological notes on Celastrina ebenina (Lycaenidae). W. H. Wagner, Jr. 
and Amos H. Showalter 


INDEX 


260 


264 


272 
277 


284 


289 


301 — 


Volume 31 1977 Number 1 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


30 March 1977 


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Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA 


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J OURNAL OF 


Tue LEPIDOPTERISTS’ SOCIETY 


Volume 31 1977 Number 1 


STUDIES ON- THE CATOCALA (NOCTUIDAE) OF 
SOUTHERN NEW ENGLAND. V. THE RECORDS OF 
SIDNEY A. HESSEL FROM WASHINGTON, 
CONNECTICUT, 1961-1973 


THEODORE D. SARGENT 
Department of Zoology, University of Massachusetts, Amherst, Massachusetts 01003 


With the death of Sidney A. Hessel on 11 November 1974, lepidopter- 
ists lost one of their most enthusiastic and inspiring colleagues. This sad 
event also closed the pages on an unprecedented compilation of records 
on the moths of a single genus at a single location, for Hessel had faithfully 
noted all of the Catocala specimens taken at two light sources near his 
home on virtually every night of 12 seasons between 1961 and 1973. 

These records, portions of which have been previously published 
(Sargent & Hessel, 1970; Sargent, 1976), are summarized and analyzed 
here, particularly with a view to assessing (1) the variability in Catocala 
populations from year to year, (2) the extent of seasonal separation 
among the various species, and (3) the degree of stability in hindwing 
diversity across seasons. I also hope to demonstrate the usefulness of 
such records for the development and testing of hypotheses relating 
to the ecology of these moths. Specifically, I will propose a mechanism 
for the maintenance of stability in the frequencies of certain hindwing 
patterns, drawing upon applicable data from Hessel’s records. 

I hope that this paper will illustrate the value of complete and de- 
tailed records that extend over several seasons and thus will encourage 
others to gather similar data at their own locations. 


METHODS 


Washington is located in the Litchfield Hills of west-central Con- 
necticut. The collecting site itself was at the bottom of a narrow north- 
south valley through which an all-season stream flowed southward. The 


NS) 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TasLe 1. Numbers of individuals (N) of all Catocala species taken over 12 
seasons (1961-65, 1967-73) at Washington, Connecticut and the corresponding per- 
centages (%) of the Catocala sample. 


Species N % Species N % 
palaeogama 1795 17.45 cara 82 0.80 
residua ONG 11.83 ilia 67 0.65 
habilis 964 9.37 crataegi AT 0.46 
amica 790 7.68 dejecta 46 0.45 
concumbens 676 Gio parta 45 0.44 
ultronia 632 6.14 subnata 44 0.43 
grynea 466 4.53 unijuga 42. 0.41 
neogama 446 4.34 flebilis 40 0.39 
antinympha 44] 4,29 blandula 40 0.39 
retecta 403 3.92 coccinata 39 0.38 
serena 353 3.43 praeclara 38 0.37 
epione 264 2bt relicta 23 0.22 
obscura 224 2.18 similis 21 0.20 
andromedae PAID 2.06 amatrix 7 0.07 
judith PAIL 05) innubens 6 0.06 
mira 201 1.95 briseis 3 0.03 
micronympha 159 1.55 piatrix 1 0.01 
badia 127 25 vidua iL 0.01 
gracilis 114 If. 1taL cerogama ue 0.01 


site was surrounded by hills, mostly of mixed deciduous woodlands, but 
including patches of earlier seral stages that result from the periodic 
establishment and abandonment of farms and pastures. 

Most of the moths were obtained in a Robinson mercury vapor 
light-trap that was operated from dusk to dawn. The contents of this 
trap were checked each morning, and the number of specimens of each 
Catocala species was recorded. The majority of the specimens was re- 
leased near the trap iocation after examination, so some individuals may 
have been captured and recorded on more than one occasion. However, 
studies of color-marked Catocala have shown that very few specimens 
are recaptured under such circumstances (Sargent, 1976). A few records 
were obtained at a 15-watt fluorescent black-light, which was checked 
periodically during the evening, and these records were combined with 
the Robinson trap data in Hessel’s daily compilations. Both light sources 
were in operation from mid-March to mid-November each year (except 
for occasional 1-3 day absences). 

The species of Catocala were identified as keyed and described in 
Forbes (1954), except that gracilis and sordida were not always dis- 
tinguished; these species are considered together (as gracilis) through- 
out the present report. 

A total of 10,288 Catocala specimens of 38 species was recorded over 


VoLUME 31, NuMBER 1 3 


2400 

2000 
n 
_ 

~=¢ 1600 
= 
= 
2 
a 
= 

1200 
ue 
O 
far 
Lhd 
= 

> 800 
z 

400 


61 62 63 64 65 66> = 167 68 69 70 71 Tia. 73 


YEARS 


Fig. 1. Total number of Catocala taken each year at two light sources at Wash- 
ington, Connecticut. 


the 12 seasons, 1961-65 and 1967-73. The numbers of each species taken, 
ranked in decreasing order of abundance over the 12 seasons, are given 
ie fable! 


RESULTS AND DISCUSSION 


Annual Variations 


Analysis of Hessel’s records revealed considerable variation in the 
Catocala samples from year to year, despite essentially identical collect- 
ing procedures. These variations included changes in (1) the total 
abundance of all Catocala, (2) the relative abundance of particular 
species, and (3) the overall pattern of species abundance. 

The size of the Catocala sample fluctuated markedly from year to 
year, ranging from a low of 306 specimens in 1963 to a high of 2337 
specimens in 1971 (Fig. 1). No long-range trend of increasing or de- 
creasing Catocala abundance could be discerned against the erratic 
fluctuations in annual abundance. 

In addition to changes in total abundance, there was also considerable 


4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TaBLE 2. The highest and lowest annual totals, with the corresponding percent- 
ages of total Catocala taken, for those species having at least 100 records over 12 
seasons at Washington, Connecticut. 


Annual Totals 


High Low 
Species N % N % 

palaeogama 690 29.53 12 3.92 
residua 345 24.43 q 1.89 
habilis 159 13.81 23 6.22 
amica 281 12.02 10 Byte 
concumbens 130 5.56 12 0.85 
ultronia 220 9.4] 14 4.58 
grynea We S05 8 2.61 
neogama 37 Veo 4 0.86 
antinympha 182 Hel 13 2.45 
retecta 83 6.51 9 DA 
serena 124 5.31 0 — 

epione 69 5.41 5) 1.07 
obscura 28 1.20 3 0.98 
andromedae 40 3.14 2) 0.64 
judith | 5} 4.07 0 _ 

mira 76 BAD) 2; ()) SUF 
micronympha 43 3.05 i 0.21 
badia hil 1.16 D) 0.38 


variation in the relative abundance of particular species from year to 
year. The highest and lowest annual totals of those species for which 
there were at least 100 records over the 12 years of collecting are given 
in Table 2. These data suggest that the more common species in the 
overall totals were more erratic in terms of annual abundance than were 
the less common species. This suggestion is supported by comparisons 
of the relative annual frequencies of certain more and less common 
species (Fig. 2). It is apparent that the most abundant species overall 
exhibited explosive increases in numbers from time to time, whereas 
the less common species maintained rather constant frequencies over 
the years. These differences in relative abundance across years suggest 
differences in the mechanisms by which populations of various species 
are regulated, and this possibility certainly warrants further study. 

The 12-year totals of the Catocala species from this location (Table 1) 


> 


Fig. 2. Fluctuations in abundance from year to year of several Catocala species 
at Washington, Connecticut. Abundance is expressed as a percentage of the total 
Catocala recorded each year. The species considered range in status from abundant 
(A) to common (B) to uncommon (C). 


VoLuME 31, NuMBER 1 


PERCENT OF TOTAL CATOCALA PERCENT OF TOTAL CATOCALA 


PERCENT OF TOTAL CATOCALA 


40 


*— palaeogama (1795) A 
o———o habilis (964) 


YEARS 


40 
B 
e— concumbens (676) 
8 °——o antinympha (441) 
30 
25 
20 
i ° 
15 . 
10 
° 
5- p 
So. 
oO 
T T ae T 
70 7) 72 73 
YEARS 
40- 
(¢ 
35-4 
—e andromedae (212) 
o——o badia (127) 
30 
25 
20-4 
15 
10 
544 
St mieyG. & ° : : o 
eae T Tha Ph ee ic T ] ] i} | i I | | 
61 62 63 64 65 66 67 68 69 70 7) 72 73 


6 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


reveal a distribution of species abundance that is characteristic of most 
field samples of this sort, ie., a few very common and many uncommon 
species. In this case, the five most common species comprised over 50% 
of the records, whereas the 20 most uncommon species comprised less 
than 7% of the total sample. However, the extent to which this pattern 
was developed did vary from year to year. For example, the most com- 
mon species in 1962 (palaeogama) comprised 35% of the sample for 
that year, but the most common species in 1969 (concumbens) com- 
prised only 10% of the sample. At the other extreme, 15 species were 
recorded fewer than five times in 1963 (nine species occurred only once), 
whereas only six species were recorded fewer than five times in 1962 
(only one species occurred only once). 

These examples of annual variation in samples from a single location 
should illustrate the futility of making long-term assessments of Catocala 
populations on the basis of limited collecting. For even these records of 
Hessel, as extensive and complete as any known for the Catocala, will 
permit few conclusions regarding the status, or trends in the status, of 
the species at his location. This finding, however, is perhaps one of the 
most valuable to emerge from his records. As I have said elsewhere 
(Sargent, 1976), “Perhaps the lesson here is to view most general as- 
sessments of status in the Catocala as tentative.” 


Seasonal Occurrence 


One of the most interesting questions regarding the Catocala con- 
cems the nature of the isolating mechanisms that prevent hybridization 
among the many species which occur together at any one place. This 
problem has been discussed in detail elsewhere (Sargent, 1976), and it 
seems likely that many factors coact to isolate the various sympatric 
species. These factors include differences in daily and seasonal activity 
periods, and in courtship and mating behaviors. Here we will be con- 
cerned with only one of these factors—differences in seasonal occurrence. 

Hessel’s daily records, which cover the entire Catocala season for many 
years, are particularly useful for analyses of such differences; for his 
records, especially when summed across the years, provide the large 
sample sizes essential for the detection of relatively small seasonal offsets. 

Hessel took adult Catocala over a four-month period (July—October), 
but most of his records fell between mid-July and mid-September ( Fig. 
3). A total of 33 species was taken during the second half of August, and 
as many as 21 species were recorded on a single night during that period 
(Sargent & Hessel, 1970). Clearly, many species had overlapping flight 
seasons, 


VoLUME 31, NuMBER 1 7 


NO. SPECIES 16 30 32 33 29 20 16 


3000 


2500 


< 
—_ 
< 
Y 2000 
O 
= 
< 
U 
te 
O 
a4 1500 
tL! 
co 
= 
=) 
z 
—_ 
=< 1000 
O 
~— 
500 


JUL JUL AUG AUG SEPT SEPT OcT 
1-15 16-31 1-15 16-31 1-15 16-30 1> 


SUCCESSIVE HALF-MONTH PERIODS 


Fig. 3. The total number of Catocala recorded during successive half-month 
periods of the season at Washington, Connecticut, summed over 12 years. The 
number of species taken during each half-month period is given at the top of the 
graph. 


However, if one compares the median dates of capture of those spe- 
cies for which there were 24 or more records (Fig. 4), some interesting 
seasonal offsets between certain species become apparent. Thus, for 
example, approximately a month separates the median dates of capture of 
dejecta (3 August) and retecta (2 September), and serena (11 August) 
and habilis (14 September). In these cases, three-quarters of the records 
of the earlier species occurred before the first quarter of records for the 
later species. Other closely related species pairs exhibiting marked dif- 
ferences in median capture dates include blandula (13 July) and mira 
(1 August), swbnata (10 August) and neogama (6 September), residua 
(19 August) and obscura (6 September), and concumbens (25 August) 


8 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


JULY SEPT 
SPECIES 2, 6 ,10,14,18,22,26,30, 3, 7,11,15,19,23,27,31, 4 , 8, 12,16,20,24,28, 2, 6 ,10,14,18,22,26, 30 


blandula 
coccinata 
micronympha 
crataegi 
mira 

amica 
badia 
gracilis 
antinympha 
praeclara 
andromedae 
epione 
palaeogama 
dejecta 
judith 
ultronia 
grynea 
subnata 
serena 

ilia 
residua 
unijuga 
concumbens 
flebilis 
retecta 
neogama 
obscura 
parta 

cara 


habilis 


Fig. 4. Seasonal occurrence of Catocala species at Washington, Connecticut 
based on records summed over 12 seasons. The lines run from the earliest to the 
latest dates of capture, and quartile dates are indicated by a dot (median) and 
dashes (first quarter, third quarter). Only species for which there were 24 or 
more records are considered, and these are arranged in descending order on the 
basis of a seasonal sequence from early to late. 


and cara (13 September). Assuming that most matings occur near the 
beginning of the flight season of a species, it seems likely that seasonal 
offsets such as these must contribute to the reproductive isolation of the 
species involved. 

On the other hand, certain other pairs of closely related species had 
nearly identical median dates of capture. Among such pairs were badia 
(1 August) and antinympha (2 August), gracilis (2 August) and an- 
dromedae (2 August), and praeclara (2 August) and grynea (6 August). 
Clearly such species pairs must depend on isolating mechanisms other 
than seasonal separation. 

Complete understanding of the complex of factors that isolate all of 
the Catocala species at any one location must await much more study. 
But these records of Hessel suggest that seasonal separation is one of the 
factors involved in certain cases. 


VoLUME 31, NuMBER 1 9 


TaBLE 3. Distribution of Catocala in five hindwing groups at Washington, 
Connecticut (1961-65, 1967-73). 


Hindwing Group Species Number % 
1 relicta 23 0.22 
2 epione, judith, flebilis, obscura, residua, 
retecta, dejecta, vidua, andromedae 2618 25.45 
3 piatrix, antinympha, badia, habilis, serena, 


palaeogama, subnata, neogama, 
cerogama, gracilis, crataegi, 
mira, blandula, grynea, praeclara, 


similis, micronympha, amica 6048 58.79 
4 innubens, ilia, parta, briseis, unijuga, 

coccinata, ultronia 834 8.11 
5 cara, concumbens, amatrix 765 7.44 


Hindwing Diversity 


The forewings of many Catocala species are strikingly variable (poly- 
morphic), but the hindwings are essentially invariable (monomorphic) 
within any species. However, hindwing diversity across species is sub- 
stantial, and this matter has been the subject of considerable prior study 
(Sargent, 1969, 1973, 1976; Sargent & Owen, 1975). The records of 
Hessel provide an opportunity to analyze the occurrence of various 
hindwing types at a single location in considerable detail. This analysis 
in turn prompts some speculation regarding the apparent maintenance 
of stability in hindwing diversity at this location. 

Dr. Denis Owen and I recently analyzed the frequencies of various 
hindwing types occurring in large Catocala samples taken at mercury 
vapor lights at four localities in eastern North America (Sargent & Owen, 
1975). For purposes of our analysis, the hindwing patterns were arbi- 
trarily divided into five groups: (1) black and white, banded; (2) black, 
unbanded (on upper surface); (3) yellow to yellow-orange and black, 
banded; (4) orange-red to red and black, banded; and (5) pink and 
black, banded. The frequency distribution of these hindwing types 
was remarkably similar at each of the localities we considered, despite 
marked differences in species composition. These frequencies closely 
resembled those obtained at Washington, Connecticut, as compiled from 
Hessel’s total records (Table 3). 

This apparent stability in hindwing diversity at different locations was 
more simply expressed by combining the hindwing groups into an achro- 
matic assemblage (groups 1 and 2) and a chromatic assemblage (groups 
3, 4, and 5). This division emphasizes the most obvious hindwing di- 
chotomy in the Catocala, i.e., the presence or absence of color. And the 


10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


frequency of occurrence of these two hindwing types was nearly con- 
stant across localities, with achromatic individuals consistently compris- 
ing ca. 20% of the total Catocala samples (Sargent & Owen, 1975). 

We interpreted the apparent stability in hindwing diversity at various 
locations as a result of selective predation, especially by birds, and 
argued that the observed frequencies function to “confuse” predators, 
presumably by introducing the element of anomaly (the unexpected) 
into the overall predator-prey system (Sargent & Owen, 1975). Thus, 
for example, achromatic hindwings might serve as effective startle de- 
vices only if they comprised no more than ca. 20% of the total Catocala 
hindwings encountered. At higher frequencies, predators might come 
to expect such a hindwing pattern, and predation would increase until 
the frequency was again returned to 20% of the total. Presumably, such 
selection pressure would eventually result in stabilization of the different 
hindwing types at optimal frequencies with respect to predation. 

In further analyzing Hessel’s records, I will consider only the achro- 
matic and chromatic hindwing groups, since these are the most easily 
defined and perhaps most meaningful categories with respect to Catocala 
hindwing diversity. And since most of the species with achromatic hind- 
wings feed as larvae on the Juglandaceae (hickories, Carya, and walnuts, 
Juglans), particular attention will be devoted to the species that utilize 
those foodplants. 

The percentage of specimens with achromatic hindwings at Washing- 
ton, Connecticut ranged from 16.76% in 1973 to 36.24% in 1961 and 
averaged 25.67% over the 12 years. However, a more striking constancy 
of achromatic hindwings can be demonstrated when only those Catocala 
whose larvae feed on the Juglandaceae are considered. This analysis 
excludes only two achromatic species from Hessel’s totals (relicta, a 
Salicaceae feeder; and andromedae, an Ericaceae feeder), leaving eight 
achromatic (epione, judith, flebilis, obscura, residua, retecta, dejecta, 
and vidua) and six chromatic species (piatrix, habilis, serena, pala- 
eogama, subnata, and neogama). All of the chromatic species in this 
case have yellow-orange and black, banded hindwings. 

The percentage of individuals with achromatic hindwings among 
these Juglandaceae-feeding Catocala was remarkably constant from year 
to year, despite considerable variation in the species composition and 
the number of individuals taken each year (Fig. 5). Such stability in 
the occurrence of achromatic hindwings suggests the operation of a con- 
trol mechanism related in some way to predation. 

Two possibilities immediately come to mind: (1) predators con- 
sistently select Catocala such that trapped samples will reveal a constant 


VoLUME 31, NuMBER 1 4 


100 
Us 
50 


25 


PERCENT ACHROMATIC 


750 


500 


NUMBER OF INDIVIDUALS 


250 


YEARS 


we achromatic chromatic 


Fig. 5. Number of Juglandaceae-feeding Catocala taken each year at Washing- 
ton, Connecticut with achromatic and chromatic hindwings distinguished (bottom) ; 
and the corresponding percentages of individuals with achromatic hindwings (top). 


frequency of achromatic hindwings; and (2) the moths themselves, in 
response to long-term predator selection, have evolved the means of 
maintaining a constant frequency of achromatic hindwings. Both pos- 
sibilities pose difficulties, but it seems particularly unlikely that the 
stability of achromatic hindwings in trapped samples is entirely a product 
of immediate predator selection; for this would assume that few moths 
are trapped prior to their exposure to intense predator selection, and such 
an assumption seems clearly unreasonable. 

Thus, the possibility that the moths themselves are maintaining a 
constant frequency of achromatic hindwings must be examined. This 
possibility is rendered particularly perplexing in view of the fact that 


NUMBER RECORDED 


NUMBER RECORDED 


NUMBER RECORDED 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


700 


e— palaeogama 


o——o residua 


600 


5¢0 


400 


300 


200 


100 


YEARS 
125 
e—— serena B 
o—o judith 
100 
75 
50 
25 
. 
i} 
ih T 
61 62 63 64 65 66 
YEARS 
1004 
*——* neogama i 
o o retecta 
° 
80 
60 
° 
40 
o 
o 
20- o 
= T T T iT T as 
6) 62 63 64 65 66 


VoLUME 31, NuMBER 1 13 


none of the species involved are polymorphic with respect to hindwing 
types, and, consequently, mechanisms that would maintain a balanced 
polymorphism within a species (Ford, 1964) cannot be operating. How, 
then, is a stable relationship between two hindwing types to be achieved 
in a complex of species, each monomorphic with respect to hindwing 
type, and each highly variable with respect to abundance from year to 
year? 

One theoretical possibility in a system wherein each species regulates 
its own density by assessing and responding to the density of every 
other species present. Such density regulation might be envisioned for 
a single species (Wynne-Edwards, 1962) but seems entirely implausible 
for an assemblage of species, given the complex social behaviors required 
in such a control system. 

It may be, however, that an overall stability in the relationship between 
achromatic and chromatic hindwings could be achieved on the basis of 
simpler interactions between or among certain species. If, for example, 
the achromatic and chromatic species were paired and the members of 
each pair exhibited parallel fluctuations in annual abundance, then a 
stable relationship between the two hindwing types would result. 

That such pairings of species may exist is suggested by the nearly 
identical fluctuations in annual abundance of certain achromatic and 
chromatic species (Fig. 6). These similarities suggest that the two 
species involved in each case are responding to environmental variables 
in the same fashion and thus may have identical, or nearly identical, 
ecological niches. This suggestion, however, seems to raise problems 
with respect to the so-called competitive exclusion principle, i.e., the 
ecological dictum that two species cannot share the same niche, since 
competition between them should eventually exclude the less well- 
adapted species (see discussion in any ecology text, e.g., Ricklefs, 1973). 
This principle clearly assumes that the two species compete for some 
limiting resource, usually food. The Catocala, however, may be limited 
by predation rather than the availability of food, and in that case the 
competitive exclusion principle would not apply. 

The fact that many Catocala species may utilize the same hostplant 
suggests that food is not generally limiting for these moths. Many of the 


< 


Fig. 6. Numbers of individuals of three pairs of Juglandaceae-feeding Catocala 
species taken each year at Washington, Connecticut. The species are paired on the 
basis of similarities in abundance across years, and each pair includes a species 
with achromatic hindwings (open dots) and a species with chromatic hindwings 


(solid dots). 


14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Juglandaceae feeders under consideration here, for example, are known 
to feed on shagbark hickory (Carya ovata) (Sargent, 1976). On the 
other hand, the fact that the ova, larvae, and adults of most Catocala are 
highly cryptic implies that predation has long been substantial on these 
insects. There is some evidence for heavy bird predation on the adults, 
since beak-damaged individuals may comprise as much as 4% of trapped 
specimens (Sargent, 1973). Such beak-damaged individuals have es- 
caped from their predators, but presumably many more individuals are 
actually eaten by birds. I have shown that naive blue jays (Cyanocitta 
cristata (L.)) will quickly learn to capture Catocala, rarely losing indi- 
viduals after 12-15 experiences with these moths (Sargent, 1973). 

It seems likely that the simultaneous presence of two species with dif- 
ferent hindwing types would be advantageous to both species with 
respect to the effectiveness of their hindwings as startle devices, for 
birds are clearly less effective predators when confronted with the novel 
or unexpected in their prey (Sargent, 1976). And since novelty and 
anomaly are functions of scarcity, the advantage of any one hindwing 
type should increase as the numbers of the other hindwing type increase. 

Thus, it follows that two species with different types of hindwings 
might share the same niche, and the advantage of each species with 
respect to predation would increase as the other species increased in 
abundance. In such a situation, neither species should act to exclude 
the other from the niche, and each should become as abundant as other 
limitations (climate, parasites, etc.) permit. Given that the two species 
are Closely related and are adapting to the same niche, it should not be 
surprising to find them occurring in approximately equal numbers, as 
seems to be the case in several instances at Washington, Connecticut 
(Fig. 6). 

The system envisioned here would result in stable relationships be- 
tween pairs of species with achromatic and chromatic hindwings and 
would not require intrinsic mechanisms for the assessment or adjustment 
of population densities. One species would need only to adapt to a niche 
already occupied by a species with a different type of hindwing. If 
such pairs of species comprised a substantial portion of the total of 
species under consideration, then a stable overall relationship between 
the different hindwing frequencies would be expected. 

There remains the question of why hindwing diversity, if it is such an 
advantage with respect to predation, has not developed within any 
species. The answer must be that there is an even greater advantage 
associated with hindwing monomorphism at the species level. This sug- 
gests that the hindwings function as specific recognition devices, per- 


VOLUME 31, NuMBER 1 15 


haps serving as releasers during courtship and mating behaviors, and 
thus act to isolate various species. If the hindwings do serve as isolating 
mechanisms, it seems possible that sympatric speciation on the basis 
of hindwing differentiation might occur on occasion in the Catocala. 
Sympatric speciation might then account for the phenological similarities 
we have seen in certain pairs of species with different tvpes of hind- 
Wings. 

These ideas regarding the maintenance of stability in hindwing diver- 
sity, though often quite speculative, are based on data that Hessel 
acquired over many years at Washington, Connecticut. I hope that 
other workers will be stimulated to test these ideas by acquiring addi- 
tional data and conducting further studies on the Catocala at their 
locations. Whether such studies support or refute the ideas developed 
here, the results can only advance our understanding of these moths. 
And in this way, the records of Hessel will make their most important 
contribution. 


SUMMARY 


The late Sidney A. Hessel of Washington, Connecticut recorded all 
of the Catocala taken at two light sources near his home over 12 seasons 
(1961-65, 1967-73). Totals of 38 species and 10,288 individuals were 
recorded. 

The Catocala populations at this location varied considerably from 
year to year. These annual variations included changes in the total 
number of Catocala taken, the relative abundance of particular species, 
and the overall pattern of species abundance. The more common species 
exhibited more erratic fluctuations in annual abundance than the less 
common species. It is concluded that limited collecting will not permit 
long-term assessments of status and trends in Catocala populations. 

The Catocala season at Washington extended from July—October, and 
most of the species had overlapping flight seasons. However, detailed 
analyses, including comparisons of the median dates of capture of 
various species, suggested that certain closely related species might be 
isolated in part by seasonal offsets. 

The frequency distribution of various hindwing types at Washington 
is summarized. The percentage of individuals with achromatic hind- 
wings, particularly within the group of Juglandaceae-feeding species, 
remained remarkably stable over the years. A possible mechanism for 
the maintenance of that stability is proposed, based on observations of 
nearly identical fluctuations in annual abundance of certain pairs of 
species that included one member with achromatic hindwings and one 


16 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


member with chromatic hindwings. It is suggested that the species in 
each of these pairs may share the same ecological niche, and may have 
arisen sympatrically. 


LITERATURE CITED 


Forses, W. T. M. 1954. Lepidoptera of New York and neighboring states. ILL. 
Noctuidae. Cornell Univ. Agri. Exp. Sta. Memoir 329. 433 p. 

Forp, E. B. 1964. Ecological genetics. Methuen, London. 335 p. 

Rickters, R.E. 1973. Ecology. Chiron Press, Newton, Mass. 861 p. 

SARGENT, T. D. 1969. A suggestion regarding hindwing diversity among moths 
of the genus Catocala (Noctuidae). J. Lepid. Soc. 23: 261-264. 

1973. Studies on the Catocala (Noctuidae) of southern New England. 

IV. A preliminary analysis of beak-damaged specimens, with discussion of 

anomaly as a potential antipredator function of hindwing diversity. J. Lepid. 

Soc. 27: 175-192. 

1976. Legion of night: The underwing moths. Univ. Mass. Press, 

Amherst, Mass. 222 p. 

& S. A. Hessen. 1970. Studies on the Catocala (Noctuidae) of southern 

New England. I. Abundance and seasonal occurrence of the species, 1961— 

1969. J. Lepid.-Soc. 24: 105-117. 

& D. F. Owen. 1975. Apparent stability in hindwing diversity in samples 
of moths of varying species composition. Oikos 26: 205-210. 

WynneE-Epwarps, V. C. 1962. Animal dispersion in relation to social behaviour. 
Oliver & Boyd, Edinburgh. 653 p. 


MELITAEA SAXATILIS MOD. “SASSANIDES” (NYMPHALIDAE) IN IRAN: 
CONFIRMATION OF AN OLD RECORD 


On 5 July 1974 I took eight adult specimens of Melitaea saxatilis mod. “sassanides” 
(Higgins) in Alborz, Mount Damavand, northern Iran. The butterfly was re- 
stricted to the height of 4000 m, near the third mountaineer’s shelter where a steep 
rock slope was covered by a few scattered species of Cruciferae, Labiatae and 
grasses. The adults were feeding on the few Labiatae flowers that existed. No early 
stages were found. 

Higgins (1941) in his “An illustrated catalogue of the Palearctic Melitaea”’ (Trans. 
Roy. Ent. Soc. London 91: 175-365) mentioned that the only specimens he saw 
were amongst the ex. coll. Grum-Grshimaile collection at the British Museum. 
They were collected on 29 June 1894 and no additional record has ever been 
published. Personal contact with Dr. Higgins and the literature confirm this 
claim. Unfortunately, due to the change of the weather and the dangerous location 
of the butterfly habitat, I was not able to collect a sample of the vegetation or 
investigate farther. 


JaAvAD Hasnemr Tarresut, 4 Whiteheads Lane, Bradford-On-Avon, Wiltshire, 
England. 


VoLUME 31, NUMBER 1 1 or 


DISTRIBUTION AND BIOLOGY OF A PLEISTOCENE RELICT: 
OCHLODES YUMA (HESPERIIDAE ) 


JAMEs A. Scott!, OAKLEY SHIELDS”, AND Scorr L. EL.is?® 


The purpose of this paper is to summarize our knowledge of the 
distribution, life history and behavior of Ochlodes yuma (Edwards), 
a little-known western United States skipper. 

Larval foodplant. Phragmites communis Trin., the Common Reed, is 
a large (ca. 2m), cosmopolitan, perennial grass forming canelike thickets 
in wet places, with a wind-dispersed fruit and spreading rhizome (Mason, 
1957; Polunin, 1960, p. 98). It occurs in Europe, Asia, Africa, the 
Americas, and Australia but is absent from many islands (Ridley, 1923). 
It may be the most widely distributed flowering plant in the world 
(Polunin, 1960, p. 98; Sculthorpe, 1967, p. 366). In western United States 
it occurs along watercourses, irrigation canals, freshwater springs, and 
alkaline or even sulphurous seeps. 

C. Don MacNeill, J. M. Burns, and. J. F. and T. C. Emmel raised O. 
yuma larvae on P. communis leaves in the Central Valley of California 
(Arnaud, 1960; Emmel & Emmel, 1973). J. Scott observed oviposition on 
leaves at the base of the plant in San Juan Co., Utah. J. F. Emmel 
and C. Sekerman found ova and larval shelters with leaf edges fastened 
together to form a tube at Surprise Canyon, Inyo County, California. J. F. 
Emmel found many last instar larvae in larval shelters at Mesquite Spring, 
Inyo Co., Calif. O. yuma is extraordinarily restricted to P. communis; it 
is almost always found in or within a few meters of stands of P. communis. 
We know of only one record away from P. communis, a male from Home- 
wood Canyon, Inyo Co., California, 0.5 mile from P. communis. 

Habitats with O. yuma have only one thing in common: the presence 
P. communis. In the Central Valley of California colonies occur along 
estuaries, sloughs, and canals. Colonies occur along the Colorado and 
other rivers in the Great Basin. In desert parts of the Great Basin, colonies 
are to be found at springs, on alkaline salt-encrusted flats with sufficient 
subsurface water to support Phragmites, and in semi-irrigated streamside 
marshes. One colony (near Mina, Mineral Co., Nevada) was at a sulfurous 
spring, and another (Surprise Canyon, Inyo Co., California) was at a seep 
with Phragmites on a hillside. Agricultural activity seems to have 
increased the habitat for O. yuma along the Colorado River drainage in 


Colorado. 


1 Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, Colorado 80523. 
2 Department of Entomology, University of California, Davis, California 95616. 
3 Ecology Consultants, Inc., P.O. Box 1057, Fort Collins, Colorado 80521. 


18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Distribution records. Dates are given only for those records not included 
in Fig. 2. 


ARIZONA. Coconino Co.: Little Colorado River at Cameron, 4100’, K. Roever, and 
Tilden, 1957; 1 mi. W of Tuba City, 4500’, K. Roever; Indian Gardens, Grand Canyon, 
Tilden, 1957; Pima Co.: Tucson, Tilden, 1957 (this record may be dubious, as Mr. 
Kilian Roever has not found it there ). 

CauirorNniA. Calaveras Co.: Sand Flats, Tilden, 1957; Contra Costa Co.: Antioch, 
F. H. Chermock, J. Scott, and Tilden, 1957; Bethel Island, J. Scott; Jersey Island, 
N. La Due; Inyo Co.: near Aberdeen, 12 mi. N of Independence, 3840’, S. L. Ellis 
and S. A. Johnson; Antelope Spring, J. S. Buckett; Darwin Falls, J. W. Tilden, L. M. 
Martin, S. S. Nicolay, R. Holland; Hank Lubkin Ranch, Cartago, C. Henne; Homewood 
Canyon, Argus Range, 3600—4000’, J. F. Emmel & O. Shields; Hunter Canyon, Saline 
Valley, Los Angeles County Museum; 4 mi. NE of Independence just W Owens River, 
S. L. Ellis & S. A. Johnson; 2 mi. N Lone Pine, J. S. Buckett; Limekiln Spring, 4000’, 
Surprise Canyon, Panamint Range, J. F. Emmel & O. Shields; Lone Pine Ranger 
Station road, 5 mi. W Lone Pine, 6500’, S. L. Ellis & S. A. Johnson; Mesquite Spring 
Campground, J. F. Emmel; Olancha, Comstock, 1927; 1 mi. N of Shoshone on Hwy. 
127, 1630’, J. F. Emmel & O. Shields; Deep Springs, Tilden, 1957; Whitney Portal nr. 
Lone Pine, S. L. Ellis; Haiwee, Tilden, 1957; Owens Lake, Tilden, 1957; Wyman 
Creek Canyon, White Mts., 6000’, J. F. Emmel & O. Shields; Mono Co.: Farrington 
Camp, Tilden, 1957; Mammoth Camp, Tilden, 1957; Sacramento Co.: Bannon Island, 
F. H. Chermock; South Stone Lake, A. M. Shapiro; Beach Lake, A. M. Shapiro; 
Jackson Slough Road, Brannan Island, J. Scott; Sherman Island, C. D. Ferris, W. 
Howe, R. Davis; Twitchel Island, N. La Due; Elkhorn Slough, C. D. Ferris; San 
Bernardino Co.: Topock Marsh, 15 mi. SSE Needles, 500’, K. Roever; San Joaquin 
Co.: Bishop Tract, J. Scott; Empire Tract, J. Scott; Solano Co.: Suisun Slough, A. M. 
Shapiro; Stanislaus Co.: Modesto, Tilden, 1957. 

Coxtorapo. Delta Co.: Austin, 5000’, S. L. Ellis; Columbine Ranch Rd., 3 mi. SW 
Hotchkiss, 5750’, S. L. Ellis; Federal Fish Hatchery, SE of Lazear, N. Fk. Gunnison 
River, 5300’, S. L. Ellis; Leroux Creek, #4 Ditch takeout, 5700’, S. L. Ellis; Mesa Co.: 
1 mi. NE jet. I-70 & Hwy. 65, J. Scott; 5 mi. S Debeque, between Debeque & Cameo, 
Colorado River, J. Scott; Unaweep Canyon nr. Gateway, 6300’, S. L. Ellis, J. Scott; 
Moffat Co.: Echo Park, Dinosaur National Mon., 5300’, J. F. Emmel, O. Shields, S. L. 
Ellis; Montrose Co.: Hwy. 90, 10 road mi. NE Naturita, S. L. Ellis & O. Shields; W. 
Paradox Creek, nr. Paradox, 5400’, S. L. Ellis, S. A. Johnson; Rio Blanco Co.: White 
River, cotypes of scudderi. 

Nevapa. Clark Co.: Cold Creek, Spring Mts., 6200’, A. Austin; Corn Creek, J. F. 
Leser; Corn Creek Station, Desert Big Game Refuge Hdq., 3000’, K. Roever, O. 
Shields, P. Herlan; Moapa, 1600’, K. Roever; Logandale, P. Herlan, J. F. Leser; 
Overton, P. Herlan; Rogers Spring, 8 & 12 mi. S of Overton, P. Herlan; Stewart 
Springs, ca. 1 mi. W of Overton arm of Lake Mead, P. Herlan; Tule Springs, ca. 10 mi. 
N of Las Vegas city limits, A. Austin, K. Roever; Whitney Mesa, J. F. Leser; Elko Co.: 
21 mi. S of Bear Creek Summit (August), P. Herlan; Esmeralda Co.: Lida Summit, 
P. Herlan; Lander Co.: Humboldt River NE of Battle Mtn. (August 5), J. Scott; 
Lincoln Co.: 2 mi. N of Caliente, J. F. Emmel & O. Shields; Mineral Co.: at the 
mouth of Cottonwood Canyon, 5.5 mi. SW of Hawthorne, P. Herlan; 4 mi. S of Mina, 
K. Roever; Whiskey Flats on the Pole Line Rd. 15 mi. S of Hawthorne, P. Herlan; 
Nye Co.: Beatty, J. Scott; 5 mi. N of Beatty, 3500’, K. Roever; 5.8 mi. NE of Currant, 
J. Scott. 

Urau. Emery Co.: San Rafael River, jct. I-70, J. Scott; Garfield Co.: Calf Creek, 
12 mi. S of Boulder, K. Roever; 1 mi. W Henrieville, K. Roever; Grand Co.: 12.5 mi. 
NE jct. Hwy. 128 & Castleton road, Hwy. 128, 3800’, S. L. Ellis, O. Shields; Kane 
Co.: N of Glendale, J. F. Emmel & O. Shields; 2 mi. S of Kanab, 4800’, K. Roever; 


VoLUME 31, NuMBER 1 19 


TABLE 1. (Continued) 


San Juan Co.: NE ject. Hwy. 160 & Hatch Wash, J. Scott; Uintah Co.: Green River 
at Split Mtn., C. J. & B. V. Durden; nr. Jensen P. O., O. A. Paterson (figured by 
Holland, 1931, plate 53), in Carnegie Museum; Merkley Park, R. E. Stanford; Wash- 
ington Co.: ca. 1 road mi. E of Park Headquarters on Hwy. 15, Zion Nat. Park, 
along Clear Creek, 4500’, J. F. Emmel & O. Shields. 


Distribution. Tilden (1957) presented some records. We now know 
that O. yuma occurs in central California, Nevada, Utah, western Colo- 
rado, and northern Arizona (Table 1 & Fig. 1). 

We have found no geographic variation, and treat Ochlodes scudderi 
Skinner as a synonym of O. yuma. There is some individual variation in 
the width of the dorsal forewing dark border, and in size. Types and type 


Fig. 1. Distribution records of O. yuma in western U.S. 


20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


CENTRAL CALIFORNIA 


E. CALIFORNIAY 


NUMBER S, NEVADA 


OF 
RECORDS 


i i 5 ARIZONA 
| 2) UAT ee 


E, UTAH, 
COLORADO 


O Ne, OCWO by CO On 


JUNE JULY AUG Sera 


Fig. 2. Histograms of daily collection records (ignoring numbers seen or collected ), 
grouped into four-day intervals, from late May—early October. 


localities of yuma (Inyo Co., Calif.) and scudderi (Rio Blanco Co., Colo- 
rado) were treated by Tilden (1961) and Brown (1957). 

Colonies usually are very isolated from each other. Like Speyeria 
nokomis nokomis (Edwards), another western United States butterfly 
found at isolated springs, the current colonies seem to be relicts of a 
formerly widespread Pleistocene distribution. All of the records we have 
are within the drainage basin of the Colorado River during the Pleisto- 
cene, except the records from the Central Valley of California, where it 
may have been introduced from the Great Basin during this century. It 
has only recently been discovered to occur in the Central Valley (Tilden, 
1957). 

Colonies are often very small; in several cases the area of plants was 
only about 30 x 10 m, and at Mesquite Spring, Inyo Co., California, the 
isolated Phragmites patch was only 1 X 5 m in size. Most of the colonies 
in the Great Basin are many miles from other Phragmites patches. The 
persistence of these isolated colonies is amazing; it has been thousands of 
years since wetter Pleistocene conditions may have permitted more 
extensive populations to exist. 

Time of emergence. There are two broods in California and southern 
Nevada (Fig. 2). In the eastern part of the range there is only one brood 


VoLuME 31, NuMBER 1 21 


(Fig. 2); in Colorado peak numbers along the Colorado River are several 
weeks later than at sites farther from the river. Records are too few from 
southern Utah and Arizona to determine the number of broods. Males 
slightly precede females in emergence by a few days as in most butterflies. 

Behavior. O. yuma is a perching species (Scott, 1974), defined as a 
mate-iocating strategy in which males rest at characteristic sites and 
investigate passing objects in search of females that fly to these sites to 
mate. O. yuma males rest usually on P. communis leaves 1-2 m above 
ground, in a low spot among the P. communis or, when the plants grow on 
a river bank, on leaves or sometimes boulders on the bank side of the 
plants. Males sometimes patrol among the plants. Males investigate 
passing objects, usually other males, then usually rest in the vicinity of 
their previous resting site. Males show perching behavior at all times of 
day. We found a copulating pair in Moffat Co., Colorado, at 1340 
(24-hr. standard time). 

Adults have been observed feeding on flowers of yellow Chrysothamnus 
nauseosus (Pursh) Britton, Grindelia sp., and Helianthus sp., reddish 
purple Polygonum pennsylvanicum, Cirsium sp., and Asclepias sp., rose- 
purple Arctium minus Schk., and bluish Aster sp. 

Parasites. A tachinid larval parasite from Contra Costa Co., California 
was identified as Spathidexia dunningi (Coquillet) (Arnaud, 1960). 


ACKNOWLEDGMENTS 


We thank A. T. Austin, F. H. Chermock (now deceased), C. J. Durden, 
Ceo ethers. |r. Emmel, C: Henne, P. J. Herlan, N: La Due, S. 5S. 
Nicolay, K. Roever, and R. Stanford for distribution records. 


LITERATURE CITED 


ARNAuvD, P. H., Jr. 1960. A review of the genus Spathidexia Townsend ( Diptera: 
Tachinidae). Wasmann J. Biol. 18: 1-36. 

Brown, F. M. 1957. The type-locality for Ochlodes yuma. Lepid. News 11: 
153-154. 

Comstock, J. A. 1927. Butterflies of California. Published by the author. 

EmMeEL, T. C. & J. F. Emmet. 1973. The butterflies of southern California. Natural 
History Museum, Los Angeles, Calif. 148 p. 

Hotuanp, W. J. 1931. The butterfly book. Garden City, New York. 424 p. 

Mason, H. L. 1957. A flora of the marshes of California. Univ. Calif. Press, 
Berkeley & Los Angeles. 878 p. 

Potuni, N. 1960. Introduction to plant geography. McGraw-Hill Book Co. New 
York. 640 p. 

Ripotey, H.N. 1923. The distribution of plants. Ann. Bot. 37: 1-29. 

Scott, J. A. 1974. Mate-locating behavior of butterflies. Amer. Midl. Nat. 9L: 
103-117. 

, S. L. Exuis, & D. Err. 1968. New records, range extensions, and field 

data for Colorado butterflies and skippers. J. Lepid. Soc. 22: 159-171. 


bo 
bo 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ScuLTHorPE, C. D. 1967. The biology of aquatic vascular plants. Edward Arnold, 
London. 610 p. 

TiwvEN, J. W. 1957. Taxonomic history and distribution of Ochlodes yuma. Lepid. 
News 11: 151-152. 

. 1961. Studies in the genus Ochlodes Scudder. I. The type material of the 

North American species (Lepidoptera: Hesperiidae). Entom. News 72: 37-45. 


BIZARRE CAPTURE OF A BUTTERFLY BY AN AMBUSH BUG 


Pyle (1973, J. Lepid. Soc. 27: 305-307) reported the communal feeding of am- 
bush bugs (Hemiptera: Phymatidae: Phymata sp.) upon a single adult silver 
bordered fritillary, Boloria selene Denis & Schiffermiiller (Nymphalidae). He stated 
that due to the small size of the ambush bugs relative to the butterfly that the 
“means of actual capture. . .baffled me.” Personal observation of a similar situation 
has revealed a possible mode of capture. 

On 11 November 1968 at the Brackenridge Field Laboratory of the University of 
Texas at Austin (within Austin), I observed the capture of a dogface butterfly, 
Colias (Zerene) cesonia Stoll (Pieridae), by an ambush bug, Phymata fasciata 
(Gray), on an inflorescence of cowpen daisy, Verbesina encelioides Cav. (Gray) 
(Compositae). C. cesonia is common in this area at this season and frequently 
visits inflorescences of V. encelioides for nectar. One particular butterfly was ob- 
served to visit several inflorescences in rapid sequence (very little time is spent at 
a single inflorescence). Upon approaching one inflorescence, the butterfly quickly 
dipped down to the plant but did not rise immediately to fly to another. Instead, 
a rapid beating of the wings ensued with the body of the butterfly remaining 
stationary. Shortly, the butterfly ceased movement but later began beating its wings 
again. 

Investigation of the inflorescence revealed that the butterfly was being held by 
its proboscis which was caught fast in one of the foretibia of the bug. This impasse 
(butterfly unable to escape, bug unable to consume a meal) continued for at least 
fifteen minutes, after which time observations ceased. Possibly the bug could have 
maneuvered its beak into position to pierce the body of the butterfly. Great strength 
and elasticity of the proboscis as illustrated by C. cesonia would indicate that the 
proboscis probably would not break (permitting freedom but in a mutilated condi- 
tion). Body length of the bug was about 10 mm while that of C. cesonia averages 
about 23 mm. 


Raymonp W. Neck, Texas Parks and Wildlife Department, John H. Reagan 
Building, Austin, Texas 78701. 


VoLUME 31, NuMBER 1 ae: 


HYBRIDIZATION OF CALLOSAMIA (SATURNIIDAE) 


RICHARD S. PEIGLER 
303 Shannon Drive, Greenville, South Carolina 29615 


The genus Callosamia Packard contains three closely related species 
that generally do not hybridize in nature because of effective temporal 
isolation. These are C. promethea (Drury), C. angulifera (Walker), 
and C. securifera (Maassen). Although no wild hybrids have been 
found and I can see only scant evidence of introgression, the species can 
be easily crossed in captivity. The cross angulifera 6 < promethea 2 has 
been described and figured by Haskins & Haskins (1958) and Remington 
(1958), but I find nothing published on other crosses in this genus. 
Several lepidopterists in the northeastern states of the U.S.A. have made 
the above cross, and the two following crosses have been reared to 
adults at least once: promethea ¢ X angulifera 2 and securifera ¢ X 
angulifera 2. The rarity of hybrids with C. securifera is due to the un- 
availability of stock of that species. 

For three years I have crossed the species of Callosamia, with varied 
success. The purpose of this paper is to describe and figure some of the 
stages of the hybrids and to discuss techniques that may aid the reader 
in making crosses with Lepidoptera. 

To compare my hybrids described and pictured here with stages of 
the parent species, the reader is referred to Jones (1909), Packard 
(1914), Peigler (1976), and the excellent color plates and text in Fergu- 
son (1972). The C. securifera larva figured in color by Dominick (1972) 
shows larger tubercles than almost all those in several broods that I have 
reared from Florida and South Carolina. 


MATERIALS AND METHODS 


Because C. promethea is rare or absent throughout the South, I ob- 
tained cocoons from northern states. Except where noted, all C. angu- 
lifera stock used was from Clemson, South Carolina and was mostly wild 
males taken at lights, although some were reared from ova. All C. secu- 
rifera stock was from Berkeley Co., South Carolina, mostly from wild 
cocoons, although again some were reared from ova. 

After adults emerged from cocoons and their wings were dry, they 
were transferred to a shoebox in the refrigerator. The box contained a 
wet paper towel to provide humidity. The lower temperature prevented 
fluttering and prolonged adult life. Females that had mated were kept 


24 : JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in envelopes with wings folded back to minimize wing damage during 
oviposition. 

Adults were hand-paired by a method very different from that de- 
scribed in Collins & Weast (1961). I held moths by the thorax below the 
wings, but no squeezing of the abdomen was done, as it is not necessary 
for the female genitalia to protrude. A gentle rubbing together of the 
posterior ends of the moths usually caused the male to clasp on within 
a few minutes. Transferring to a foothold was necessary, and if the pair 
tried to pull apart, I found that blowing strongly on them gave the needed 
calming effect. Also, it sometimes helped to clip off metathoracic legs 
of a female that was trying to coax the male to release her. 

Ova were kept in petri dishes. Upon eclosion, the larvae were put 
into large cloth bags on limbs of growing foodplants, as ventilation is a 
must for larvae in this genus. This method of rearing gave larger adults 
but prevented close observation of the early instars. Therefore, I only 
describe mature larvae, since the data on early instars are fragmentary.’ 
Only in large hybrid broods were any larvae killed for preservation, but 
those that died of disease or other causes were put into alcohol or the 
freezer. The late Dr. R. B. Dominick kindly freeze-dried larvae of 
crosses 2 and 3. All hybrid larvae were reared in Greenville or Clemson, 
South Carolina on tuliptree (Liriodendron tulipifera LL.) or sweetbay 
(Magnolia virginiana L.). Cocoons were spun in the bags and, there- 
fore, under natural conditions, although some were spun in folds of the 
bags rather than on branches among leaves. 


DEscRIPTIONS oF Hysrips 


1. C. angulifera ¢ x promethea ° 


The ova, supplied by Dale F. Schweitzer, were a mixture from two 
females reared from cocoons collected on wild black cherry (Prunus sero- 
tina Ehrh.) in Medford, New Jersey. Both females were induced to emit 
pheromone after dark by artificial light in Strafford, Pennsylvania and 
attracted wild C. angulifera males; the matings were natural, not hand- 
paired. The percent hatch was very high. Larvae were reared on tulip- 
tree. A total of 33 cocoons was obtained, which produced 19 males 
and 14 females. Seven females emerged singly the first spring, but 26 
cocoons overwintered again (probably because they were kept in the 
refrigerator the first winter). The emergence pattern the second spring 


1 My larval descriptions do not involve the various black markings on the head, prolegs, and anal 


plate because my impression is that they are unreliable because of much variation within the pure 
species, especially C. angulifera. 


VoLUME 31, NuMBER 1 95 


Figs. 1-6. Larvae of hybrid Callosamia. 1, angulifera § x promethea Q. 2, 
promethea 8 xX securifera 2. 3, angulifera $ xX securifera Q. 4, securifera g xX 
angulifera @. 5, (angulifera 6 xX securifera 2) 6 X securifera 2. 6, securifera § 
x (securifera 6 X< angulifera 9 )@. (Figs. 1-5 on tuliptree, fig. 6 on sweetbay; all 
larvae in last instar except smaller one in Fig. 5. ) 


was spread over 43 days. Excepting two early males, all females emerged 
singly, then all males emerged singly or in twos. 


Larva (Fig. 1): Very uniform in all characters. Red and yellow scoli cylindrical 
or slightly swollen. Black scoli intermediate in size. Yellow subspiracular abdominal 
stripe absent.” 

Cocoon: All very dark brown, but a few golden before weathering. All but eight 
with good peduncles. 

Male (Fig. 9): Size and outline same as C. promethea, yet quite variable. Apex 


2 This lateral yellow stripe, absent in C. promethea and C. promethea hybrids, occurs in C. 
angulifera, C. securifera, and hybrids between them. A larva hanging head-down on a tuliptree or 
sweetbay leaf midrib does not seriously disrupt the pattern of the leaf’s underside because of this 
stripe, even if viewed at high angles from either side. 


26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of forewing very pointed and falcate in most specimens. Color intermediate and 
uniform among individuals, but a few with more golden suffusion beyond postmedian 
line. Discal spots present but reduced, better developed in forewing. Underside 
almost exactly like C. promethea except for discal marks. A few with sparse scaling on 
thorax and abdomen. 

Male genitalia (Fig. 19): Callosamia promethea characters dominant. Costal lobe 
of valve almost as narrow as in C. angulifera, median lobe intermediate, saccular lobe 
wider than in C. promethea but heavily sclerotized and toothed slightly. Posterior 
opening of anellus V-shaped. In two examples seen, aedeagi identical and inter- 
mediate. Cornuti on vesica almost as large as in C. promethea. 

Female: Color reddish but lighter than C. promethea and none orange in color. 
Highly variable in blackish suffusion in all areas of wings. Discal marks all well 
developed in all wings of all specimens. Underside with less contrast than C. 
promethea, but marginal lines thicker than in C. angulifera. A few with less than 
normal number of ova. 


2. C. promethea 3 X securifera ? 


Two broods of this cross were reared. The male used for the first 
brood was one I reared from Auburn, Pennsylvania. Hatch was 89%. 
Although newly hatched larvae were offered wild black cherry, they 
chose tuliptree. Of 55 cocoons obtained, over 40 contained dead larvae 
that failed to pupate. Only three males were obtained, the other adults 
being too weak to pull themselves out of cocoons. Emergences were at 
the end of July. 

The following year this cross was made again with a male from Pine 
Grove, Pennsylvania (reared by Wm. H. Houtz, Jr. on Lindera benzoin 
(L.) Blume). Percent hatch was very high again and about 35 cocoons 
were obtained; the larvae were reared on tuliptree. Two did not pupate 
successfully, and I cut open anterior ends of all cocoons to facilitate 
emergences. In late July and early August, 23 males and eight females 
emerged in a highly clustered pattern. Many adults had poorly formed 
wings, some hindwings being scaleless and transparent. Scaling on the 
body was sparse in all adults of both broods. 

A few larvae in the second brood showed a distorted pattern, having 
segments and scoli out of line. One of these made a cocoon and a male 
emerged that differed strikingly from the other males by closely re- 
sembling pure C. securifera with golden suffusion in the postmedian 
area, larger discal marks, and lighter underside. One cannot ignore the 
possibility that a genetic correlation existed between the anomalous larva 
and unique imago. 

Females emitted pheromone at “C. promethea time” on the first day 
and during the flight times of both parent species on the second day. 
A hybrid male in a cage was attracted to, and mated with, a calling 
sister 3 hr before dark. The ova produced no larvae. 


Larva (Fig. 2): Intermediate but homogeneous in appearance. Colored scoli very 


bo 
~l 


VoLUME 31, NUMBER 1 


11 12 


Figs. 7-12. Cocoons and adults of hybrid Callosamia. 7, angulifera 8  securifera 
2, cocoons. 8, promethea 6 X securifera 9, cocoons. 9, angulifera 8  promethea 
@, male. 10, securifera 6 xX (securifera 6 x angulifera 2)2, male. 11-12, 
promethea 8 X securifera 2, male, female. 


short and tapered in first brood, cylindrical in other brood. Color bluish as in C. 
promethea; black scoli intermediate. Lateral yellow abdominal stripes lacking. 

Cocoon (Fig. 8): Intermediate in size and compactness. Light brown with gray or 
red cast, never silvery. About 61% had strong peduncles, 25% made weak attach- 
ments, and 14% made none. 

Male (Fig. 11): Outline like C. promethea. Color very dark but not as black as 
C. promethea. Discal marks weakly developed in forewing, usually absent in hindwing. 
Very minimal golden suffusion in postmedian area. Apices of forewings more pointed 
than either parent species. Underside intermediate with weak red suffusion past 
postmedian line. 

Male genitalia (Fig. 20): Callosamia promethea characters very dominant. 
Median lobe of valve as large and long as in C. promethea, costal lobe intermediate. 
Anellus opening variable. In one specimen, cornuti on vesica large; in another 
specimen, cornuti half that size. Aedeagi almost as large as in C. promethea. 


28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Female (Fig. 12): Undersized but full of ova. Color dull orange, being between 
females of parents. Discal marks developed in all wings. Postmedian line less 
undulating (as in some C. promethea) and more proximal than in both parental 
species. Underside very similar to C. angulifera. 

Ova: Chorion thin, causing collapse. White color changing to translucent yellow 
when dry. Size same as C. promethea, smaller than C. securifera. 


3. C. angulifera 3 X securifera ? 


About five-sixths of the ova hatched. Most of the brood were reared 
on tuliptree, but some were reared on sweetbay. The latter grew slower 
but attained equal proportions as adults. Of 52 cocoons, only a few died. 
Adults totalled 24 males and 23 females. The emergence pattern ran 
from 15 July to 25 August with heaviest emergences early in this range. 
The first 12 to emerge were females, the last ten all males, and between 
was a mixture. Six cocoons overwintered and adults emerged in early 
May; five were females. 


Larva (Fig. 3): Homogeneous in all aspects. Epidermal color and minute black 
scoli very close to C. angulifera, but colored scoli like C. securifera. Lateral stripe 
prominent. 

Cocoon (Fig. 7): Most golden brown, a few dark brown like father species. 
Intermediate in size but closer to C. securifera. About 33% made strong stem attach- 
ments, 45% made weak ones, and 22% made no attempt to attach. 

Male (Fig. 13): All intermediate but the series quite variable. Callosamia securifera 
characters dominant but larger than in that species. Color dark brown and present, 
but reduced, discal marks (because they are summer form). Underside of forewing 
like C. securifera; hindwing more intermediate with less contrast than C. angulifera 
but with dark brown median area as in C. angulifera. Abdominal terga and anal 
margin of hindwing maroon as in C. securifera. 

Male genitalia (Fig. 21): Median lobe of valve short like in C. angulifera. Smaller, 
shorter, more rounded saccular lobe than in reciprocal cross. Anellus opening rounded. 
Vesica with pair of large cornuti. 

Female (Fig. 14): Extremely variable, some assignable to spring form, others 
summer form, others intermediate. All with traits of both parent species, with C. 
angulifera characters predominating. Color orange with varying degrees of black 
suffusion. Underside much more like C. securifera, but more contrast between median 
and postmedian areas. 

Ova: Size of C. securifera, but larger than C. angulifera. A few with weak chorion. 


4. C. (angulifera 6 X securifera 2) F> 


A male and female from cross 3 were hand-paired, and six or seven 
of the 125 ova hatched. Tuliptree was used for food, and two cocoons 
were obtained that yielded females the following May. One was under- 
sized and weak with thin scaling on the wings; the description below is 
based on the other specimen. 


Larva: Surprisingly like C. promethea. Thick red thoracic and yellow scoli cylin- 
drical. Black scoli larger than in most C. securifera, approaching C. promethea, 
dorsal ones largest. Abdominal yellow stripes present. 

Cocoon: No peduncles. Color of silk and size intermediate. 


VoLuME 31, NuMBER I 99 


aes , 18 


Figs. 13-18. Adults of hybrid Callosamia. 13-14, angulifera 8 x securifera 9, 
male, female (ventral views). 15-16, securifera 6 x angulifera 2, male, female. 
17-18, angulifera 8 x (angulifera 6 X securifera 2)2, male (ventral view), 
female. 


Female: One specimen. Would easily pass for pure C. angulifera if examined 
closely dorsally or ventrally. One minor C. securifera trait noted—a double distal 
edge of discal mark on underside of forewing. 

Ova: Size like C. securifera. Chorion thin, collapsing and color becoming trans- 
lucent yellow as ova dry, a few remaining white. 


5. C. securifera 6 X angulifera 2 


Only 21 ova hatched of 128 laid. Larvae were reared on tuliptree, a 
few being started on sweetbay and transferred to tuliptree in third in- 
star because of higher mortality on sweetbay. Twelve cocoons were 
obtained and six males and five females emerged in May. One female 
was observed to emit pheromone during C. promethea flight time, ceased 


30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


at dark, and resumed for another hour under a 15-watt lamp with re- 
flector. 


Larva (Fig. 4): All but one very close to reciprocal cross but colored scoli slightly 
thinner. Aberrant one (Fig. 4, foreground) very unusual. Color grayer overall. 
Colored scoli shorter, tapered, and much thicker. Thoracic ones dull brick orange 
similar to faded ones in freeze-dried specimens. Black scoli large like in C. promethea. 
( Aberrant larva produced female imago closely matching sisters; mother of cross 7). 

Cocoon: Of 12 cocoons, half with strong, long peduncle, four with partial, two 
with none. Color duller gray brown than those of cross 3 and smoother silk. Color 
and size intermediate, with father species dominant. 

Male (Fig. 15): One male (figured) quite unlike brothers, having much golden 
suffusion. Other males alike, closely resembling males of reciprocal cross, but 
smaller overall and a bit redder in median area of underside of hindwings. Discal 
marks not prominent. Abdominal terga and anal margins of hindwings maroon. 

Male genitalia: Costal lobe of valve very much like C. angulifera, median lobe 
long, saccular lobe wide. Callosamia angulifera traits dominant. Teeth of uncus 
shorter. Anellus with V-shaped opening. Aedeagus exact length and thickness as in 
cross 8, but two cornuti much smaller. 

Female (Fig. 16): Very close to C. angulifera and cross 4. Extremely large discal 
marks. Underside mostly like C. angulifera, but postmedian line much like C. 
securifera (unlike C. angulifera). Lateral ornamentation of abdomen more like 
C. securifera. Antennae intermediate. Wing outline rather variable in view of all 
other similarities. 

Ova: Intermediate size. Some with weak chorion, others normal appearing. 


6. C. angulifera 6 x (angulifera 3 X securifera 2) 9 


This backcross was done three times using females from cross 3, but 
fertility of ova and viability of larvae and pupae were low. Hatching 
ranged from about 10-60%. A total of six cocoons and three adults were 
obtained; one pair was the spring form and another male a summer form. 


Larva: Most intermediate or closer to C. angulifera. One with black scoli split into 
pairs, metathoracic scoli with thick double (disjunct) black base. Black circled area 
on anal prolegs with black line running through middle. 

Cocoon: Dark brown and small like C. angulifera, but five of six with silken 
attachment to branch. 

Male (Fig. 17): Spring form (figured) indistinguishable from C. angulifera, but 
size and wing shape more like C. securifera. Summer form closely resembling males 
of crosses 3, 5, and 8. Wing scaling slightly sparser. 

Male genitalia: Like pure C. angulifera except for a few minor trends: rounded 
anellus, slightly longer median lobe of valve, and vesica with larger cornuti. Aedeagus 
as wide as in C. securifera. 

Female (Fig. 18): All characters like C. angulifera but smaller and browner. 
Double edge of discal marks on underside barely discernable. Underside areas with 
much less contrast along post median line than C. angulifera. 

Ova: Like C. angulifera, perhaps a bit larger. 


7. C. securifera 8 X (securifera 8 x angulifera 2) 


The mother of this hybrid brood was the aberrant larva with larger 
tubercles mentioned in cross 5, The ova gave 8% hatch, and the larvae 


VoLUME 31, NuMBER 1 on 


21 


Figs. 19-22. Male genitalia of hybrid Callosamia. 19, angulifera § x promethea 
Q. 20, promethea $ xX securifera 9. 21, angulifera $8 x securifera 2. 22, 
(securifera 6 X angulifera 2)é x (angulifera 6 x securifera 2) 9. 


were reared on sweetbay. Only four cocoons were obtained and one 
adult. 


Larva (Fig. 6): Brood included largest larvae I have ever seen of genus. Black 
scoli not as large as in mother but larger than in most C. securifera. Colored thoracic 
scoli somewhat light, short, and thick. 

Cocoon: A bit smaller and darker than C. securifera and peduncle wanting. Very 
much like C. securifera nonetheless. 

Male (Fig. 10): Pattern and colors exactly like summer form of C. securifera. 
Hindwings less rounded, more like C. promethea. Larger than in C. securifera. 

Male genitalia: Pure C. securifera in all respects. Median lobe of valve slender 
and longer than costal lobe. Costal lobe squared off. Aedeagus longer than in C. 
angulifera. Two large cornuti on vesica. 


8. (securifera 6 X angulifera 2) 3 X (angulifera é Xx 
securifera @ ) @ 
Only eight larvae were obtained from 190 ova laid by one female. Two 


larvae reached maturity, all others dying in the earliest instars. All were 
on tuliptree. Two males emerged in mid-July. 


32 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Larva: No characters seen to distinguish from pure C. angulifera, but possibly less 
yellowish epidermal color. 

Cocoon: No peduncles. Intermediate in color. Almost as large as in C. securifera. 

Male: Closely resembling males in cross 3 and the summer male of cross 6. 
Especially red on undersides in postmedian area. Discal marks on forewing average, on 
hindwing feeble. One male with maroon abdomen. 

Male genitalia (Fig. 22): Costal and median lobes of valve more like C. angulifera, 
saccular lobe more like C. securifera, less pointed, more rounded. Anellus rounded. 
Uncus comparatively small. Noticeable bend in aedeagus, which is thicker than in 
C. angulifera. Large cornuti on vesica. 


9. C. (angulifera 3 x securifera 2) é X securifera 2 


Twenty-three of 118 ova hatched, and the larvae were reared on 
sweetbay until the last two instars, when they were transferred to tulip- 
tree. All larvae died of a disease except one, which made a cocoon and 
died at pupation. About a third of the brood was “super-tuberculate” in 
that there was a dorsal pair of reduced colored tubercles on the first 
abdominal segment. This peculiar character apparently occurs occasion- 
ally in the pure species. It has been reported in X-radiated C. prome- 
thea stock by Haskins (1934) and is incorrectly given as a constant 
character of C. securifera by Ferguson (1972), although I find only 
four C. securifera larvae of about 30 in the Wedge Plantation Collection 
show it. That collection also has a few C. angulifera larvae (from SE 
Pennsylvania) with this trait. 

Larva (Fig. 5): See discussion above for most unusual characteristic. Black scoli 
the size of those in C. securifera or slightly larger. Scarlet scoli slightly swollen or 
cylindrical, thicker than in C. securifera and C. angulifera. 


Cocoon: One specimen. Smaller and darker than C. securifera but with firm 
attachment to stem. Silk golden. 


10. C. (securifera ¢ X angulifera 2) 6 x [angulifera 6 x 
(angulifera 3 X securifera 2 )2)? 

I killed the female (Fig. 18) after 72 ova were laid. One egg hatched 
and the larva was reared to maturity on tuliptree. Unfortunately, it was 
killed accidentally in the last instar. 

Larva: Colored scoli with very thick black bases, especially metathoracic pair. 


Black scoli size of those in C. securifera. Color like C. angulifera, and yellow lateral 
stripe present. 


DISCUSSION 


One inherent problem in the hybridization project was the large 
genitalia of C. promethea when compared with the other two species. 
Best results can be obtained with reared C. promethea intentionally 
made undersized by crowding or poor food during larval life. Females of 


VoLUME 31, NuMBER 1 33 


C. angulifera, C. securifera, and their hybrids can be ruptured and 
killed by mating with a male C. promethea. Conversely, it is difficult 
for C. angulifera, C. securifera, or hybrid males to clasp onto C. prome- 
thea females. 

A large number of crosses were made that produced no larvae from 
the ova. In most cases, matings seemed successful, lasting over 4% hr to 
several hours, and females oviposited freely. These crosses included 
duplicates and reciprocals of some of the above crosses and seven that 
involved all three species. An example of the latter is C. (securifera 6 x 
angulifera 2) & X promethea 2, in which the female (from Cedar 
Rapids, Iowa) laid 157 ova. 

If a hybrid that combines all three species is reared, probably one 
parent will be pure C. promethea since all my tests suggest hybrids 
which involve C. promethea are sterile. Ova of crosses between C. 
angulifera and C. securifera often give a high percentage of eclosion, 
and resultant adults of both sexes are partially fertile, as shown by six 
of my crosses. Hybrid females of the allied genus Hyalophora rarely 
contain ova (Collins & Weast, 1961 and pers. obs.), but all Callosamia 
hybrid females do. Most mortality in hybrid broods occurs in ova and 
earliest instar larvae. 

Some observations were made on certain behavioral traits that are 
probably polygenic. The attachment of the cocoon (almost always in 
C. promethea and C. securifera, rarely in C. angulifera) seems to be a 
dominant trait. Unlike the other two species, females of C. angulifera 
do not oviposit freely. However, all hybrid females that I mated ovi- 
posited freely. If the males used in cross 2 came from univoltine popula- 
tions, it would appear that this trait was not expressed in the offspring. 
Larvae of C. promethea and C. securifera regurgitate a green fluid when 
handled, but I have not observed C. angulifera larvae to do so. This 
trait of C. angulifera occurred in hybrid larvae of crosses 3-6 and 8. 

Examples of my hybrids can be found in the following private and 
museum collections: Dale E. Pforr, Canada; Michael M. Collins, Cali- 
fornia; Dr. Claude Lemaire, France; Wedge Plantation Collection; Los 
Angeles County Museum of Natural History; United States National 
Museum. A future paper will describe and figure my hybrids of the 
cross C. angulifera 6 xX Samia cynthia (Drury) @. 


ACKNOWLEDGMENTS 


I am most grateful to Dr. G. R. Camer, insect pathologist at Clemson 
University, for making all the photographs in this paper. He often took 
time from his busy schedule to photograph hybrid larvae. Dr. Claude 


34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Lemaire, a saturniid authority and personal friend, very kindly made 
the genitalic preparations. Dr. T. R. Adkins of Clemson University 
allowed me to bag tuliptrees in his yard for two summers. 


LITERATURE CITED 


Coxiiins, M. M. & R. D. Weasr. 1961. Wild silk moths of the United States. 
Collins Radio Corp. 138 p. 

Dominick, R. B. 1972. Practical freeze-drying and vacuum dehydration of cater- 
pillars. J. Lepid. Soc. 26: 69-79. 

Fercuson, D. C. 1972. Bombycoidea, Saturniidae (in part). In R. B. Dominick 
et al., The moths of America north of Mexico, fase. 20.2B: 155-269, 22 pls. 
Haskins, C. P. 1934. Preliminary note of morphological variations occurring in 
X-rayed stock of the attacine moth Callosamia promethea Dru. J. N. Y. Ent. Soc. 

42: 145-154. 

. & E. F. Haskins. 1958. Note on the inheritance of behavior patterns for 
food selection and cocoon spinning in F; hybrids of Callosamia promethea x C. 
angulifera. Behaviour 13: 89-95. 

Jones, F. M. 1909. Additional notes on Callosamia carolina. Ent. News 20: 49-52. 

Pacxarp, A. S. 1914. Monograph of the bombycine moths of North America, 
part 3 (ed., T. D. A. Cockerell). Mem. Natl. Acad. Sci. 12: 1-516. 

PeicLer, R. S. 1976. Wing color variation in Callosamia (Saturniidae). J. Lepid. 
Soc. 30: 114-115. 

Remincton, C. L. 1958. Genetics of populations of Lepidoptera. Proc. Tenth 
Internatl. Congress Ent. 2: 787-805. 


COCYTIUS DUPONCHEL (SPHINGIDAE): SECOND UNITED STATES 
CAPTURE 


On 30 September 1975 while operating a UV light along Route 29 near Immokalee, 
Collier Co., Florida, the author took a large male sphingid that under the light ap- 
peared to be the resident species, Cocytius antaeus (Drury). Upon mounting the 
specimen, it tentatively was indentified as Cocytius duponchel (Poey). The speci- 
men was sent to William Sieker of Madison, Wisconsin who confirmed the identifica- 
tion as C. duponchel. 

This constitutes only the second U.S.A. record for this neotropical species. The 
other capture of C. duponchel within the U.S.A. is from Edwards Co., Texas in May 
1902 (Hodges 1971, Moths of North America, Fascicle 21, Sphingoidea, 25). The 
specimen has been deposited in the Florida State Collection of Arthropods in 
Gainesville, Florida. 


JAMeEs P. TurrLe, 2691 West Temperance Road, Temperance, Michigan 48182. 


VoLUME 31, NuMBER 1 35 


AGGREGATION BEHAVIOR OF CHLOSYNE LACINIA 
LARVAE (NYMPHALIDAE) 


Nancy Stamp! 
Department of Zoology, Arizona State University, Tempe, Arizona 85281 


The objective of this study is to describe the larval aggregation 
behavior of Chlosyne lacinia crocale (Edwards) in terms of a daily 
activity pattern and the relation of larval size to the tendency to ag- 
gregate. Although other authors mention these topics, (Edwards, 1893; 
Bush, 1969; Drumond et al., 1970), they provide little quantitative sup- 
port for their conclusions. 

Chlosyne lacinia crocale ranges from the southwestern United States 
to Mexico (Emmel & Emmel, 1973). From March—November, adult 
females deposit multiple broods on leaves of the common sunflower, 
Helianthus annuus L., and related species (Drummond et al., 1970; Neck, 
1973; Gorodenski, 1970). There are five instars, each lasting 3-7 days, 
with an average generation time in the field of 35-40 days (Edwards, 
1893; Drummond e¢ al., 1970). Hatching synchronously, the larvae feed 
gregariously under a silk web on the underside of a leaf through the 
first instar (Edwards, 1893; Bush, 1969). After a leaf is devoured by a 
group, individuals move single file to a new leaf using and reinforcing 
silk thread trails (Bush, 1969). Coinciding with the appearance of dis- 
tinct larval color patterns, dispersal by single individuals to different 
leaves and plants occurs in the fourth and fifth instars (Drummond et al., 
1970). 


METHODS 


Larvae were observed on sunflower plants (Helianthus annuus) along 
fence rows in Tempe, Arizona. Observations were made between 0700 
and 1800 hours from 10 October-15 November 1975. The mean minimal 
temperature for this period was 12.1°C, and the mean maximal tempera- 
ture was 27.7°C. A record was made of the type of plant part used, 
height on plant of larvae, number of larvae per group, and length of 
larvae, in mm. Testor’s red enamel paint was used to mark the larvae. 

To study the tendency to aggregate, 10 portions of sunflower plants 
were placed in containers with water in the laboratory at 24°C. Each 
portion had two main stems of about equal diameter, length, and number 
of healthy leaves, buds, and flowers. On five of these plants, a group of 


1 Present address: Department of Zoology, University of Maryland, College Park, Maryland 20742. 


36 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


n=577 n=678 n=606 n=-485 n-571 


100 
Percent 
of 
total 
apiae 28 Single 
[]Grouped 
50 
0730- 1000- 1200- 1400- 1600- 
0959 1159 1359 1559 1759 
Hours 
30 
avg. 
no. wee 
in 
group 
18 
0730- 1000 - 1200- 1400- 1600- 
0959 1159 1359 1559 1759 
Hours 


Fig. 1. Daily activity patterns of Chlosyne lacinia. Upper—the proportion of 
solitary and aggregated larvae by 2-hour intervals. Lower—average size of group. 


five larvae was placed at the divergence of the two main stems. On each 
of the other five plants, five larvae were placed singly at diverging stem 
points. The number of larvae in groups was recorded every half 
hour for 3.5 hours. This experiment was conducted four times with large 
larvae (5-10 mm in length). 


RESULTS 


Small, light-colored larvae (<2 mm in length) usually clustered on the 
underside of large leaves. Individuals often fed at the same site on a 
leaf. These groups fed on one large leaf for several days and usually 
molted before moving to a new leaf. 


VoLUME 31, NuMBER 1 37 


single grouped single grouped single grouped 


too Mees 122769 n=148 n=2764 n=148 n=2668 


. 
\ 


30 


Percent of larvae 


Plant part used Height on plant Length of larvae 


Leaves J <06m <2mm MB 4-10mm 
Stems — 0.6-1.2 m 2-4 mm >10mm [J 
Buds & flowers [] he? ne E 


Fig. 2. Comparisons of single larvae and larvae in groups. 


Larger, darker larvae (5-10 mm) clustered on one or both sides of a 
leaf, devoured it in a few hours, and, leaving only the network of the 
leaf intact and scattered dark fecal pellets, departed for a new leaf. 

Tendency to aggregate was dependent on time of day (2 X 5 con- 
tingency table with x? = 33.35 and p < 0.001). As a day progressed, the 
percentage of aggregated larvae decreased to 91.5% and then increased 
to 98.2% after 1600 hours (Fig. 1). Correspondingly, the average number 
of larvae per group was lowest after noon and rose sharply by 1600 hours. 
Throughout the daylight hours, grouped larvae fed exclusively on leaves 
and spent most of their time on them (Fig. 2). The percentage of ag- 
gregated larvae on stems was highest between 1400 and 1559 hours, 
when larval movement was greatest. Most (97.0%) of the grouped 
larvae were on leaves higher than 0.6 m from the ground. 

Through the day, increasing numbers of single, active larvae were 
observed, but after 1600 hours, the number of solitary larvae decreased 


38 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


100 
Single larvae 


Sig at start 
a n=100 
> = 
i aa 
‘N 
N 
‘\ 


50 i 


Percent 


Grouped larvae 


at start 
n=100 


0.5 1.0 15 2.0.. 225 3.0 B55 
Hours 


Fig. 3. Testing aggregation tendency. Percent of single larvae are indicated by 
solid and dashed lines. Larvae placed singly on sunflower stems tend to move until 
they reach other individuals. Larvae placed in groups tend to remain in groups after 
some initial scattering. See text for additional explanation. 


(Fig. 1). Solitary larvae were encountered more frequently than ag- 
gregated larvae on stems, buds, and flowers (Fig. 2). Although the 
majority of single larvae were found on plant parts higher than 0.6 m 
from the ground, 26.4% of the solitary larvae were closer to the ground 
(<0.6 m) as a result of their tendency to wander. 

Most (60.0%) of the aggregated caterpillars were <4 mm in length, 
whereas only 2.0% of the solitary larvae were this small (Fig. 2). Of the 
single larvae, 50.3% were 4-10 mm and 47.0% were >10 mm. 

An experiment was conducted to test whether the larvae tend to 
aggregate (Fig. 3). The percentage of solitary larvae on plants origi- 
nally provided with single larvae was compared with the percentage on 
plants stocked with group larvae. In the first group of plants, percentage 
of solitary larvae decreased significantly with time (2 X 7 contingency 


VoLUME 31, NuMBER 1 39 


table with y? = 26.65 and p < 0.001). Percentage of solitary larvae in 
the second group of plants remained fairly constant (2 * 7 contingency 
table with y? = 5.55 and 0.50 > p > 0.20), with a mean of 41% single 
larvae. 

Several aggregations on sunflower plants were marked to determine 
whether these groups remained together, but the results were incon- 
clusive. One group of 20 larvae remained together for three days and 
did not mix with a second group of 67 individuals 0.6 m higher on the 
plant. However, another marked group of 18 larvae was scattered singly 
or in pairs over its host plant on the second day. An unmarked group 
was also scattered, and two pairs of larvae consisted of marked and 
unmarked individuals. 


Discussion 


Larvae of C. lacinia tend to form groups, with individuals leaving 
and wandering in the afternoon, then rejoining groups by early evening. 
The large larvae are more likely to wander and be found as isolated 
individuals. 

What are the advantages associated with the formation of aggrega- 
tions? Larvae in groups, especially small individuals (<4 mm), exhibited 
synchrony in feeding, molting, and moving to a new leaf. By moving in 
a group, individuals may gain an advantage in exploiting the food source. 
Ghent (1960) found that small larvae of the jack pine sawfly (Neodiprion 
pratti banksianae Roh.) had difficulty boring into pine needles, but once 
one individual was successful, others could easily join in feeding at 
that site. 

As larvae grow larger, the feeding advantage linked with being in a 
group decreases. The larger caterpillar can more easily chew into a leaf, 
and competition for food becomes more intense. Large larvae (5-10 
mm) consume leaves at a much faster rate than small caterpillars. Con- 
sequently, large larvae move to new feeding sites more often, which 
requires an increased expenditure of time and energy. If a larva locates 
a leaf and devours it alone, it will maximize eating time and minimize 
time and energy expended on movement. 

Though probably a function of food competition, dispersal may also 
reduce disease, parasitism, and predation in the fourth and fifth instars 
(Drummond e¢ al., 1970). Parasitism is especially high in the third, 
fourth, and fifth instars, but there is some evidence of differential 
parasitism among the three larval color morphs (Drummond et al., 1970). 
Perhaps to avoid pupal parasitism, fifth instars migrate singly to pupation. 

Costs and benefits of group membership change radically with age 


40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


and size of larvae, which leads to age-linked changes in the tendency 
to aggregate. 


ACKNOWLEDGMENTS 


I thank Dr. J. Alcock for reviewing this manuscript and his encourage- 
ment to pursue this project. Also, I appreciate the suggestions and help 
in identification of specimens from Dr. F. Hasbrouck. 


LITERATURE CITED 


Busy, G. L. 1969. Trail laying by larvae of Chlosyne lacinia. Ann. Ent. Soc. Amer. 
62: 674-675. 

DrumMmonp, B. A., III, G. L. Buso, & T. C. Emmeu. 1970. The biology and 
laboratory culture of Chlosyne lacinia Geyer (Nymphalidae). J. Lepid. Soc. 
JA: 135-142. 

Epwarps, W. H. 1893. Notes on a polymorphic butterfly, Synchloe lacinia, Geyer 
(in Hub. Zutr. ), with description of its preparatory stages. Can. Ent. 25: 286-291. 

EMMEL, T. C. & J. F. Emmen. 1973. The butterflies of southern California. 
Natural History Museum of Los Angeles County, Los Angeles. 148 p. 
GuEenT, A. W. 1960. A study of the group-feeding behaviour of larvae of the 
jack pine sawfly, Neodiprion pratti banksianae Roh. Behaviour 16: 110-148. 
GoroveNskI, S. A. 1970. The genetics of three larval color forms in Chlosyne 
lacinia and the phenotypic frequencies of this polymorphism in natural popula- 
tions. M.S. Thesis. Ariz. State Univ. 43 p. 

Neck, R. W. 1973. Foodplant ecology of the butterfly Chlosyne lacinia (Geyer) 
(Nymphalidae). I. Larval foodplants. J. Lepid. Soc. 27; 22-33. 


A MELANIC FORM OF PHIGALIA STRIGATARIA (GEOMETRIDAE ) 


A dark geometrid moth was caught by the author at black light at Lebanon, New 
Jersey on 5 April 1972. A genitalic slide proved it to be a melanic form of Phigalia 
strigataria Minot. On p. 128 of “A revision of the New World Bistonini,” Frederick 
H. Rindge states that he never saw a melanic specimen of P. strigataria. This ap- 
parently is the first verified one. I gave the specimen with genitalia slide to the 
American Museum of Natural History, New York. 

Compared with Phigalia titea form “deplorans” no difference can be detected. 
Forewing length of this strigataria from apex to base is 16 mm. Small sizes of 
“deplorans” are also found, but most are larger. Colors of both are the same. 


JosEpH Mutter, R.D. 1, Lebanon, New Jersey 08833. 


VOLUME 31, NuMBER 1 4] 


A REVIEW OF NORTH AMERICAN RHODOPHAEA 
(PHYCITINAE: PYRALIDAE), WITH DESCRIPTION 
OF SIX NEW SPECIES 


PauL A. OPLER 


Office of Endangered Species, U.S. Fish and Wildlife Service, Department of the 
Interior, Washington, D.C. 20240 


As conceived by Heinrich (1956) the Holarctic phycitine genus Rho- 
dophaea was represented in North America by two species—R. caliginella 
(Hulst), an oak-feeder, and R. supposita (Heinrich), which feeds on 
Rosaceae. Six species are described here; all feed on members of the 
family Fagaceae in California or adjoining portions of Arizona and 
Nevada. Heinrich’s R. swpposita, described from British Columbia, is 
synonymized with the Palaearctic R. suavella (Zincken), and probably 
represents an introduction by man. The type of genus Rhodophaea, 
R. advenella (Zincken), and three other species, (including R. suavella), 
occur in the Palaearctic. 

During an ecological study of microlepidoptera feeding on Fagaceae in 
California (Opler, 1974), six new Rhodophaea were reared, and larval 
shelters probably representing other new species were found. 

Forewing color patterns of North American Rhodophaea are similar 
and the identification of any specimen should be based on the examina- 
tion of dissected genitalia. Variation in color pattern among individuals 
reared from one host population may approach the variation shown be- 
tween species. Both male and female genitalia display relatively little 
variation within reared series, and offer good specific characters. 

Members of the genus Rhodophaea are extremely similar to those of 
Acrobasis Zeller. Heinrich distinguished Rhodophaea chiefly on the 
shape of the basal antennal segment. Thus, as now conceived, Rhodo- 
phaea possibly is constituted of several species groups independently 
derived from Acrobasis or its ancestors several times in the Tertiary. 


Taxonomic Characters 
Forewing color pattern. The scale color markings used are (Figure 
1): (1) small elongate patch at base of inner margin, usually dark 
(inner basal patch); (2) basal portion, exclusive of (1), usually pale 
(basal area); (3) post-basal bar extending from costal margin toward 
inner margin, usually dark contrasting with pale basal area, and otten 
edged outwardly with brown, tan or reddish (basal bar); (4) a sub- 


42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


girl 


AOL, § 


I 


Fig. 1. Diagrammatic illustration of Rhodophaea forewing color pattern features 
employed in descriptions, 1. inner basal patch, 2. basal area, 3. basal bar, 4. s-m patch, 
5. p-m patch, 6. spots, 7. band, 8. p-m line, 9. patches. 


medial triangular patch of black scales along costal margin, distal to bar 
(s-m patch); (5) postmedial pale area along costal margin (p-m patch); 
(6) two black spots in lower portion of p-m patch (spots); (7) a diffuse, 
diagonal, dark band extending from costal margin inwardly toward, but 
not reaching, inner margin (band); (8) a sinuous pale postmedian line 
(p-m line); (9) and a row of small black elongate patches just inward 
to outer margin (marginal patches). 

Genitalia. In male Rhodophaea the inner surface of the valva pos- 
sesses protuberances which are constant in configuration for each species, 
and are the prime species character used for that sex. In female 
Rhodophaea the number and configuration of the signa bursae are most 
diagnostic. 

Hosts. New World Rhodophaea are known to feed on only one or two 
closely related hosts, e.g., Rhodophaea caliginella on Quercus agrifolia 
and QO. wislizenii. More than one Rhodophaea may be found on the 
phenotypically and genetically complex Quercus dumosa-turbinella 
populations of mainland California. 


43 


VoLUME 31, NuMBER 1 


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44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Type deposition. Holotypes and allotypes of species described herein 
are deposited in the California Academy of Science, San Francisco, on 
indefinite loan from the Essig Museum of Entomology, University of 
California, Berkeley. 


Taxonomic Treatment 


Type of the genus: Phycis advenella Zincken, 1818. 


The type of Rhodophaea is quite distinct from known American con- 
geners. The apical process of the gnathos is divided, whereas it is un- 
divided in New World species. The valvae of male R. advenella each 
display one long inwardly directed bar-like projection on their inner 
face (Figure 16). The corpus bursae of female R. advenella lacks 
pronounced signa, but is covered with minute microspicules (Fig. 20). 
The ductus bursa is very short when compared to those of New World 
species. Furthermore, the male of R. advenella possesses a mid-ventral 
hair-tuft on the eighth abdominal segment which is lacking in American 
Rhodophaea. 


Rhodophaea caliginella (Hulst) 
Fie. 2 


Synonymy is given by Heinrich (1956) and Opler (1974). Lectotype designation 
is given by Opler (1974). 

Diagnosis. A large gray moth usually separable from other Rhodophaea by the 
dull black inner basal patch. 

Male. Length of forewing 9.5-11 mm (reared). Inner basal patch dull black; 
basal area of gray white-tipped scales; basal bar a small medial black patch edged 
outwardly with tan; s-m patch a narrow black triangle; p-m patch narrow, pale, 
composed of scales with basal half gray, distal half white; spots both distinct, lower 
the largest; band distinct, of black or dark gray white-tipped scales; p-m line distinct 
throughout its length, of dark gray white-tipped scales; marginal patches a broken 
line of five black elongate lines, each composed of shining black narrowly white-tipped 
scales. 

Genitalia. As in Fig. 8, valva with a small indistinct raised area on medial portion 
of inner face. 

Female. Forewing 9-11 mm (reared). As in male except inner basal patch not as 
distinct; basal bar with tan scales not as extensive; spots upper absent; band merging 
along inner margin with s-m patch. 

Genitalia. As in Fig. 21, corpus bursa with a small, indented, circular, scobinate 
patch; distinct signa lacking. Distal two-thirds of corpus bursae with pebbly texture 
and some minute spine-like projections. 

Hosts. Quercus agrifolia Nee, Q. wislizenii A. DC. and Q. wislizenii frutescens 
Engelm. 

Distribution. Known to occur only in California, but is probably also present in 
Baja California Norte. Distribution is coincident with that of its hosts, from Shasta 
County south to San Diego County in the coast range, Sierra Nevada foothills (where 
it is sparsely distributed), the transverse ranges, and the peninsular ranges. The 
moth is apparently absent from Santa Cruz Island, Santa Barbara County, where 
suitable hosts occur (Opler, 1970). Quercus agrifolia populations on the more 
remote Santa Rosa Island have not been sampled. 


VoLuME 31, NuMBER 1 45 


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PO SOLS | ae 
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C e . ws he - ie 
. ’ ‘ e Pr) ere 3 ' 
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‘ ae =e 
’ . Yr 
5 - 


Figs. 8-9. Rhodophaea, male genitalia, left valva not shown, aedeagus and juxta 
removed, A. juxta, B. aedeagus, 8. R. caliginella, 9. R. cruza. Scale bar = 1 mm. 


46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 10-11. Rhodophaea, male genitalia, left valva not shown, aedeagus and juxta 
removed, A. juxta, B. aedeagus, 10. R. kofa, 11. R. durata. Scale bar = 1 mm. 


The Arizona specimen referred to this species by Heinrich (1956) may have been 
mislabelled. The specimen is R. caliginella, but neither host of the moth occurs in 
Arizona, and furthermore, the only documented Arizonan Rhodophaea is R. kofa, 
described below from Yuma County’s Kofa Mountains, an area remote from the 


VOLUME 31, NUMBER 1 AT 


State’s principal oak-dominated habitats. Sampling of most Arizona Quercus in 1969 
indicated an absence of Rhodophaea throughout most of that state. 


Rhodophaea cruza Opler, new species 
Fig. 3 


Diagnosis. This is the only Rhodophaea with a purple-red inner basal patch and 
purple red scaling distal to the basal bar. 

Male. Length of forewing 10-10.5 mm (reared). Forewing pattern as for R. 
caliginella male except inner basal patch of purplish red scales; basal bar a broad 
purplish red band for posterior two-thirds edged inwardly with a narrow black patch; 
s-m patch dark at costal margin, included scales becoming narrowly white-tipped 
distally and posteriorly; p-m patch with anterior spot absent. 

Genitalia. As in Fig. 9, valvae with a low medial linear ridge on inner face 
extending from near base distally to half distance to tip. Juxta with small lateral 
projections, 

Female. Length of forewing 9.5 (reared). Maculation as in male except s-m patch 
and band more extensive; both p-m patch spots absent; p-m line with adjacent reddish 
scales. 

Genitalia. As in Fig. 23. Corpus bursa with single triangular signum composed of 
pointed projections, and a circular scobinate patch. 

Type material. Holotype: male, California, Canada de Ja Cuesta, Santa Cruz 
Island, Santa Barbara County, 15 March 1969, reared from Quercus dumosa, J. Powell 
lot 69C40, emerged 30 June 1969, P. Opler prep. 78, P. A. Opler collector. Allotype: 
female, same data except P. Opler prep. 81. Paratypes: 3 646, 4 2Q same data 
except some emerged 29 June and 4 July. 

Host. Quercus dumosa Nutt. 

Distribution. Santa Cruz Island, Santa Barbara County, California. 

Discussion. Present evidence indicates this moth is endemic to Santa Cruz Island, 
yet future study may well alter this assignment. On Santa Catalina Island, Los 
Angeles County, a single Rhodophaea shelter was found on Quercus dumosa, so that 
R. cruza could eventually prove to be endemic to more than one California island. 
On the mainland, larvae and shelters were found on Quercus dumosa throughout 
much of its range, but only one reared individual exists (a male from Los Angeles 
County ). Color pattern and genitalic features relate this individual to R. cruza, but it 
differs to such a degree that assignment to that species is unsure. 


Rhodophaea kofa Opler, new species 
Fig. 4 


Diagnosis. The extensiveness of white-tipped scales gives this moth a distinctive 
“salt and pepper” appearance. 

Male. Length of forewing 7-8.5 mm (reared). Forewing as for R. caliginella male 
except inner basal patch of white-tipped black scales; basal bar mostly tan, inwardly 
edged with narrowly white-tipped black scales; s-m patch very restricted in size; 
p-m patch very extensive along costal margin; spots both present with narrowly 
white-tipped black scales; band restricted, not extending to inner margin. 

Genitalia. As in Fig. 10. Valva with a stout diagonal ridged swelling at base of 
inner face. 

Female. Length of forewing 8.5 mm (reared). Forewing as in male except 
marginal patches extending to costal margin. 

Genitalia. As in Fig. 18. Similar to that of R. cruza, corpus bursa with singular 
triangular signum of pointed projections, and with a circular scobinate patch, denser 
and more extensive than that of R. cruza. 

Type material. Holotype: male, Arizona, Palm Canyon, Kofa Mountains, Yuma 


48 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 12-13. Rhodophaea, male genitalia, left valva not shown, aedeagus and juxta 
removed, A. juxta, B. aedeagus, 12. R. fria, 13. R. neva. Scale bar = 1 mm. 


County, 8 April 1963, reared from Quercus turbinella « ajoensis hybrid, J. Powell lot 
63D11, emerged 12 May 1963, P. Opler prep. 42, C. A. Tauber (Toschi) and J. Powell 
collectors. Allotype: female, same data except emerged 10 May 1963, P. A. Opler 
prep. 82. Paratypes: 3 64, 8 2@ same data except some emerged various dates 
between 19 April and 31 May 1963. 


VoLUME 31, NuMBER 1 49 


Host. Quercus turbinella < ajoensis. 
Distribution. R. kofa is known only in the Kofa Mountains, Yuma County, Arizona. 


Rhodophaea durata Opler, new species 
Fig. 5 


Diagnosis. This moth is extremely similar to R. caliginella but is sevarable by the 
combination of a pale brown or dark gray inner basal patch and the brown scales on 
the distal portion of the basal bar. 

Male. Length of forewing 7.5-9.5 mm (reared). Forewing as for R. caliginella 
male except inner basal patch of pale brown scales; basal bar of pale brown scales 
not reaching either costal or inner margins, inwardly edged with narrow black band; 
s-m patch of all black scales, limited in extent; seven small marginal patches. 

Genitalia. As in Fig. 11. Similar to that of R. cruza. Juxta lacking lateral projections. 

Female. Length of forewing 9 mm (reared). Forewing as for male except inner 
basal patch dark gray with a few reddish brown scales intermixed; basal bar with 
reddish brown instead of pale brown scales; s-m patch with dark more extensive, 
merging broadly with band; p-m patch distinct but reduced in extent. 

Type maierial. Holotype: male, California, Alpine Lake, 1100’, Marin County, 
25 April 1958, reared from Quercus durata, J. Powell lot 58D10, emerged 14 June 
1958, P. Opler prep. 75, J. A. Powell collector. Allotype: female, same locality as 
holotype, 15 April 1972, reared from Quercus durata, J. Powell lot 72D10, emerged 
27 June 1972, abdomen lost, J. Powell collector. Paratypes: 9 ¢4, 1 @ same 
locality date, 3 ¢ ¢ from 1958 collection, 5 ¢ 4, 1 2 (abdomen lost) from 1972 
collection, 1 ¢ 17 April 1970 reared from.Quercus durata, J. Powell lot 70D48, 
emerged 3 June 1970, J. A. Powell collector. 

Host. Quercus durata Jeps. 

Distribution. This moth is known only in the slopes of Mt. Tamalpais, Marin 
County. The range of the moth is presumed to be more extensive, since larval shelters 
were found on Q. durata at Cedar Mountain, Alameda County, and nine miles west of 
Atascadero, San Luis Obispo County. At other localities where the host was searched 
no Rhodophaea shelters were found, and R. durata has an even patchier distribution 
than its host. 


Rhodophaea fria Opler, new species 
Fig. 6 


Diagnosis. This moth is distinguished by its gray inner basal patch and the fact 
that the p-m line is indistinct near the inner margin. 

Male. Length of forewing 9.5-10 mm (reared). Forewing as for R. caliginella 
male except inner basal patch gray; basal bar with outer portion brown; s-m patch 
small, restricted; band present, dark on costal margin, paler toward inner margin; 
p-m line indistinct near inner margin, very narrow. 

Genitalia. As in Fig. 12. Valva with a bulbous, irregular projection on medial 
portion of inner face. Juxta with darkened sclerotized areas not extending to base. 

Female. Length of forewing 9.5 mm (reared). Forewing as for male except s-m 
patch more extensive. 

Genitalia. As in Fig. 17. Corpus bursa with two signa consisting of approximately 
rectangular patches of pointed projections arranged in rows, and with a small 
quadrangular patch of scobinations. 

Type material. Holotype: male, California, 4 mi. SE Clayton Contra Costa 
County, 26 May 1968, reared from Quercus douglasii, J. Powell lot 68E64, emerged 
27 June 1968, P. Opler prep. 35, P. A. Opler collector. Allotype: female, same data 
except emerged 10 July 1968. Paratypes: 5 66,5 92“, same data as holotype, 
emerged 14 June—5 July 1968; 3 @¢, Russelmann Park, Contra Costa County, 


50 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


ifs) 


Figs. 14-15. Rhodophaea, male genitalia, left valva not shown, aedeagus and juxta 
removed, A. juxta, B. aedeagus, 14. R. yuba, 15. R. suavella. Scale bar = 1 mm. 


VOLUME 31, NuMBER 1 51 


Fig. 16. Rhodophaea advenella, male genitalia, left valva removed, A. juxta, B. 
aedeagus. Scale bar = 1 mm. 


California, 11 May 1968, reared from Quercus douglasii, J. Powell lot 68E23, emerged 
10-14 June 1968; 1 ¢ 8 miles southeast Clayton, Contra Costa County, California, 
26 May 1968, reared from Quercus douglasii, J. Powell lot 68E53, emerged 27 June 
1968, P. A. Opler collector. 

Host. Quercus douglasii H. T. 

Distribution. In addition to adults reared from Q. douglasii at localities on the 
eastern slope of Mt. Diablo, Contra Costa County, shelters were found on this host 
at Folsom Dam (Sacramento County), 6 mi. SW Mariposa, Mariposa County, and at 
Keene and 6 mi. S Monolith, both Kern County. 

Discussion. This species is the only defined Nearctic Rhodophaea which feeds on 
a deciduous host. The leaves which surround its silken larval shelters are tied to 
host twigs prior to leaf abscission in fall. The partially grown larvae thus overwinter 
in situ and complete their development on newly produced leaves in the spring. 


Rhodophaea neva Opler, new species 
Fig. 7 

Diagnosis. This moth is separable by the combination of a dark basal area, broad 
basal bar, and vague indistinct patches. 

Male. Length of forewing 8.5-9 mm (reared). Forewing scaling of allotype male 
rubbed presumed to be as in female. 

Genitalia. As in Fig. 13. Valva with a ridged projection on basal portion of inner 
face surmounted by a tubercular process. 

Female. Length of forewing 8.5 mm (reared). Forewing as for R. caliginella 
males except inner basal patch of reddish brown scales; basal area dark, pale only 


JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


l 


Figs. 17-18. Rhodophaea, female genitalia, 17. R. fria, 18. R. kofa. Scale bar = 


mn. 


VoLuME 31, NuMBER 1 53 


Figs. 19-20. Rhodophaea, female genitalia, 19. R. neva, 20. R. advenella. Scale 
bar — | mm. 


54 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 21-23. Rhodophaea, female genitalia, 21. R. caliginella, 22. R. suavella, 23. 
R. cruza. 


adjacent to basal bar; basal bar broad, of reddish brown scales, inner black portion 
for posterior %; band especially black at costal margin becoming diffuse inwardly; 
scattered pale brown scales along inner margin. Marginal patches vague, not distinct. 

Genitalia. As in Fig. 19. Corpus bursa with two signa similar in configuration to 


VoLuME 31, NuMBER 1 55 


those of R. fria, but with one distinctly larger. The scobinate patch is between the 
signa, rather than to one side as in R. fria, and is triangular in outline. 

Type material. Holotype: female, Nevada, summit Kingsbury Grade, Douglas 
County, 30 June 1968, reared from Chrysolepis sempervirens, J. Powell lot 68F108, 
emerged 9 July 1968, P. A. Opler collector. Allotype: male, same data except 17 May 
1969, J. Powell lot 69E88, emerged 29 June 1969. Paratypes: 2 2 2, same data as 
holotype, emerged 6 and 9 July 1968. 

Host. Chrysolepis sempervirens ( Kell.) Hjelm. 

Distribution. This moth is known only from the type locality, but it probably 
occurs at other localities where its host occurs in the Carson Range. No evidence of 
this species’ presence was found at other localities where the host was sampled. 
Sampling of Chrysolepis chrysophylla (Doug. ex. Hook.) Hjelm. failed to disclose 
evidence of Rhodophaea. 


Rhodophaea yuba Opler, new species 


Diagnosis. This moth has a more uniform pale gray appearance than its congeners, 
and is further distinguished by its indistinct band and pale gray tan scales on the 
distal margin of the basal bar. 

Male. Length of forewing 10.5 mm (reared). Forewing as for R. caliginella male 
except inner basal patch of pale tan scales; basal bar indistinct, pale gray tan, nar- 
rowly edged inwardly with black; s-m patch restricted; p-m patch with dull white 
scales: band present but indistinct; p-m line distinct but narrow, becoming indistinct 
toward costal margin; marginal patches a distinct narrow broken line, becoming 
indistinct at inner margin. 

Genitalia. As in Fig. 14. Valva with a low irregular ridge-like protuberance on 
medial portion of inner face. Juxta truncate ventrally. 

Type material. Holotype: male, California, Yuba Pass summit, 6708 feet eleva- 
tion, Sierra County 19 April 1968, reared from Quercus vaccinifolia, J. Powell lot 
68D152, emerged 10 June 1968, P. A. Opler collector. 

Host. Quercus vaccinifolia Kell. 

Distribution. This moth is known only from the type locality. R. ywba may range 
widely through the range of its host, which was sampled at few localities. 


Rhodophaea suavella (Zincken ) 


Phycis suavella Zincken, 1818. 

Myelois suavella: Herrich-Schaffer, 1849. 

Eurhodope suavella: Meyrick, 1927. 

Rhodophaea suavella: Ragonot, 1893. 

Rhodophaea supposita Heinrich, 1956 (New Synonymy). 


Diagnosis. This moth is distinguished from other Rhodophaea by its dark appear- 
ance, absence of an s-m patch, and more distal placement of the basal bar which 
lacks black scaling. 

Male. Length of forewing 9-10 mm (reared). Maculation as in R. caliginella 
except inner basal patch brown; basal area dark, not clearly differentiated; basal band 
positioned more distally on wing, narrow white, not edged inwardly with black; s-m 
patch absent; p-m patch strongly reduced; band present but indistinct on posterior 
half of wing. 

Genitalia. As in Fig. 15. Valva with long projection on inner face surmounted by 
a more heavily sclerotized circular setiferous area. 

Female. Length of forewing 8.5 mm (reared). Forewing as for male. 

Genitalia. As in Fig. 22. Corpus bursa lacking distinct signa, but with an indented 
scobinate patch on medial portion, and a small area of minute thorn-like projections 
on proximal portion. 


56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Hosts. Various Rosaceae: Cotoneaster (British Columbia), Crataegus spp. (En- 
gland), Prunus (France ). 

Distribution. In North America known only from Vancouver, British Columbia, 
Canada. In the Palaearctic in southern England, central and southern Europe, as 
well as the Near East. 


Future Taxonomic Work 


During the 1967-1970 study of California Oak Microlepidoptera, 
Rhodophaea larvae and shelters were discovered on five hosts from 
which no reared material is currently available (Opler, 1970). These 
were Lithocarpus densiflora, Quercus chrysolepis, Q. dunnii, Q. engel- 
manni and Q. garryana. Moths feeding on Lithocarpus can be expected 
to represent an undescribed species, while at least one and possibly 
other undescribed species are represented by moths which feed on the 
other four. The specific identity of Rhodophaea associated with Quer- 
cus dumosa throughout its range must be studied in detail. 


ACKNOWLEDGMENTS 


J. A. Powell contributed significantly to all phases of this study, and 
reviewed an early draft of the manuscript. Facilities at the Smithsonian 
Institution were made available through the kindness of W. D. Duck- 
worth, who made arrangements for the adult photographs. During this 
study's investigative phase (1967-1970) supporting funds were provided 
by National Science Foundation grants GB 4014 and GB 6813X (Princi- 
pal Investigator, J. A. Powell). 


LITERATURE CITED 


Hernricu, C. 1956. American moths of the subfamily Phycitinae. U.S. Natl. Mus. 
Bull. 207, 581 p. 

Orter, P. A. 1970. Biology, ecology, and host specificity of Microlepidoptera 
associated with Quercus agrifolia (Fagaceae). Doctoral thesis, University of 
California, Berkeley. 

1974. Biology, ecology, and host specificity of Microlepidoptera associated 

with Quercus agrifolia (Fagaceae). Univ. California Publ. Entomol. 75: 1-83, 

anolse 


VoLuUME 31, NuMBER 1 od 


A NEW NORTH AMERICAN SPECIES OF APAMEA 
FORMERLY CONFUSED WITH A. VERBASCOIDES 
(GUENEE) (NOCTUIDAE) 


DoucLas C. FERGUSON 


Systematic Entomology Laboratory, IIBIII, Agr. Res. Serv., USDA 
c/o U.S. National Museum, Smithsonian Institution, Washington, D.C. 20560 


While “sugaring” for moths in early August 1961, near Lake 
Kejimkujik, Nova Scotia, in what has since become Kejimkujik National 
Park, I collected five specimens of a species of Apamea Ochsenheimer 
(formerly Septis Hubner) that might have been mistaken for verba- 
scoides (Guenée) except that they did not appear entirely normal. 
Even when sitting on the tree trunks the moths seemed smaller and 
brighter than verbascoides, with the gray and reddish areas of the fore- 
wings more prominent and contrasting. The species considered to be 
the true verbascoides is not uncommon in that region and was even 
collected in the same place on the same field trip. Subsequent investiga- 
tion revealed that the five unusual specimens represent an undescribed 
species with distinguishing characters in the genitalia of both sexes. 
Study of material from other collections disclosed that the new species 
occurs also in eastern Massachusetts, the vicinity of New York City, and 
in the pine barrens area of New Jersey. In New Jersey it seems to be 
much commoner than verbascoides. 

Although it has thus been apparent for some time that two species 
are involved, I delayed publishing on this problem because of uncertainty 
as to the proper application of the one available name, Xylophasia verba- 
scoides Guenée (1852: 141), based on specimens collected by Edward 
Doubleday in the state of New York (probably at Trenton Falls, Oneida 
Co.). The original description does not give the exact locality or the 
number or sex of the type specimens; no holotype was specified. Through 
the kindness of Mr. D. S. Fletcher, I have been able to examine one male 
and two female specimens in the collection of the British Museum 
(Natural History) that are believed to be syntypes of verbascoides. As 
expected, these clearly represent the larger, commoner, more widely 
distributed species. None of the supposed syntypes bears an original 
type label or any label with the name verbascoides, and it may therefore 
be argued that there is no proof of their authenticity as types. However, 
there is no doubt that the original description fits this species or its 
sibling herein described, and Guenée clearly stated the source of the 
specimens as “Amérique du Nord, Etat de New-Yorck. Coll. Doubleday.” 


58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The three examples in question in the British Museum are labelled as 
Doubleday specimens, and it is reasonable to assume that these are the 
specimens Guenée described, although we cannot be sure that he saw 
all of them. Assuming that he did, I hereby designate the male of the 
syntypic series as the lectotype. It bears the following labels: “699” 
(on green paper); “Xyloph. Cucullioides Gn.”; “U.S. America. Double- 
day. 46—110.”; “Type” (red-bordered circular label). A lectotype 
label has been added. Guenée did not describe a Xylophasia cucul- 
lioides, and the meaning of this label is uncertain. There is no cucul- 
lioides in Apamea or in any closely related genus. However, Guenée 
made a point of comparing verbascoides with species of Cucullia, deriv- 
ing the name itself from that of the European Cucullia verbasci (L.), 
and he may have changed his mind about what to call it; or perhaps he 
or Doubleday simply made a lapsus calami in labelling. Other species 
that appear most closely related are Apamea nigrior (Smith) of the 
Northeast, which is a much duller brown species, and A. cuculliformis 
(Grote) of western North America, easily distinguished by its larger 
size, paler coloring, and reduced dark markings on the forewing. 

Apamea verbascoides occurs across Canada from Newfoundland to 
Saskatchewan and in the northern United States from Maine at least to 
Wisconsin, and southward to Pennsylvania, southern New Jersey, and at 
higher elevations in the Appalachians to North Carolina. It has one 
brood which flies from late June to early August. To my knowledge 
nothing has been reported on the early stages or host plants. The ex- 
ample figured by Holland (1903, pl. 19, fig. 43) is a female of the true 
verbascoides. In the illustrations published by Barnes & McDunnough 
(1913, pl. 14), figures 1 and 2 are of verbascoides, but figure 3 repre- 
sents the unnamed species, which I describe as follows: 


Apamea inebriata Ferguson, new species 


Description. Upperside of forewing like that of A. verbascoides except for the 
following differences: areas of brown coloring of a brighter, more red-brown shade, 
this being especially evident through the middle of the wing longitudinally; costal 
area with more gray scales; pale zone along inner margin from near base to postmedial 
line more solidly gray and, especially in fresh females, much more contrasting than in 
verbascoides; basal dash weak or absent and, if present, brown, diffuse, merely a 
continuation of the deep red-brown shade that runs through middle of wing, not a 
distinct, tapered, sharp-pointed black streak as in verbascoides; small patch of white 
scales at juncture (or apparent juncture ) of Mz and Cu: at outer end of cell, character- 
istic of verbascoides, nearly always absent or much reduced, this point being marked 
only with a few light yellowish-brown scales or not at all. Although not 100% 
reliable, this is a very useful character in fresh specimens. 

Upperside of hindwing, undersurface, and vestiture of head and body about as in 
verbascoides. No visible differences in antennae, palpi, or other external structures. 

Length of forewing: males, 17-19 mm; females, 15-18 mm; holotype male, 18 mm; 


VoLUME 31, NuMBER 1 59 


3 


Figs. 1-4. Apamea spp. 1, A. inebriata ¢, holotype; 2, A. inebriata 2, allotype; 
3, A. verbascoides 6, Lake Kejimkujik, Queens Co., Nova Scotia, 8 Aug. 1961; 4, A. 
verbascoides 9, Halifax watershed area, Halifax Co., Nova Scotia, 28 July 1957. 
About natural size. 


allotype female, 16 mm. Mean wing length: males (of 11), 18.09 mm; females (of 
11), 16.95 mm. (For verbascoides these measurements are as follows: males, 
17.5-20.5 mm; females, 16.5-19.5 mm; lectotype male, 20 mm. Mean wing length: 
males (of 34), 19.12 mm; females (of 33), 18.27 mm. Although both species 
are variable, it is thus apparent that the forewing of inebriata averages more than one 
mm shorter than that of verbascoides. When series of the two species are compared, 
the size difference is readily noticeable without measurement. 

The most obvious characters in the male genitalia that distinguish this species from 
verbascoides are found in the vesica and in the form of the basocostal lobe of the 
valve. On the inner face of the valve the basocostal corner takes the form of a large, 
protuberant, rounded lobe in verbascoides, overlying the point of articulation of 
costa and tegumen; in inebriata the lobe is reduced to less than half this size, is 
somewhat irregular, and does not completely overlie the articulation of costa and 
tegumen. Also, the lobes are nearly symmetrical in verbascoides, more often asym- 
metrical in inebriata, the one on the left valve being shallowly notched, the one on 
the right valve entire. The sclerotized structures of the aedoeagus and vesica are 
quite similar in the two species but differently situated, and the vesica itself is 
differently shaped. The everted vesica consists of two divergent lobes, one being 
the actual ductus ejaculatorius and the other a blind pouch. The lobe leading to the 
ductus ejaculatorius bears, dorsally, a cluster of several cornuti similarly situated in the 
two species. The other lobe bears one large tooth with a broadly expanded base, in 
verbascoides located at about the same relative position as the cluster of small cornuti 
on the opposite lobe but ventrally; in inebriata this large tooth occupies an apical 
position at the very end of the blind lobe, and the lobe itself is only half as long. 
There are one or two additional cornuti of intermediate size attached to the wall of 
the aedoeagus by a straplike connection; in inebriata these are smaller, and the 
straplike connection is only half as long as in verbascoides; also, they are located in a 
left sublateral position instead of being midventral as in verbascoides. To the right 
of this, at the end of the aedoeagus, lies a disk bearing 8 to 20 much smaller cornuti; 
this is almost exactly ventral in inebriata but lies in a right sublateral position in 
verbascoides. 


60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


"Sa 


Ne 

Sere tes 
he 
vl 


Figs. 5-11. Genitalia of Apamea spp. 5, A. inebriata 8, genitalia of holotype, 
aedoeagus omitted; 6, A. inebriata ¢, aedoeagus of holotype; 7, A. inebriata ¢, 
aedoeagus of paratype from Martha’s Vineyard, Massachusetts; 8, A. verbascoides 6, 
aedoeagus, West Dover, Halifax Co., Nova Scotia; 9, A. verbascoides 6, right valve 
of same specimen; 10, A. verbascoides 2, Lake Kejimkujik, Queens Co., Nova Scotia; 
11, A. inebriata 9, paratype, Lake Kejimkujik, Nova Scotia. 


VoLUME 31, NuMBER 1 61 


In the female genitalia the only important difference seems to be in the size of the 
ductus bursae. This structure is rather rigidly sclerotized, heavily rugose, and finely 
but densely scobinate in both. In inebriata, however, it is shorter and more slender, 
being no more than two-thirds as thick as that of verbascoides. 

Types. Holotype: ¢, Lake Kejimkujik, Queens Co., Nova Scotia, August 7, 1961 
(at bait), D. C. Ferguson, U.S. National Museum Type No. 74,001 (fiz. 1). Allotype: 
2, same data but taken August 6 (fig. 2). Paratypes: 1 ¢, 2 9 9, same locality, 
August 5, 6, 11, 1961, D. C. Ferguson; 1 ¢, Concord, Massachusetts, July 21, 1913, 
William Reiff; 1 ¢, Martha’s Vineyard, Massachusetts, July 29, 1948, F. M. Jones; 
I g¢, CN.Y./7-8-02,” Coll. A. C. Weeks; 1 2, “N. York”; 1 4, Jerseyville, 3 mi. E 
of Freehold, New Jersey, July 9, 1961, M. Shulgin; 1 @, Freehold, New Jersey, July 
17, 1955, M. Shulgin; 8 ¢ 6, 3 9 2, Lakehurst, New Jersey, June 26—-July 27, 1955, 
J. G. Franclemont; 18 ¢ 6,13 2 9, Lakehurst, New Jersey, June 29-30, July 1, 1937, 
July 1-25, July 4, 1910, July 12, July 17, July 17-30, July 19, July 25, Fred’k. 
Lemmer; 1 @, Lakehurst, New Jersey, July 10, 1928, T. D. Mayfield; 2 92 9, 
Pitman, New Jersey, July 11, 1910; 1 ¢@, Elizabeth, New Jersey, “8-6-08"; 1 9, 
Morris Co., New Jersey, May 26, 1937 [wrong date?]; 1 2, Jersey City, no date. 
Holotype and allotype in collection of U.S. National Museum; paratypes in U.S. 
National Museum, American Museum of Natural History, Peabody Museum of 
Natural History at Yale University, Franclemont collection at Cornell University, 
Canadian National Collection, Nova Scotia Museum, and British Museum (Natural 
History ). 

Distribution. The localities given for the types represent the entire known distribu- 
tion, which seems limited to areas near the coast. The type-locality, 30 miles from 
Annapolis Basin, on the Bay of Fundy, is the farthest inland; the U.S. localities all 
appear to be less than 20 miles from tidewater. 

Early stages. Unknown. 

Remarks. The habitat where inebriata was collected in Nova Scotia is different 
from that of the New Jersey pine barrens, being Canadian Zone woodland, although 
of a rather southern type bordering on Transition Zone. It is characterized by a 
mixture of second growth white and red pine, red spruce, hemlock, balsam fir, beech, 
red oak, sugar and red maple, white and yellow birch, aspen, ash, hop hornbeam, 
and a wide variety of shrubs, mainly those that favor acid soil conditions in that 
region. The moths were taken along old logging roads on the eastern side of Lake 
Kejimkujik close to the boundary between Annapolis and Queens counties, and about 
one-third of a mile north of the Grafton Lake fish hatchery. Apamea inebriata is a 
new addition to the list of northeastern endemics with curiously limited, or disjunct, 
coastal distributions. I believe that these are relicts of a Pleistocene coastal plain 
fauna which, in part, survived glaciation on large, emergent island refugia off New 
England, Nova Scotia, and Newfoundland in a climate tempered by proximity to 
the Gulf Stream. This was the subject of my paper, The Grand Banks Refugium, 
presented at the 1974 annual meeting of The Lepidopterists’ Society. 


ACKNOWLEDGMENTS 


I am indebted to D. S. Fletcher of the British Museum (Natural His- 
tory) for the opportunity to see and photograph the types of verbasco- 
ides, to J. G. Franclemont of Cornell University, F. H. Rindge of the 
American Museum of Natural History, and J. D. LaFontaine of the 
Biosystematics Research Institute, Agriculture Canada, for the privilege 
of examining additional material, and to E. L. Todd of the Systematic 
Entomology Laboratory for a very helpful review of this paper. 


62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


LITERATURE CITED 


Barnes, W. & J. H. McDunnoucu. 1913. Contributions to the natural history of 
the Lepidoptera of North America 2(1): pls. 1-21. 

GuENEE, A. 1852. Species Général des Lépidopteres, Noctuélites 1: I-XCVI + 
1-407 pp. 

Hoxuanp, W. J. 1903 and later editions. The moth book. Doubleday, Page & Co., 
New York. 479 p., 48 pls. 


STAPHYLUS AZTECA, NEW RECORD FOR THE UNITED 
STATES (HESPERIIDAE ) 


Staphylus azteca (Scudder, 1872) 


Type locality. Tehuantepec, Oaxaca, Mexico. 

Distribution. Mexico: Mante, Tamaulipas, June 1967 (H. A. Freeman); Ciudad 
Valles (Hotel Covadonga), San Luis Potosi, June, July, August 1966-1975 (H. A. 
Freeman); Tamazunchale, San Luis Potosi, June 1967 (H. A. Freeman); San Blas, 
Nayarit, September 1966 (W. S. McAlpine); Comala and Colima, Colima, March, 
April 1967 (Robert Wind); and Catemaco, Veracruz, August 1967 (H. A. Freeman). 
Guatemala, Salvador, and Costa Rica (various records in the British Museum and 
American Museum of Natural History, New York). 


On 2 June 1940 I collected a female specimen of Staphylus at Alpine, Brewster 
County, Texas, approximately 11 mi. N of town. This specimen was sent to Mr. 
E. L. Bell at the American Museum of Natural History for determination. In 
response Mr. Bell wrote that this specimen being a female was not possible to 
identify as it was not like any they had in the museum and could possibly represent 
a new species. After searching for males for many years in that general area north 
of Alpine with no results I finally gave up and left the specimen unidentified in my 
Texas collection. Recently I re-examined the specimen and compared it with my 
Mexican Staphylus and found it to be azteca (Scudder). This represents a new Hes- 
periidae record for the United States. 


Hucu Avery FREEMAN, 1605 Lewis Drive, Garland, Texas 75041. 


VOLUME 31, NuMBER 1 63 


ADDITIONAL MATERIAL OF SCOPARIA HUACHUCALIS 
MUNROE, WITH DESCRIPTION OF MALE GENITALIA 
(PYRALIDAE, SCOPARIINAE) 


EUGENE MUNROE 


Biosystematics Research Institute, Agriculture Canada, 
Ottawa, Ontario KIA O0C6 


Scoparia huachucalis Munroe was described from a single damaged 
female in the Los Angeles County Museum. (Munroe, 1972: 46, “1973” 
[1974]: pl. 2, fig. 42, pl. H, fig. 2). Subsequently, through the kindness 
of Mr. Julian Donahue, I have been able to examine a short series of 
better specimens from Madera Canyon, Santa Rita Mountains, Arizona, 
6000 ft., 21 May 1940, Lloyd M. Martin. The maculation (Figs. 1, 2) 
agrees well with that of the holotype, but the reniform spot of the fore- 
wing is filled in with light brown and the hind wing is lightly infu- 
scated. 3 

The characters of the male genitalia (Fig. 3a, b) are as follows: 
Uncus long, narrowly hood-shaped, lightly setose laterally and dorsally. 
Gnathos narrow, as long as uncus, with short basal section and long, 
weakly sinuate, slender, strongly sclerotized distal section, finely spinu- 
lose dorsally near tip, and with extreme apex decurved and acute. Juxta 
small, with evenly convex ventral margin, acute lateral angles, and con- 
cave dorsolateral margins, converging to the long, acute dorsal extremity. 
Vinculum with ventral part produced into a short rounded _ saccus. 
Valve about three times as long as greatest width, weakly expanded 
distally, tip symmetrically rounded; costa narrowly inflated; sacculus 
scarcely inflated, but with distinct free distal spine from ventral margin 
at about three-fourths from base to tip of valve. Penis curved, about 
five times as long as wide, with a single short, straight, spinelike cornutus. 

The original description of the female genitalia states that the ductus 
bursae of the female genitalia is slender and membranous. Actually the 
proximal part of the ductus (Fig. 4a) is weakly sclerotized and tapers for 
some distance, then there is a short membranous zone leading to a 
sclerotized and contorted zone, bearing a few spinules; the remainder 
of the ductus is wider and membranous, leading to the oval to round 
bursa; the spinules of the latter (Fig. 4b) are arranged in two large 
patches, facing each other on opposite sides; the spinules of one patch 
are more widely spaced and larger than those of the other; the large 
spinules have stellate bases. These features are visible in the preparation 


64 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1, 2. Scoparia huachucalis Munroe, Madera Canyon specimens. 1, male; 2, 
female. 


of the holotype genitalia and in the figure in The Moths of America 
North of Mexico, but are more obvious in the new preparation. 


ACKNOWLEDGMENTS 


I thank Mr. Julian Donahue, Los Angeles County Museum of Natural 
History, for the loan of material as already detailed. Slides of genitalia 


«. BAS 


male genitalia 


? 


imen 


Madera Canyon spec 


? 


Munroe 
penis. 


is 
b, 


) 


s removed 


ni 


. Scoparia huachucal 


1 with pe 


b 


vt 


ON hai Hil hee 


la. 


, female genital 


d proximal part of ductus bursae; b, distal part of ductus 


cimen 


Munroe, Madera Canyon spe 


is 


huachucal 


aria 
ium an 


tor, ost 


ae and bursa. 


i 


a, ovipos 


b 


urs 


Fig. 4. Scop 


66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


were made by Mr. Douglas Kritsch, Biosystematics Research Institute. 
Photographs were made by Mr. Tom Stovell, Graphics Unit, Research 
Branch, Agriculture Canada, with the assistance of Mr. Kritsch. The 
plates were mounted by Mr. R. Bennet, Graphics Unit. 


LITERATURE CITED 


Munroe, E., in Dominick, R. B., et al. 1972. The moths of America north of 
Mexico, fasc. 13.1A, Pyraloidea (in part). London. 

Munrog, E., in Dominick, R. B., ef al. “1973” [1974]. Ibid., fasc. 13.1C, Pyraloidea 
(in part). 


VoLUME 31, NuMBER 1 67 


GENERAL NOTES 


EFFECTS OF 1933 HURRICANES ON BUTTERFLIES 
OF CENTRAL AND SOUTHERN TEXAS 


Hurricanes are massive tropical storms that may be accompanied by tremendous 
winds, precipitation, and/or surge tides. Such storms are normal occurrences in 
southem Texas, having been observed since the beginning of historical records 
(Schlesselman, 1945, Trans. Tex. Acad. Sci. 38: 173-182). Two hurricanes struck 
the Brownsville area in 1933 (5 August and 4 September) causing widespread dam- 
age and torrential rains from there southward into Mexico. On 6 July a hurricane 
struck the Mexican coast approximately equidistant from Tampico, Tamaulipas, and 
Brownsville. Rainfall from this hurricane in the Brownsville area broke a lengthy 
drought (July precipitation was 4.50 in. vs the normal of 1.68 in.). Rainfall from 
November 1932—June 1933 inclusive totaled 8.59 in., compared with a normal 
rainfall of 13.66 in. for November—June. Precipitation at Brownsville during July— 
September 1933 was 26.14 in., compared with an average of 9.44 in. (normal 
annual precipitation is only 26.75 in.). Rainfall in central Texas was unaffected by 
these hurricanes; San Antonio precipitation during July-September 1933 was 7.94 
in. (exactly normal). 

Rainfall surges following droughts typically cause large increases in the numbers 
of many lepidopteran species, some of which subsequently migrate in tremendous 
numbers. The September storm was very severe, whereas the August storm was 
relatively minor (names were not applied to tropical storms until 1953). All three 
storms had profound effects on the biota of southern and central Texas. Three literature 
reports describe tropical butterfly occurrences that seemingly resulted from these 
storms, although the observers did not connect their observations with the hurricanes. 
These three accounts deal with, respectively: 1) initial population movements in 
direct response to environmental disturbances; 2) establishment, albeit temporarily, 
of a breeding population in the same year as the storms; and 3) apparent overwinter 
survival and continued reestablishment of a population during the following year. 

H. B. Parks (remarks published by Engelhardt, 1934, Bull. Brooklyn Ent. Soc. 
29: 16) collected numerous specimens of Anteos chlorinde nivivera Fruhstorfer and 
Anteos maerula lacordairei (Boisduval), both Pieridae, at San Antonio, Bexar Co. 
during the last week of August and first week of September 1933. Parks. states, 
“these huge butterflies were a glorious sight. They arrived in large numbers and 
stayed with us about a week.” San Antonio was apparently the center of the flight, 
but specimens were also seen or reported at Kerrville, Austin, and San Antonio. 
Northward movement of these species was apparently in direct response to the 
August storm. During this same period, and subsequently, he collected the following 
species: Hesperiidae: Chiomara asychis georgina (Reakirt); Nymphalidae: Siproeta 
steneles biplagiata (Fruhstorfer) and Marpesia petreus (Cramer); Pieridae: Phoebis 
philea (Johanssen); and Heliconiidae: Heliconius charitonius vasquezae Comstock 
& Brown and Dryas julia moderata (Stichel). By listing these species, Parks was 
indicating that these were unusual captures for the San Antonio area. This classifica- 
tion is verified by my observations in the area. The two heliconians are seen in 
most years in low numbers but normally not until late summer or fall, although early 
summer occurrence of both species was observed in 1968 (unpub. data). Siprocta 
steneles biplagiata has been observed sporadically in the Austin area. 

The second report involves a species that had not previously been seen in Texas 
or the rest of the United States and probably has not been observed since. A 
“perfect specimen” of Dryadula phaetusa (L.) was collected on 19 December 1933 
at Sarita, Kenedy Co. by H. Glazbrook (1934, Ent. News 45: 251-252). Dryadula 
phaetusa apparently was able to disperse northward in late 1933 after the storms 
and breed at Sarita. Larvae of this species feed on passionflower ( Passifloraceae; 


68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Passiflora). Passiflora foetida LL. var. gossypiella (Hamilton) Mast probably occurs 
in the more mesic wooded habitats of this area. No severe winter weather had 
been experienced in this area by mid-December 1933. The nearest weather records 
are for Corpus Christi, Nueces Co. (88 km to the NNE). Ironically, the coldest tem- 
perature for late 1933 was 43°F on 19 December. Dryadula phaetusa is known from 
the temperate and tropical regions of southern Mexico but extends up the coasts an 
unknown distance toward the United States (Hoffman, 1940, An. Inst. Biol. Mex. 11: 
639-739 ). 

The third report involves the occurrence, which may or may not be associated 
with these hurricanes, of another tropical butterfly in central Texas. Anartia jatro- 
phae luteipicta Fruhstorfer (Nymphalidae) was reported near Sutherland Springs, 
Wilson Co. in late October 1934 (Parks, 1935, Bull. Brooklyn Ent. Soc. 30: 83). 
The existence of newly emerged specimens among those collected indicates that a breed- 
ing colony had been established. Doubt about the association between the occurrence of 
this species and these hurricanes results from the observation of this butterfly west 
of San Antonio on 9 November 1931 by A. J. Boyles. However, Parks (op. cit.) 
states that “a very careful search has been made [since 1931] without success [until 
1934].” Therefore, occurrence of A. j. luteipicta in central Texas is believed to be 
the result of a colony established in late 1933 (or early 1934) after northward 
dispersal associated with the hurricanes of August and September 1933. Normally, 
A. j. luteipicta would not be expected to survive the winter cold of central Texas. 
However, the winter of 1933-1934 was exceptionally warm; the coldest temperature 
recorded at San Antonio (45 km NWW of Sutherland Springs) was 29°F. A popula- 
tion was probably established in late 1933 with successful overwintering and 
survival until the following winter. Survival through the 1934-1935 winter is 
unlikely (low temperature 18°F). Survival between the 1931 sighting and the 
putative 1933 establishment is unlikely since colder weather occurred in both 1931— 
1932 (low temperature 24°F on 13 March) and 1932-1933 (low temperature 12°F). 

In September 1967, Hurricane Beulah, after striking land at the mouth of the 
Rio Grande River near Brownsville, Cameron Co., Texas, brought extremely heavy 
rains to southern Texas. As a result of this storm, a fairly large number of lepidoptera 
species previously unknown in Texas (and the United States) have been reported in 
southern Texas (Doyle, 1970, J. Lepid. Soc. 24: 212; Heitzman, 1970, Mid-Cont. 
Lepid. Ser. 12: 10-11; Heitzman & Heitzman, 1972, J. Res. Lepid. 10: 284-286; 
Kendall, 1970, J. Lepid. Soc. 24: 59-61, 266; Kendall, 1972, Ibid. 26: 49-56). 
At least some of these species have established permanent populations in this area 
(see later collections reported by Tilden, 1974, J. Lepid. Soc. 28: 22-25). 


RaymMonp W. Necx, Texas Parks and Wildlife Department, John H. Reagan 
Building, Austin, Texas 78701. 


COLONY OF PIERIS NAPI OLERACEA (PIERIDAE) IN INDIANA 


According to Blatchley (1891, Ann. Rep. Indiana State Geol. 17: 365-408), Pieris 
napi oleracea—aestiva Harris was collected by Mr. A. B. Ulrey in Kosciusko Co., 
Indiana in the summer of 1890. When collecting extensively in Kosciusko Co. and 
other northeastern counties in the mid-1930’s and from 1964-1970, I failed to locate 
P. napi. On 12 July 1971, John Campbell, a high school student in my collecting 
party, collected one in the Pigon River State Fish and Game Area, Mongo, LaGrange 
Co., Indiana. My identification of this speciman as Pieris napi oleracea Harris was 
confirmed by Mr. Harry K. Clench of the Carnegie Museum. John and I returned 
to Mongo on 24 August 1971 and collected 23 more specimens. Since that time P. n. 
oleracea has been found annually in the Mongo tamarack bog, the largest bog of its 
type in Indiana and only 7 miles from the Michigan border. 


VoLUME 31, NuMBER 1 69 


Figs. 1-2. 1, left, habitat showing Barbarea sp., one of favorite larval foodplants of 
Pieris napi oleracea, Pigeon River State Fish and Game Area, Mongo, Indiana; 2, right, 
close up of P. n. oleracea, second summer brood ( photographs by David Eiler). 


Recently this species has suffered a great restriction in habitat, partly because of 
competition with Pieris rapae and partly because of habitat destruction by man. In 
the north central and eastern states P. napi is found only in the Transition and 
Canadian zones, not extending south of the Catskill Mountains in New York ( Klots, 
1951, A field guide to the butterflies, Houghton Mifflin, Boston, 349 p.). Old 
records are unreliable since P. napi was often confused with Pieris virginiensis, a 
single brooded species of more southern (Transition Zone) distribution. Pieris rapae, 
a species accidentally introduced from Europe into Quebec about 1860, has spread 
rapidly throughout most of North America (Howe, 1975, The butterflies of North 
America, Doubleday & Co., Inc., 633 p.). Pieris rapae continues to invade P. napi 
territory. Pieris n. oleracea formerly occurred in northern Illnois, but it is now ap- 
parently extinct in the state (Irwin & Downey, 1973, Illinois Nat. Hist. Survey, 
Biological Notes 81: 1-60). Thus, additional notes and records of this species in 
Indiana should be of interest to entomologists. 

Not only is P. napi geographically variable, but the generations of P. n. oleracea 
vary from one brood to another. On 19 May 1975 I collected the spring form in 
the Mongo tamarack bog. It had the veins prominently marked. On 1 July 1975 I 
collected 18 of the summer form (second brood) and observed dozens more flying 
in the bog. Members of this large colony flew up and down the bog, concentrating 
on a very limited patch of water cress (Barbarea) (Cruciferae) that nearly choked 
the narrow stream (Fig. 1). Jewelweed (Impatiens), shrubby cinquefoil ( Dasiphora 
fruticosa), the beautiful but dangerous poison sumac (Rhus vernix), narrow-leaved 
and broad-leaved cattail (Typha), and other plants were growing in this area among 
the scattered tamarack trees ( Larix laricina). Euphydryas phaeton was flying nearby, 
not far away from its larval foodplant, turtlehead (Chelone glabra). Most of the 
July P. n. oleracea were plain white, except for the blackish basal dustings and traces 
of black on the apical border; a few had washes of yellow on the underside of the 
hind wing. A specimen collected on 20 August 1974 had a faint dark spot in cell 
M: of the front wing, a rare variety. In a letter to me, Mr. Clench describes the spring 


70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


brood as having sharp, almost black, thin lines on the veins of the hind wing under- 
side, whereas the summer brood has no black striping on the veins (Fig. 2). In both 
broods the underside ground color on the hind wing and apex of the front wing is 
yellowish. Of the 18 collected on 1 July 1975 only three were females. The females 
are more seclusive and thus better protected for reproduction. Occasionally a few P. 
n. oleracea left the bog, flying into an adjoining open field. A few P. rapae likewise 
flew into the bog, mingling with P. n. oleracea. 

I found two separate colonies of P. n. oleracea in the bog, one at the eastern end 
of the Pigeon River camp grounds and the other at the western end. The two 
colonies were about one quarter of a mile apart. Collecting in the bog can be 
somewhat dangerous, not only because of the ever-present poison sumac, but because 
of the sponge-like soil, which makes the collector feel as if he were walking on a 
spring coil mattress. When I occasionally lost my footing, I simply fell forward and 
the vegetation easily supported my weight. Suprisingly, by 10 July 1975, I found 
only three P. n. oleracea in the bog, but by 31 July their numbers had greatly in- 
creased. Again I collected 18 and saw dozens more. The highlight of the day came 
when I found a pair in copula. It was 1235 EST and the temperature was 90°F. 
The pair was settled low in the grass and did not fly, even when I placed the net 
over them. When I touched the male through the net, it lifted the female upward. 
In the Pieridae I have found that only the males are the flight partners. After making 
these observations, as a conservation measure, I released the mating pair, which 
remained in copula. 

On 18 August 1975, Dr. David L. Ejiler and I collected in the two colonies at 
Mongo. I netted 25 P. n. oleracea but released six of them. Dr. Eiler caught seven, 
the first he had ever seen in their natural habitat. Dozens more were seen flying 
and feeding in the bog, and a few in the open field. In the bog itself, the P. n. 
oleracea far outnumbered the P. rapae. 

These data show that there were at least two broods of P. n. oleracea in the Mongo 
bog in 1975 and that the species seems to be well established in LaGrange Co., 
Indiana. 


Ernest M. Suutzt, 402 North Wayne Street, North Manchester, Indiana 46962. 


AN ECOLOGICAL NOTE ON POLITES SABULETI SABULETI 
AT THE NORTHERN LIMIT OF ITS RANGE (HESPERIIDAE)? 


The Sabuleti Skipper, Polites sabuleti sabuleti (Boisduval), is reported here for 
the first time as occurring in southern Canada, at Penticton in the Province of British 
Columbia. This locality represents the most northerly distribution of this skipper, the 
known geographic range of which extends from Washington State to Baja California, 
the Great Basin east to Colorado, and through western Arizona south into Mexico 
(MacNeill 1975, Family Hesperiidae, p. 423-578 in Howe, ed., The butterflies of 
North America, Doubleday & Co., Inc., New York). Its presence as an abundant and 
apparently established species at Penticton was discovered only recently. Previous 
faunal studies in southwestern Canada have omitted any mention of this skipper 
(Gregory 1975, Checklist of the butterflies and skippers of Canada, Lyman Ent. 
Mus. & Res. Lab. Mem. 3, 44 p.). Its range, therefore, seems to have been extended 
into southern British Columbia at some time during the quite recent past. 

Field observations indicate that this skipper is bivoltine at Penticton. The flight 
period of the adults extends from late May to early July and occurs again during 


' This paper was presented at the joint meeting of the Washington State Entomological Society 
and the Oregon Entomological Society in Pullman, Washington on 25 September 1976. 


VoLUME 31, NuMBER 1 ya 


the latter part of August and early September. The bivoltine condition prevails in 
Washington State as well (MacNeill, op. cit.). 

At Penticton, the observed nectar sources visited by adults of this skipper comprise 
two groups of plants in particular; namely, phlox and knapweed. During the flight 
period of the first generation, the major nectar source is Phlox longifolia Nutt. 
This plant possesses pink or lavender-coloured blossoms from May into June, and it 
is found abundantly as far north in the dry interior as Peachland (Lyons 1974, Trees, 
shrubs and flowers to know in British Columbia, J. M. Dent & Sons (Canada) Ltd., 
Toronto, Vancouver). During the flight period of the second generation, the 
predominant flowering plant is one of several species of knapweed. At an observation 
site near the Penticton Industrial Park, the knapweed present is diffuse knapweed, 
Centaurea diffusa Lam. This Eurasiatic plant is common in southern British 
Columbia, and individual plants possess either white or purplish flowers (Frankton 
& Mulligan 1970, Weeds of Canada, Canada Department of Agriculture Publ. 948). 

The Sabuleti Skipper seems to have pioneered this territory successfully, presuming 
that it truly was absent during the time of previous faunal studies. A rapid flying, 
small species of skipper, it is difficult to capture. However, it has recently been 
numerous in and around developed and inhabited areas, making it difficult to over- 
look. To accomplish a pioneering extension of range, the adults of this species had 
to be able to find adequate nectar sources during two periods of the growing season. 
Without a substantial and acceptable nectar source in the fall, second generation 
adults presumably would starve. Diffuse knapweed is the only abundant nectar 
source that this skipper has been observed to visit during the fall in the vicinity of 
Penticton. The presence of this particular species of introduced knapweed, there- 
fore, may have been directly associated with the success of northward pioneering 
and apparent establishment by this skipper in the dry interior region of British 
Columbia. ; 


ACKNOWLEDGMENT 


I thank the staff of the Lyman Entomological Museum & Research Laboratory, 
Ste.-Anne-de-Bellevue, Quebec, Canada, for making available work facilities and 
a reference collection of skippers during June 1976, at which time I determined the 
identity of the skipper species involved in this study. Voucher specimens have been 
deposited at that institution. 


J. ALLAN GarLAND. 2—1491 Government Street, Penticton, British Columbia V2A 
4V9, Canada. 


LARVAL HIBERNATION OF GEOMETRIDAE IN EASTERN UNITED STATES 


Over the past few years, I often encountered partially grown gray stick-like geo- 
metrid larvae in autumn well after leaf fall at various localities in Pennsylvania, New 
Jersey, and Massachusetts. Similarly, such larvae were also found in early spring 
before leaf development. They were found hanging by silken threads from low limbs 
or shrubs or on my clothing in fall and crawling actively in spring. 

I placed such larvae in small glass jars with tight lids and left them outside for 
the winter. Absorbent paper was placed in the jar for the larve to rest on. The larvae 
usually spun silken pads and attached themselves to the paper. The paper was 0oc- 
casionally dampened, often by placing a small amount of melting snow in the jar. 
The larvae usually survived until spring but most died in March, apparently from 
desiccation or excessive exposure to sunlight resulting in part from neglect. 

The few surviving ones were fed in spring whatever deciduous trees or shrubs 
that were readily available. In all cases mixtures of Rosa spp., Pyrus spp., Prunus 


72 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


spp., and Quercus spp. formed the bulk of the diet. From seven larvae so fed, the 
following results were obtained. One of two apparently identical larvae from a 
small series taken on a sugar maple (Acer saccharum) from 28 October—1 November 
1974 at Sunderland, Franklin Co., Massachusetts eclosed in late May 1975 as a 
melanic of a Hypagyrtis sp., probably H. unipunctata (Haworth). The other larva 
died in May after the last molt and is preserved at the Peabody Museum of Natural 
History at Yale University. 

Another moth of this genus was reared from a larva found on white oak (Quercus 
alba) at Glendale, Camden Co., New Jersey on 30 November 1975. The larva fed 
on the usual plants and also molted once in addition to pupation. Eclosion was in 
June 1976. The specimen is a female of the powdery brown form common in and 
near the New Jersey Pine Barrens. Many other identical appearing larvae were 
collected earlier in November in Cape May Co., New Jersey mostly on white oak. 
These died from mold in February. I tentatively identified the reared moth as Hypa- 
gyrtis pustularia Hubner. 

A larva found on my clothes at Leverett, Franklin Co., Massachusetts 10 November 
1873 eclosed the following May as Protoboarmia porcelaria Guenée. The larva molted 
once or twice before the pupal molt. It fed in the spring on all of the food plants 
mentioned. A similar, but more mature larva was found on 25 March 1974 at Batsto, 
Burlington Co., New Jersey. It was crawling on the twigs of Leucothoe racemosa on 
which the flower racemes were just beginning to grow. These proved to be suitable 
as food in the laboratory. This larva was parasitized by a tachinid, and its identity 
was not definitely established. 

On 22 April 1974 large gray geometrid larva with short lateral filaments was 
found on an apple limb at Amherst, Massachusetts. The tree had partially expanded 
leaves at the time, which were the bulk of the diet of this larva in the laboratory. 
It eclosed in late May as a Campaea perlata Guenée. It did not molt prior to pupation. 

Finally, on 20 April 1975 a large, rather slender gray larva with a protuberance on 
one thoracic segment was found crawling in a blueberry ( Vaccinium vaccillans ) patch 
on the barrens at Batsto, New Jersey. The spring was quite late, and the only plants 
with any new growth in the immediate area were Leucothoe racemosa, whose racemes 
had partially expanded in late February. This larva was reared to adult, chiefly 
on Pyrus spp. Prunus serotina and Vaccinium corymbosum were also offered and 
eaten freely. It molted once. The moth eclosed very late in May as Euchlaena 
astylusaria Walker, a female of the tan form. 

Unfortunately, detailed descriptions are not available for the above larvae. All 
were marked solely with gray, black, and brownish shades. All rested in the typical 
stick-mimicking posture of many Geometridae. 

Besides the larval hibernation, I have noticed another similarity between these 
species, namely that all have two generations in the above areas, and adults of the 
spring flight period are larger in size than those of the later emergence. This is espe- 
cially pronounced in females of Campaea perlata. It seems likely that this phe- 
nomenon is the direct result of feeding on superior quality spring foliage. 


Date F. Scuweirzer, Peabody Museum of Natural History, Yale University, New 
Haven, Connecticut 06520. 


VoLUME 31, NuMBER 1 he 


NOTES AND NEWS 


Recent Letters 


Dear Dr. Godfrey: 


I noted with great interest Dr. Ferris’s review of The Butterflies of North America 
by William H. Howe in the recent issue of the Journal (vol. 30(2): 138-143). As a 
devoted collector and student of Holarctic Rhopalocera, I purchased the volume 
shortly after it appeared, although strong misgivings had been expressed to me by 
several American friends who, like Dr. Ferris, are to be considered leading profes- 
sional lepidopterists. As far as I can judge (from collecting Nearctic butterflies only 
during the first six and most inexpert years of my 30 as a lepidopterist), their and 
Dr. Ferris’s criticisms are beyond dispute and extremely well-founded. 

In very large part the shortcomings of this volume could have been avoided; 
rather few seem to be attributable to lax editorship and the inevitable unevenness 
that results from the varying competence of 21 contributors. In a particular field where 
I am better informed (worldwide Parnassiinae being one of my areas of specializa- 
tion), Dr. Ferris’s judgment, though basically sound, has perhaps been more severe 
than necessary. The current status of Parnassius Latreille is so totally chaotic the 
world over, due to hopeless oversplitting, that Jon and Sigrid Shepard have under- 
standably and even properly gone in for some solid “lumping.” Almost un- 
doubtedly they have overdone this. But, like most workers in this group, they have 
uncritically accepted most of the voluminous work of the two most recent author- 
ities among Parnassiologists, F. Bryk and C. Eisner, who are heavily responsible for 
the oversplitting; for instance, by now 200 odd subspecies of Palaearctic P. apollo 
L. have been described. 

Undoubtedly, however, Dr. Ferris’s strictures must be considered as entirely fair 
on Howe’s book as a whole, and we are left with the problem of making the best 
of it. I feel strongly that we cannot content ourselves with criticism. For most of 
us, the book has been too heavy a personal investment, and, much more important, 
it is likely to remain the only major modern work in print on Nearctic Rhopalocera 
for amateur and professional alike. Nor, unfortunately, do I believe that there will 
soon be a new edition in which errors could be corrected and omissions rectified. 
Admittedly, this is a mere guess, for the present volume does not carry any indication 
of how large an edition was printed; but in view of its bulk, high cost, and inadequa- 
cies, the present edition seems unlikely to be exhausted soon and replaced by a 
new one. Conditions are undoubtedly very different for the highly portable, in- 
expensive, highly praised, and by now well-established Field Guides on butterflies. 
These, deservedly, sell extremely well—often even getting into a publisher's best- 
seller list—and therefore run fairly rapidly through several editions. 

Thus, Klots’s superb, even uniquely excellent, Field Guide to the Butterflies of 
North America, East of the Great Plains was, and perhaps even still is, a distinct 
publishing success, though probably not so great as that of Roger Tory Peterson's 
Field Guide to the Birds, which initiated the series, and which, as the Chief Editor 
of Houghton Mifflin told me in the late forties, had become the best-selling book in 
American publishing history, after the Bible. Dr. Higgins told me when his very 
good Field Guide to the Butterflies of Britain and Europe first appeared that the 
British publishing house had run off 400,000 (sic!) copies of the color plates to 
provide for English and foreign language editions. The newest English edition, 
just out, is thus the third in six years. It is a sad reflection, incidentally, that the 
color plates in Higgins & Riley, which costs less than one-fifth of the Howe book, 
should be of greatly superior quality, and life-size to scale throughout. 

Therefore, I wish to suggest that the Lepidopterists’ Society assume the burden 
and responsibility of raising the value of Howe’s book. To begin with, and taking 


74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Dr. Ferris’s review as a point of departure, I recommend that a succinct, page by 
page, corrections and addenda to Howe be published in the Journal, quite particu- 
larly bringing the status of subspecific taxa in the various groups up to date. If 
Dr. Ferris cannot be persuaded to undertake this task, it should not be too difficult 
for the Editorial Board of the Journal to appoint a panel of experts for it, perhaps 
including at least some of the specialists who contributed to the Howe volume. 
Thereafter the Journal might publish at regular (annual?) intervals, or as the number 
of discoveries warrants, a brief annotated list of addenda that will keep the book up 
to date. Society members may recall that early in its history the News, and then the 
Journal, published regularly a worldwide section called “Recent Literature on 
Lepidoptera.” This highly ambitious and most valuable service, which provided 
brief abstracts of all new lepidoptera literature, was discontinued with vol. 20, no. 2, 
in 1966, no doubt because it proved too space-consuming for the publication and 
too time-consuming for the contributors. Although the Society is happily and suc- 
cessfully international, still, its area of greatest expertise and prime scientific 
responsibility, as well as the main interest of most of its readers, remains the 
Nearctic. The service of bringing Howe’s volume regularly up to date would 
therefore, on a more manageable and modest scale, resume the abstracting service 
of earlier times. 

Nor is this entirely an innovation for the Journal. Cyril F. dos Passos himself 
published addenda and corrigenda to his Synonymic List of the Nearctic Rhopalocera 
(1964) in the Journal (19: 192; 23: 115-125; 24: 26-38). This uniquely valuable 
and excellent work has thus maintained its great value, and it is to be hoped that 
the Journal will continue to publish addenda for the new taxa described. Conceiv- 
ably, the Journal editors may be able to publish such additions to either of the works 
in such a form that they can be readily inserted into the individual owner’s copy, 
selling such separata for a reasonably low charge. But even failing this, one can 
readily bring one’s own copy up to date by making marginal notes or, even better, 
by interleaving the volume (small and thin interleaf sheets, gummed along one edge, 
are available for just such purpose at many university bookstores ). 


Hans J. EpsTEIN 


Ep. Note: Mr. Epstein’s comments and suggestions regarding The Butterflies of 
North America are most notable and challenging. I too am interested in the accuracy 
of all lepidopterological information. However, the responsibility for rectifying 
mistakes in non-Society publications rests squarely on the shoulders of their pub- 
lishers, editors, and authors. With the exception of the following letter by Dr. 
Ferris, I suggest that any additional comments and corrections be sent directly to 
the author(s) of the book in question. They may then use them more advantageously 
for compiling an Addenda et Corrigenda to be published either in the Journal or 
elsewhere than if the Society was to take the initiative. 


G. L. GopFREY 


Dear Dr. Godfrey: 


Regarding my recent review of Howe’s The Butterflies of North America (J. 
Lepid. Soc. 30(2): 138-143, 1976), Mr. H. A. Freeman has kindly pointed out two 
oversights on my part. The chromosome number of Megathymus coloradensis is 27 
and that of M. yuccae is 26. Thus two distinct species are involved and Killian 
Roever, who prepared this section of the book, was in error in placing coloradensis 
as a subspecies of yuccae. 


VoLUME 31, NuMBER 1 75 


In the Agathymus section, chisosensis was placed as a subspecies of neumoegeni. 
A. chisosensis belongs to a different species group as its chromosome count is 18 
while that of neumoegeni is 10. 

Partial chromosome numbers of the Megathymidae are given in Freeman’s review 
of the family (J. Lepid. Soc. Supp. 1, 23: 1-59, 1969). 


CLIFFORD D. FERRIS 


Dear Dr. Godfrey, 


With reference to the note, “Aberrant Chlosyne lacinia Nymphalidae) from central 
Texas” (Neck 1975, J. Lepid. Soc. 29: 259): if Mr. Neck examines the forelegs of 
his abdomen-less specimen, he should have little difficulty in determining its sex. 
The fore-legs in both sexes of the Nymphalidae are useless for walking, but that of 
the female bears some likeness to a leg, whilst that of the male has degenerated into 
little more than a brush. 

American authors seem strangely reluctant to mention the use of the fore-leg as 
a means of sexing butterflies. British authors, using any form of key for classification, 
invariably mention the condition of the fore-leg as one of the basic couplets... . 
Imms (A General Text Book of Entomology) .. . 


D. G. SEVASTOPULO 


Dear Sir: 


The suggestion by Professor Ehrlich (vol. 30, p. 149) that P. xuthus may have 
reached Hawaii by natural dispersal may well be correct, although I think the 
intervention of some form of human agency is more probable. However, the analogy 
of the Lycaenid Vaga is misleading. “Vaga’ ogasawaraensis of the Bonin Is. is a 
Celastrina species very closely allied to, and clearly derived from, C. swugitanii 
(Matsumura) of Japan, which in turn is closely allied to C. argiolus (L.). None of 
these species are at all closely allied to the Hawaiian Vaga blackburni, which, to 
judge by its male genitalia, has its nearest relatives in a group of Papuan species which 
includes “Candalides’ meeki Bethune-Baker, “Holochila’ owgarra Bethune-Baker, 
“Lycaenopsis’ manokwariensis Joicey & Noakes, “L.” pullus Joicey & Noakes and 
several other unnamed species, for which a new genus is required and will be named 
in the review of the Lycaenopsis section currently being undertaken by Akito Kawazoé 
and myself. 

It is certain that the ancestor of Vaga must have reached Hawaii by transoceanic 
dispersal, but it seems likely that the route followed was a more southerly one than 
that taken by P. xuthus. 


Ti2Ny BLIoeT 


Corrigendum 


The species name myrtale was misspelled “myratle” in the title of Dr. Ichiro 
Nakamura’s recent article (J. Lepid. Soc. 30: 305-309). I thank Dr. Nakamura 
for pointing out my error. 


Editor 


76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


OBITUARY 


EUGENE S. MILJANOWSKI (1908-1976 ) 


Dr. Eugene S. [Yevgeniy Semionovitsh] Miljanowski died of a heart attack on 
29 May 1976 in his home in Sukhumi, where he spent the last 43 years of his life. 
He was one of the most remarkable persons in Soviet lepidopterology and was widely 
known for his important contributions to the faunistic study and zoogeographic in- 
terpretation of the macrolepidoptera of the Caucasus, especially in the Abkhazian 
Autonomous Soviet Socialist Republic. 

Eugene S. Miljanowski was born on 12 January 1908 in Warsaw, into the family 
of a Russian officer. In 1916 his family settled in Poltava (Ukraine), where he 
took his first steps in his profession and life-long passion—entomology. It is here 
he met Alexandr S. Danilevski, and the ensuing friendship, first based on a common 
interest in butterflies and moths, persisted till Danilevski’s death in 1969 (see A. 
Diakonoff 1970, J. Lepid. Soc. 24: 70-72). After finishing high school and Horti- 
cultural Technical School in 1927, he was appointed in the Poltava Agricultural 
Experiment Station as an assistant with the special task to study the macrolepidop- 
teran fauna of the Poltava district. In 1930-1933 he co-operated with the Lubny 
Medical and Etheric-Oil Plant Experiment Station. From 1933 on, his permanent 
residence spot was Sukhumi, where he was appointed as an assistant and then ad- 
vanced to the head position of the Plant Protection Division, Sukhumi Etheric-Oil 
Plants Research Station. He worked there until his retirement in 1976, several months 
before his death. Starting in 1946 he was pluralistically employed at the Natural 
History Division, Abkhazian Lore Museum, where his efforts resulted in a remarkable 
increase in the collections and scientific level of the staff activity. 

Because of strong myopia he was not fit for military service and was unable to 
take a direct part in World War II. At this time he studied the possible use of 
etheric oils and parts of etheric plants for wound cures. This research resulted in 
obtaining some original, therapeutically active preparations for wound microflora 
control and supporting quick healing of wounds. For efforts in this field he was 
awarded the medals “For Defence of Caucasus” (1945) and “For Valiant Labor in 
the Great Patriotic War’ (1947). 

His decision to leave Ukraine for Caucasus he explained by the fact that Western 
Caucasus and its Black Sea Coast in particular were then unstudied by lepidopterists 
so he found this area far more worthy of study and discovery-promising. Alone, 
sometimes in company with visiting entomologists, or as a member of botanical, zoo- 


VoLUME 31, NuMBER 1 fe 


logical and geological expeditions he collected in practically all accessible—and often 
not so accessible—parts of Abkhazia, from the swampy lowlands to the high mon- 
tane areas. He was really tireless in field work. During the weeks, after days of 
hard climbing and walking he was still able to spend nights collecting with a light 
trap, having only an hour or two of drowsiness with head on sleeve. In the field 
he ignored comfort and his luggage was limited to a minimal omnia mea me cum 
porto. The most important results of these year-to-year excursions was the develop- 
ment of the most complete regional collection of butterflies and moths that ever 
existed in Caucasus. This was bequeathed to the Zoological Institute of the U.S.S.R. 
Academy of Sciences, Leningrad. 

Miljanowski published more than 50 scientific articles on the macrolepidoptera 
of Abkhazia and other parts of the Caucasus, and more than one hundred papers 
(including popular articles) on plant protection, nature conservation (with special 
reference to insect protection), insect behavior and ecology, botany, herpetology, 
batrachology, and speleology. For the past 10 years he was President of the Spe- 
leology Section at the Sukhumi Tourist Club. He was a man with a wide range of 
interests in different aspects of human activity. He was an expert in Ancient Greek 
mythology, and often recited his own hexameters on recent affairs where Hellenistic 
persons were employed. His second passion was classic music, especially opera. He 
collected many hundreds of records, and I often heard him singing his favorite parts 
of operas in the field. He was always ready to teach. As a man of generous soul 
he was easy to contact, and his home was always filled with people. 

Superficially Miljanowski represented a naturalist of an older generation profess- 
ing the Staudinger-Seitz taxonomic doctrine. At the same time, under this “Tory” 
mask, a man of unordinary approach and world vision was covered. One example 
will depict his original way of thinking. A puzzling peculiarity of the Black Sea 
coast of the Caucasus and the Crimea as a habitat is their relative faunal wretched- 
ness, though these areas with their ‘subtropical’ climate would seem to be richer in 
Lepidoptera than the adjoining mountains and plains of Ciscaucasia. Contrary to 
recognized theory of geological-geographical isolation of N. J. Kuznetsov (1929), 
Miljanowski proved the ecological background of this phenomenon. The reason 
appeared to be surprisingly simple: these coastal areas cannot be colonized by boreal 
species because they are too warm for their hibernation and winter diapause (the lack 
of freezing ); on the other hand, relatively low winter temperatures prevent coloniza- 
tion of the coast by southern species (Miljanowski 1956, Zool. Journ. (Moscow) 35: 
1170-1176). 

A central position in his scientific work occupies “Macrolepidoptera of Abkhazia: 
Ecology and Economic Importance” which he defended in 1961 as a doctoral thesis 
in Tbilisi Agricultural Institute. Together with more recent publications, this work 
promises to be a concise guide to the butterflies and moths of Abkhazia and adjoining 
areas of Georgia and Russia, and it may be hoped that in the near future it will be 
prepared for publication. It will be the best possible monument to the man whose 
knowledge, advice and suggestions were widely used by his numerous pupils, col- 
leagues, and friends. His name will never be omitted in the Caucasian lepidopterology 
and he will persist in our memory. 


Yurt P. NEKruteNnko, Ukrainian Institute for Plant Protection, 252627 Kiev 127, 
Ukraine, U.S.S.R. 


78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


BOOK REVIEWS 


Motus OF SOUTHERN AFRICA, by E. C. G. Pinhey. 1975. Tafelberg Publishers Ltd., 
28 Wales Street, Cape Town, South Africa, 273 p., 19 figures (line drawings), 63 
color plates. Price $35.95 (U.S.). 


At the risk of offending the ardent rhopalocerists who have grown accustomed in 
recent years to the availability of exquisite butterfly books, complete with color plates, 
from nearly all comers of the world, it is refreshing to see, occasionally, a regionalized 
treatment of moths. E. C. G. Pinhey’s Moths of Southern Africa is designed to 
stimulate interest in the moth fanua of southem Africa and is a broad introduction 
to the subject, not a definitive treatment of this entire fauna. Pinhey includes 1181 
of the more common, colorful, or economically important moth species, mainly from 
the defined region, in the color plates (plus a line drawing of an additional species ) 
and provides brief descriptions of them in the text. A hostplant index, glossary, 
substantial number of cited references, and line drawings supplement the color 
plates and textual information. 

Using a balance of technical and nontechnical terminology, Pinhey introduces the 
moths by reviewing their basic morphological, ecological, and behavioral character- 
istics. The written portion of the introduction certainly should stir the interests of 
budding lepidopterists. However, I was surprised to find virtually no discussion on 
the genitalia that are so very useful in indentification and classification, yet two 
pages of space are devoted to line drawings of these important structures. 

Taken in their entirety, the line drawings, which make up a substantial proportion 
of the introductory material, are satisfactory. The beginning student might occasion- 
ally be confused by tke structural terminology in the text and symbols used in some 
figures because a few problems are caused by combinations of printing errors and 
inconsistencies in symbol application. For example, Figure 5 is referred to as “fig. 
3” on page 7, and “ar,” the symbol for areole in the figure legend, is “Ar” in Figures 
5 c, e, and f. In other figures “RS” denotes both the reniform stigma and the 
radial sector. 

The author has chapters on “Collecting and preparing insects” and “Identification 
and classification.” The former is rather brief, though references are provided that 
may be consulted for information on specific procedures. The latter also is brief, 
but interestingly written for its intended audience. However, the identification key 
to suborders and superfamilies ideally belongs here, not in the subsequent chapter 
entitled “Swifts and Longhorns”! Pinhey should have given figure references in the 
couplets of his keys, especially since his book is meant to interest and aid persons 
who are not professional and/or experienced lepidopterists, not discourage them. 
The remaining keys generally go to the family level (a few extend to the subfamily 
level)—beyond that the color figures and brief species accounts serve as the 
identification aids. 

The printing quality of the color plates is excellent with the exception of a few 
cases where the colors of the plate backgrounds and moths are so similar that the 
specimens are scarcely visible. There is a problem with the organization of the plates 
that lessens their effectiveness and utility for identifying specimens. Namely, fami- 
lies, congeners, and, in some cases, the same species frequently are depicted on 
different plates separated by several pages of text and other plates. For example, the 
Limacodidae occur on plates 1, 2, 5, and 7 and adult Lasiocampidae on plates 
1, 22, 24-26, and 29-30. The author should have spent less time coining almost 
useless, common names and more time on the overall organization of his book and 
embellishment of the introductory chapters. 

Considering the large number of species and color plates in Moths of Southern 
Africa, the price is not unreasonable. The book should prove rather useful to those 


VoLuME 31, NuMBER | 79 


for whom it was designed. The book’s true value will be realized if a host of minds is 
inspired to continue the study of heterocerous species in southern Africa or elsewhere. 


Grorce L. Goprrey, Section of Faunistic Surveys and Insect Identification, IIli- 
nois Natural History Survey, Natural Resources Building, Urbana, Illinois 61801. 


LEcIon oF NiGHT—THE UNDERWING Morus, by Theodore D. Sargent. 1976. Univer- 
sity of Massachusetts Press, Amherst, Massachusetts, 222 + xiii p., 8 color plates. 
Price $15.00 (U.S.). 


My introduction to the underwing moths was dramatic. As a high school student 
I had just taken up collecting Lepidoptera and had read with great interest the 
article on “sugaring” in W. J. Holland’s The Moth Book. I had just sugared a path 
along a road in the northern Wisconsin forest. In the fading light of the evening 
I had discerned the outline of a beautifully banded underwing moth. I was forever 
hooked on the Catocala. 

So it was with unexpected pleasure that I read Theodore D. Sargent’s Legion of 
Night—The Underwing Moths. He has captured the romance of these unusual and 
beautiful insects. 

Statistically the book contains two hundred twenty-two pages, and has eight 
colored plates depicting one hundred twenty-six specimens in color consisting of 
seventy-one species of underwing moths found in the eastern United States. In 
addition there are numerous black and white colored plates and many fine drawings. 

Sargent’s book gives a complete survey of the Catocala of the eastern United States, 
a summary of the current biological information on these species, and introduction 
to the scientific investigations which are being conducted on these moths. 

The species accounts give the range, within the scope of the geographical area 
covered by this book, the seasonal occurrence, larval food plants, and the interesting 
behavioral aspects of the adults of each species. In this respect it is quite unique. 
None of the work done on the Catocala in the past has gone into these aspects. In 
this respect it is most helpful to the field entomologist. So few works on Lepidoptera 
seem to be concerned with this aspect of collecting. Every species of Catocala one 
might encounter in the eastern United States is covered completely. 

Dr. Sargent gives an interesting history of the entomologists who described the 
species found in the area or wrote about these moths in his chapter “Of Men and 
Names.” Thus, we get an insight into the entomological lives of such men as 
Coleman T. Robinson, Augustus Radcliffe Grote, Achille Guenée, Ferdinand Hein- 
rich, Herman Strecker, William Henry Edwards, the Reverend George D. Hulst, and 
Francis Walker. The book contains excerpts from the published letters of several 
of these entomologists, particularly Strecker, Grote, and Hulst. None of these men 
held the others in any great esteem, and their sniping at each other makes for 
interesting reading. Apparently they were unaware of the laws on libel and slander. 
In any event there is no record of any civil actions arising out of these feuds. 

Each species is accurately described, and similar species distinguished. The 
relative abundance or scarcity of the particular species is fully discussed. Of great 
importance to the collector is something about the larval habits and the habits of 
the adults in coming to light or to bait, or to various types of traps. The book 
contains much statistical material on abundance of the various species and something 
about the predators of the larva and adults, particularly bird and bat damage to the 
adults. 

I would make only one criticism of the book. On the theory nobody is perfect, 
I would liked to have seen the plates with a lighter background. Since so many of 


80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the Catocala have at least dark forewings, the specimens do not stand out against 
the black background, and in this respect I think a paler background would have 
produced much better colored plates. 

In any event this is a most worthwhile contribution to the science of Lepidoptera. 
This volume should be in the library of every moth collector. 


WiiuiAM E. Srexer, 119 Monona Avenue, Madison, Wisconsin 53703. 


eae a 


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Via a 


EDITORIAL STAFF OF THE JOURNAL 
GrorcE L. Goprrey, Editor 


Illinois Natural History Survey, Natural Resources Building 
Urbana, Illinois 61801 U.S.A. 


WitiiaM H. ALLEN, Associate Editor JAMeEs G. STERNBURG, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study 
of Lepidoptera. Contributors should prepare manuscripts according to the following 
instructions. 

Text: Manuscripts should be submitted in duplicate, and must be typewritten, 
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11 
inch paper. Titles should be explicit and descriptive of the article’s content, including 
the family name of the subject, but must be kept as short as possible. The first men- 
tion of a plant or animal in the text should include the full scientific name, with 
authors of zoological names. Insect measurements should be given in metric units; 
times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). 
Underline only where italics are intended. References to footnotes should be num- 
bered consecutively, and the footnotes typed on a separate sheet. 

Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically 
under the heading LireraTure Cirep, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, 
London. 209 p. 

196la. Some contributions to population genetics resulting from the 
study of the Lepidoptera. Adv. Genet. 10: 165-216. 


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

Illustrations: All photographs and drawings should be mounted on stiff, white 
backing, arranged in the desired format, allowing (with particular regard to lettering ) 
for reduction to their final width (usually 4% inches). Illustrations larger than 8% 
x 11 inches are not acceptable and should be reduced photographically to that size 
or smaller. The author’s name, figure numbers as cited in the text, and an indication 
of the aricle’s title should be printed on the back of each mounted plate. Figures, 
both line drawings and halftones (photographs), should be numbered consecutively 
in Arabic numerals. The term “plate” should not be employed. Figure legends must 
be typewritten, double-spaced, on a separate sheet (not attached to the illustrations), 
headed ExpPLANATION OF FicuRES, with a separate paragraph devoted to each page of 
illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings 
for tables should not be capitalized. Tabular material should be kept to a minimum 
and must be typed on separate sheets, and placed following the main text, with the 
approximate desired position indicated in the text. Vertical rules should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer's errors. Excessive author’s changes at this time will be charged 
to authors at the rate of 75¢ per line. A purchase order for reprints will accompany 
the proofs. 

Correspondence: Address all matters relating to the Journal to the editor. Short 
manuscripts such as new state records, current events, and notices should be sent to 
the editor of the News: Ron Leuschner, 1900 John Street, Manhattan Beach, Cali- 
fornia 90266. 


ALLEN PRESS, INC. NE) LAWRENGE, KANSAS 
uUuS.x 


CONTENTS 


STUDIES ON THE CATOCALA (NocTUmDAE) OF SOUTHERN NEw ENGLAND. 
V. Tse Recorps oF Siwney A. HessEL FROM WASHINGTON, CON- 
NECTICUT, 1961-1973. Theodore D. Sargent _____._ > ae 


DISTRIBUTION AND BIOLOGY OF A PLEISTOCENE RELICT: OCHLODES 
yumM4 (HesperupaE). James A. Scott, Oakley Shields, and Scott 
LL.  Bbig es Se ie 


HYBRIDIZATION OF CALLOSAMIA (SATURNIDAE). Richard S. Peigler 


AGGREGATION BEHAVIOR OF CHLOSYNE LACINIA LARVAE (NYMPHALI- 
DAE). Nancy Stamp 0 00 ee 


A Review or NortH AMERICAN RHODOPHAEA (PHYCITINAE: PYRALI-— 


DAE), WITH DescripTION OF Six New Species. Paul A. Opler 


A New NortrH AMERICAN SPECIES OF APAMEA FORMERLY CONFUSED 
witH A. VERBASCOIDES (GUENEE) (NocrumwaE). Douglas C. 
Persusorn 200000000 sa 


ADDITIONAL MATERIAL OF SCOPARIA HUACHUCALIS MUNROE, WITH DE- 
SCRIPTION OF MALE GENITALIA (PYRALIDAE, SCOPARIINAE). Eu- 
gene Mainroe)? 0 Ns 


GENERAL NOTES 


Melitaea saxatilis mod. “sassanides” (Nymphalidae) in Iran: confirmation 
of an old record. Javad Hashemi Tafreshi 
Bizarre capture of a butterfly by an ambush bug. Raymond W. Neck ___ 
Cocytius duponchel (Sphingidae): second United States capture. James 
Bi Death 20 0 NSN ee YM Pas Ne rr 
A melanic form of Phigalia strigataria (Geometridae). Joseph Muller __— 
Staphylus azteca, new record for the United States (Hesperiidae). Hugh 
Avery Freeman 2 ONG gs Ny Sr 
Effects of 1933 hurricanes on butterflies of central and southern Texas. Ray- 
mond W Neck 220 ii No) A 
Colony of Pieris napi oleracea (Pieridae) in Indiana. Ernest M. Shull Ke 
An ecological note on Polites sabuleti sabuleti at the northern limit of its 
range (Hesperiidae). J. Allan Garland 2 so eee 
Larval hibemation of Geometridae in eastern United States. Dale F 
Schuetiser: 20080 en 2a 


63 


Volume 31 1977 Number 2 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
Publié par LA SOCIETE DES LEPIDOPTERISTES 

3 Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 

4 Publicade por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


30 June 1977 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


S. S. Nico.ay, President CarLos R. BEUTELSPACHER, 

MreiAmM Roruscuiip, Ist Vice President Vice President 

THEODORE D. SArcENT, Vice President Joun M. Sniper, Treasurer 
JuLtian P. DonauHueE, Secretary 


Members at large: 


J. T. BREWER D. F. Harpwick R. A. ARNOLD 
K. S. BROWN J. B. ZrecLER E. D. CasHatr , 
K. W. Puivip F. S. CHEw R. E. STANFORD 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and 
formally constituted in December, 1950, is “to promote the science of lepidopterology 
in all its branches, .... to issue a periodical and other publications on Lepidoptera, 
to facilitate the exchange of specimens and ideas by both the professional worker and 
the amateur in the field; to secure cooperation in all measures’ directed towards 
these aims. 

Membership in the Society is open to all persons interested in the study of 
Lepidoptera. All members receive the Journal and the News of the Lepidopterists’ 
Society. Institutions may subscribe to the Journal but may not become members. 
Prospective members should send to the Treasurer full dues for the current year, 
together with their full name, address, and special lepidopterological interests. In 
alternate years a list of members of the Society is issued, with addresses and special 
interests. There are four numbers in each volume of the Journal, scheduled for 
February, May, August and November, and six numbers of the News each year. 


Active members—annual dues $13.00 
Student members—annual dues $10.00 
Sustaining members—annual dues $20.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $18.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
John M. Snider, 3520 Mulldae Ave., San Pedro, Calif. 90732 U.S.A. 


Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA 


by Cyrit F. pos Passos 


Price: Society members, $5.00 U.S.; non-members, $7.50 U.S. Paper covers, revisions 
of the Melitaeinae and Lycaenidae supplied separately. 


Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni- 
versity of Louisville, Louisville, KY 40208, U.S.A. 


The Lepidopterists’ Society is a non-profit, scientific organization. The known 
office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second 
class postage paid at Lawrence, Kansas, U.S.A. 66044. 


—S— ee 


JOURNAL OF 


Tue Leripoprerists’ Society 


Volume 31 1977 Number 2 


IMMATURE STAGES AND ECOLOGICAL OBSERVATIONS OF 
EOPARARGYRACTIS PLEVIE (PYRALIDAE: NYMPHULINAE) 


SANDY B. FIANCE! AND ROBERT E. MOELLER? 


Eoparargyractis plevie was described by Dyar (1917) from an adult 
female. Lange (1956) hypothesized that the unknown larvae in this 
genus feed on the rock-encrusting periphyton of small lakes. Our ob- 
servations from Mirror Lake, New Hampshire, indicate that, in fact, the 
larvae of E. plevie feed on several species of aquatic macrophytes. 

Collections of the plant species Lobelia dortmanna L., Isoetes tucker- 
mani A. Br., and I. muricata Dur. in 1974 and 1975 frequently included 
larvae of this species. Leaves of infested macrophytes were usually 
damaged, suggesting that the larvae were feeding on the macrophytes 
themselves, rather than on any sparse periphyton they might have sup- 
ported. Behavioral observations in the laboratory confirmed this in- 
terpretation. Apparently there are no previous reports of insects feeding 
on these species. 

The following report (1) describes the previously unknown larva and 
pupa of Eoparargyractis plevie; (2) establishes that the pupal stage is 
passed underwater and possesses what may be a stridulatory apparatus; 
(3) provides the first information on larval feeding habits in the genus; 
and (4) is the first discussion of insect feeding on aquatic rosette plants. 


Description of Immature Stages 


Larva: Head (Fig. 4C) 0.96 mm wide. Right mandible (Fig. 3C) with 5 
cusps, slightly curved along outer margin and with a simple peg-like tooth basal 
to the first cusp. Ventral margin of mandible with a broadly produced flange having 
ing two setae inserted at its base, the proximal one less than 1% length of the distal. 
Labrum (Fig. 2) with three highly modified setae on each side of midline (Fig. 
3A, 3B). Head capsule translucent yellow, pigmented black only along adfrontal 
region and around ocelli. Total body length 11 mm. Prothoracic coxae nearly 


1 Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853. 
Tie of Ecology and Systematics, Langmuir Lab, Comell University, Ithaca, New York 


JOURNAL OF THE LEPIDOPTERISTS’ SocIE'Ty 


A, habitus, lateral view; B, dorsal view of 


plevie: 


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1 


larva of E., 


terminal abdominal segments. 


Mature 


Fig, 


bo 


VoLUME 31, NUMBER §3 


Fig. 2. Labrum of larva of E. plevie (outer surface to the left and inner surface 
to the right of diagonal line). 


contiguous; metathoracic coxae widely separated. Dorsal abdominal setae *4 to 
equal length of antero-dorsal abdominal gills. Integument spiculate, with coarse, 
very short setae arising from knob-like projections, setae becoming longer toward 
caudal end. Setation of head as in Fig. 4C. Integument in living condition un- 
pigmented, light green interior showing through. Gills as in Fig. 1, unbranched 
with one exception, or arising from a common base, present on last two thoracic 
and all abdominal segments. Last gill cluster on margin of tenth abdominal segment 
having 5 branches. Gills variable in number and exact position, those of segments 
8-10 beset with fine hairs around their entire periphery, and forming a flap-like 
extension of the last abdominal segment (Fig. 1B). Crochets a uniserial biordinal 
circle on prolegs 1-4 and a uniserial biordinal mesoseries on anal proleg. Setation 
of first thoracic and third abdominal segments in Figs. 4A & B respectively. 

Material examined: New Hamepsuire, Grafton Co.: W. Thornton, Mirror Lake; 
8 September 1975, Robert E. Moeller; 10 larvae killed immediately; same locality: 
22 October, 1 larva reared, killed 11 December 1975. New Yorx, Herkimer Co.: 
Webb Township, Upper Sylvan Pond; 10 June 1975, Leo M. Demong; 3 larvae. 
New York, Herkimer Co.: Webb Township, Panther Lake; 8 October 1975, Leo 
M. Demong; 2 larvae. 

Pupa: Cocoon (Fig. 7) spun on leaf at rosette base (Lobelia dortmanna), 
often omamented with bits of vegetation. Inner silken sheath complete on all sides, 
no gap or holes along leaf. Total length of pupa 6-7 mm. Unpigmented. Four 
respiratory tubercles on abdominal segments 2 & 3 (Fig. 6A). Cremaster as in Figs. 
5B, 6B. Metathoracic tarsi heavily sclerotized, each bearing a long tooth at tip 
(Fig. 54). Two series of raised ridges present on abdominal sternites closest to 
tips of metathoracic legs (Fig. 5B). 

Hinton (1948) reviewed a number of mechanisms of sound production in 
lepidopterous pupae. He concluded that stridulation probably frightens predators 
and parasites, thus serving as a defense mechanism. According to Hinton, an 
elongated proboscis is rubbed against raised ridges of the fifth abdominal sternite 
producing a hissing sound in Gangara thyrsis F. E. plevie has a similar morpholog- 
ical arrangement (Fig. 5). Although sound production by the pupae of EF. plevic 
has not been demonstrated, stridulation by this species is suggested by the two 
series of raised ridges on the abdominal sternites near the heavily sclerotized teeth 
at the tip of the metathoracic tarsi. In view of the aquatic habit of the pupae of 


84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ee 


Figs. 3, 4. Larva of E. plevie: 3A & B, enlarged views of modified setae borne 
on labrum; C, view of inner surface of right mandible. 4, setation diagrams of 
first thoracic segment (A) and third abdominal segment with small circles indicat- 
ing positions of gills (B); frontal view of right half of head, excluding modified 
setae of labrum (C); setation of left prothoracic leg (D). 


E. plevie, an alternative hypothesis involving the defense of localized resources 
(a rosette ) seems in order. 

Material examined: New Hampsuree, Grafton Co.: W. Thornton, Mirror Lake; 
larvae collected October and November 1975, Robert E. Moeller. Reared on Lobelia 
dortmanna, 5 cocoons recovered January 1976 containing 4 pupal exuvia, | pupa. 


VoLUME 31, NUMBER 2 


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Figs. 5-7. Pupal stage of E. plevie: 5, ventral view of pupa, habitus (A) and 
cremaster (B); 6, dorsal view of pupa, habitus (A) and cremaster (B); 7, cocoon 
attached to leaf of Lobelia dortmanna, containing pupa and exuvia of last larval 
instar. 


86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


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Fig. 8. Temperature during 1975 at a depth of 2 m in Mirror Lake, New Hamp- 
shire (curve) and temporal incidence of severely damaged, dead, or vanished 
Lobelia dortmanna as a proportion of the experimental population (histogram ), 


Distribution: Maine, Massachusetts, Nova Scotia, Quebec; NEw RECORDS from 
New Hampshire and New York. Larval and pupal voucher specimens are deposited 
at the Canadian National Collection and the Cornell University Collection. 


Ecological Observations 


In Mirror Lake, New Hampshire, larvae are frequently encountered 
during late summer on Lobelia dortmanna, Isoetes tuckermani, and 
I. muricata. These small (1-5 cm high) plants consist of a rosette of 
stiff, narrow leaves at the base of which the larvae construct a protective 
purse of silk and detritus. Larvae cut through and ingest the thin 
epidermis that surrounds the intercellular air spaces of the largely hollow 
leaves. Damage to the plant can be severe; badly injured leaves die and 
separate from the plant, and growing leaves may be deformed by the 
silken tube attachments. Defoliated plants have been encountered, as 
well as decayed plants that had died as a result of the defoliation. Al- 
though no more than a single larva has been found on any Lobelia 
dortmamna plant, several may occur on a single rosette of Isoetes tucker- 
mani. Muskrats (Gaevskaya, 1966) and ducks (Fassett, 1969) have 
been reported to feed on rosette species, and muskrats occasionally feed 
on L. dortmanna and other plant species in Mirror Lake. E. plevie seems 
to be the first reported insect herbivore on these plants. 

The period of most intensive feeding in Mirror Lake was estimated 
from a plant growth experiment carried out on Lobelia dortmanna during 
the summer of 1975. Seventy-five individual rosettes over a depth range 
of 0.5-1.5 m were marked, and the appearance of new leaves was followed 


VOLUME 31, NUMBER 2 87 


at 4-week intervals over the growing season (May-September). Fig. 8 
(histogram ) indicates the temporal incidence of severely damaged, dead, 
or vanished plants. Much of this mortality was due to E. plevie, but ad- 
ditional factors cannot be discounted. The early summer peak probably 
corresponds to feeding of late-instar larvae before pupation, and the late 
summer peak with feeding of the newly hatched larvae of the next gen- 
eration. Although these results suggest a very great impact of the insect 
on the host plant population, it appears that the experimental plants 
suffered heavier infestation than the population as whole, perhaps as the 
result of greater spacing between experimental plants. 

Mirror Lake contains approximately 20 species of aquatic macrophytes 
(Moeller, 1975). Larvae of E. plevie have been found on the three species 
already listed, and could occur on others that have been less intensively 
examined. They do not occur on Utricularia purpurea Walt., Nuphar 
variegatum Engelm., or Nitella flexilis (L.) Ag. The known distribu- 
tion of E. plevie (Munroe, 1972) coincides with regions of soft-water, 
oligotrophic or dystrophic lakes in which Lobelia dortmanna and Isoetes 
species are common, and to which many of them, including L. dortmanna, 
are restricted (Fassett, 1930; Swindale & Curtis, 1957). Larvae of E. 
plevie from New York came from an oligotrophic Adirondack lake where 
Isoetes sp. is abundant. This insect’s distribution may be determined 
by the restriction of its host plants, Lobelia and Isoetes, and possibly 
other rosette plants, to unproductive lakes in regions of granitic bedrock 
(northern New England and eastern Canada) or sandy outwash (Cape 
Cod and Martha’s Vineyard). 


Kite’ Cycle 


Larvae have been collected in Mirror Lake from August through mid- 
November. The absence of plant damage after the middle of September 
indicates that the larvae have become quiescent, but if disturbed, they 
will move about. Forbes (1923) described overwintering larvae in the 
closely related genera Nymphula and Parargyractis. E. plevie also over- 
winters as a larva. The pupal stage is passed underwater, cocoons being 
constructed at the bases of Lobelia dortmanna rosettes in laboratory- 
reared specimens. Pupation and emergence have not yet been observed 
on Mirror Lake, but the observed pattern of damage to L. dortmanna is 
consistent with the July-August flight period known elsewhere (Munroe, 
1972). 

Larvae collected 22 October and reared at 20-25°C on Lobelia dort- 
manna emerged 8 weeks later on about 20 December. Larvae collected 
12 November emerged about 10 January, again after about 8 weeks. 


88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


These development times are consistent with a flight period of late July 
if the overwintering larvae resume development above a 16—18°C thresh- 
old at the end of May (Fig. 8, showing temperature at a depth of 2 m, 
representative of the 0.5-3 m recorded range of the insect). The absence 
of significant new damage to Mirror Lake L. dortmanna after mid- 
September indicates a slowdown in feeding and development below 
18-20°C. 


ACKNOWLEDGMENTS 


A number of our associates at Cornell have provided substantial aid 
in this publication. Dr. John G. Franclemont generously provided the 
initial adult determination, constant encouragement and advice and 
critically read the manuscript. Tim McCabe and Richard Brown con- 
tributed many ideas and technical expertise. We are also indebted to 
Dr. C. O. Berg and Dr. George Eickwort for considerably improving 
the manuscript and figures. Leo Demong kindly provided larval speci- 
mens from upstate New York. We gratefully acknowledge the partial 
financial assistance of the Department of Entomology, Cornell University 
and the National Science Foundation through Dr. Gene E. Likens and 
the Hubbard Brook Ecosystem Project. 


LITERATURE CITED 


Dyar, H. G. 1917. Notes on North American Nymphulinae. Insecutor Inscitiae 
Menstruus 5(4—6): 75-79. 

Fassett, N. C. 1930. The plants of some northeastern Wisconsin lakes. Trans. 
Wisconsin Acad. Sci. 25: 157-168. 

1969. A manual of aquatic plants. Revision appendix by E. C. Ogden. 
Univ. Wisconsin Press, Madison, 405 p. 

Forses, W. T. M. 1923. The Lepidoptera of New York and neighboring states. 
Cornell University Agric. Expt. Sta., Mem. 68: 1-729 p. (Nymphulinae p. 
574-581 ). 

GarvskayA, N. S. 1966. Rol’ vysshikh vodnykh rastenii v pitanii zhivotnykh 
presnykh vodoemov. Nauka, Moscow. Translated as “The role of higher aquatic 
plants in the nutrition of the animals of fresh-water basins,” D. G. Maitland 
Muller (trans.), National Lending Library for Science and Technology, 
Boston Spa, Yorkshire, England, 1969, 629 p. 

Hinron, H. E. 1948. Sound production in lepidopterous pupae. Entomologist 
81: 254-269. 

Lance, W. H., Jr. 1956. A generic revision of the aquatic moths of North 
America: (Lepidoptera: Pyralidae, Nymphulinae). Wasmann J. Biol. 14: 59- 
144. 

Moe.tuier, R. E. 1975. Hydrophyte biomass and community structure in a small, 
oligotrophic New Hampshire lake. Verh. Internat. Verein. Limnol. 19: 1004— 
1012. 

Munrog, FE. 1972. The moths of America North of Mexico. Classey Ltd & R. B. 
Publications Inc. London. Fascicle 131A. 134 p. 

SwInDALE, D. N. & J. T. Curtis. 1957. Phytosociology of the large submerged 
plants in Wisconsin lakes. Ecology 38: 397-407. 


VoLUME 31, NUMBER 2 89 


SIX NEW SPECIES OF HESPERIIDAE FROM MEXICO 


Hucu Avery FREEMAN 
1605 Lewis Drive, Garland, Texas 75041 


During the process of conducting research on the Hesperiidae of 
Mexico, several undescribed species recently have been found, six of 
which are described in this article. 


Pyrrhopyge hoffmanni Freeman, new species 
Bigs) O54. li? 


Male (Upper side). Primaries black with an orange-yellow spot near base in 
space lb, the upper half of the spot is orange and the lower half yellow. There is a 
circular spot in space lb situated directly below the spot in space 2. The cell is 
linear and situated inward from the oval shaped spot in space 2. There is a linear 
spot in space 3 and another in space 4. The four apical spots form an even curve 
starting with one in space 6 and extending to space 9. All spots are sordid white. 
Fringes white, black at end of veins. . 

Secondaries black, with the outer margin somewhat crenulate. Fringes white, 
black at ends of veins. 

Male (Under side). Primaries black. There is a narrow orange streak in space 
la. In space 1b and the cell near the base there is an orange area. The costa from 
the base to near mid wing is bright orange. The discal and apical hyaline spots are 
well developed and are clear white. The veins are somewhat lighter than the ground 
color. 

Secondaries black with a bright orange basal area covering somewhat less than 
one half of the wing. There is a distinct black cell spot and one specimen has another 
black spot below the cell spot. There is a narrow black line at the base of the wing. 
The veins are somewhat lighter than the ground color. 

Thorax above black with an orange spot on each side below the head, beneath 
black and orange striped. Abdomen black above, orange and black striped be- 
neath. Head black, white spotted. Palpi black, with distinct white outer areas and 
white spotted cheeks. Legs black and orange. Antennae, both shaft and club, 
solid black. 

Wing measurements. Primaries: base to apex, 25 mm; apex to outer angle, 
17 mm; outer angle to base, 19 mm. Secondaries: base to end of vein 3, 15 mm; 
center of costa to anal angle, 18 mm. Total expanse, 48 mm. 

Female. Very similar to the male. The only difference is the less crenulate 
secondaries and the larger size. 

Wing measurements. Primaries: base to apex, 32 mm; apex to outer angle, 
22 mm; outer angle to base, 22 mm. Secondaries: base to end of vein 3, 22 mm; 
center of costa to anal angle, 22 mm. Total expanse, 55 mm. 

Type material. Holotype, male, Tenosique, Tabasco, Mexico, 3 September 1962 
(E. C. Welling collector), will be placed in the American Museum of Natural 
History, New York. Allotype, female, Chimalapa, Oaxaca, Mexico, September 1965 
(T. Escalante collector), is in my collection. There are three male paratypes from 
Tenosique, Tabasco, one collected 18 August 1962, one 26 August 1962, and the 
other 7 September 1962, and one male paratype from Middlesex, Stann Ck. District, 
British Honduras, 24 March 1965. All paratypes were collected by E. C. Welling. 


This new species is named for the late C. C. Hoffmann, who did so 
much to increase our knowledge of the Mexican Rhopalocera. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


90 


VoLUME 31, NUMBER 2 9] 


Pyrrhopyge hoffmanni belongs in the maculosa group of Evans (1951). 
His concept of there being but two species in this group is completely in 
error. Actually with the discovery of hoffmanni there are now six species 
present, four occur in Mexico, mulleri Bell, erythrosticta Godman & 
Salvin (Figs. 1 & 2), hoffmanni and araxes Hewitson and its subspecies 
arizonae Godman & Salvin. Maculosa Hewitson and cossaea Druce are 
found in Colombia. Mulleri, erythrosticta, and hoffmanni fly in the same 
general area in Tabasco, Veracruz and Oaxaca. Besides differences in 
the genitalia hoffmanni can be separated easily from mulleri by the fol- 
lowing differences: (1) mulleri lacks hyaline spots on the primaries; 
(2) mulleri lacks the cell spot on the lower surface of the secondaries 
which is present in the orange basal area of hoffmanni; and (3) hoff- 
manni is slightly smaller than mulleri. Hoffmanni differs from eryth- 
rosticta in the following ways: (1) the discal spots on the primaries of 
erythrosticta in spaces 1b, 2, and the cell form a straight line, whereas in 
hoffmanni the spot in space 2 is displaced outward from the other spots, 
not forming a straight line; (2) the basal orange area on the lower 
surface of the secondaries is much more extensive in erythrosticta cover- 
ing approximately two thirds of the wing, while in hoffmanni it covers 
less than one half of the wing; (3) there is no cell spot in the orange area 
of erythrosticta which is present in hoffmanni; (4) the orange-red spot 
near the base of the primaries on the upper side in space lb is solid 
deep orange-red in erythrosticta, while in hoffmanni the upper half is 
orange and the lower half is yellow; and (5) erythrosticta lacks the orange 
spots on the thorax just below the head which are present in hoffmanni. 


Epargyreus deleoni Freeman, new species 
Riess 5. 6. 1S 


Male (Upper side). Primaries light brown, with the discal spots yellowish- 
orange; spot in lb linear, midway between the outer margin spot in space 2; spot in 
space 2 broader at the top than bottom, overlapping midway the cell spot; cell 
spot broader at the top than bottom; spot in space 3 small and linear. There are 
two minute linear spots over the outer edge of the cell spot. There is one minute 
apical spot. Basal and discal areas including space la heavily overscaled with 
golden-yellow scales. Costal fold well developed. Fringes yellowish-brown. 

Secondaries yellowish-brown, heavily overscaled with golden-yellow scales ovet 


< 


Figs. 1, 2. Pyrrhopyge erythrosticta Godman & Salvin. Male. Middlesex, Stann 
Ck., Dist., British Honduras, 24 August 1965 (E. C. Welling; H. A. F.). 

Figs. 3, 4. Pyrrhopyge hoffmanni, n. sp. Holotype, male, Tenosique, Tabasco, 
Mexico, 3 September (E. C. Welling; A. M. N. H.). 

Figs. 5, 6. Epargyreus deleoni, n. sp. Holotype, male, X-Can, Quintana Roo, 
Mexico, 13 June 1969 (E. C. Welling; A. M. N. H.). 

Figs. 7, 8. Typhedanus salas, n. sp. Holotype, male, Piste, Yucatan, Mexico, 26 
August 1968 (E. C. Welling; A. M. N. H.). 


JOURNAL OF THE LEPIDOPTERISTS SocIETY 


ae, LP ry 


VoLuME 31, NUMBER 2 93 


the basal and discal areas to near the outer margin. Fringes yellowish-brown, brown 
at ends of the veins. 

Male (Under side). Primaries, light brown, with a purplish wash over the 
apical and outer cellular areas. Space la light tan. The hyaline spots are yellowish- 
brown. 

Secondaries dark chocolate brown over basal and discal areas. Outer margin gray- 
ish with a purplish sheen. Discal spots narrow, beginning at space 1b and extending 
in a straight line to space 7. The spot in space 3 extends slightly outward. The 
spots extending through cell and above, narrow and linear. All spots silvery-white. 
There is an indistinct band beginning at space 1b and running in an irregular 
ene just outside the silvery, discal band, terminating at the upper edge of the 
cell. 

Thorax golden-brown above, dark brown beneath. Abdomen golden-brown above, 
lighter brown beneath, with indistinct segmental striping. Head golden-brown. Palpi 
chestnut brown. Legs tan. Antennae, shaft light brown above and below, club 
slightly darker. 

Wing measurements. Primaries: base to apex, 27 mm; apex to outer angle, 
18 mm; outer angle to base, 17 mm. Secondaries: base to end of vein 3, 15 mm; 
center of costa to anal angle, 21 mm. Total expanse, 51 mm. 

Female (Upper side). Primaries brown, with some golden-yellow overscaling 
near base and midway to discal band. Discal spots yellowish-brown, similar to 
those in the male except larger and there is a distinct costal spot above the cell spot 
and there are three apical spots, lower one is minute and displaced outward from the 
other two. 

Secondaries brown, heavily overscaled with golden-yellow scales over the basal 
and to near the discal areas. Fringes sordid white and brown at ends of veins. 

Female (Under side). Very similar to the male except there is a broad grayish 
marginal area on both the primaries and secondaries. 

Thorax golden-brown above, dark brown beneath. Abdomen brown above, 
grayish-white striped beneath. Head golden brown. Palpi sordid yellowish-white. 
Legs and antennae same as in the male. 

Wing measurements. Primaries: base to apex, 31.5 mm; apex to outer angle, 
20 mm; outer angle to base, 20 mm. Secondaries: base to end of vein 3, 19 mm; 
center of costa to anal angle, 24 mm. Total expanse, 61 mm. 

Type material. Holotype, male, X-Can, Quintana Roo, Mexico, 13 June 1969 
(E. C. Welling collector), will be placed in the American Museum of Natural 
History. Allotype, female, X-Can, Quintana Roo, Mexico, 7 June 1967 (E. C. Welling 
collector), is in my collection. There is one female paratype, Chichen Itza, Yucatan, 
Mexico, 1 November 1930 (F. M. Gaige collector), will remain for the present in my 
collection. 


This new species is named for my good friend Lorenzo DeLeon of 
Cuidad Valles, S. L. P., Mexico, the golf professional at Hotel Covadonga. 
Superticially above this new species slightly resembles E. windi Free- 


— 


Figs. 9, 10. Polythrix guatemalaensis, n. sp. Allotype, female, X-Can, Quintana 
Roo, Mexico, 26 July 1962 (E. C. Welling; H. A. F.). 

Figs. 11, 12. Codatractus yucatanus, n. sp. Holotype, female, Piste, Yucatan, 
Mexico, 1 September 1967 (E. C. Welling; A. M. N. H.). 

Figs. 13, 14. Bungalotis milleri, n. sp. Holotype, male, Candelaria Loxicha, 
Oaxaca, Mexico, 22 September 1968 (E. C. Welling; A. M. N. H.). 

Figs. 15, 16. Bungalotis milleri, n. sp. Allotype, female, San Quintin, Chiapas, 
Mexico, 16 September 1970 (Robert Wind; Allyn Museum). 


94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 17. Male genitalia. Pyrrhopyge hoffmanni, n. sp. Paratype, Tenosique, 
Tabasco, Mexico, 18 August 1962 (E. C. Welling; H. A. F.). 

Fig. 18. Male genitalia. Epargyreus deleoni, n. sp. Holotype, X-Can, Quintana 
Roo, Mexico, 13 June 1969 (E. C. Welling; A. M. N. H.). 

Fig. 19. Male genitalia. Typhedanus salas, n. sp. Paratype, Piste, Yucatan, 
Mexico, 29 August 1968 (E. C. Welling; H. A. F.). 

Fig. 20. Male genitalia. Polythrix guatemalaensis, n. sp. Holotype, Sayaaxche, 
E] Petan, Guatemala, 23 August 1963 (E. C. Welling; A. M. N. H.). 


VoLuME 31, NUMBER 2 95 


man, however, on the lower surface it does not resemble any other spe- 
cies of Epargyreus due to the unusual arrangement of the silvery-white 


discal band. 


Typhedanus salas Freeman, new species 
Pigse:( 5.819 


Male (Upper side). Primaries dark brown. There is a dark, straight band of 
spots between the base and the discal band, extending from space 1b to near the 
costa. There is a dark discal band which is broken outward at vein 3 and is ir- 
regularly continuous with the dark apical band. Fringes dark brown. 

Secondaries dark brown, with the slightest indication of a discal band. There 
is a prominent radiating hair tuft arising from near the base of space lc, which is 
yellowish in coloration. The remainder of the wing dark brown. Fringes sordid 
yellowish-white, uncheckered. 

Male (Under side). Primaries similar to above except space la is yellowish. 

Secondaries similar to above except the discal bands are darker. 

Thorax dark brown above and beneath. Abdomen dark brown above and _ be- 
neath. Head dark brown. Palpi light brown. Legs dark brown. Antennae, shaft 
dark brown, club dark brown with the apiculus orange. 

Wing measurements. Primaries: base to apex, 22 mm; apex to outer angle, 
15 mm; outer angle to base, 15 mm. Secondaries: base. to end of vein 3, 14 mm; 
center of costa to anal angle, 18 mm. Total expanse, 39 mm. 

Female. Very similar to the male, the only difference is on the lower surface 
of the secondaries near the anal angle where it is a slight degree lighter brown. 

Wing measurements. Primaries: base to apex, 26 mm; apex to outer angle, 
17 mm; outer angle to base, 19 mm. Secondaries: base to end of vein 3, 18 mm; 
center of costa to anal angle, 17 mm. Total expanse, 47 mm. 

Type material. Holotype, male, Piste, Yucatan, Mexico, 26 August 1968, will 
be placed in the American Museum of Natural History. Allotype, female, same 
location, 16 August 1968, is in my collection. There are nine male paratypes and 
one female paratype from the same area in my collection. All specimens were 
collected by E. C. Welling. 


This new species is named for Felipe Salas, Assistant Manager of 
Hotel Covadonga, Ciudad Valles, S. L. P., Mexico, who helped me with 
my collecting in that area of Mexico. 

This new species resembles Typhedanus ampyx Godman & Salvin ex- 
cept it lacks all of the bright yellow coloration on both surfaces of the 
secondaries, is somewhat smaller and there are differences in the geni- 
talia. Both species fly together in Yucatan. 


Polythrix guatemalaensis Freeman, new species 
Figs. 9, 10, 20 


Male (Upper side). Primaries dark brown. There is a rather compact discal 
band of four yellow, hyaline spots. The spot in space 2 is large and square. There 


< 


Fig. 21. Female genitalia. Codatractus yucatanus, n. sp. Paratype, Piste, Yucatan, 
Mexico, 2 August 1967 (E. C. Welling; H. A. F.). 

Fig. 22. Male genitalia. Bungalotis milleri, n. sp. Paratype, Candelaria Loxicha, 
Oaxaca, Mexico, 5 August 1969 (E. C. Welling; H. A. F.). 


96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


is a small spot in space 1b directly under the outer edge of the spot in space 2. In 
space 3 there is a small spot directly over the outer edge of the spot in space 2, and the 
inner edge of the cell spot, and the inner edge of the spot in space 2 form an even 
straight line. There are three apical spots, one in space 6 is larger than the one 
in space 8, while the one in space 7 is small. The inner edge of the three spots 
forms an even curve. The costal fold is well developed. Fringes dark brown, con- 
colorous with rest of wing. 

Secondaries dark brown. The lower half of each wing rather badly torn thus not 
indicating the tail length. Fringes dark brown. 

Male (Under side). Similar to above except slightly lighter in coloration and 
there is a dark area just beneath the apical spots. The veins are slightly lighter 
than the ground color. 

Secondaries brown. There is a dark cell spot and another one just above it. The 
discal row of spots from 1B to the cell are dark black and very prominent. The 
veins are slightly lighter than the ground color. 

Thorax brown above with some yellowish hair-like scales intermixed, beneath 
brown. Abdomen brown above and beneath. Head brown with some yellowish 
hair-like scales intermixed. Palpi yellowish-brown. Legs brown. Antennae, shaft 
and club, brown above, yellowish beneath. 

Wing measurements. Primaries: base to apex, 21 mm; apex to outer angle, 15 
mm; outer angle to base, 19 mm. Secondaries: base to end of vein 3, 13 mm; cen- 
ter of costa to anal angle uncertain due to damage of both wings. Total expanse, 
4] mm. 

Female (Upper side). Primaries very similar to male except there is a distinct 
costal spot directly over the cell spot. Fringes tan, lighter than the rest of the wing. 

Secondaries dark brown. Tails broad and fairly short (5 mm), evenly curved 
outward. Fringes sordid white, slightly darker at ends of veins. 

Female (Under side). Very similar to the male. 

Thorax, abdomen, head, palpi, legs, and antennae similar to male. 

Wing measurements. Primaries: base to apex, 22 mm; apex to outer angle, 15 
mm; outer angle to base, 15 mm. Secondaries: base to end of vein 3, 14 mm; center 
of costa to anal angle (end of tail), 23 mm. Total expanse, 43 mm. 

Type material. Holotype, male, Sayaaxche, E] Petan, Guatemala, 23 August 
1963, will be placed in the American Museum of Natural History. Allotype, female, 
X-Can, Quintana Roo, Mexico, 26 July 1962, will remain for the present in my 
collection. Both specimens were collected by E. C. Welling. 


This new species resembles Polythrix procerus (Ploetz) on the upper 
side except in guatemalaensis the discal spots are slightly darker and 
form a more compact band. On the lower surface they do not resemble 
each other at all due to the very dark cell spot and discal band of 
guatemalaensis on the secondaries. The nearest related species appears 
to be P. callias (Mabille) from Bolivia as there is some similarity in the 
dark macular bands on the lower surface of the secondaries. The geni- 
talia are distinct. 


Codatractus yucatanus Freeman, new species 
ie Seale ea 
Female (Upper side). Primaries uniform dark brown, immaculate except for two 


indistinct apical spots in spaces 8 and 9, Fringes indistinctly checkered dark and 
light brown. 


VOLUME 31, NUMBER 2 97 


Secondaries uniform dark brown, immaculate. Tails approximately 14 mm _ in 
length. Fringes indistinctly checkered and light brown. 

Female (Under side). Primaries varying shades of brown, somewhat resembling 
Codatractus carlos Evans, except there are no hyaline spots present except two 
minute apical ones in spaces 8 and 9. The entire coloration is darker than carlos 
and the cellular light area is more extensive. 

Secondaries dark brown with heavy black basal and discal markings. The white 
discal area is similar to that in C. alcaeus (Hew.) and not extending as far upward 
as in carlos. There is a dark bar at the upper end of the white discal area. The 
veins are slightly lighter than the ground color. 

Thorax dark brown above, light, yellowish-brown beneath. Abdomen dark brown 
above, striped yellowish and brown beneath. Head brown, with a white line at the 
base of the eyes. Palpi sordid yellowish-white. Legs light brown. Antennae, shaft 
and club dark brown above, beneath shaft minutely striped yellow and brown, 
club bright yellow. 

Wing measurements. Primaries: base to apex, 31 mm; apex to outer angle, 20 
mm; outer angle to base, 23 mm. Secondaries: base to end of vein 3, 21 mm; 
center of costa to anal angle (end of tail), 35 mm. Total expanse, 58 mm. 

Type material. Holotype, female, Piste, Yucatan, Mexico, 1 September 1967, 
will be placed in the American Museum of Natural History. There are eight fe- 
male paratypes from the same location, collected during June, July, August and 
September 1967 and 1968, in my collection. All specimens were collected by E. C. 
Welling. 


On the upper surface this species does not resemble any other member 
of the genus Codatractus due to the absence of hyaline spots. On the 
lower surface it resembles both carlos and alcaeus based entirely on 
ground color. 


Bungalotis milleri Freeman, new species 
Bigs. Wen ta los 6s-22 


Male (Upper side). Primaries bright orange-fulvous. No hyaline spots. There 
is a dark brownish-black cell spot. At the end of the cell there is a dark bar and 
another dark bar midway between the end of the cell and the apical area forming 
a portion of an evenly curved row of dark apical spots. In spaces la and lb there 
are two circular dark spots, directly above in space 2 there is a dark marking re- 
sembling the letter “B,” above this spot is a dark bar midway between the bar at 
the end of the cell and the one forming the lower portion of the apical spots. There 
is a distinct costal fold. Fringes dark brown. 

Secondaries, costa uniform dark brown with no brilliant blue in side light, re- 
mainder of wing bright orange-fulvous. There is a dark cell spot and a row of 
rather indistinct discal spots. Fringes dark brown. 

Male (Upper side). Primaries, costa, apical area and outer margin to midway 
of space 1 dark brown, remainder of wing bright orange-fulvous. The spots reappear 
but are much less distinctive. 

Secondaries dark brown with some orange-fulvous scales between the dark discal 
markings which are much more pronounced on this surface of the wing. 

Thorax above bright orange-fulvous, brown below. Abdomen above bright 
orange-fulvous, below approximately the same. Head orange-fulvous. Palpi and 
cheeks dark tawny with some lighter scales beneath the eyes. Legs light brown. 
Antennae, shaft both above and below dark brown, club brown, with the terminal 
end of the apiculus light orange both above and below. 

Wing measurements. Primaries: base to apex, 30 mm; apex to outer angle, 


98 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


20 mm; outer angle to base, 23 mm. Secondaries: base to end of vein 3, 20.5 mm; 
center of costa to anal angle, 26 mm. Total expanse, 60 mm. 

Female (Upper side). Primaries dark brownish-black, with a central band of 
white, hyaline spots from space lb to cell, with a detached but approximate spot 
in space 8. There is a minute, linear spot in space 4 midway between the spot in 
space 3 and the outer margin, and a minute dot in space 5. There are three apical 
spots forming a straight line pointing toward the upper fifth of the outer margin. 
Fringes concolorous with rest of wing. 

Secondaries, unmarked, dark brownish-black. Fringes concolorous with rest of 
wing. 

Female (Under side). Very similar to above except space lb is bright yellowish. 

Secondaries dark brownish-black with some gray discal and subbasal spots. There 
are two spots below the costa that are somewhat lighter in coloration than the others. 

Thorax and abdomen dark brown above and below. Head dark brown. Palpi 
and cheeks dark brown, with the slightest indication of some lighter scales below 
the eyes. Legs dark brown. Antennae, shaft and club, dark brown, with the 
apiculus slightly lighter. 

Wing measurements. Primaries: base to apex, 40 mm; apex to outer angle, 25 
mm; outer angle to base, 34 mm. Secondaries: base to end of vein 3, 30 mm; 
center of costa to anal angle, 27 mm. Total expanse, 67 mm. 

Type material. Holotype, male, Candelaria Loxicha, Oaxaca, Mexico, 22 Sep- 
tember 1968, will be placed in the American Museum of Natural History. This 
specimen and two male paratypes from the same location were collected by E. C. 
Welling. Allotype, female, San Quintin, Chiapas, Mexico, 16 September 1970, col- 
lected by Robert Wind, is in the Allyn Museum of Entomology, Sarasota, Florida. 
The two male paratypes from Candelaria Loxicha are in my collection. There is a 
male paratype in the American Museum of Natural History from Rancho San Carlos, 
Chiapas, collected August 1968 by Peter Hubbell. There are single male and fe- 
male paratypes collected 15 November 1971 by Peter Hubbell at Catemaco, 
Veracruz, Mexico, in the collection of Dr. W. W. McGuire, San Antonio, Texas. 


This new species is named for Dr. Lee Miller, Allyn Museum of En- 
tomology, Sarasota, Florida, for his outstanding work on the Rhopalocera. 

This new species does not fit any of the known species of Bungalotis 
in that the costa of the secondaries of the males is not shot with blue 
from the side light thus placing it in the borax Evans (1952) complex, 
but the markings and palpi do not fit other members of that group and 
the genitalia are distinct even though there are some similarities to 
astylos (Cramer) which has the blue shot on the costa of the secondaries 
and the cheeks are distinctly white at the base in both sexes. 


ACKNOWLEDGMENTS 


The author wishes to thank the National Geographic Society for fur- 
nishing research grants which made it possible for this research to be 
conducted during the past four summers. I would also like to thank Dr. 
Frederick H. Rindge, Curator, Department of Entomology, the American 
Museum of Natural History, E. C. Welling and Dr. W. W. McGuire for 
the loan of specimens for determination. The photographs of the adults 
used in this article were made by Melvin Cannon, ICT Coordinator, 
Hillcrest High School, Dallas, Texas. 


VoLUME 31, NuMBER 2 99 


LITERATURE CITED 


Evans, W. H. 1951. A catalogue of the American Hesperiidae indicating the 
classification and nomenclature adopted in the British Museum. Part I. In- 
troduction and Group A, Pyrrhopyginae. London: British Museum, 92 p., 
pls. 1-9. 

1952. A catalogue of the American Hesperiidae indicating the classification 
and nomenclature adopted in the British Museum. Part II. Pyrginae, Sec. 1. 
London: British Museum, 178 p., pls. 10—25. 


ABERRANT ERYNNIS TRITUS TATITUS (HESPERIIDAE ) 


Although they occur with some frequency in many genera, aberrant specimens ap- 
pear uncommon in the North American Pyrginae. Burns, in his monograph Evolution 
in Skipper Butterflies of the Genus Erynnis ( Univ. of Calif. Publ. in Entomol., Vol. 
37, Berkeley, 1964) made no mention of such in this genus. 

In July 1976 while I was collecting in Grant Co., New Mexico, I took an unusual 
Erynnis specimen which I thought at first was a male horatius (Scudder & Burgess). 
Upon genitalic examination, it was determined to be an aberrant E. tristus tatius 
(Edwards), a fairly common species in this region. The specimen was collected 
along the Gila River at Riverside, 4250’ (1295 m), Grant Co., New Mexico on 
4 July 76. 

Fig. 1 shows the dorsal and ventral views of the aberrant specimen while Fig. 
illustrates a typical tatius taken on the H-Y Ranch, Mule Creek area, Grant Co., 
N.M., on 10 August 75. In the aberrant specimen, the ventral HW _ white spots 
are reduced in size and replaced by pale brown with a central light dot. The wing 
fringes are brown rather than white. The forewings resemble normal tatius. A sim- 
ilar aberrant female was taken S of Silver City, Grant Co., N.M. on 26 August 76. 

While I have collected many hundreds of Erynnis, including some unusually small 
and large examples, this is the first aberration that I have encountered. 


2 


Cuirrorp D. Ferris, Bioengineering Program, University of Wyoming, Laramie, 
Wyoming 82071. 


1 


Figs. 1-2. Erynnis tristus tatius. 1, aberrant, dorsal left, ventral right; 2, normal, 
dorsal left, ventral right. 


100 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


STUDIES ON THE BIOLOGY OF PARIDES IPHIDAMAS 
(PAPILIONINAE: TROIDINI) IN COSTA RICA 


ALLEN M. YounG 


Invertebrate Division, Milwaukee Public Museum, Milwaukee, Wisconsin 53233 


The tropical butterfly Parides iphidamas iphidamas (Fabricius) oc- 
curs, along with several congeners (arcas mylotes Bates, arcas mycale 
Goodwin & Salvin, childrenae Gray, and erithalion Druce) in the Pre- 
montane Wet and Lowland Tropical Wet Forest life zones (Holdridge, 
1967) on the Caribbean drainage of the Cordillera Central in Costa 
Rica. It is not uncommon to find butterflies of these Parides visiting the 
same patch of Cephaelis tomentosa Aubl (Vahl) (Rubiaceae) flowers 
each day. At higher elevations in Costa Rica, such as the Montane Wet 
Forest life zone, generally P. arcas mylotes prevails, and although P. 
erithalion and P. childrenae are relatively rare in premontane wet forest, 
they are more abundant in Lowland Tropical Wet Forest. 

The life cycle and natural history of P. arcas mylotes from Costa 
Rican Premontane Wet Forest have been reported (Young, 1973). The 
present paper summarized similar information for P. iphidamas at the 
same locality. Being a very widespread and familar species throughout 
much of Central America, it is likely that this species has been studied 
in various places, but data from Costa Rica are not available. Rothschild 
& Jordan (1906) report that P. iphidamas iphidamas ranges from southern 
Mexico to Panama, and no information is given for the biology of this 
subspecies and the others. This paper is the second in a series on the 
biology of Parides species in Costa Rica, and it emphasizes differences in 
early stages and behavior between P. iphidamas and P. arcas mylotes. 


METHODS 


Studies were conducted from 10 January through 28 February 1976 
on the properties of Compania Agricola Myristica S.A. (C.A.M.S.A.) and 
Compania Agricola Huntro S.A. (C.A.H.S.A.). The two companies have 
adjacent, extensive land holdings in northeastern Costa Rica. This lo- 
cality was called “Finca Tirimbina” in Young (1973), and it is ca. 8 km 
from La Virgen (elev. 220 m), Heredia Province. The region (premon- 
tane wet forest) experiences heavy rainfall throughout most of the year 
and a short, erratic dry period in February and March. Of the land 
encompassed by both companies (a total of ca. 1500 acres), ca. 60% is 
still undisturbed primary forest, whereas the remainder is secondary 
forest and cultivated habitats. Owing, however, to plans to convert most 


VoLUME 31, NUMBER 2 101 


of the primary and secondary forest habitats into cultivated areas, the 
habitats of Parides are disappearing very fast in this region. 

Parides butterflies were studied by searching for adults in a variety 
of different habitats and recording where they feed and oviposit. 

To study the early stages of P. iphidamas, several eggs were collected 
at various times and confined, along with fresh cuttings of the host plant, 
in clear plastic bags kept tightly shut. The technique is essentially the 
same used to study P. arcas mylotes (Young, 1973). Although P. iphi- 
damas has been reared before but incorrectly called P. arcas mycale 
(Young, 1972), the present study focused on the description of the 
early stages, for comparison with those of P. arcas mylotes (Young, 
1973). In addition to finding eggs in the wild, two females were con- 
fined in separate bags with host plant cuttings to obtain eggs. Both 
females were caught from the same flower patch and within two weeks 
of each other during the latter half of January 1976. Parides females 
will generally oviposit readily in captivity, and they can be kept alive on 
sugar-water solutions for several weeks (Young, 1972). The eggs ob- 
tained were reared to the adult stage. 


RESULTS 
Habitat, Adult Feeding, and Larval Host Plant 


Parides iphidamas, in contrast with P. arcas mylotes and P. childrenae, 
flies most frequently in open secondary habitats, and along the edges of 
primary forest. At a flower patch (Cephaelis tomentosa) observations 
of male Parides for 3 hr on sunny mornings resulted in the scoring of 
8-35 visits of P. iphidamas relative to only 3-10 of P. arcas mylotes and 
1-3 of P. childrenae. When C. tomentosa is abundantly in bloom in 
shaded forest, but within a few meters of the forest edge, several species 
of Parides visit the conspicuous red flowers. For example, during the late 
afternoon of 17 January 1976, six visits by both sexes of P. iphidamas, 
four by P. arcas mylotes, and two by P. childrenae were scored within 
20 min (16:10-16:30 hrs) at a small patch of C. tomentosa in an old 
secondary forest of Goethalsia meiantha (Tiliacae) trees. 

Other than C. tomentosa, Parides has not been seen visiting other 
inflorescences. However, probably other food sources exist. Cephaelis 
tomentosa produces it’s large red inflorescences throughtout the year, 
providing a predictable food source for these and other buttertlies, 
in addition to hummingbirds. However, there is considerable asynchrony 
of flower production within a patch, such that as some flowers wither 
following several days of bloom, others come into bloom. A far less 
abundant nectar source for Parides at this locality is Impatiens sultani 


102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(Balsaminaceae), which occurs as small patches in heavily shaded forest 
openings where the ground is almost of mud consistency. The pink or 
red inflorescences attract Parides in mountain forest (Young, 1973), 
where this plant is very abundant. At lowland tropical wet forest lo- 
calities, the bright red inflorescences of Hamelia patens (Rubiaceae) at- 
tract Parides species (Young, 1971). 

At “Tirimbina,” Parides butterflies share C. tomentosa nectaries with 
some Heliconius species (e.g., H. hecale and H. cydno). In the absence 
of precise data from field mark-recapture experiments, it appears that 
males of P. iphidamas and other Parides are more predictable visitors 
than females at a given flower patch on a day-to-day basis. For example, 
at one small forest edge patch of C. tomentosa, the same male P. iphi- 
damas visited for six successive days, whereas several different females 
passed through the patch only one time each during the same period. I 
have witnessed similar patterns at other patches of C. tomentosa. On a 
given day, the same male would reappear 5-30 times at a flower patch, 
whereas a female would appear only once or twice. 

Whereas P. arcas mylotes flies along forest edges (Young, 1973), P. 
iphidamas is more abundant in this ecotonal habitat. All 12 oviposition 
acts took place at forest edges where the larval host plant, Aristolochia 
constricta (Aristolochiaceae), occurs as large mature vines exposed to 
direct sunlight most of the day. Oviposition takes place throughout the 
day in clear weather; generally only one egg is laid, but occasionally 
two or even three eggs are laid separately on a single visit. The egg is al- 
ways placed on the ventral side of an older leaf on a mature vine. The 
larval host plant is very abundant along the edges of primary forest, in 
forest openings, and in open, secondary habitats. Very small seedlings 
of this species occur in heavily shaded understory of forest remnants, al- 
though mature vines are not common in these places. Aristolochia con- 
stricta is microsympatric with A. pilosa, but the latter is generally rare 
at this locality, and it is not used as a host plant by Parides butterflies. 

In a previous study, it was estimated that Parides females are capable 
of producing a few hundred eggs (Young, 1972). But the two females 
held captive in the present study only produced 22 eggs within a few 
days. One female, a young adult as judged by the condition of the wings, 
produced eight eggs between 14 and 16 January 1976, and the second 
female, judged as middle-aged, produced 14 eggs on 5-6 February 1976. 
Additional eggs could have been obtained, but one female was sacrificed 
and the other released. All of the eggs appeared viable in terms of size 
and coloration. 


VOLUME 31, NuMBER 2 103 


Fig. 1. Early stages of P. iphidamas. Left column, top to bottom: egg, first 


instar, second instar, and third instar caterpillars. Right, top: third instar feeding 
on A. constricta in the wild; right, bottom: fourth instar. 


Life Cycle and Behavior of Caterpillars 


Ege (Fig. 1). Brown to honey-orange, spherical, clearly visible vertical fur- 
rows at 10 magnification, 1.0 mm diameter. Hatches in 6-7 days (13 eggs). 


First instar (Fig. 1). Thoracic and abdominal areas rusty-orange, head capsule 


104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


eee 


‘hie, 


Fig. 2. Top: fifth instar caterpillar. Bottom: frontal and lateral aspects of the 
pupa. 


black. Thoracic and abdominal areas tuberculate. Tubercles white or rusty-orange 
with latter color replaced by purple in all later instars; otherwise colors and tubercle 
number and positions the same. Tubercle pattern as follows (all instars): thoracic 
area bears 3 pairs (dorsal, sub-dorsal, lateral), abdominal area bears 2 pairs (dorsal 
and lateral). Prothorcic dorsals and sub-dorsals white, laterals rusty-orange; meso- 
and metathoracic dorsals and laterals white, subdorsals rusty-orange. First abdom- 


VoLUME 31, NUMBER 2 105 


Fig. 3. Adult P. iphidamas obtained from laboratory rearing: female above, 
male below. 


inal, both sets rusty-orange, laterals of second abdominal becoming white. Pattern 
reversed for third abdominal segment. Both sets of fourth and fifth abdominal seg- 
ments rusty-orange, both white on sixth, only dorsals white on seventh. Both 
white on eighth and ninth abdominal segments; tubercles of ninth very reduced. 


106 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Black setae prominent on tips of all tubercles but disappear in later instars. Grows 
from 2.5 to 4.0 mm in 5 days; eats egg shell to varying degrees (11 individuals ). 

Second instar (Fig. 1). As first instar, but with rusty-orange replaced with 
purple. Prominent white “collar” evident just behind head, present in previous in- 
star but reduced. Body more cylindrical with tubercles shorter relative to thickness 
of body. Both first and second instars feed on young leaves of host plant in the 
field and laboratory. Grows from 4.0 to 9.0 mm in 5 days (10 individuals). 

Third instar (Fig. 1). Identical in color and form to previous instar. First 
three instars quickly evert bright yellow osmeteria when disturbed, and this ability 
diminishes drastically in later instars. First three instars feed from ventral side of 
young or old leaves in wild. Grows from 9.0 to 23.0 mm in 5 days (8 individuals ). 

Fourth instar (Fig. 1). Thoracic and abdominal areas cuticle has glossy sheen. 
One striking color pattern difference retained in fifth instar: white of lateral 
tubercles of third abdominal segment extends to meet expended white basal area of 
fourth abdominal segment, giving appearance of thick, broken line connecting the 
segments. Similar expansion of sub-dorsal tubercle colors on metathoracic area. 
Grows from 23.0 to 34.0 mm in 5 days (8 individuals ). 

_ Fifth instar (Fig. 2). Same as fourth instar. Both fourth and fifth instars eat 
woody stems of host plant in addition to leaves. Both accept A. maxima in the 
laboratory, but for two tested, both died after feeding several days. Grows from 
34.0 to 56.0 mm in 6 days (8 individuals ). 

Pupa (Fig. 2). Angulate, bluish-green ventrally, yellow-green dorsally; 31.0 
mm long, 10.0 mm wide (dorsal-ventral), 15.0 mm thick. Head capsule strongly 
forked. Darkens to adult colors day prior to eclosion. Lasts 28-31 days (7 individ- 
uals). Adult ready for flight 2 hr after eclosion. 

Adult (Fig. 3). Good descriptions in Godman & Salvin (1879). Easily dis- 
tinguished from P. arcas mylotes in the wild by marginal dots on wings being 
white in P. iphidamas and red in the former. Females readily identified by presence 
of variable red spot in cell between veins M: and RS ventrally on hindwings; spot 
always absent in P. arcas mylotes examined at this locality. 

Total development time 60-63 days (7 individuals ). 


DIscussIONn 


In premontane tropical wet forest regions such as “Tirimbina” and the 
surrounding area, P. iphidamas and P. arcas mylotes both exploit Aristo- 
lochia constricta as a larval host plant. A. constricta is by far the most 
abundant of the four or five species of the genus that occur here. It oc- 
curs as large patches in open secondary habitats and as isolated small 
patches or young single vines in heavily shaded understories of forest 
remnants that dot the area. Individual vines in forest understory possess 
thinner leaves than mature vines in open areas, and their leaves are gen- 
erally darker green. Although four species of Parides visit C. tomentosa 
inflorescences in heavily shaded forest understory (old secondary forest) 
and along forested roads and paths, there is a strong preference for 
P. iphidamas to oviposit on mature vines in open areas, whereas P. arcas 
mylotes oviposits primarily on very small seedlings of A. constricta in 
forest understory. It was noted elsewhere (Young, 1973) that females 
of P. arcas mylotes, presumably mated, make frequent excursions into 
shaded forest in search of oviposition sites. This is not the case with P. 


VoLUME 31, NUMBER 2 107 


iphidamas. Although females of both species and also those of P. 
childrenae, the latter being a far more elusive species than the other 
two, may often be found together in the same places, there is a definite 
preference in P. iphidamas for oviposition in open places. In the wild, 
it is generally easier to follow ovipositing females of P. iphidamas than 
those of P. arcas mylotes, and, at least in part, this probably involves 
a behavioral difference between these species for oviposition site selec- 
tion. Since P. arcas appears to be a species that oviposits in forests, 
where seedlings and individual vines of the host plant are probably 
more widely dispersed than in secondary habitats, the females may 
spend greater amounts of time on a daily basis searching for oviposition 
sites than do P. iphidamas females searching in secondary and forest edge 
habitats. 

Elsewhere (Young, 1973), the developmental time of P. arcas mylotes 
was reported to be ca. 53 days, compared with 60-63 days for P. iphi- 
damas in the present study; the host plants were the same in the two 
studies and rearing conditions very similar. It is expected, however, 
that an ecological statistic such as developmental time will be very 
sensitive to subtle environment factors, making comparisons difficult. 
Certainly the size of adults will be altered by differences in develop- 
mental time within and between species. Considerable differences in 
developmental time occur in Parides even when reared under similar 
conditions (Young, 1972; 1973). 

Although caterpillars of P. iphidamas and P. arcas mylotes are similar 
in color and size, there are some consistent and useful differences be- 
tween them that assist in field identification. Since the two species be- 
long to different groups within the genus, differences in early stages 
are expected. The fourth and fifth instars of P. arcas mylotes are colored 
with a variegated pattern of brownish-purple and velvety black, whereas 
those of P. iphidamas are glossy purple; the caterpillars of P. arcas 
mylotes lack the broken white band of P. iphidamas caterpillars. The 
caterpillar of P. arcas mylotes lacks the white coloration of the lateral 
tubercles of the third abdominal segment that comprises the lower por- 
tion of the lateral white band in P. iphidamas caterpillars. Furthermore, 
the caterpillar of P. arcas mylotes lacks the white thoracic tubercles of P. 
iphidamas (compare Fig. 3 in Young, 1973 with Fig. 1 in present paper). 
Other differences occur in the pupa stage for the two species: when 
placed side by side, it is evident that the pupa of P. iphidamas is wider 
and more greenish-yellow than that of P. arcas mylotes, which is narrow 
and bluish-green. 

Aside from the clear differences in the males of P. iphidamas and P. 


108 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


arcas mylotes, it is interesting that the males of former species possess 
a very thick white fold of inner margin on the hindwing. This structure 
(Fig. 3) is black and reduced in males of P. arcas mylotes. 


SUMMARY 


Various aspects of the biology of the tropical butterfly Parides iphi- 
damas were studied at one locality in the premontane tropical wet forest 
life zone of northeastern Costa Rica. Here, this species occurs with sev- 
eral other of the genus, including the similar-appearing P. arcas mylotes. 
Although both P. iphidamas and P. arcas mylotes exploit Aristolochia con- 
stricta ( Aristolochiaceae ) as a larval host plant, the former species shows 
a preference to oviposit on mature vines in open secondary habitats and 
the latter species in forest understory. These species, along with 
others of the genus, show the same preference to visit Cephaelis tomen- 
tosa inflorescences in forest understory. Emphasis in interpreting the 
form and features of early stages is placed on a comparison between 
P. iphidamas and P. arcas mylotes caterpillars and pupae for aiding 
field identifications. 


ACKNOWLEDGMENTS 


This research was supported by National Science Foundation grant 
GB-33060. The cooperation of Dr. J. Robert Hunter of C.A.M.S.A. and 
C.A.H.S.A. with logistical matters and field station facilities is greatly ap- 
preciated. I thank Dr. Lee D. Miller (Allyn Museum of Entomology) for 
confirming the identification of the species. 


LITERATURE CITED 


GopMan, F. D. & O. Satvin. 1879. Biologia Centrali-Americana. Insecta. Lepi- 
doptera-Rhopalocera. Taylor & Francis, London. 

Houprince, L. R. 1967. Life zone ecology. Tropical Science Center, San Jose, 
Costa Rica. 

RotuscuHitp, W. & K. Jorpan. 1906. A_ revision of the American Papilios. 
Novitates Zool. 8: 27-753. 

Younc, A. M. 1971. Mimetic associations in natural populations of tropical 
papilionid butterflies (Lepidoptera: Papilionidae). J. New York Ent. Soc. 79: 
210-224. 


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

1973. Notes on the life cycle and natural history of Parides arcas mylotes 
(Papilionidae) in Costa Rican premontane wet forest. Psyche 80: 1-22. 


VoLUME 31, NuMBER 2 109 


OBITUARY 


RICHARD B. DOMINICK, M.D. (1919-1976) 


Dr. Richard B. Dominick, Dick to his friends, died suddenly on 4 May 1976. He 
appeared healthy in the morning and was dead by the evening; he was spared the 
difficulty of a lingering illness. 

Dick was an energetic, many faceted man. Academically, he was an alumus of 
Yale University and the Columbia University Medical School. Among his activities 
and interests at different times were crew, piloting in WWII and subsequently, 
boy scout leadership, hunting, fishing, newspaper writing, literature, the Church, 
medicine, support of the Peabody Museum’s research program, the Charleston Mu- 
seum, photography, the Wedge, Lepidoptera, conservation and preservation of nat- 
ural areas and people. 

I first met Dick in 1968 at the Wedge with Doug Ferguson and Jack Franclemont 
when the Moths of America North of Mexico began to assume form. Since then 
much of Dick’s energy, drive, and talent was directed to making the project a 
reality. Dick was not a taxonomist, and he readily proclaimed it, but he was de- 
termined to play an active role in the series. This he did in several ways: he 
learned color photography and photographic developing to the point that he pro- 
duced the excellent positives used for the colored plates; he set deadlines and drove 
the editor-authors to try to meet them; he made the project financially viable; he 
provided a setting in which work could be done; and more. The project was left 
at a stage where it could stand on its merits and momentum and move forward. 

Dick was genuinely friendly, helpful, and interested in people. To spend time at 
the Wedge (the name of his home and property on the Santee River in Charleston 
County, South Carolina), where many of life’s vagaries seemed to vanish, was al- 
ways a pleasant experience. Dick believed in individuality at a time when many 
seemed lost in conformity. He usually got up by 3 a.m. and then went to the lab, 
an extremely well-equipped facility, where he worked on some aspect of Lepidoptera, 
Meals were taken at varying times. He would literally fall asleep standing up by 
8 or 9 p.m. He developed an excellent collection of those butterflies and moths found 
on the Wedge by bait and light trapping and aerial collecting. He reared many 
species, particularly satumiids and sphingids and became interested in freeze- 
drying as a method of preserving larvae and pupae. 


110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


I cannot long think of Dick without considering the interplay between him and 
his wife, Tatiana. She is also a person of immense energy, vitality, and competence, 
an extremely suitable counterpart to Dick’s sometimes boisterous activities. The 
two were always thoughtful and protective of each other and were jointly responsive 
to a multitude of demands. 

Dick is survived by his wife, his children Julia and Oliver, his stepchildren Eliza- 
beth, Stephanie, Victoria, and Christopher, and a grandson Alexander. 


Ronatp W. Honces, Systematic Entomology Laboratory, Agriculture Research 
Center, USDA, Beltsville, Maryland 20705. 


VoLUME 31, NUMBER 2 fil 


NOTES ON THE BEHAVIOR OF ASTEROCAMPA LEILIA 
(NYMPHALIDAE) IN SOUTHERN ARIZONA 


GrEoRGE T. AUSTIN 


Department of Biological Sciences, University of Nevada, Las Vegas, 
Las Vegas, Nevada 89154 


While conducting studies on the Santa Rita Experimental Range, 
Pima Co., Arizona, in 1970 and 1971, I obtained data on the behavior 
of Asterocampa leilia Edwards, especially with regard to temperature 
and territoriality. Although these data are largely incomplete, they are 
reported now because my studies in southern Arizona are not being 
continued and these aspects of life history are unknown for this species 
and poorly known for butterflies in general. 


METHODS 


Relative abundance was determined by counting all butterflies as they 
were encountered within 5 m of me as I walked through the study area. 
Time budget studies were made over stopwatch-timed intervals on ca. 
10 individuals. Concurrent air shade temperatures (T,) were obtained. 
Microhabitats were distinguished as full sun, partial shade with an in- 
terspersion of sun and shade, and full shade. Posture was expressed as 
wings fully spread at 180°, wings partially spread, or wings closed 
tightly together. Orientation with respect to the sun was also noted. Too 
few data were obtained to ascertain any diurnal changes in behavior 
patterns that may have been present. Most data on interactions with 
other fauna were gathered during time budget observations. Data were 
collected on clear, windless days. Additional miscellaneous lite history 
data were obtained incidentally. 


Study Area 


The study area was located on the western slope of the Santa Rita 
Mountains, ca. 10 km SE of Sahuarita, at an elevation of 1150 m. The 
area was relatively flat, sloped slightly to the northwest, and was dis- 
sected by a maze of small and large desert washes. The community, in- 
cluding the study area, was described as desert-grassland (Lowe, 1964) 
which has been invaded by considerable woody growth mainly as a re- 
sult of protection from fire (Humphrey, 1958). No grazing has oc- 
curred in much of the area for the last several years. The dominant 
vegetation included mesquite (Prosopis glandulosa), paloverde (Cer- 
cidium microphyllum), hackberry (Celtis pallida), and cholla cacti 


112 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(Opuntia fulgida and O. spinesior), with an understory of several grass 
species, some small woody bushes, succulents, and herbs. Photographs 
of the area were published by Anderson & Anderson (1973). 

Rainfall and temperature were typical for this part of Arizona. Rain 
fell principally in winter (December—February) and summer (July- 
August). Daytime temperatures during midsummer commonly exceeded 
30°C whereas morning lows were often near 20°C. 


Territoriality 


Males of A. leilia flew at and chased nearly anything that came close 
to them. I have records of 179 attacks by males, of which 64 were intra- 
specific, 78 were directed at other butterfly species that ranged in size 
from Microtia dymas Edwards to Battus philenor (L.), one was di- 
rected at the moth Celerio lineata (Fabricius), 24 at Odonata, 11 at 
Hymenoptera, including one at an ant, and one at a lizard (Cnemidoph- 
orus sp.). I never saw one fly directly at a bird, although perched A. 
leilia usually flew when a bird passed overhead. 

No butterflies were marked, but three males I studied were recog- 
nizable by distinctive tears in their wings. These three were apparently 
resident for at least 8, 14, and 17 days after I first saw them. Addi- 
tionally, males which were observed at various times throughout the 
course of a day remained within a definite area at all times. These used 
very few perch sites, of which one or two seemed preferred and were 
returned to time after time following flights. Favored perch sites were 
on the ground, in a wash, and usually near the center of the area used 
by the individual. 

The majority (63%) of flights were initiated by disturbance caused 
by another organism passing over or near a perched individual. Many 
of these flights terminated in a patrol within a well-defined area that did 
not change diurnally or over several days. Furthermore, many flights 
were initiated without any obvious stimulus when the butterfly would 
similarly patrol this area. The patrolled areas of neighboring individuals 
overlapped little. Many intraspecific interactons occurred near the bound- 
aries of the patrolled areas when one male entered the area occupied 
by another. Invariably, the intruding individual was chased by the resi- 
dent male. Nearly all of these areas were along washes. 

This strong site tenacity and associated behavior indicate true ter- 
ritoriality. The occurrence of territoriality in butterflies has been the 
subject of some debate. Considerable evidence for territoriality in males 
of two Nymphalids was presented by Baker (1972). Scott (1974), 
however, argued that territoriality was absent or rare among butterflies. 


VoLUME 31, NUMBER 2 1S 


TABLE 1. Average length of intraspecific and interspecific interactions by 
Asterocampa leilia males. 


Mean number of seconds 


Interaction piesa anaes 
with ie 4076 Re a0 Total 
Asterocampa leilia USO oe 15.6 (14) 16.8 (23) 
Other butterflies 6.5 (26) 5.1 (21) 5.9 (47) 
Other insects - - 5.9 (26) 
1 Seconds. 


2 Number of flights. 


He claimed that site tenacity was poor and that the apparent pugnacity 
of perched males was in reality a mate-seeking behavior. 

My data on A. leilia indicate that although males were disturbed by 
nearly any passing object, a distinct difference existed between responses 
to and contacts with intraspecific and interspecifc objects. Intraspecific 
interactions averaged nearly three times longer than interspecific inter- 
actions (Table 1). Thus, males were able to quickly distinguish a conspe- 
cific. Initially, all contacts were investigative. If the individual was not 
an A. leilia, it was on occasion chased briefly, but usually not out of 
the defined area. Conspecific males were pursued to the boundaries 
of the area. Four contacts with known female A. leilia lasted an average 
of 35 s. The results of these contacts were unknown, but after one, the 
male could not be found within his known territory, although he was 
present later in the day. The pair may have disappeared into the vege- 
tation. On one occasion, I found a male and female perched near each 
other in the middle of a Celtis pallida. 

Fourteen territories were examined in some detail. All except one 
were along washes. They contained an average of 5.7 (range 4-10) 
shrubs and small trees of which an average of 1.6 (range 1-3) were 
C. pallida. This tree was the only one common to all territories. The 
defended area averaged 0.07 ha. (range 0.03-0.13 ha). 

The territorial behavior of A. leilia shows both similarities and dif- 
ferences to that of two other Nymphalids (Baker, 1972). A. leilia is 
similar to Aglais urticae (L.) in that the male defended an area which 
included an oviposition site, whereas Inachis io (L.) males de- 
fended areas along female flight paths on route to oviposition sites. 
Asterocampa leilia, however, was similar to Inachis io in that there was 
but one male per territory. In Aglais urticae, territories were often oc- 
cupied by several males. Asterocampa leilia apparently defended ter- 
ritories for much of the day compared with only ca. 4 h per day by 
Inachis io and 1.5 h per day by Aglais urticae. The latter two species 


114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Time budget and number and length of flights for male Asterocampa 
leilia as a function of ambient temperature. 


Ambient Temperature (°C) 


20-25 25-30 30-35 35-40 
% time perched 92.6 91.8 81.0 98.6 
% time flying 7.4 8.2 19.0 1.4 
Total time (s) PAILS 15426 10812 3636 
Mean number of flights per hour 3.2 25.2, 39.0 2.2 
Mean length of flights (s) : 81.0 LiL 7-6 25.5 


spent much of the morning feeding; I have observed feeding by adult 
Asterocampa leilia but once (on Coyote, Canis latrans, feces containing 
much Opuntia fruit). 


Temperature-related Behavior 


Behavior of Asterocampa was greatly affected by temperature. Amount 
of time spent flying was ca. 8% at low T,, increased to 19% at moderate 
T, and decreased to less than 2% at high T, (Table 2). At a T, of 20- 
25°C, few flights were made (Table 2), but these were long in duration 
and usually very fast and erratic. These were usually the first flights 
of the day. Later and at warmer T,, flights were more frequent and of 
shorter duration. These were usually the slower stroke-glide type of 
flight characteristic of the species. At high T,, flights were again in- 
frequent and of relatively short duration (Table 2). Pursuits also ap- 
parently decreased in length with increasing T, as shown in Table 1, al- 
though the sample size is small. 

As temperatures increased, there was a gradual shift in use of perch 
sites from those completely exposed to the sun at low T, to those com- 
pletely shaded at high T, (Table 3). The positioning of the wings also 
varied overall and within the three microhabitats used for perching, 
with T,. At low T,, the wings were nearly always spread when the 
butterflies were perched; above 30°C, the wings were nearly always 


TABLE 3. Microhabitat usage by male Asterocampa leilia as a function of ambient 
temperature. 


Usage (% Total Time) 


Microhabitat 20-—25°C 25—-30°C 30—35°C 35—40°C 
Sun 100.0 Tay Al ONS 17 0.0 
Partial shade 0.0 24.2, Son 0.0 


Shade 0.0 1.5 41.5 100.0 


VoLUME 31, NUMBER 2 115 


TABLE 4. Wing position by male Asterocampa leilia as a function of ambient 
temperature. 


% Total Time 


Position 20—25°C 25-—30°C 30-35°C 35-40°C 
Spread 99.2 51.6 3.9 0.0 
Partially spread 0.0 25.3 4.6 0.0 
Folded 0.8 23.1 91.5 100.0 


folded dorsally (Table 4). At moderate T,, the wings were spread 
more when perched in full shade than when perched in partial shade 
or in full sun (Table 5). 

At each T,, even when perched in the shade, the males nearly always 
perched with the head facing away from the sun. This was usually the 
orientation at which they alighted following a flight. If not, they so 
aligned themselves almost immediately. 

There are few other quantitative data that illustrate the various aspects 
of behavioral thermoregulation by butterflies as outlined by Clench 
(1966). Asterocampa leilia reacted behaviorally to temperature in a man- 
ner similar to that described for the nymphalids Argynnis paphia (L.) 
by Vielmetter (1958) and Precis villida (Fabricius) by Heinrich (1972). 

Asterocampa leilia is a dorsal basker as is typical for nymphalids in 
general (Clench, 1966). This species showed all the characteristic be- 
havioral patterns of dorsal baskers. At cool T,, the wings were opened 
fully so that they were maximally exposed to the sun. With increasing 
T,, the degree of opening was varied to adjust the amount of heat gain. 
This was concurrently augmented by selection of an appropriate micro- 
habitat to further control the rate of heat gain. Still additional control 
was attained by slowly opening and closing the wings while perched. 
This behavior was noted 0.8% and 2.2% of the total perching time at a 
T, of 25-30°C and 30-35°C, respectively, but not at any other T,. Heat 
loss or reduction of solar heat gain at high T,, was facilitated by selecting 
the coolest microhabitat and reducing activity to a minimum. 


TABLE 5. Percent of time wings are folded by male Asterocampa leilia in three 
microhabitats as a function of ambient temperature. 


% Time Folded 


Mrerahalitat 20-25°C 25-30°C 30-35°C 35-40°C 
Sun 0.8 223 100.0 as 
Partial shade = 27.0 91.8 = 
Shade = 0.0 87.1 100.0 


116 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 6. Percent of Asterocampa leilia in counts of butterflies on the Santa Rita 
Experimental Range. 


Asterocampa leilia (%) 


Year May June July Aug. Sept. Oct. Nov. 
1970 8.9 ee Zoro DAB yO Oe, 0.0 — 


1971 el 5.0 51.6 29.1 A4l.] - 38.1 


Normal activity in A. leilia occurred in the T, range of 25-35°C, 
slightly higher than the generalized range given by Clench (1966) and 
that for an alpine species (Erebia epipsodea Butler) by Brussard & 
Ehrlich (1970). Asterocampa leilia was actually active in a temperature 
niche considerably warmer than this for much of the day, since the 
temperatures given herein were obtained in the shade. Exposed air 
temperature taken with a silver-bulbed mercury thermometer average 
ca. 5°C greater than shade T,. The radiational heat load on a relatively 
dark-colored insect would be even greater. Considerable interspecific 
differences have been shown in the active thoracic temperatures of butter- 
flies (Heinrich, 1972). A. leilia apparently flies with and is able to 
tolerate high thoracic temperatures, which is of definite adaptive value 
in the southwestern deserts. 


General Life History 


On the Santa Rita Experimental Range, adult A. leilia were active 
from early May—mid-November. Fresh individuals were noted through- 
out the flight period. Peak abundance (both actual and relative) was 
from early July-September (Table 6). The daily flight period was long. 
The first flying individuals undisturbed by me were observed as early 
as 30 min. after sunrise but more usually 90 min. after sunrise. These 
early morning flights, as noted above, were usually very fast and er- 
ratic. The last active individuals were noted just before sunset. 

The general behavior of males was described in detail above. Females 
were observed infrequently. They appeared to fly along washes until 
intercepted by a male. Females were usually encountered by examining 
Celtis pallida bushes, where they were found perched in the shade on 
the lower sides of branches. In most cases, they remained in this posi- 
tion for considerable periods, up to at least 2 h. 

Oviposition was noted four times, all on C. pallida and between 0930 
and 1200. Eggs were laid in clusters of 10, 11, 13, and 15. Three clusters 
were on the upper side of leaves, and one was on the lower side. In each 
case, hatching took place on the seventh day following oviposition. 


VoLUME 31, NUMBER 2 WE 


Thirty-six of the 49 eggs hatched; the remaining eggs showed no signs 
of development. On one occasion, I found a female perched in a C. 
pallida. She perched for 82 s, flew slowly with a male among the 
branches of the C. pallida for 16 s, then laid a cluster of 10 pale yellow 
eggs on the lower side of a leaf in 109 s. After this, she again perched 
for 455 s before flying out of sight. 

The behavior of males perching with the head facing away from the 
sun, as noted earlier, may also function in quickly detecting the approach 
of an object from behind. Such an object would cast a shadow which 
passes by the perched individual before the object itself. Many times I 
noted males initiating flight in response to a shadow before the object 
itself was in view. 

I never observed A. leilia visiting flowers or mud puddles. Other 
Asterocampa were reported to visit mud puddles (Klots, 1951). 


SUMMARY 


Territorial and temperature-related behavior of A. leilia were investi- 
gated in southern Arizona. Males appeared to be truly territorial. They 
investigated nearly all passing objects and chased conspecific males to 
the boundary of a well-defined area. These territories were described. 
Behavior by males was temperature dependent, with a shift from ex- 
posed to shaded microhabitats as temperature increased. Concurrent 
changes in other behavior patterns also occurred. Other miscellaneous 
life history notes were presented. 


ACKNOWLEDGMENTS 


This study was conducted incidental to field work supported by the 
US/IBP Desert Biome Program under National Science Foundation 


Grant GB 15886 at the University of Arizona. I thank Donald Thomas 
and Kenneth Moor for critical comments on the manuscript. 


LITERATURE CITED 


ANDERSON, A. H. & A. ANDERSON. 1973. The Cactus Wren. Univ. of Arizona 
Press, Tucson. xiv + 226 p. 

Baker, R. R. 1972. Territorial behaviour of the nymphalid butterflies, Aglais 
urticae (L.) and Inachis io (L.) J. Anim. Ecol. 41: 453-469. 

Brussarp, P. F. & P. R. Enruicn. 1970. Adult behavior and population structure 
in Erebia epipsodea (Lepidoptera: Satyrinae). Ecology 51: 880-885. 

CiencuH, H. K. 1966. Behavorial thermoregulation in butterflies. Ecology 47: 
1021-1034. 

HernricH, B. 1972. Thoracic temperatures of butterflies in the field near the 
equator. Comp. Biochem. Physiol. 43A: 459-467. 

Humpurey, R. R. 1958. The desert grassland. A history of vegetational change 
and an analysis of causes. Botan. Rev. 24: 193-252. 


118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Kuots, A. B. 1951. A field guide to the butterflies. Houghton-Mifflin Co., Bos- 
ton. xvi + 349 p. 

Lowe, C. H. (ed.). 1964. The vertebrates of Arizona. Univ. of Arizona Press, 
Tucson. viii + 259 p. 

Scotr, J. A. 1974. Mate-locating behavior of butterflies. Am. Midl. Nat. 91: 
103-117. 

VIELMETTER, W. 1958. Physiologie des Verhaltens sur Sonnerstrahlung bei den 
Tagfalter Argynnis paphia L.—I. Untersuchungen im Freiland. J. Insect 
Physiol. 2: 13-37. 


UNIFORM GENITALIA AMONG WING COLOR MORPHS 
OF OLETHREUTID MOTHS 


Most traits are said to be uniform among lepidopteran wing color morphs (Ford 
1975, Ecological Genetics, ed. 4, 442 p., John Wiley & Sons, New York). This 
uniformity presumably includes structure, specifically genitalia. Important as they 
usually are taxonomically, genitalia are seldom mentioned in literature on wing color 
polymorphism (Robinson 1971, Lepidoptera Genetics, 687 p., Pergamon Press, 
New York). Because wing color polymorphism in well studied examples is con- 
trolled by only one or a few genes, structural uniformity is expected and hence not 
likely to be reported. 

In olethreutids, wing color polymorphism and its genetics have been little studied. 
The occurrence of wing color morphs is problematic in many little known species 
in this family. Empirical evidence for genitalic uniformity among putative wing 
color morphs could be taxonomically helpful. From Opler’s (1971, J. Lepidop. Soc. 
25: 115-123) discussion of two species of Epinotia having wing color morphs, 
uniform genitalia can be inferred. I report here explicitly on this point in two ad- 
ditional species. 

Sciaphila duplex (Walsingham) (subfamily Olethreutinae), feeding on Populus 
tremuloides (McGregor 1967, J. Econ. Ent. 60: 1213-1216), has two wing color 
morphs (Heinrich 1926, U.S. Nat. Mus. Bull. 132, 216 p.), one of which is melanic 
in both sexes. The melanic morph numbered 5 of 54 specimens from Michigan, 
Ontario, and Minnesota. Genitalia comparison between the morphs was based on 2 
or more genitalia slide preparations of each sex (9 preparations in all). 

Epinotia solandriana (Linnaeus) (subfamily Eucosminae), feeding chiefly on 
Betula, has 4 main wing color morphs (Lindquist and MacLeod 1967, Can. Ent. 99: 
1110-1114), each in both sexes. These morphs may not be sharply discontinuous. 
Using specimens from Ontario, Wisconsin, and Michigan, I compared genitalia 
among these 4 morphs with 1-7 preparations of each sex (18 preparations in all). 
Comparisons were made under a light microscope at 60-90, magnifications nor- 
mally used in genitalia study. There were no genitalic differences between color 
morphs in either sex of either species. This result confirms expectation and strengthens 
the usefulness of genitalia for ascertaining presence or absence of wing color poly- 
morphism in olethreutids. 

I thank curators of collections at Michigan State University, University of 
Wisconsin, and University of Minnesota for loans of material. 


Wittiam E. Miter, North Central Forest Experiment Station, USDA Forest 
Service, Folwell Avenue, St. Paul, Minnesota 55108. 


VoLUME 31, NUMBER 2 119 


CUDONIGERA: A NEW GENUS FOR MOTHS FORMERLY 
ASSIGNED TO CHORISTONEURA 
HOUSTONANA (TORTRICIDAE) 


Jerry A. POWELL AND N. S. Opsraztsoy! 


Department of Entomological Sciences, University of California, 
Berkeley, California 94720 


During 1952-1966 Dr. N. S. Obraztsov worked as a Research Fellow at 
the American Museum of Natural History, New York, on a generic 
classification of Nearctic Tortricinae. In the process he progressively 
expanded the scope of his work, both geographically, into the Neotrop- 
ical fauna, and in the taxonomic level of treatment, which led him to 
several detailed reviews at the species level. Unfortunately, these 
studies diluted his concentration on the generic revision, and at the time 
of his sudden death in 1966, only parts of the comprehensive study had 
been completed. A summary of Obraztsov’s life and varied work on 
Lepidoptera has been given by Diakonoff (1966). 

The generic treatment, which is intended to accompany and augment 
that of Obraztsov (1954-1957) treating the Palearctic fauna, is being 
continued by Powell. This has involved incorporation of considerable 
Mexican Nearctic material accumulated during the past decade, en- 
abling clarification of relationships among North American genera. Par- 
ticularly in the Cnephasiini, the New World fauna appears to consist of 
Neotropical elements, with the depauperate Nearctic representation 
showing little relationship to the Palearctic, and this tribe was developed 
only preliminarily in Obraztsov’s manuscripts. By contrast, the Tortricini 
and Archipini consist primarily of Holarctic or Nearctic genera which ap- 
pear to be more clear-cut, and Obraztsov had completed study of most 
of them. 

The genus Cudonigera was envisioned by Obraztsov as monobasic, 
related to the Holarctic genus Choristoneura Lederer. Current research 
by Mutuura and Munroe at Ottawa indicates that the populations re- 
ferred to C. houstonana (Grote) should be considered a complex of al- 
lopatric species. The genus is proposed to make the name available for 
use in their studies. 


Cudonigera Obraztsov and Powell, new genus 


Type species: Tortrix houstonana Grote, 1873. 


Tortrix (in part); Grote, 1873, Bull. Buffalo Soc. Nat. Sci. 1: 15. Fernald, 1882, 
1 Deceased in 1966. This study was in part conducted through support from National Science 
Foundation grants to Obraztsov, in 1959-1965. 


120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-5, Cudonigera houstonana (Grote): 1, head, lateral aspect; 2, wing vena- 
tion; 3, male genitalia, ventral aspect, aedeagus removed, a aedeagus lateral aspect; 
4, 5, female genitalia with structures of VIIJ—X segments in venterolateral aspect in 
4, ventral aspect in 5, showing lateral rotation of papillae anales. 


VoLUME 31, NuMBER 2 121 


Trans. Amer. Ent. Soc. 10: 17. Grote, 1882, New check list of North Amer. 
moths: 58. Fernald, “1902”/1903/, Bull. U.S. Natl. Mus. 52: 482. 

Lozotaenia (in part); Walsingham, 1879, Illustr. typical specimens of Lepid. 
Heterocera 4: 13. 

Cacoecia (in part); Meyrick, 1912, in Wagner, Lepid. Catalogus 10: 21; 1913, in 
Wytsman, Genera Insectorum, fasc. 149: 25. Barnes & McDunnough, 1917, 
Check list Lepid. Boreal Amer.: 177. 

Archips (in part); McDunnough, 1939, Mem. Southern Calif. Acad. Sci. 2(1): 56. 

Choristoneura (in part); Freeman, 1958, Can. Ent. 90, suppl. 7: 38. Powell, 1964, 
WeCalia Publ, Ent. 32: 185. 


Adult: Head (Fig. 1) densely appressed scaled, face smoother. Antennae in 
male shortly ciliated and with setae; in female only with setae. Labial palpi ascend- 
ing, densely appressed scaled; second segment length about 0.8 eye diameter, 
slightly dilated apicad; third segment about 0.3 as long as second, blunt, exposed. 

Forewing (Fig. 2) elongate-rectangular, moderately broad; costa gently arched; 
apex obtuse; termen rather straight, sometimes slightly convex, tornus broadly 
rotundate; dorsum gently convex, more curved basad. No costal fold in male. 
Twelve veins, all separate; Sc slightly curved, almost straight; R: from just before 
middle of discal cell; Re twice as near to Rs as to Ri; Rs and Rg slightly diverging 
costad; Rs to costa, Rs to termen; upper internal vein rudimentary, from between Ri 
and Re; Mz nearer to Mz than Ms; to Cui; Cu: from lower angle of discal cell; Cuz 
from shortly before two-thirds; A: vestigial, distinct tornad; basal fork of A.,. slightly 
longer than one-third of entire vein. 

Hindwing (Fig. 2) rotundate-subtrapezoidal; costa slightly sinuate, convex at 
middle; apex rotundate; termen flat or slightly concave below apex; tornus and dorsum 
forming a strongly convex arch. Eight veins; S almost straight; R and M; connate 
or short stalked; Mz gently bent downward basad, remote from Ms; Mz close to Cu, 
separate; Cu: from lower angle of discal cell; Cue from two-thirds. No cubital 
pecten. 

Male genitalia (Fig. 3): Mensis ventralis represented by two narrow, sclerotized 
folds of intersegmental membrane. Tegumen strong, wide, with broad, flat shoulders; 
pedunculi broad, narrowed basad, and bent inward at extreme base; saccus ro- 
tundate. Gnathos strong, with a long, narrow middle process. Socii minute, rudi- 
mentary. Valvae weak, short, rotundate; costa not sclerotized; sacculus moderately 
broad, slightly longer than lower edge of valva, with a free tip; valvula finely 
striated; pulvinus soft, interior; no processus basales. Uncus broad, spatulate, di- 
lated apicad, concave at caudal margin. Fultura superior a slightly arched, transverse 
bar between upper internal angles of valvae, with a short, blunt projection at 
middle. Fultura inferior subcordate, haired laterad at upper margin; caulis short, 
joined to aedeagus slightly before its middle. Aedeagus slightly curved; cornuti few, 
rather short and thick, deciduous. 

Female genitalia (Figs. 4, 5): Papillae anales rotated 90° outward, forming a 
blade-like ovipositor, hidden below eighth abdominal tergite which is enlarged, 
strongly sclerotized, forming a helmet-shaped cover that extends caudad beyond 
genitalic opening. Sinus vaginalis wide; sterigma represented by lamella post- 
vaginalis only, broad in middle, narrowed laterad. Antrum elongate, slightly 
sclerotized, rotundate cephalad, with two lateral colliculi caudad. Ductus bursae 
coincident with antrum. Bursa copulatrix with corpus ovate and cervix bursae 
long; cestum narrow, longitudinal, band-like, dilated at corpus bursae, not reaching 
antrum. Ductus seminalis opening into antrum. Signum a strong, curved thorn; its 
basal sclerotization formed as a serrate, scobinate plate; no capitulum. 

Final instar larva: Sharing Archipini characters as defined by MacKay (1962: 
29). Head: adfrontal sutures sinuate and adfrontals not attenuated posteriorly. 
Thorax: meso- and metathorax with SV group bearing one seta; dorsal pinacula 
not elongated posteriorly; SD: dorsal to SD; rather than anterodorsal. Abdomen: 


122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Dz pinacula on anterior segments with mesal margin below lateral margin of D,; 
pinacula; SV groups on segments 1, 2, 7, 8, 9 with 3, 3, 2, 2, 2 setae; D,’s on 
anal shield distinctly closer to corresponding SD,’s than to each other; crotchets 
variably biordinal, 34-44 on abdominal, 28-36 on anal proleg; anal fork well de- 
veloped, 3-8 tines. (Based on examination of larvae from California and the de- 
scription given by Heinrichs (1971) of Kansas specimens. ) 

Remarks: This genus evidently is a New World derivative of Choristo- 
neura Lederer, from which it differs in having a smoother scaled head, 
the antennae not serrated in male, shorter, broader labial palpi, and a 
slightly longer basal fork of the forewing vein Ay,3. The male genitalia 
of Cudonigera have a broader tegumen with large shoulders; the uncus 
is shorter and broader than in all known Choristoneura species. The 
gnathos is differently shaped; its lateral arms are shorter and broader, 
with the middle process narrower and longer. The valvae are shorter 
than in Choristoneura. The female genitalia are unique, with a hyper- 
trophic development of the eighth abdominal tergite which forms a kind 
of helmet-shaped cover over coriaceous papillae anales (cudo, a helmet 
made of skin; -gera, bearing). 

The distinctive features, particularly the rudimentary socii, elongate- 
narrow joined portion of the gnathos, and blade-like ovipositor situated 
beneath the hood-like development of the tergite, are characters shared 
by no species of Choristoneura. The short valvae and the larval char- 
acters suggest a reiationship with Group 1 of MacKay (1962: 36), in- 
cluding C. conflictana (Wlk.) and C. fractivittana (Clem.), rather than 
with the conifer-feeding Choristoneura (fumiferana and related species ). 


Cudonigera houstonana (Grote), new combination 


Tortrix houstonana Grote, 1873, Bull. Buffalo Soc. Nat. Sci.; 15. Fernald, 1882, 
Trans. Amer. Ent. Soc. 10: 17. Grote, 1882, New check list of North Amer. 
moths: 58. Fernald, “1902’/1903/, Bull. U.S. Natl. Mus. 52: 482. 

Cacoecia houstonana; Meyrick, 1912, in Wagner, Lepid. Catalogus 10: 21; 1913, 
in Wytsman, Genera Insectorum, fasc. 149: 25. Barnes & McDunnough, 1917, 
Check list Lepid. Boreal Amer.: 177. 

Archips houstonana; McDunnough, 1939, Mem. Southern Calif. Acad. Sci. 2: 56. 

Choristoneura houstonana; Freeman, 1958, Can. Ent., 90, suppl. 7: 38. Powell, 
1964, U. Calif. Publ. Ent. 32: 185 (biol.). Heinrichs & Thompson, 1968, 
Can. Ent. 100: 750 (biol. ). 

Lozotaenia retana Walsingham, 1879, Illustr. typical specimens Lepid. Heterocera 
Aes 13 

Tortrix retana; Grote, 1881, Papilio 1: 9 (synonymy). 


Types: of houstonana, Texas; location of type specimen unknown; of 
retana, male, Bosque County, Texas, October 5, 1874 (Belfrage), in 
British Museum (Natural History ). 

Taxonomic discussion: Aspects of the geographical variation have 
been discussed elsewhere (Powell, 1964: 186). Populations referred to 


VoLUME 31, NUMBER 2 12: 


Oo 


the name houstonana occur in Massachusetts, Kansas and are widely 
scattered in the western United States, associated with the island-like 
distribution of the larval foodplant, Juniperus. As noted, considerable di- 
versity in size, forewing pattern and hindwing color exists, in part repre- 
sented by samples in collections that are too fragmentary to permit ade- 
quate assessment. Presumably the color variation is related to cryptic 
resemblance of the resting moths on their coniferous hosts, but degrees 
of reproductive isolation may have been attained among some popula- 
tions. Morphological differences were not observed during our investi- 
gations (genitalia preparations examined; 9 4, Calif., Colo., Texas, 


Mass.; 4 2, Calif., Texas). 


LITERATURE CITED 


DraxonorF, A. 1966. Nicholas Sergeevich Obraztsov, 1906-1966. J. Lepid. Soc. 
20: 255-266. 

Hetnricus, E. A. 1971. External morphology of larvae of Choristoneura hou- 
stonana (Lepidoptera: Tortricidae). Can. Ent. 103: 12-17. 

MacKay, M. R. 1962. Larvae of the North American Tortricinae (Lepidoptera: 
Tortricidae). Can. Ent. Suppl. 28, 177 p. 

Oxsraztsov, N. S. 1954-1957. Die Gattungen der Palaearktischen Tortricidae I. 
Unterfamilien Tortricinae und Sparganothinae. Tijd. voor Ent. 97: 141-231; 
98: 147-228; 99: 107-154; 100: 309-347. 

PowEL., J. A. 1964. Biological and taxonomic studies on Tortricine moths, with 
reference to the species in California. U. Calif. Publ. Ent. 32: 1-317. 


124 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


THE STATUS OF THE GLYPHIPTERIGIDAE AND A 
REASSESSMENT OF RELATIONSHIPS IN YPONOMEUTOID 
FAMILIES AND DITRYSIAN SUPERFAMILIES! 


Joun B. HEppNER* 


Department of Entomology and Nematology, IFAS, 
University of Florida, Gainesville, Florida 32611 


Current studies of the North American Glyphipterigidae have revealed 
major fundamental morphological and behavioral characters which dem- 
onstrate that the inclusion of the choreutid and glyphipterigid groups 
within a single family is untenable. The discordant characters involved 
have been shown in the past by other workers to be so fundamentally 
and evolutionarily conservative in Lepidoptera phylogeny that it is not 
even possible to consider the two groups to have evolved within the 
same superfamily. 

Glyphipterigid moths have long been considered of unusual interest 
because of apparent affinities to the Yponomeutidae and the Sesiidae, as 
well as to the Tortricidae. Most early workers considered them as dis- 
tinct groups: the choreutids were placed with the tortricids and the 
glyphipterigids sensw stricto were placed among the tineoid moths. 
This segregation was rarely altered until Meyrick (1914) combined them 
into one family. Meyrick’s classification was based largely on general 
facies—the two groups share a number of superficial characters—and 
not fundamental relationships. He also relied strongly on wing venation 
and did not use genitalia, internal morphology or larval characters. He 
formed a conglomeration of what now are no less than nine distinct 
families in several superfamilies, although he realized the true affinities 
of many of the included genera in later years. Current revisionary studies 
on the choreutids and glyphipterigids, using modern systematic tech- 
niques, are revealing the true affinities of these moths. The results of 
these studies to date have confirmed the polyphyletic nature of the 
Glyphipterigidae sensu lato, first indicated by Brock (1967). 


Glyphipterigid Discordancies 


Brock (1971) revealed certain previously unused characters of ditry- 
sian internal morphology of which the sternal abdomino-thoracic artic- 
ulation provides a significant character for Lepidoptera phylogeny and 


1 Florida Agricultural Experiment Station Journal Series No. 341. 

* Research Associate, Florida State Collection of Arthropods. Mail address: Department of 
EneOlsey, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 

560. 


VoLuME 31, NuMBER 2 125 


the affinities of families. Whereas most genitalic characters are evolu- 
tionarily plastic at the species level in most groups, due to the selective 
pressures for reproductive isolation, it is clear that characters not likely 
to be involved in active selection should remain relatively stable and, 
consequently, useful in assessing the relationships of higher categories. 
The abdominal articulation in Lepidoptera appears to be such a stable 
character.® 

Two types of sternal articulation are found in adult Ditrysia: the 
Tineoidea type, having elongated sternal rods internally in the second 
sternal sclerite and with apodemal projections into the thoracic lumen, 
and the Tortricoidea type, having simple apodemes. There are minor 
variations in these types, but only the two major conformations of abdom- 
inal articulation are found in Lepidoptera. Having examined 16 genera 
of choreutids and 9 genera of glyphipterigids, including all 50+ Nearctic 
species assignable to these two groups, and many species of Pantropic 
and Palearctic origins, I have found no discrepancy in the abdominal 
articulation of any in terms of assignment to either group. All the 
choreutids have tortricoid apodemes, and all the glyphiptergids have 
tineoid rods. This articulation discordancy, consequently, indicates that 
the two groups have not evolved from a recent common ancestor. 

Another lepidopteran character considered evolutionarily conserva- 
tive at the family level is the chaetotaxy of larvae, with particular 
interest here involving the lateral pre-spiracular setal group of the larval 
prothorax (Werner, 1958; MacKay, 1963; Peterson, 1965; Common, 1975). 
The glyphipterigid sensu stricto larvae have a bisetose pre-spiracular 
setal group on the prothorax. The choreutids have a trisetose pre- 
spiracular setal group. The polyphyly of the Glyphipterigidae sensu 
lato is here again demonstrated by a character used in the Lepidoptera. 

A third fundamental character useful in the higher classification of 
Lepidoptera is pupal behavior at adult ecdysis and again the two groups 
show no recent common ancestry. The glyphipterigids do not protrude 
the pupa at adult ecdysis, and the choreutids do protrude the pupa. The 
protrusion or non-protrusion behavior is characteristic of superfamilies in 
the Ditrysia. It should be noted that this behavior involves the presence 
or absence of genetic components that form the pupal exterior spination, 
which is usually necessary for the pupa to be able to protrude from the 
cocoon. Some yponomeutids protrude only the head. 

Table 1 summarizes the three fundamental characters noted above 
for each of the families and superfamilies comprising the microlepidop- 

’The sexual dimorphism in abdominal articulation noted by Hodges (1974) in certain 


Oecophoridae is one of degree only and while some tortricoid tendencies occur, these do not 
form a simple apodemal articulation but retain the tineoid rod conformation. 


126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Characters of Ditrysian Microlepidoptera. 


Larval 
Abdominal Superfamily L-group  Protruded 

Articulation Setae upa 
tineoid rods Tineoidea (2 in Scardia) 3 yes 
tineoid rods Gelechioidea 3 no 
tineoid rods Copromorphoidea - Copromorphidae WA, no 
Carposinidae 2; no 

E,permeniidae 2 no 

Glyphipterigidae uy no 

tineoid rods Yponomeutoidea - Douglasiidae 8 yes 
Argyresthiidae 3 yes 

Yponomeutidae 8 yes 

Plutellidae 8 yes 

Acrolepiidae 8 yes 

Heliodinidae 8 yes 

apodemes Sesioidea - Immidae 3 yes 
Sesiidae 3 yes 

Choreutidae 3 yes 

apodemes Tortricoidea 8 yes 
apodemes Cossoidea 3 yes 
apodemes Castnioidea 3 yes 
apodemes Zy gaenoidea 3 yes 
apodemes Pyraloidea 2 yes 


terous Ditrysia. Taken together the three characters provide strong evi- 
dence that the glyphipterigids and choreutids have not evolved from a 
recent common ancestor and, consequently, are distinct families be- 
longing to different superfamilies in our present concept of these higher 
categories. 


Affinities and Rearrangements 


Rearrangements I propose for a new classification of the lower Ditrysia 
are as follows: Glyphipterigidae sensu stricto and Epermeniidae trans- 
ferred from Yponomeutoidea to Copromorphoidea, and Choreutidae and 
Sesiidae restricted to Sesioidea, with the Copromorphoidea being shifted 
between the Gelechioidea and the Yponomeutoidea, while the Tortri- 
coidea are placed after the Sesioidea in a linear arrangement altered from 
that proposed by Common (1970). There are also two Nearctic genera 
placed in Glyphipterigidae sensu lato that will be transferred to Copro- 
morphidae in a future paper: one of the genera was already assigned 
to Carposinidae by MacKay (1972) based on larval characters, but larval 
differences from true carposinids apparently apply to Copromorphidae. 

Meyrick (1928) was the first to combine Copromorphidae and Car- 
posinidae as a new superfamily, the Copromorphoidea (plus Alucitidae), 
but the Glyphipterigidae and Epermeniidae were not associated with the 


VoLUuME 31, NUMBER 2 127 


superfamily. The discordances noted above show that the characters 
of the glyphipterigids sensu stricto conform to Copromorphoidea. Their 
naked haustellum and bisetose larva excludes them from the Gelechioidea. 
Their bisetose larva and the non-protruded pupa excludes them from the 
Yponomeutoidea. 

The Epermeniidae have the same three major character states as the 
glyphipterigids, which also places the family outside of Gelechioidea and 
Yponomeutoidea. There is some doubt about the bisetose pre-spiracular 
condition of epermeniid larvae since MacKay (1972) noted larvae of an 
Epermenia species to be bisetose, but Forbes (1923) noted another to be 
trisetose. Common (1970) states that epermeniid larvae are bisetose. 
My own examination of reared epermeniid larvae in the National Mu- 
seum of Natural History, Smithsonian Institution, Washington, D.C., pro- 
duced only bisetose larvae. It may be possible that both bi- and tri- 
setose larvae occur in the family as in Tineidae where Scardia larvae 
are bisetose (Hinton, 1956) while other tineids are trisetose. The bi- 
setose condition appears to be an apomorphic development prevalent in 
endophagous larvae, although as seen in Table 1, this character is gen- 
erally conservative enough evolutionarily to serve as a useful character for 
higher classification. Not all endophagous larvae, however, are bisetose; 
for example, the trisetose endophagous Sesiidae (MacKay, 1968) among 
others. 

The epermeniids are placed between Carposinidae and Glyphipterigi- 
dae because of genitalic features showing affinities to Carposinidae, e.g., 
the uncus, and because of advanced wing venation and other characters 
showing a close relationship to the glyphipterigids. Some epermeniids 
superticially resemble glyphipterigids, for example, the Palearctic Eper- 
menia pontificella Hiibner. As with the superficial resemblance of some 
choreutids with glyphipterigids, the Epermeniidae also have wing mac- 
ulation that could be the result of convergent adaptive strategies as 
diurnal moths, although it is unclear whether all epermeniids are diurnal. 
Choreutids and glyphipterigids, as also some similar heliodinids, are di- 
urnal in adult activity. 

A distinctive feature of the Copromorphidae and the Carposinidae is 
the anal pectin of the hind wings, but not all species in these families 
have this feature (Common, 1970). The raised scale tufts of the fore- 
wings also are not found in all species of the families, which otherwise 
is a distinctive character for both families. Both characters would appear 
to be apomorphic in these two families and, thus, the lack of either in 
epermeniids and glyphipterigids should not exclude them trom the super- 
family. Some Gelechiidae and Oecophoridae (e.g., Tonica spp.) also 
have raised scale tufts on the forewings. The Epermeniidae often have 


128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


a scale tuft on the dorsal forewing margin that may indicate an affinity 
with the raised scale tufts of copromorphids and carposinids, although 
it may be a peculiarity of many epermeniids. 

An interesting cohesive character of the Copromorphoidea is the en- 
larged spiracles of the prothorax and abdominal segment 8. Spiracles 
of the 8th abdominal segment also are more dorso-caudally positioned 
than is usual in Lepidoptera larvae (Common, 1970). While spiracle 
size is close to normal in Carposinidae (MacKay, 1972), a striking en- 
hancement of this character has been illustrated and described by Moriuti 
(1960) and Kodama (1961) in the larva of the Japanese species, Gly- 
phipterix semiflavana Issiki. The larvae have the spiracles of the 8th 
abdominal segment not only dorso-caudally positioned but elevated on 
what look like scoli. The larva of a new Glyphipterix species from Flor- 
ida has protruding and enlarged spiracles as in the Japanese species. 
Larvae of the glyphipterigid genus Machlotica also have this unusual 
spiracle enlargement. I have examined reared Epermenia larvae, and 
these also showed the protruding spiracles. I have not seen larvae of 
Copromorphidae and follow Common (1970) in his notes for the family. 
The character may be apomorphic in endophagous larvae, having some 
unknown adaptive function. MacKay (1959) noted that tortricid larvae 
with more caudally positioned spiracles of the 8th abdominal segment 
invariably were borers, although this apparently does not hold for sesiid 
larvae (MacKay, 1968). Inasmuch as all Copromorphoidea larvae known 
thus far have enlarged spiracles to greater or lesser degree, but more 
than usual for Lepidoptera larvae, it appears to indicate a common an- 
cestor for the four families. The unusual spiracle development of eper- 
meniid and glyphipterigid larvae indicates that these two families are 
closely related. MacKay (1972) also noted other chaetotaxic characters 
which show affinities of epermeniids to Carposinidae. 

The Copromorphoidea, as arranged in Table 1, have a reduction in 
wing venation from Copromorphidae to Epermeniidae, while retaining 
a chorda in Glyphipterigidae and vestiges thereof in Epermeniidae. The 
presence of the chorda has in the past retained the glyphipterigids and 
epermeniids in the Yponomeutoidea—the same can be noted for the 
choreutids—but the wing venation of these two families can be accepted 
as specializations within the Copromorphoidea. 

The Douglasiidae are an anomalous family with little known about 
their biologies. The larvae are stated to be trisetose (Common, 1970), 
which I have confirmed in larvae of Tinagma balteolella (Fisher von 
Roeslerstamm). The pupa apparently is protruded at adult ecdysis, al- 
though this is unclear from published information. I retain them in 
Yponomeutoidea pending further investigation on their immature stages. 


VoLUME 31, NuMBER 2 129 


The family appears to be the most primitive yponomeutoid in relation 
to such characters as wing venation, a reduced uncus, and no socii. The 
remaining yponomeutoid families appear to form a monophyletic super- 
family and require no further notation in the context of this paper. The 
most recent research of European and Japanese workers is followed by 
the separation of Argyresthiidae, Plutellidae, and Acrolepiidae from 
Yponomeutidae. The superfamily progresses to the Heliodinidae, which 
would appear to be the most specialized yponomeutoid family. 

The Choreutidae and Sesiidae have usually been considered in the 
Yponomeutoidea, especially due to their similar wing venation, which is 
also very similar between the two families, although very specialized in 
the sesiids. In fact, in the “choreutid” genus Sagalassa the two families 
nearly merge, with many species in the genus having hyaline wing areas 
as in Sesiidae. Larvae in at least one Neotropical species, Sagalassa 
olivacea (Busck), appear to be indistinguishable from true sesiid larvae 
(Duckworth & Eichlin, pers. comm.). The naked haustellum and other 
characters of Sagalassa indicate a close relation to Sesiidae, but with af- 
finities to Imma. Since the Sesiidae also have tortricoid apodemes at 
the abdominal articulation and are otherwise closely related to the 
Choreutidae, although extremely specialized, I follow Brock (1971) in 
assigning both to a separate superfamily, the Sesioidea. Although very 
specialized, the Sesiidae retain ancestral features (e.g., genitalic charac- 
ters) that allow their placement before Choreutidae in a linear arrange- 
ment of primitive to advanced. 

The Pantropical genus Imma, in the past included in the glyphi- 
pterigids, may be assigned to Immidae, new family (type-genus: [mma 
Walker [1859] ), the most primitive family of the Sesioidea. A thorough 
revision for a clarification of the true affinities of Immidae is needed. 

Forster (1954) was the first to combine the Sesiidae and Glyphip- 
terigidae into one superfamily which he called Glyphipterygoidea, but 
he included the Glyphipterigidae sensu stricto. Meyrick (1928) had 
anticipated Forster by segregating the two families from Yponomeutoidea 
to Glyphipterygoidea—which was not followed by other workers—but 
he included Heliodinidae and Heliozelidae. Turner (1947) also had 
relationships mixed among several families, yet it is noteworthy that he 
seems to have been the first to note a possible relationship between 
Sesiidae, Glyphipterigidae sensu lato, and the Copromorphidae. Nicu- 
lescu (1964) also noted a relationship to Copromorphidae. Brock (1971) 
used the name Aegerioidea, but since Sesiidae is senior to Aegeriidae 
through the relative genus pertaining to each name, Sesioidea is the cor- 
rect superfamily name. 

The Sesioidea remain distinct from the Tortricoidea through larval 


130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


characters, wing venation, labial palpi, head vestiture, and genitalic 
features. Among these characters in the Choreutidae are many tortricoid 
affinities. The largely tropical genus Hilarographa, heretotore considered 
choreutid, has remarkable genitalic resemblance to Chlidanotinae tortri- 
cids from Australia and New Guinea (Diakonoff, XV International Con- 
gress of Entomology, August 1976, Washington, D.C.) and, together with 
the related Idiothauma and Mictopsichia, will be transferred to Tortri- 
cidae in the near future. 

The Choreutidae have a peculiar feature in their scaled haustellum, 
which is characteristic of gelechioids but not of sesiids or tortricoids 
(the three genera to be transferred to Tortricidae have naked haustel- 
lums, as do Sagalassa and Imma species). The state of haustellum scal- 
ing is usually useful at the superfamily level in Lepidoptera classification 
in terms of cohesive groups of families either having a scaled or a naked 
haustellum. As with other characters, isolated groups are found not to 
conform to some major character while otherwise having all the charac- 
teristics of the particular taxa they are related to. I believe the situation 
is the same with choreutids in their character complex between Sesiidae, 
Yponomeutidae, and Tortricidae. The Pyralidae also are the only pyra- 
loid family having a scaled haustellum. As with the choreutids, the 
haustellum scaling appears to represent the retention of an ancestral 
character to some related group (e.g., choreutid relatives in the gele- 
chioids? ) or an apomorphy. 

Figure | illustrates my understanding of the evolution of the Ditrysia 
by evidence presented herein and arranged linearly, but I do not wish 
to discuss all the details involved as this has been extensively covered 
by other authors (see Common, 1975). The taxa shown in the figure 
have lineage heights in relation to the general amount of evolutionary 
change (as a rate vector) that the group has undergone from ancestral 
forms: for example, Yponomeutidae evolved from an ancestral ypono- 
meutoid but at a slower rate than Douglasiidae and, thus, the latter 
family is placed on a higher rate vector although the douglasiids have 
other characters which indicate they are more primitive yponomeutoids. 


Superfamily Relationships 

For a linear arrangement of the ditrysian microlepidopterous super- 
families, I follow Common (1970) as modified by the studies of Brock 
(1971). Thus, the Tineoidea and Gelechioidea are considered the most 
primitive due to their articulation and wing venation. However, such 
an understanding of their phylogenetic ancestral relationships does not 
preclude the many specializations found within the Gelechioidea, being 
a result of differential rates of evolution in the various included families. 


VoLuME 31, NuMBER 2 13] 


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primitive Lepidoptera 


Amount of Divergence from Related Taxa (linear arrancement ) 


Fig. 1. Evolution of Ditrysia. 


In contrast to Common (1970), I place the Copromorphoidea after 
Gelechioidea due to their tineoid abdominal articulation and the non- 
protruding pupal behavior, which is not tortricoid. Copromorphids have 
an abdominal articulation resembling the apodemal type, yet retain the 
tineoid sternal rods: the apodemal resemblance is actually enhanced due 
to the stoutness of the rods. The trisetose larvae and protruding pupal 


132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


behavior of Yponomeutoidea indicates closer affinities to Sesioidea than 
to Gelechioidea, thus, placing them after Copromorphoidea. 

The apodemal nature of the abdominal articulation of Tortricoidea is 
a derived condition and demonstrates closer affinities to the higher 
Ditrysia, which all have the apodemal articulation, than to the Tineoidea, 
as followed by Common (1970). Larval studies by MacKay (1959) have 
also shown that most tortricids are more advanced than Tineoidea. As 
noted above, the mixed character complexes of the Sesioidea indicate 
ancestral relationships to both Yponomeutoidea and Tortricoidea, placing 
them in the middle in a linear arrangement. The Cossoidea I consider 
having evolved at a very slow rate of evolution in relation to the related 
Tortricoidea and, while more primitive in many ways compared to 
tortricoids, they are more advanced than ancestral tortricoids, thus al- 
lowing a more convenient placement after Tortricoidea for a linear ar- 
rangement. The remaining superfamilies are arranged after Common 
(1970) except for the Alucitidae. The alucitids were placed in the 
Copromorphoidea by Meyrick (1928) and Common (1970) but the tor- 
tricoid abdominal articulation would better place them in the Pyraloidea 
(Brock, 1971), which also have bisetose larvae and non-protruding 


pupae. 
CONCLUSIONS 


The long maintained assimilation of the Glyphipterigidae and Choreu- 
tidae as one family was due to their overall resemblance. Evaluation of 
more fundamental characters, as noted above, has elucidated the dis- 
cordances in considering the two groups as one family in relation to the 
desire to maintain only monophyletic groupings of related taxa. Actually, 
the two groups evolved from distinct ancestral lines and must be con- 
sidered distinct families. 

Evaluation of related families indicates that the Epermeniidae are 
much more closely related to Glyphipterigidae than previously con- 
sidered, with both showing common ancestry with the Copromorphidae 
and Carposinidae. Thus, the four families are here considered in one 
superfamily, the Copromorphoidea. Fundamental characters also dis- 
tinguish the Sesiidae and Choreutidae as Sesioidea (together, probably, 
also with Immidae), not Yponomeutoidea, and their placement be- 
tween the yponomeutoids and the tortricoids appears sound. Immidae 
will be discussed further in a forthcoming paper. 


ACKNOWLEDGMENTS 


For discussions of the conclusions presented in this paper and review 
of the manuscript, I wish to thank J. F. G. Clarke (Smithsonian Institu- 


VoLuME 31, NuMBER 2 133 


tion), D. R. Davis (Smithsonian Institution), A. Diakonoff ( Rijksmuseum 
van Natuurlijke Historie, Leiden), W. D. Duckworth (Smithsonian Insti- 
tution), T. D. Eichlin (California Dept. of Agriculture, Sacramento ), 
D. H. Habeck (University of Florida), R. W. Hodges (Systematic En- 
tomology Laboratory, USDA), N. P. Kristensen (University of Copen- 
hagen), P. A. Opler (Office of Endangered Species, U.S. Dept. of In- 
terior), J. A. Powell (University of California, Berkeley), and J. D. 
Bradley, G. S. Robinson, and K. Sattler of the British Museum ( Natural 
History), London. The Department of Entomology and Nematology, 
Institute of Food and Agricultural Sciences, University of Florida, pro- 
vided support for my studies of the Nearctic Choreutidae and Glyphi- 
pterigidae as part of my doctoral program, under the guidance of D. H. 
Habeck. I wish to also thank W. D. Duckworth for providing facilities 
and the Smithsonian Institution for providing a predoctoral fellowship for 
research at the National Museum of Natural History, Washington, D.C. 
Funds provided, in part, by the Smithsonian Institution and the Na- 
tional Science Foundation (Dissertation Improvement Grant DEB 76- 
12550) also materially enhanced research presented herein by supporting 
an extended stay at the British Museum (Natural History). 


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—. 1971. A contribution towards an understanding of the morphology and 
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Common, I. F. B. 1970. Lepidoptera (moths and butterflies). Pp. 765-866, in 
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. 1975. Evolution and classification of the Lepidoptera. Ann. Rev. Ent. 
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Forses, W. T. M. 1923. The Lepidoptera of New York and neighboring states. 
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Forster, W. 1954. Biologie der Schmetterlinge. In W. Forster and T. A. Wohl- 
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KopaMa, T. 1961. The larvae of Glyphipterygidae (Lepidoptera) in Japan (1). 
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. 1963. Evolution and adaptation of larval characters in the Tortricoidea. 

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1968. The North American Aegeriidae (Lepidoptera): a revision based 

on late-instar larvae. Ent. Soc. Can. Mem. 58: 1-112. 


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. 1972. Larval sketches of some microlepidoptera, chiefly North American. 
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Meyrick, E. 1914. Lepidoptera Heterocera. Fam. Glyphipterygidae. In P. Wyts- 
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1928. <A revised handbook of British Lepidoptera. [1968 facsimile]. 
Classey, Hampton. 941 p. 

Morivutl, S. 1960. Description of the larva and pupa of Glyphipteryx semiflavana 
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Peterson, A. 1965. Larvae of insects. An introduction to Nearctic species. Part 
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Turner, A. J. 1947. A review of the phylogeny and classification of the Lepidop- 
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WerneER, K. 1958. Die Larvalsystematik einiger Kleinschmetterlingsfamilien. ( Hy- 
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dae). Abh Larvalsyst. Insekten 2: 1-145. 


AN “ALBINIC” PIERIS SISYMBRII (PIERIDAE) FROM 
THE CALIFORNIA SIERRAS 


“Albinic” or “depigmentized” forms largely or wholly lacking melanin pigment 
from the wings are known in several pierid butterflies. Within the genus Pieris a 
weakly melanized form is known from P. protodice Bdv. & LeC. (Shapiro 1970, 
Wasmann J. Biol. 28: 245-256) and Gardiner (1962, Ent. Gaz. 13: 97-100; 1963, 
J. Res. Lep. 2: 127-136) has reported a form from P. brassicae L. in which the 
normally black scales lack pigment altogether, producing a translucent “shadow” 
pattern. In both of these cases the genetics is known. Crowe (1967, J. Lepid. 
Soc. 21: 121) reported a female P. sisymbrii Bdv. from Harney Co., Oregon which 
seems to resemble Gardiner’s form of P. brassicae in totally lacking melanin on the 
wings. Although the accompanying photograph does not show a “shadow” pattern, 
it is mentioned in the text. On 23 May 1975 a very similar male with “shadow” pat- 
tern was taken flying among normal individuals on Washington Road, off state High- 
way 20 in Nevada Co., California. As in Crowe’s specimen, the normally dark wing- 
veins contrast strongly with the ground color and the body, legs and antennae are 
normally pigmented. This male was kept alive for two days but no virgin females were 
available and I could not induce wild females to mate. The Washington Road popu- 
lation is unusual in that it is an isolated colony on the highest-elevation outcrop of 
serpentine soil (elev. 5000 ft) in the central west-slope Sierra. The vegetation on 
this atypical site is digger pine-manzanita-scrub oak, contrasting strongly with 
nearby stands of mixed montane coniferous forest on non-serpentine soils; the 
nearest known sisymbrii colony is seven miles away. This is the first aberrant in- 
dividual I have seen among about 750 wild P. sisymbrii in about 20 California 
populations. 


Arruur M. SHapriro, Department of Zoology, University of California, Davis, 
California 95616. 


VoLUME 31, NuMBER 2 135 


TWO NEW SPECIES OF PETROVA MOTHS FROM PINE 
IN SOUTHEAST ASIA (TORTRICIDAE, OLETHREUTINAE ) 


WILLIAM E.. MILLER 


USDA Forest Service, North Central Forest Experiment Station, 
Folwell Avenue, St. Paul, Minnesota 55108 


The genus Petrova is holarctic in distribution and its known hosts are 
conifers, pines in particular. The larvae are usually shoot borers. For- 
est entomological investigations in India and Thailand have revealed 
two previously unknown members of Petrova whose descriptions follow. 
One of them has adapted to an introduced North American pine in 
southeast Asia. 

In this paper the letter n denotes number of specimens underlying an 
observation or conclusion. Color names accompanied by a parenthetical 
number indicate standard colors in the system of Kelly & Judd (1965). 

For specimens and information I thank Chavewan Hutacherern, Thai- 
land Royal Forest Department, Bangkok; M. J. Chacko and T. Sankaran, 
Commonwealth Institute of Biological Control (CIBC), Indian Station, 
Bangalore; and Fujio Kobayashi, Government Forest Experiment Sta- 
tion, Momoyama, Kyoto, Japan. 


Petrova khasiensis Miller, new species 
Figs. la—c 


Male. (2 n) Head: Labial palpus short, second segment 0.9 eye diameter, 
apical segment 0.4 length of second segment; scaling spreading, partly obscuring 
apical segment, orange brown, darker toward base. Scaling of front and crown 
lighter hued than labial palpus. Thorax: Dorsal scaling, including tegulum, shining 
brown, some scales white tipped. Venter and prothoracic and mesothoracic legs 
shining brown, metathoracic legs lighter, tibiae and tarsi white banded. Forewing: 
Length 7.0-8.5 mm, 2.6 width; apex acute, termen slightly convex, tornal angle 
distinct. Upper side (Fig. la) with indistinct basal patch, shining gray from base 
to middle with sprinkling of strong brown, strong brown beyond middle except for 
four paired white marks on costa, which continue irregularly and intermittently to 
dorsum as silver and white bands and form a rudimentary ocellus; about 15 black 
scales sprinkled in ocellar area. Fringe gray, tinged with orange brown. Underside 
gray, mottled with white in costal area. Hindwing: Wider than forewing, termen 
concave, curved to dorsum, tornal angle not discernible. Shining gray throughout. 
Fringe pale gray. Underside gray, lighter than underside of forewing. Abdomen: 
Dorsal scaling shining light brown, ventral shining white. Genitalia (2 n) with 
valva moderately constricted, clasper a broad ridge (Fig. 1b); uncus slightly de- 
veloped. 

Female. (6 n) As described for male with following exceptions. Forewing: 
Length 8.0-9.5 mm, 2.4xX width. Abdomen: Genitalia (4 n) with sterigma a 
shield-like plate (Fig. 1c), a sclerotized incomplete ring near middle of ductus, two 
unequal sized thorn-like signa. Posterior apophyses shorter than anterior. 

Types. Holotype, male (Fig. la) and allotype, female: Upper Shillong, Assam 


136 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


\ 
\ 
| 
! 
) 
| 
| 
| 
! 


Fig. 1. Petrova khasiensis: a. holotype, male; b. male valva; c. female sterigma. 

Fig. 2. Petrova salweenesis: a. holotype, male; b. male valva; c. female sterigma. 

Fig. 3. Petrova cristata: a. female from near Kyoto, Japan, forewing length 8.0 
mm; b. male valva; c. female sterigma. 


(now Meghalaya), March 1963, CIBC, male genitalia slide 173 and female genitalia 
slide 170 (R. B. Moore) and wing slide II.24.76 (W. E. Miller), both types in US 
National Museum of Natural History. Two paratypes: same data as holotype ex- 
cept male genitalia slide 376301 (W. E. Miller) in US National Museum of Natural 
History, and female genitalia slide 412763 (W. E. Miller) in collection of CIBC 
Indian Station, Bangalore. Four additional females examined, one with same data 
as holotype, two with same except August 1968. 

Host. Pinus khasya Royle. 

Geographic distribution. Khasi Hills of northeastern India. 


Petrova salweenensis Miller, new species 
Figs. 2a—c 


Male. (4 n). As in khasiensis with following exceptions. Thorax: Dorsal scal- 
ing lighter and sprinkled with strong brown. Forewing: Length 6.0-7.5 mm; upper 
side (Fig. 2a) more heavily sprinkled with strong brown from base to middle, basal 
patch distinct. Abdomen: Genitalia (3 n) with valva greatly constricted (Fig. 2b), 
uncus moderately developed. 


VoLUME 31, NUMBER 2 137 


Female. (5 n). As described for male with following exceptions. Forewing: 
Length 6.0-8.5 mm. Abdomen: Genitalia (5 n) as in khasiensis except sterigma 
more elongate (Fig. 2c). 

Types. Holotype, male (Fig. 2a): Baw Luang, Thailand, March 20, 1972, 
D. Chaiglom; allotype, female, Chiangmai, Thailand, March 10, 1975, Chavewan, 
female genitalia slide 22476a (W. E. Miller), both types in US National Museum of 
Natural History. Two paratypes: Baw Luang, Thailand, March 1976, Chavewan, 
male genitalia slide 376305, and female genitalia slide 376304 (W. E. Miller) in 
US National Museum and Thailand Royal Forest Department, Bangkok, respectively. 
Five additional specimens examined, two males and three females, same data as 


paratypes. 

Hosts. Pinus khasya, P. merkusii De Vries, and P. caribaea Morelet, the last in- 
troduced from North America. 

Geographic distribution. Northwestern Thailand. 


DIscussION 


The closest known occurrences of Petrova khasiensis and P. salweenen- 
sis to one another are 960 air km apart. Both species are similar to P. 
cristata (Walsingham) (Figs. 3a—c; illustrated by Issiki (1957) in color) 
whose nearest known occurrence is Canton, China (Obraztsov, 1964), 
1,500 air km northeast of Chiangmai. Both new species were compared 
with P. cristata reared from Pinus thunbergii Parl. near Kyoto, Japan 
(5n). The main anatomical differences are summarized as follows: 


Item khasiensis salweenensis cristata 


Forewing (5-9 n): 


Length, mm 7.0-9.5 6.0-8.5 6.0-8.0 
Ground color Strong brown Strong brown Dark brown 
(55) (5) (59) 
Basal patch Obscure Distinct Distinct 
Male genitalia (2-3 n): 

Ventral margin of Obtuse Right Right 
valval sacculus angular angular angular 
Valval constriction Moderate Great Great 
Uncus development Slight Moderate Slight 


Female genitalia (3-5 n): 
Width of sterigma > Length < Length < Length 


Strong brown is much lighter than dark brown. Color and other anatomi- 
cal differences between adult Petrova khasiensis and P. salweenensis are 
minor, but it is desirable for communication purposes to treat them as 
separate taxa unless future biological investigations prove them conspe- 
cific and reduce the weights of current diagnostic characters. 

Pine biogeography suggests how Petrova khasiensis and P. salweenensis 
may have evolved. The two species appear to be isolated from one an- 
other by disjunct distribution of the shared host, Pinus khasya (variously 
spelled such as kesiya and considered by some authors to be P. insularis 


138 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Endl.). Pinus khasya does not occur at low elevations and there are 
100-km gaps between its occurrences from the Khasi Hills of north- 
eastern India (range of Petrova khasiensis) to the highlands of Thailand 
(range of P. salweenensis) (Critchfield & Little, 1966; Mirov, 1967). 
Pinus merkusii and other pines likewise seem to be absent in these gaps. 
In Pleistocene times, the cool climate that allowed pines to migrate south- 
ward through Indochina (Mirov, 1967) must have allowed P. khasya 
to occur at low elevations, perhaps in continuous distribution. As the 
climate warmed at the end of the Pleistocene, P. khasya would have 
retreated from low elevations to form its fractured distribution pattern, 
thereby creating islands of Petrova that speciated. Under this hypothesis, 
related Petrova might occur in other subdivisions of the Pinus khasya 
range. 


LITERATURE CITED 


CritcHFIELD, W. B. & E. L. Lirrie, Jr. 1966. Geographic distribution of the 
pines of the world. U.S. Dept. Agric., For. Serv. Misc. Publ. 991, 97 p. 
Issrkr, S. 1957. Eucosmidae. P. 53-75. In Icones Heterocerorum Japonicorum in 
Coloribus Naturalibus, by T. Esaki and others. 318 p. Hoikusha, Osaka, Japan. 
(Japanese ). 

Keiiy, K. L. & D. B. Jupp. 1965. The ISCC-NBS method of designating colors 
and dictionary of color names. U.S. Dept. Comm. Natl. Bur. Stand. Circ. 553, 
158 p. 

Mirov, N. T. 1967. The genus Pinus. Ronald Press, New York. 602 p. 

Opsraztsov, N. S. 1964. Die Gattungen der palaearktischen Tortricidae. II. Die 
Untertamilie Olethreutinae, 5. Teil. Tijd. Entom. 107: 131-178. 


A RECORD OF URBANUS SIMPLICIUS (HESPERIIDAE ) FOR THE USA 


Tilden (1965, J. Lepid. Soc. 19: 53-55) summarized the differences between 
Urbanus simplicius (Stoll) and Urbanus procne (Plotz). He found that most if not 
all records of simplicius from the USA were erroneous, a result of confusion of that 
species with procne. 

I took a fresh male simplicius in Bentsen-Rio Grande Valley State Park, Hidalgo 
Co., Texas, on 13 April 1974. The specimen was collected at a large patch of 
thistle, Cirsium texanum Buckl. (Compositae), whose blossoms were attracting 
many skippers. Of the 35 species of Hesperiidae present, other interesting species 
were Urbanus doryssus Swainson, Astraptes anaphus annetta Evans, Aguna asander 
( Hewitson ), Cogia outis (Skinner), and C. hippalus (Edwards). These are apparently 
the first records from the Lower Rio Grande Valley for the two Cogia species. 

I wish to thank the Texas Parks and Wildlife Department for the issuance of a 
collecting permit for Bentsen-Rio Grande Valley State Park. 


Mike A, Rickarp, 4618 Holly, Bellaire, Texas 77401. 


VoLUME 31, NUMBER 2 139 


DATA SUGGESTING ABSENCE OF LINKAGE BETWEEN TWO 
LOCI IN THE MIMETIC BUTTERFLY HYPOLIMNAS BOLINA 
(NYMPHALIDAE) 


C. A. CLARKE AND P. M. SHEPPARD! 
Department of Genetics, University of Liverpool, England 


A large number of female forms of the polymorphic Batesian mimic 
Hypolimnas bolina (L.) have been described, but the situation is com- 
plicated because essentially similar phenotypes have been given dif- 
ferent names in different geographical areas. In fact, the forms can be 
described in terms of four basic phenotypes and their combinations, 
together with minor modifications of pattern. We have used for the 
sake of clarity the varietal names for the four main forms given by 
Poulton (1924). In the present paper we discuss the genetics of three 
of these, the mimic f. ewploeoides and two of the three nonmimetic 
forms, f. nerina and f. naresi. 

In a previous paper (Clarke & Sheppard, 1975) we showed that this 
polymorphism, sex-controlled to the female, is determined by two loci, 
one with two allelomorphs (E and e) and the other with three (P, P", 
and p). It is possible that the locus with three alleles consist of two very 
closely linked loci with epistatic interactions between the allelomorphs. 

The data used in the previous investigation gave a crossover value 
between the two loci E and P of 45.2%, which was not significantly dif- 
ferent from independent assortment. Since all the evidence, including 
that from Ephestia kuehniella Z. (Traut & Rathjens, 1973), suggests the 
absence of chiasmata in female Lepidoptera, and in our broods the 
double heterozygote was the female, it seemed unlikely that the two 
loci are on the same chromosome. However, despite the absence of 
chiasmata in E. kuehniella, Robinson (1971) reports a brood in which 
crossing over had apparently occurred in the female of this species, and 
it therefore seemed important to investigate the possible linkage between 
E and P further. The present paper reports five broods in which the 
progeny were all female (a phenomenon surprisingly common in FH. 
bolina (Clarke et al., 1975)). One of the backcross broods (14228) in 
which the female was a double heterozygote produced 126 offspring 
and therefore could give information on linkage. 


MATERIAL AND METHODS 
Genotypes and corresponding phenotypes. Clarke & Sheppard (1975) 
list all the genotypes and their corresponding phenotypes. One of these, 


1Ep. Nore: Professor P. M. Sheppard died 17 October 1976. His obituary will appear in the 
next issue. 


140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-4. Four female offspring of brood 14228. 1. f. euploeoides-nerina ( geno- 
type EeP"p), wing span 85 mm. 2. f. euploeoides (Eepp), wing span 89 mm. 3. 
f. nerina (eeP"p), wing span 95 mm. 4. f. naresi (eepp), wing span 78 mm. 


euploeoides-nerina (sometimes called aphrodite, Fig. 1) was synthesised 
on many occasions by crossing both homozygous and heterozygous eu- 
ploeoides (EEpp and Eepp, Fig. 2) with nerina, again when both homo- 
zygous and heterozygous (eeP”P” and eeP"p, Fig. 3). However, we 
never succeeded in showing that a wild euploeoides-nerina was in fact 
carrying both the allelomorph E and the allelomorph P”, rather than 
being of some as yet unknown genotype. 

In the present experiment the original female sent to us was a wild 
gravid euploeoides-nerina from Sarawak. Her progeny were mated to 
males from a stock of hybrid origin. These were of the genotype eepp, 
which in the female produces the form naresi (Fig. 4), the bottom 
recessive. 

Breeding methods. The butterflies were bred in heated greenhouses in 
Liverpool using the methods described in Clarke & Sheppard (1975). 


VoLUME 31, NuMBER 2 141 


TaBLE 1. Broods giving linkage data in H. bolina. 


Offspring 
Provenance and Provenance 
Brood No. Form of Mother of Father oS ee 
14067 Wild Sarawak Wild Sarawak 0 ir 11 euploeoides-nerina 
euploeoides-nerina 6 nerina 
14181 14067 hybrid 0 19 11 euploeoides-nerina 
euploeoides-nerina  eepp 8 nerina 
14187 14067 hybrid 0 4 3 euploeoides-nerina 
euploeoides-nerina  eepp 1 nerina 
14228 14181 hybrid Oe 26 34 euploeoides-nerina 
euploeoides-nerina eepp 28 euploeoides 
32 nerina 
32 naresi 
14229 14181 hybrid 0 10 2 euploeoides-nerina 
euploeoides-nerina eepp 2 euploeoides 
2 nerina 
A narest 


RESULTS 


Table 1 gives the progeny of the original female and subsequent 
broods. The first matings at Liverpool were those of two of her euploe- 
oides-nerina offspring, which were mated to males of hybrid stock 
homozygous eepp. Two of the resulting euploeoides-nerina progeny 
were again backcrossed to hybrid males eepp. 

The original female euploeoides-nerina, or her wild mate, or both, 
were probably homozygous for nerina (P"P"). Thus, brood 14067 ap- 
pears to be a backcross with respect to E. If neither of the parents was 
homozygous P*P", then one would expect at least one in three of the 
progeny to be neither euploeoides-nerina nor nerina. Since no other 
phenotype (euploeoides or naresi) appeared, it is likely that at least one 
of the parents was in fact homozygous P”P”. On the other hand, if the 
wild brood were an F2 for E, then the ratio should be at least three 
nerina to four non-nerina (euploeoides or naresi) among those insects 
that are not euploeoides-nerina. 

That the wild male was certainly carrying nerina is shown by the next 
generation. His ewploeoides-nerina daughter (EeP"P") mated to a 
naresi male (eepp) (brood 14181) produced no euploeoides or naresi, 
showing that she was homozygous nerina; her offspring must therefore 
have been heterozygous (EeP"p and eeP"p). This was confirmed by 
the second backcross (brood 14228), which segregated euploeoides- 
nerina (EeP"p), euploeoides (Eepp), nerina (eeP"p), and naresi (eepp ) 


142 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in a good 1:1:1:1 ratio. The small brood 14229 is also consistent with 
this ratio. Thus, the series of matings not only shows that a female 
euploeoides-nerina from the wild was in fact produced by the combina- 
tion of the allelomorphs EF and P", but the broods also strongly support 
the view that the two loci segregate independently and there is no need 
to invoke crossing over in the female. 

Since two of the forms (nerina and naresi) are nonmimetic, eu- 
ploeoides-nerina is a poor mimic, and only euploeoides is a good one, 
it seems unlikely that the polymorphism is maintained by frequency- 
dependent selection due to the mimicry alone, particularly in the absence 
of close linkage. It therefore seemed important to look at other possible 
selective effects of the allelomorphs, such as viability and speed of 
development. The present second backcross broods can give us no 
information on the viability of the euploeoides and nerina homozygotes. 
However, they also give no evidence, under laboratory conditions, of 
differential viability between the four genotypes that could be tested. 

There was a suggestion of differential speed of development, with an 
apparent excess of naresi emerging early and butterflies carrying eu- 
ploeoides (EeP"p and Eepp) coming out late. However, an analysis of 
variance revealed no significant differences with respect to the allelo- 
morphs and their interactions, the comparison “euploeoides” (EeP"p and 
Eepp ) to non-euploeoides (eeP"p and eepp) giving 0.1 > p > 0.05, nerina 
(EeP"p and eeP"p) to non-nerina (Eepp and eepp) p > 0.1, and the 
interaction 0.1 > p > 0.05. 

Because two of the probabilities were quite low we investigated similar 
large broods from Clarke & Sheppard (1975). As in the present brood, 
an insignificant excess of females carrying euploeoides came out late 
(p > 0.1). Clearly, it would be worth investigating the matter further. 
We could not combine the analyses because under the different cond- 
itions of raising the broods the variances in the emergence times were 
heterogeneous. 


SUMMARY 


The complicated polymorphism in the mimetic butterfly Hypolimnas 
bolina can be described in terms of the four main phenotypes and their 
combinations. Genetic analysis from broods bred in the laboratory 
reveals that these are controlled by two unlinked loci, a matter that was 
previously in doubt. 

No differences in the viability of the four genotypes tested could be 
detected. Although there was a suggestion that these allelomorphs af- 
fected speed of development, the differences were insignificant from 
the amount of data so far available. 


VoLUME 31, NUMBER 2 143 


ACKNOWLEDGMENTS 


We are grateful to the Science Research Council, the Nuffield Founda- 
tion, and the Royal Society for support. We would also like to thank Mr. 
Stephen Kueh for butterfly material from Sarawak. 


LITERATURE CITED 


CLARKE, C. A. & P. M. SHEPPARD. 1975. The genetics of the mimetic butterfly 
Hypolimnas bolina (L.). Phil. Trans. R. Soc. Lond. B. 272: 229-265. 

CLARKE, C. A., P. M. SHEPPARD, & V. Scaui. 1975. All-female broods in the butter- 
fly Hypolimnas bolina (.). Proc. R. Soc. Lond. B. 189: 29-37. 

Poutton, E. B. 1924. Mimicry in the butterflies of Fiji considered in relation to 
the Euploeine and Danaine invasions of Polynesia and to the female forms of 
Hypolimnas bolina L. in the Pacific. Trans. Ent. Soc. Lond., p. 564-691. 

ROBINSON, R. 1971. Lepidoptera genetics. Pergamon Press, Oxford. 

Traut, W. & B. RATHJENS. 1973. Das W-Chromosom vom Ephestia kuehniella 
(Lepidoptera) und die Ableitung des Geschlechtschromatins. Chromosoma 
(Berlin) 41: 437-466. 


CAPTURE OF PAPILIO ANDROGEUS (PAPILIONIDAE) IN SOUTHERN 
FLORIDA, A NEW RECORD FOR THE USA 


A female Papilio androgeus (Cramer) was captured on 22 March 1976 in Broward 
Co., Florida (Figs. 1-2). The badly worn specimen was captured in an overgrown 
orange grove adjacent to Flamingo Road, near State Road 84, west of Ft. Lauder- 
dale. I captured the specimen while she rested on weeds beneath the orange trees. 
According to Barcant (1970, Butterflies of Trinidad and Tobago, London) the 
main foodplant of the species is the orange tree, Citrus sinensis (Osbeck). 

Irving Finkelstein and Ray Suydam, reported seeing two males of P. androgeus 
in the same orange grove on 23 March and 27 March, respectively. Charles V. 
Covell, Jr. saw one male of the same species about 1 mi. W of the above location 


on 23 May 1976. 


ArTHUR JoE Patrerson, 749 Peachtree Street, Apt. C-2, Atlanta, Georgia 30308. 


Figs. 1-2. 1, left, female Papilio androgeus, upper surface; 2, right, same speci- 
men, under surface. 


144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


GENERAL NOTES 


DETERMINATION OF SEX IN FOUR SPECIES OF 
GIANT SILKWORM MOTH LARVAE (SATURNIIDAE) 


We consider the ability to determine the sex of giant silkworm moth larvae to be 
of applied value from two standpoints. First, we could rear only (or primarily) in- 
dividuals of the sex required for special research purposes. Second, we could in- 
sure that small colonies (< 12 individuals), maintained from year to year as breed- 
ing stock, contain sufficient numbers of each sex. Except for information reported 
for the domestic silkmoth, Bombyx mori Linnaeus (Tazima, 1964, The genetics of 
the silkworm, Prentice-Hall: Englewood Cliffs, New Jersey), we have found very 
little information concerning sex determinations in lepidopterous larvae. A recent 
paper by Hinks & Byers (1973, Can. J. Zool. 51: 1235-1241) reported structures that 
appear to be reliable indicators of sex in certain noctuid larvae. In female noctuid 
larvae these characters consist of four pits or other modifications of the integument 
associated with the developing female genitalia and occur between the ventral and 
subventral setae on the 8th and 9th abdominal segments; in male noctuid larvae they 
consist of a pit or other modification of the integument associated with the develop- 
ing male genitalia (Herold’s Organ) and occur on the venter of the 9th abdominal 
segment. Similar, but less detailed, information has been reported for the genus 
Malacosoma (Stehr & Cook, 1968, Bull. U.S. Nat. Mus., 276: pp 46-47). 

During 1975, while maintaining large research colonies of several giant silkworm 
moth species, we decided to examine some of our colonized larvae to determine 
whether they exhibited similar sex-related characters. We selected random samples 
of four giant silkworm moth species (Table 1) and categorized individuals as male 
or female on the basis of the characters reported by Hinks & Byers. The larvae were 
then segregated according to sex and reared to pupation or adulthood to confirm the 
sex of each individual. 

Larvae of Antheraea polyphemus (Cramer) were examined in both the 4th and 
5th instars. We found only the structure related to the developing male genitalia, 
visible to the naked eye in both instars as a single black pit on the venter of the 
9th abdominal segment; female characters were not observed even at 60% magnifi- 
cation. Individuals exhibiting this black pit were categorized as males; those with- 
out it were categorized as females. 

Larvae of Ewpackardia calleta (Westwood) were also examined in both the 4th 


TABLE |, Larval sex determinations for four giant silkworm moth species. 


Number Actual 
Number Categorized Number 
Larvae Male or Male or Probability of 
Species Examined Female Female! Misclassification? 
A. polyphemus 27 10 3 10 3 0 (0-0.27) 
179 Ter Q oes 0.12 (0.02-0.34 ) 
E. calleta 33 16 2 Gia 0 (0-0.18) 
ie® 179 OCOD 7) 
H. cecropia 28 iss A Wa I © 0.07 (0.0003—0.3 ) 
13 9 18).2 0 (0-028) 
C. promethea 43 I AO Bile) 0.05 (0.0002-0.21 ) 
22.9 92.9 0 (0-0.13) 


1Sex determined by examination of pupae or adults. 
* Estimates of probability of misclassification and 95% confidence interval. 


VOLUME 31, NUMBER 2 145 


Fig. 1-2. Ventral views of 5th-instar E. calleta larvae. 1, female larva showing 
location of the pits associated with the developing genitalia (FH) on the 8th and 


9th (S8 and S9) abdominal segments. 2, male larva showing the absence of these 
pits on S8 and S9. 


and 5th instars. We found only the developing female genitalia, visible to the naked 
eye in both instars as four prominent dark pits on the ventral side of the 8th and 9th 
abdominal segments (Fig. 1). We did not observe any male character (Fig. 2), 
even at 60X magnification. Individuals with the four dark pits were categorized 
as females; all others were categorized as males. 

Of the Hyalophora cecropia (Linnaeus) specimens, we examined 3rd-, 4th-, and 
Sth-instar larvae. We found only the developing female genitalia in the 4th- and 
oth-instar larvae, visible to the naked eye as four white subsurface spheres on the 
ventral side of the 8th and 9th abdominal segments. Microscopic examination of the 
integument over these white spheres did not reveal any pits or other modifications 
of the surface. Individuals with these white spheres were categorized as females; 
all others as males. 

Larvae of Callosamia promethea (Drury) were examined only in the 4th instar. 
Using the naked eye, we were unable to find any evidence of developing male or 
female genitalia. However, microscopic examination (60) revealed the presence 
of developing female genitalia in the form of two obscure, irregular, subsurface, 
dark green to black bodies on the ventral side of the 8th abdominal segment. There 
was a slight modification of the integument over these structures. Male characters 
could not be found even at 60 magnification. Individuals with the dark sub- 
surface structures were categorized as females; all others as males. 

Our observations (Table 1) demonstrate that the characters associated with the 
developing genitalia can be used to determine the sex of larvae of these four giant 
silkworm moth species. When selecting larvae, the probability that an individual 
will be misclassified depends on how distinct the characters are, the sex in which 
the characters occur, and the sex being sought. In A. polyphemus only the male 
character was observed. Thus, if one is seeking to obtain only male larvae of this 
species, the chance of selecting a group free of females is very good, particularly 
if any questionable individuals are excluded. Conversely, if one is seeking to obtain 
only female larvae of A. polyphemus, the chances are not as good because males 
with indistinct genital characters might be included with the females. Our findings 
bear this out for A. polyphemus and the other species we studied, although for E. 
calleta, H. cecropia, and C. promethea the situation is reversed. 

In addition to the colonized larvae, we examined five wild larvae of H. cecropia 


146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


that were collected in the 4th instar and examined only at that stage of their de- 
velopment. We categorized all these larvae as males, but later examination of the 
pupae showed four males and a female. These results agree with our findings for 
colonized H. cecropia larvae (Table 1). 

The findings from this study should be of interest, and perhaps of applied value, 
to lepidopterists, dealers, and researchers who colonize or study giant silkworm 
moths. 


THomMaAS A. MILLER, WILLIAM J. COOPER, AND JERRY W. HicHritt, U.S. Army 
Medical Bioengineering Research and Development Laboratory, Fort Detrick, Mary- 
land 21701. (The opinions contained herein are those of the authors and should 
not be construed as official or reflecting the views of the Department of the Army.) 


A METHOD FOR HANDLING EGGS AND FIRST INSTAR LARVAE 
OF CALLOSAMIA PROMETHEA (SATURNIIDAE) 


In an earlier paper (Miller & Cooper, 1976, J. Lepid. Soc. 30: 95-104) we re- 
ported the use of portable outdoor cages to effect the mating of various giant silk- 
worm moths, including Callosamia promethea (Drury). Since that time we have 
conducted studies to evaluate methods for the collection of eggs and the transfer 
of newly-hatched larvae to food plants. 

We routinely collect eggs from giant silkworm moths by placing fertile females 
in paper bags where they can oviposit on the inner surfaces. For larvae reared 
outdoors, we turn the paper bags inside out and place them in sleeve cages already at- 
tached to branches. For larvae reared indoors, we cut the bags into small pieces of 
paper containing the eggs masses, and these are variously attached to food plant 
cuttings. These methods are not novel and have long been used, with variations, 
by lepidopterists who colonize giant silkworm moths (Crotch, 1956, A  silkmoth 
rearers handbook, The Amateur Entomologist’s Society, London; Taschenberg & 
Roelofs, 1970, Ann. Ent. Soc. Amer. 63: 107—111; Waldbauer & Stemmburg, 1973, 
Biol. Bull. 145: 627-641; Dirig, 1975, Growing moths, N.Y. State College of Agri- 
culture & Life Sciences). For large-scale indoor colonization of giant silkworm 
moths we found that the time required to cut around the egg masses and then at- 
tach them to the food plants was unacceptable. Therefore, we developed a modi- 
fied procedure for collecting the eggs and transferring the larvae to food plants. 
This paper reports our results with C. promethea. 

We used 12 C. promethea females, each placed in a brown paper bag (lunch 
size) on the first night after mating. The following morning the female moths were 
removed and the bags, containing the eggs, were folded to their original flattened 
configuration and held for 8 days. On the 9th day 3 fresh wild cherry (Prunus) 
cuttings, each 15-20 cm long and containing 4—5 large leaves, were inserted into 
each bag. The tops of the bags were folded over about 1.5 cm and a small hole 
was made at the crease to allow the stems to protrude about 5.0 cm. The bags 
were inverted and the stems were placed in water containers. Observations of 
hatching and migration of larvae to the food plants were made by carefully opening 
the creased end of the bags and looking inside. 

The eggs hatched on the 10th day and the larvae crawled about on the inner 
surfaces of the bags; a few transferred to the wild cherry leaves, but none of these 
were observed feeding. By the end of the 11th day most of the larvae had transferred 
to the food plants and were feeding. Observations were continued through the 
13th day after oviposition, but no additional larvae transferred to the food plants 
after day 11. On the 14th day the bags were removed and cut open to record rele- 


VoLUME 31, NUMBER 2 147 


TABLE 1. Results of oviposition and larval transfer studies with Callosamia prome- 
thea (Drury). 


Number Larvae! 


Female Number Eggs Number Eggs Percent Transferring to Percent” 

Number Deposited Hatched Hatch Food Plant Transferring 

if 73 69 94.5 68 98.5 

2 WL 65 91.5 DD 84.6 

3 87 §2 94.2 45 54.8 

4 UO 69 86.6 66 95.6 

5 S7 om 100.0 49 85.9 

6 92 91 98.9 46 Si URs 

7 oi 54 94.7 50 92.5 

§ 16 15 93.7 15 100.0 

9 34 - 9 94.1 28 87.5 

10 a2 52 100.0 49, 80.7 

nia 40 39 97.5 Bil 94.8 

12 4§ 44 91.6 4] 93.1 

704 669 95.0 542 81.0 


1 By second day after hatching. 
2 Number on food plant/number hatched. 


vant data. The wild cherry cuttings, containing the Ilst-instar larvae, were placed 
in rearing cages along with other colonized C. promethea larvae and no further 
records of this group of experimental larvae were kept. 

We have concluded from the results obtained with this modified procedure 
(Table 1) that it is an effective and efficient method for handling eggs and _Ilst- 
instar larvae of C. promethea. Of the eggs that hatched (95%), 81.0% of the 
larvae migrated to the food plants within two days. We consider this percent 
transfer to be very acceptable, in view of the fact that we were able to obtain 542 
Ist-instar larvae on food plants in rearing containers with only a minimum of 
effort on our part. 

We have also found that this procedure gives acceptable results in obtaining eggs 
and Ist-instar larvae for the indoor colonization of Antheraea polyphemus (Cramer ) 
and Eupackardia calleta (Westwood), but we have not collected any detailed ex- 
perimental data for the transfer of these species to food plants. 


THomMas A. MILLER AND WILLIAM J. Cooper. U.S. Army Medical Bioengineering 
Research and Development Laboratory, Fort Detrick, Maryland 21701. (This re- 
search was not supported by government funds; the opinions contained herein are 
those of the authors and should not be construed as official or reflecting the views 
of the Department of the Army. ) 


HYPOSOTER FUGITIVUS (ICHNEUMONIDAE) PARASITIC WITHIN 
MEGALOPYGE OPERCULARIS LARVAE (MEGALOPYGIDAE ) 


The puss caterpillar, Megalopyge opercularis (J. E. Smith), is quite important 
from the medical standpoint since it is highly poisonous. 

On collecting larvae of this species from oak trees (Quercus) in New Orleans 
at the end of June 1976, some were noticed to be distinctly underdeveloped and 
quiescent. The latter were attached to leaves and measured only 7-8 mm. Most 


148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Remains of parasitized, young caterpillar of Megalopyge opercularis 
showing the hole from which the adult Hyposoter fugitivus emerged. 


other larvae found at that time measured ca. 20 mm in length. All the larvae 
were transferred to artificial diet. Three days later, two adults of Hyposoter 
fugitivus (Say) appeared in the container. On the examination of the larvae the 
parasitized ones were found. Nothing was left of the host except transparent 
cuticle attached to the leaf of the host plant. There was a 1.5 mm hole between 
the hairs from which the adult wasp emerged (Fig. 1). The wasp is parasitic on 
young puss caterpillars. Parasitism apparently resulted in the paralysis and death of 
the larva, which accounts for their underdevelopment. The parasite pupated within 
the host and emerged as adult. 

Cocoons of puss caterpillar are known to be parasitized by an ichneumonid wasp. 
However, this is the first record of an ichneumonid wasp parasitic on the larvae of 
this host. 

I am grateful to Dr. R. W. Carlson, Systematic Entomology Laboratory, IIBIII, 
for the identification of the wasp. 


KaMEL T, KHALAF, Loyola University, New Orleans, Louisiana 70118. 


WOODPECKER FEEDING ON CALLOSAMIA PROMETHEA 
(SATURNIIDAE ) COCOON 


About midday on 15 March 1975, while on a combined bird-walk and cocoon 
hunt along Bean Creek, near Morenci in Lenawee Co., Michigan, I observed a 
Downy Woodpecker, Dryobates pubescens medianus (Swainson), feeding on a live 
Callosamia promethea Drury cocoon. The day was bright, although cloudy, with 
temperatures in the forties—ideal weather for such activities. My attention was 


VoLUME 31, NUMBER 2 149 


drawn to the woodpecker as it landed in a Wild Black Cherry, Prunus serotina 
Ehrh., approximately 25 ft above the creek bank. I focused my 6-12 % 32 zoom 
binoculars on the woodpecker and observed that it had spotted the suspended 
cocoon about 2 ft away on a small lateral branch. It then flew to the cocoon, 
landed on the lower end and proceeded to peck into the cocoon. The woodpecker was 
clinging upside down while it fed on the cocoon from 1202-1209 EST, and then 
flew away. While I was observing this predator, another Downy Woodpecker 
flew into the same tree about 3 ft above the first one. It had spotted an- 
other C. promethea cocoon and studied it for about one minute before flying away 
without actually landing on the cocoon as had the first woodpecker. 

After retrieving both cocoons and cutting them open, I found that the first 
woodpecker had completely emptied the pupal contents through a hole about 2 mm 
in diameter at the thickest part of the cocoon, while the other cocoon contained a 
parasitized pupa. One can only speculate how these woodpeckers can discriminate 
between cocoons containing live or dead pupa prior to actually pecking and pene- 
trating the cocoon! Waldbauer et al. (1970, Ann. Ent. Soc. Amer. 63: 1366-1369 ) 
reported Downy and Hairy Woodpeckers, D. villosus (L.), feeding on cocoons of 
Hyalophora cecropia (L.) under field and captive conditions. Their observations 
suggest that woodpeckers may, without making a hole, identify cocoons that con- 
tain live pupae. They also reported evidence, although no field observations were 
cited, of woodpecker attacks on Antheraea polyphemus (Cramer) and C. promethea 
cocoons. 

This was my first observation of cocoon predation in 30 years of field experiences 
in Michigan, although several other saturniid cocoons have been found which show 
evidence of woodpecker attacks. It would appear that similar field observations are 
either rarely made or reported by lepidopterists. 


M. C. Nietsen, 3415 Overlea Drive, Lansing, Michigan 48917. 


150 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


OBITUARY 


ARTHUR H. NAPIER (1895-1976 ) 


Arthur H. Napier, a charter member of The Lepidopterists’ Society died 1 No- 
vember 1976 at the age of 81. Arthur was a graduate of Chestnut Hill Academy, 
Philadelphia, in 1913 and of Haverford College in 1917. He was a member of a 
number of learned societies and clubs, here and abroad, but his interest was 
focused on youth activities. He was a Merit Badge Counselor for the Philadelphia 
and Valley Forge Council of Boy Scouts of America and received the Award of 
Merit for his work in scouting. He will be remembered by the many younger 
people for his interesting Lepidoptera and other insect lectures that he gave for 
many years in the township schools near his home. Arthur was married to Elizabeth 
Doyle who died in 1966. In 1968 he married the former Eleanor M. McConnell 
who had made his lecture appointments in one of the schools where she was em- 
ployed. Besides his wife Eleanor, the others who survive him are: Arthur H. 
Napier, Jr., a son; two grandchildren, Constance Fraser and Arthur H. Napier, 3rd; 
three great-grandchildren, Elizabeth and Alison Fraser, David Groton Napier, and 
a stepson Leonard Sheppard. 


JosepH W. Avams, Morris Arboretum of the University of Pennsylvania, Phila- 
delphia, Pennsylvania 19118. 


VoLUME 31, NUMBER 2 151 


BOOK REVIEW 


REVISED CATALOGUE OF THE AFRICAN SPHINGIDAE (LEPIDOPTERA) WITH DESCRIP- 
TIONS OF THE EAST AFRICAN SPECIES, by R. H. Carcasson. 2nd Edition, 1976. E. W. 
Classey Ltd., Faringdon, England. 148 p., illus. + 17 plates. Price: $11.95 (U.S.) 
paperback. 


This publication was originally published in 1968 in the Journa! of the East 
Africa National History Society and National Museum, Nairobi, Kenya, and as far 
as I can see is an unchanged reprint, now in form of a monograph. It therefore 
contains, of course, again the systematic weaknesses pointed out already by Hodges 
(1971, The moths of America north of Mexico, fasc. 21). The introduction of the 
subfamily names “Asemanophorinae” and “Semanophorinae’—although the author 
himself admits that the correct name of the former should be Smerinthinae—is un- 
fortunate as they are contrary to the rules, notwithstanding the fact that the group- 
ing into two subfamilies is the right thing to be done. The names then should be 
with Hodges (1971): Sphinginae and Macroglossinae. 

In naming his tribes: Ambulicini, Acherontiini, Dilophonotini, Philampelini, and 
Choerocampini the author tries to keep the connection with Rothschild & Jordan 
(1903, Novit. Zool., vol. 9, suppl.). Today one would prefer to call them with Hodges 
(1971): Smerinthini, Sphingini, Dilophonotini, Philampelini, and Macroglossini. The 
breaking down of the tribes into subtribes is appropriate and even possible rightfully in 
accordance with the Code of Zoological Nomenclature which admits in article 35 “any 
supplementary categories required.” In North America, however, one usually forms 
subtribal names on the ending -iti, e.g. Sphingiti instead of Carcasson’s “Sphinges.” 

If the single Hawaiian species (Tinostoma smaragditis) taken on p. 7 into the 
subtribe “Philampeli” is a member of the subfamily “Semanophorinae” at all it is 
still an unsolved riddle. The palpus has certainly no sensory hairs as found in 
Eumorpha (= Pholus). This is now known quite thoroughly after examining more 
specimens of the species that have come into collections. It may well be that the 
genus Tinostoma (perhaps together with Sataspes which also lacks sensory hairs 
on its palpus) takes a kind of an intermediate position between the two acknowledged 
subfamilies, at least as far as the labial palpus is concerned. 

Interesting and to be commended is the line drawing of the wing pattern of a 
sphingid (p. 9) and the distributional tables for the tribes and subtribes resp. given 
in the general part (p. 2-9). They are based for the African species on Carcasson’s 
present work, for the other ones on Rothschild & Jordan (1903). 

The main part comprises p. 11-183 and is accompanied by 17 plates of adults 
(pl. 1-10), genitalia (pl. 11-17), and some immatures on pl. 16. The plates of the 
adults and immatures are all in black and white halftones that in most cases give a 
pretty good idea of the appearance of the moths in question. The genitalia are mostly 
halftones from microscopic slides, sometimes therefore difficult to interpret (alcohol 
preparations should be used exclusively for such purposes). Some of them are re- 
touched, especially those of females. Overall the photographs are too small, espe- 
cially of the aedoeagi. Pl. 17 and an unnumbered plate between p. 12 and 13 as well 
as a single drawing on p. 67 shows genitalia by use of linedrawings. The difference 
is apparent at once. The costs of publication may have been a hinderance to 
general use of linedrawings. Certainly in any case all the genitalia on the plates 
are still preferable to the ones in Rothschild & Jordan. 

To go deep into the main part would exceed the scope of this review. Only some 
remarks should be made: On p. 52 the generic name Herse has to be changed to 
Agrius, and on p. 111 the name Celerio to Hyles for well-known reasons. 

Carcasson creates in this work 12 new genera for 29 species and describes 7 new 
species. For one species a preoccupied name is replaced (Hippotion griveaudi tor 
Hippotion albolineata Griveaud). There are also four new “subspecies described 


152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


among which at least one, so it seems to me, deserves specific status based on the 
strong differences of the genitalia, as far as illustrations show ( Polyptychus (andosus ) 
amaniensis). In another “subspecies” (Hippotion rosae guichardi) the illustrations 
do not bear out what is said in the text. H. rosae rosae should be larger than 
r. guichardi, but, in the illustrations it is exactly the opposite. A scaled marker on the 
photographs would have been useful. 

Additions to the general distribution of 3 species have to be made: Deilephila 
neriti add Japan, Hawaii; Hyles lineata add Hawaii, Solomon Islands; Hippotion 
celerio add Papuasia, Polynesia (partially ). 

Hippotion isis is also represented in the Carnegie Museum and labelled “India.’ 
It does not seem to be a hybrid, much more it seems to stand in a similar relation to 
H. celerio as H. swinhoei to H. velox. The wing pattern is as in H. chloris, only 
that the color is uniformly clayish. 

The references at the end show that there are really only very few publications 
available about African sphingids, and only 4 of more recent date for Nigeria, 
Madagascar, Congo (Brazzaville) and Central and South Africa. With Carcasson’s 
work the whole of Africa is in the moment quite well covered: it also lists and pro- 
vides information on species not from East Africa. Throughout the whole work 
are valuable systematic clarifications, e.g., in the dividing of the previously “com- 
pound” genus Polyptychus into separate entities. 

A good and useful glossary follows, and after it is an index that shows where to 
look for illustrations of species not illustrated here. 

There are only very few printing errors (like “HIPPOTRION” on p. 121), as far 
as I can see. The only one of systematic importance is found on p. 5, line 22 from 
above, where one has to read “Asemanophorae” (italics mine) instead of Seman- 
ophorae. 

To sum it all up: It was a good deed of the publisher to make this worthy work 
again accessible to the entomological community. It is only in this way that we 
finally will be able to build up a sufficient knowledge of the Sphingidae of the 
world. 


J. C. E. Riorre, Bernice P. Bishop Museum, P.O. Box 6037, Honolulu, Hawaii 
96818. 


> 


EDITORIAL STAFF OF THE JOURNAL 
GerorcE L. Goprrey, Editor 


Illinois Natural History Survey, Natural Resources Building 
Urbana, Illinois 61801 U.S.A. 


WiiuiAM H. ALLEN, Associate Editor JAMes G. STERNBURG, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study 
of Lepidoptera. Contributors should prepare manuscripts according to the following 
instructions. 

Text: Manuscripts should be submitted in duplicate, and must be typewritten, 
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Underline only where italics are intended. References to footnotes should be num- 
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Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically 
under the heading Lirerature Cirep, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, 
London. 209 p. 

196la. Some contributions to population genetics resulting from the 
study of the Lepidoptera. Adv. Genet. 10: 165-216. 


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

Illustrations: All photographs and drawings should be mounted on stiff, white 
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for reduction to their final width (usually 4% inches). Illustrations larger than 8144 
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ALLEN PRESS, INC. eee LAWRENCE, KANSAS 
USA 


CONTENTS 


IMMATURE STAGES AND ECOLOGICAL OBSERVATIONS OF EOPARARGY- 
RACTIS PLEVIE (PYRALIDAE: NYMPHULINAE). Sandy B. Fiance 
and. Robert E. Moeller 2.00 5 ee 


Six New Species oF HEsPeRIDAE FROM Mexico. Hugh Avery 
Freeman 0. ee 


STUDIES ON THE BioLocy OF PARIDES IPHIDAMAS (PAPILIONINAE: 
TROWINI) IN Costa Rica. Allen M. Young __---.----- 


NoTES ON THE BEHAVIOR OF ASTEROCAMPA LEILIA (NYMPHALIDAE) 
IN SOUTHERN ARIzONA. George T. Austin ___. ae 


CUDONIGERA: A New Genus FoR Motus FORMERLY ASSIGNED TO 
CHORISTONEURA HOUSTONANA (TORTRICIDAE). Jerry A. Powell 
and N.S: Obraztsovu 20 


THE STATUS OF THE GLYPHIPTERIGIDAE AND A REASSESSMENT OF RE- 
LATIONSHIPS IN YPONOMEUTOID FAMILIES AND DITRYSIAN SUPER- 
FAMILIES. John B. Heppner 


Two New Species or PETROVA MOTHS FROM PINE IN SOUTHEAST 
Asta (TORTRICIDAE, OLETHREUTINAE). William E. Miller ___ 


DaTA SUGGESTING ABSENCE OF LINKAGE BETWEEN Two LOCI IN THE 
Mrmetic BuTTERFLY HYPOLIMNAS BOLINA (NYMPHALIDAE). C. 
A. Clarke and. P. M; Sheppard. 0 


GENERAL NOTES 


Aberrant Erynnis tristus tatius (Hesperiidae). Clifford D. Ferris 
Uniform genitalia among wing color morphs of olethreutid moths. William 


Pi Mailer? OS eh Nae Er 
An “albinic” Pieris sisymbrii ( Pieridae ) from the California Sierras. Arthur 
Mi STrapin 05 iim ah Oa Oh al vs Nr 
A record of Urbanus simplicius (Hesperiidae) for the USA. Mike A. 
Rickard ioe Ca Ban SU ta er 


Capture of Papilio androgeus (Papilionidae) in southern Florida, a new 
record for the USA. Arthur Joe Patterson _._......_ 
Determination of sex in four species of giant silkworm moth larvae (Saturni- 
idae). Thomas A. Miller, William J. Cooper, and Jerry W. Highfill _. 
A method for handling eggs and first instar larvae of Callosamia promethea 
(Saturniidae). Thomas A. Miller and William J. Cooper _......- 
Hyposoter fugitivus (Ichneumonidae) parasitic within Megalopyge oper- 
cularis larvae (Megalopygidae). Kamel T. Khalaf _...- 
Woodpecker feeding on Callosamia promethea (Saturniidae) cocoon. 
M. C. Nielsen 


OBITUARIES 


81 


89 


100 


111 


119 


124 


135 


139 


148 


151 


Volume 31 1977 Number 3 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


16 September 1977 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


J. W. Trtpen, President KENELM W. Puip, Vice President 
I. F. B. Common, Ist Vice President JuLian P. DonanueE, Secretary 
LIioNEL Hiccins, Vice President JouHn M. Sniper, Treasurer 


Members at large: 


F. S. CHEw R. A. ARNOLD J. F. EMMEL 
D. F. Harpwick E. D. CasHatr R. R. GATRELLE 
J. B. Z1EGLER R. E. STANFORD A: P. Pua 


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formally constituted in December, 1950, is “to promote the science of lepidopterology 
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Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA 


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JOURNAL OF 


Tue Lepipoprertsts’ SoctEtTy 


Volume 31 1977 Number 3 


SELECTION OF THE COCOON SPINNING SITE BY THE 
LARVAE OF HYALOPHORA CECROPIA (SATURNIIDAE)! 


A. G. ScarBroucH’, J. G. STERNBURG, AND G. P. WALDBAUER 
Department of Entomology, University of Illinois, Urbana, Illinois 61801 


The univoltine Hyalophora cecropia (L.) overwinters as a diapausing 
pupa in a silken cocoon that is usually anchored to a woody plant. Some 
are well above ground level on the branches of deciduous trees or shrubs, 
but most are near the ground among the stems of deciduous shrubs, 
among adventitious shoots at the base of a deciduous tree, or on an 
evergreen shrub. After leaf fall, cocoons on bare branches are clearly 
visible. Cocoons near ground level are usually hidden by grass, debris, or 
evergreen foliage. 

The location of a cocoon significantly affects the probability that it will 
be found by a vertebrate predator during the winter, and largely deter- 
mines the species of predator that will find it. In residential areas of 
Champaign and Urbana, Illinois, from 86.5%-90.9% of the high cocoons 
were destroyed by woodpeckers, while cocoons low in shrubs or shoots 
were almost exempt from predation (Waldbauer & Sternburg, 1967b; 
Waldbauer et al., 1970; Scarbrough, 1970). In rural habitats, not only 
are most pupae in cocoons on bare branches killed by woodpeckers, but 
those near ground level may be killed, although less frequently, by mice 
of two species, Peromyscus leucopis noveboracensis (Fisher) and P. 
maniculatus bairdii (Hoy & Kennicott), which do not occur in urban 
and suburban areas (Scarbrough et al., 1972) (see also Marsh, 1937). 

There is little literature on the behavior which leads to the selection 
of a spinning site by cecropia, no quantitative data on whether or not 
the larvae migrate from the food plant before spinning, on the extent of 
migrations, and on whether or not the migrations are affected by the 


1 Part of the Ph.D. dissertation presented by the first author to the Department. of Entomology, 
University of Illinois. This investigation was supported in part by PHS Training Grant no 
2- TO1- GM-01076 from General Medical Sciences. fet 

2 Present address: Department of Biology, Towson State College, Baltimore, MD 21204, USA. 


154 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


growth form of the host plant. We examined the behavior of cecropia 
larvae during the period which begins with the cessation of feeding and 
ends with the beginning of spinning. We are particularly concerned with 
the effects of the environment, especially plantings in urban and suburban 
residential areas, where cecropia is most abundant in central Illinois. 
Cecropia larvae feed on many species of deciduous plants, almost all 
woody and including both shrubs and trees (Ferguson, 1972; Scarbrough 
et al., 1974). Feeding larvae are relatively sedentary and seldom leave 
the host plant (unpublished observations ). Cocoons may be anchored to 
shrubs or trees which are known hosts of cecropia, but a significant 
number is found on plants, usually shrubs, which are not eaten by 
cecropia larvae (Waldbauer & Sternburg, 1967a; Scarbrough et al., 1974), 
indicating that some larvae leave the host plant before spinning. 


MATERIALS, METHODS, AND RESULTS 


The larvae: used in these experiments, progeny of locally collected 
pupae, were reared outdoors under nets on Malus pumila Mill. (apple), 
Prunus serotina Ehrh. (wild black cherry), Acer saccharinum L. (silver 
maple), or Cornus stolonifera Michx. or C. alba L. (shrubby dogwoods) as 
described by Waldbauer & Sternburg (1973). Experiments and sampling 
were done on the University of Illinois campus or residential areas in 
Champaign and Urbana, Illinois. 

Larvae transferred to trees or shrubs were always fifth instars which 
would continue to feed for at least four or five days before wandering 
off to spin their cocoons. They almost invariably completed the feeding 
phase within a few feet of the twig to which they had been transferred. 


Expt. I. Emptying the gut and moving to the spinning site. 


Feeding larvae reared on apple, silver maple or wild black cherry were 
transferred for observation to an unnetted tree of the same species, with 
no shoots at its base. They were placed singly on branch tips at the outer 
edge of the crown. Each morning they were counted and then watched 
continuously from 0500-1900 hours. We recorded relevant behavior and 
the time at which it occurred. At the end of the day larvae which had 
left the tree were replaced with new larvae. 

Mature larvae emptied the gut before moving to a cocoon spinning 
site, clinging from the underside of a branch by the thoracic legs and the 
first two or three pairs of prolegs, and allowing the end of the abdomen 
to hang free. They first passed dry, black feces, then progressively 
softer greenish feces, and finally a transparent, gelatinous liquid. When 
finished they crawled away on the upper side of the branch. 


155 


VoLUME 31, NUMBER 3 


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The times at which cecropia larvae began to empty their guts (A) and the 


Fie. I. 
times at which they began to spin the cocoon (B). 


the nearest hour after sunrise. 


156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


57cm 


Fig. 2. Diagrammatic representation of the trails of silk left by wandering 
cecropia larvae on a thin branch (a) and a wide branch or trunk (b). 


Gut emptying always occurred during daylight, most larvae beginning 
between 3 and 9 hours after sunrise, the mean being 5.7 hours after 
sunrise in both 1967 and 1968 (Fig. 1A). The process required from 
15-130 minutes, a mean and S.D. of 38 + 19. 

After emptying their guts, they quickly moved to the trunk and 
crawled downward, often diverging onto side branches, but usually 
returning to the trunk and the downward path. Some spun in the tree 
(see below), but most descended to the ground. 

They swung the head and thorax from side to side as they crawled, 
leaving a zig-zag trail consisting of a single strand of silk (Fig. 2). This 
motion is similar to the “swing-swing” reported as a component of 
cocoon construction, but not of crawling, by van der Kloot & Williams 
(1953a, b). On very small branches or rough bark the larvae obtained a 
secure hold and crawled rapidly, leaving a spread-out pattern (Fig. 2a). 
On larger branches with smooth bark they crawled slowly and left a 
compressed pattern (Fig. 2b). Ten larvae timed on a 1 cm diameter 
branch crawled at a mean rate of 78.0 cm per minute, while 10 others, 
timed on a smooth-barked branch 5 cm in diameter, crawled at a mean 
rate of 3.0 cm per minute. 


VoLUME 31, NUMBER 3 157 


Larvae descending large, smooth-barked branches or trunks, often 
paused at small branches or areas of rough bark, continuing to deposit 
silk with the swing-swing movement, and sometimes pausing long enough 
to form a thin sheet of silk. Although we did not see it happen, this 
might lead to the spinning of a cocoon. Indeed, in a sample of 323 
cocoons from small silver maples or birches (not including cocoons from 
shoots at the bases of the trees), 26% had been spun on small patches 
of rough bark or at the base of small branches on the predominantly 
smooth-barked trunk and large branches. 

Fig. 1B includes larvae that left the tree in search of a spinning site. 
Their wandering phase usually lasted from less than an hour to 8 hours; 
mean + §.D. were 3.8 + 1.8 hours in 1967 and 3.2 + 1.3 hours in 1968. 
They began to spin cocoons from 4-14 hours after sunrise, the means 
being 8.9 and 8.5 hours after sunrise in 1967 and 1968 respectively (Fig. 
1B). Spinning usually began on the same day the gut was emptied, but 
on a few unusually cool days migrating larvae rested on a woody plant 
from late afternoon until they resumed wandering the following day. 

We found, as did van der Kloot & Williams (1953a), that wandering 
larvae often pass locations which appear to be suitable for cocoon con- 
struction. This was exemplified by 40 larvae of Expt. IV that we timed 
after they had descended a tree with many adventitious shoots at its base. 
They eventually spun among these shoots, but first spent from 15-215 
minutes (x = 70) crawling among them. The wandering phase and the 
cocoon construction phase blend into each other since both involve the 
deposition of silk. 


Expt. II. Selection of the spinning site by larvae feeding in shrubs. 


We used 5 sites of two types. Two sites were single shrubs of Cornus 
stolonifera on mowed lawns and at least 10 m from any other woody 
plant. Three sites were rows of 70 or more contiguous, alternating C. 
stolonifera and C. alba on lawns. In one of these rows Viburnum sp. and 
Malus sp. were regularly interspersed. 

About once a week 5 feeding fifth instars, reared on one of the two 
species of Cornus, were released in each isolated shrub and at four widely 
separated points in each row. The larvae were marked with a distinctive 
color for each release point. The following October all cocoons were 
collected from these sites and their locations noted. The release point 
of the larva which spun each cocoon was determined by eluting in xylol 
the marker color from the exuviae in the cocoon (Scarbrough et al., 
1970). 

Sixty larvae were released on the isolated shrubs, but 2 were found 
dead. Cocoons of 51 (88%) of the survivors were found on the shrubs 


158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TaBLE 1. Location of cocoons of Hyalophora cecropia collected within a 9 m 
radius of host trees in residential areas of Champaign and Urbana, Ill. (Expt. III). 


Cocoons collected: 


1967-1968 1968-1969 1969-1970 Total 
Location No. % No. % No. % No. % 
Grass or shoots at 
trunk base 45 14.8 rom eZ 254 36.5 436 24.3 
Shrubs 148 48.8 343 A3.1 QA? 34.8 oo 40.8 


Total not in trees 193 Gow, 480 60.4 496 (io MtGs 65.2 
Total in trees 110 36.3 315 39.6 200 2501. 625 34.8 


on which they had been released. One of the 7 missing cocoons was at 
the base of a tree about 9 m away, but the other 6 were not found. 
Ninety larvae were released in the rows of shrubs. All of their cocoons 
were found and identified. Only 9% had left the row to spin on nearby 
shrubs or trees. Thirteen percent spun on the same shrub on which they 
had been released, 62% on a shrub within 4.6 m in the row, and 16% on a 
more distant shrub in the row. 

The fact that these larvae spun near the release site does not mean 
that they did not wander just before spinning. We watched some of 
them, and they crawled about at the base of the shrubs for a long time. 
The shrubs afford ample shade and the numerous closely-spaced stems 
provide a multitude of potential spinning sites. Thus, negative phototaxis 
and positive thigmotaxis probably kept the wandering larvae in the 
cover of the shrubs. If they did wander away, they were probably led 
back by a visual response to the silhouette of the shrub (see Expt. VI). 
Indeed, two marked larvae were seen returning to the isolated shrub on 
which they had fed. 


Expt. III. The proportion of larvae which leave the host tree to spin. 


We estimated this proportion by searching for wild cocoons in plant- 
ings on lawns with host trees for cecropia that were relatively isolated 
from other trees, but were near shrubs which cecropia larvae do not eat 
(Scarbrough et al., 1974). All plants and structures within 9 m of the 
tree were searched for cocoons. The number and location of the shrubs 
varied, but in most cases there were from 1-3 shrubs within 2 or 3 m of 
the tree. From past experience we knew that in similar situations few 
cocoons are found more than 9 m from the host tree. 

Less than 35% of the cocoons were on the branches of the host tree, 
but over 65% were on adventitious shoots or tall grass at the base of its 


VoLUME 31, NUMBER 3 159 


TasLE 2. Number and percent of Hyalophora cecropia larvae which spun cocoons 
in trees with two different growth forms after feeding in these trees at different 
heights above the ground (Expt. IV). 


Larvae which: 


Spun in tree Left tree 
Height of No. = SE =e 
Foliage type larvae (m) larvae No. To No. To 
Dense canopy & 3 40 he LS 33 82.5 
drooping branches 6 40 fe) 225 31 1s 
9 40 17 A425 25 ees 
Total 120 oo (27.5% ) 87 (72.5% ) 
Thin canopy & 3 AO 10 25.0 30 75.0 
horizontal branches 6 40 12 30.0 28 70.0 
9 AO 13 32:5 AT 67.5 
Total 120 35 (29.2% ) 85 (70.8% ) 


trunk, or on shrubs within 9 m (Table 1). The proportion of cocoons 
found off the trees is a little lower than might be suggested by the results 
of Expt. IV (Table 2). This probably reflects differences in sampling 
techniques. In Expt. IV we accounted for all larvae as they left the 
trees. However, counting cocoons in the sampling areas probably under- 
estimated the number of larvae which left the trees because cocoons are 
easier to find in trees than in shrubs, because some larvae probably spun 
beyond the 9 m limit, and because some wandering larvae were probably 
killed before they spun a cocoon. 

Adventitious shoots at the base of the host tree are the preferred spin- 
ning site of larvae which migrate from trees. In sample areas where such 
shoots were present, 83.3% of the cocoons found off the tree were on 
them. Where the host tree had no basal shoots, 13.1% of the cocoons 
off the tree were in tufts of grass at the base of the trunk; the remainder 
were in nearby shrubs. 


Expt. IV. Effect on cocoon site selection of the tree’s growth form and 
the height at which the larvae feed. 


We used two isolated silver maples on lawns. Both were about 12 m 
tall and had trunks extending about 2.4 m from the ground to the bases 
of the bottom branches. One had dense foliage, numerous drooping 
branches which shaded the trunk and ground all day, and adventitious 
shoots at the base of the trunk. The other had sparse foliage and rela- 
tively few branches, which were mostly horizontal; the trunk and several 
major branches received direct sunlight; there were no basal shoots. 


160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Feeding fifth instars reared on silver maple were placed in these trees 
at heights of 3, 6, or 9 m after being marked with a color indicating the - 
height. Y-shaped twigs with the larvae clinging to them were hung in 
the tree with a long cane pole. The larvae crawled to the leafy tip of a 
nearby branch (at approximately the intended height) and fed there 
until they moved to the spinning site. No more than 5 larvae were ever 
at the same height on one tree at the same time. During daylight the 
larvae were watched almost continuously, and the number with each 
color marking that left the tree was noted. 

The number of larvae which remained in the trees to spin, 27.5% on 
one and 29.2% on the other (Table 2), did not differ significantly. How- 
ever, on the tree with a dense canopy and drooping branches, the greater 
the height at which the larvae had been placed, the less likely they were 
to leave the tree (yx? = 7.02, P < 0.05), while on the tree with a sparse 
canopy and horizontal branches, their height did not significantly affect 
the probability that they would leave the tree to spin (y? = 0.56, P > 
0.975) (Table 2). 

Almost all larvae which remained in the trees spun low in the crown, 
near where the lower branches join the trunk, no matter at what height 
they had been placed. We found a similar distribution in a sample of 404 
wild cocoons collected from the branches of trees during the winter of 
1968-69. Ninety-seven percent were within a spherical volume with a 
2m radius that centered on the point where the trunk and lower branches 
join. Seventy-eight percent were within a concentric volume with only a 
1 m radius. The lower portion of these volumes included drooping 
branches. 

The results of Expt. IV raise the question of why high larvae were less 
likely than low larvae to leave the densely foliated tree, but not the 
sparsely foliated tree. Van der Kloot & Williams (1953a) postulated that 
during the wandering phase some internal event must occur before the 
larva can begin cocoon construction. The occurrence of this event may 
be a function of time or metabolic or hormonal processes. Larvae which 
reached the lower crown from the highest release point had expended 
more time and effort in wandering than the others, and, therefore, were 
presumably more internally motivated to stop wandering and spin a 
cocoon. We suggest that the summation of two factors, internal motiva- 
tion and satisfaction of the larvae’s spinning requirements, was more 
likely to end the wandering phase in the densely foliated tree than in the 
sparsely foliated tree, perhaps because the larvae are negatively photo- 
actic. 

As in Expt. III, shoots at the base of the host tree were the preferred 
spinning site. Eighty-four of 87 larvae (96.5%) spun among the shoots at 


VoLuUME 31, NuMBER 3 161 


Ui Wy _ 


\\ 


10 


1 Acer saccharinum 
2 Picea: sp: 

3 Juniperus sp. 

4 Forsythia sp. 

5 A. saccharum 

6 Sidewalk 

7 Street 

8 Fraxinus sp. 


Fig. 3. Diagram of the site of Expt. V, showing residential plantings. 


the base of the tree which had them, but only 6 of 85 larvae (6.9%) spun 
in tufts of grass at the base of the other tree. 


Expt. V. Distance from the host tree to the spinning site. 


The experimental area was a closely mowed lawn with a silver maple 
and a number of other shrubs and trees at distances of from 3 m to 23 m 
from the base of its trunk (Fig. 3). There were no adventitious shoots or 
tall grass at the base of the silver maple. Eight groups of 10 feeding fifth 
instar larvae, reared on silver maple, were placed on the lower branches 
of the silver maple at 6-day intervals. The larvae were marked as de- 
scribed above. They were watched continuously during daylight, and the 
behavior and route of each migrating larva were recorded. 

All larvae left the tree, although two eventually returned to spin on its 
branches. Seventy-five percent moved more or less directly to the closest 
spinning site, a spruce (Picea sp.) about 3 m from the base of the trunk 
(Fig. 3). Most of the others moved in the general direction of other 
shrubs, turning directly toward them when they came within about 1 m. 


162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Taste 3. Spinning sites chosen by Hyalophora cecropia larvae which left the 
host tree (A. saccharinum) (Expt. V). See Fig. 3 for code numbers and locations of 
the spinning sites. 


Meters from Larvae spinning: 
trunk of 

Spinning site host tree No. Jo 
Acer saccharinum (1) 0 ye 226 
Picea sp. (2) LL 43 50.8 
Forsythia sp. (4) 2.4 5 6 
Juniperus sp. (3) 3.0 li ON AL 
Picea sp. (9) 10.0 2 2.6 
Fraxinus sp. (8) UB 7 6 7.8 

A. saccharum (5) 9.8 0 0 
House (10) — 2 236 

Total ie 


4 These larvae left the tree and returned. 
b Does not include 3 larvae which left the experimental area. 


A few moved toward the southeast where there were no shrubs; after 
moving about 2.5 m from the trunk they tumed either north toward the 
house or south toward the closest tree. 

Over 84% of the larvae spun on one of the three woody plants closest 
to the tree, 56% on the small spruce, the closest spinning site (Table 3). 
Only 3 left the experimental area. They crawled into the street and were 
lost. 


Expt. VI. Spinning site selection by larvae in trees with no other spin- 
ning sites nearby. 


The experimental site was a closely mowed lawn with a silver maple 46 
m from the nearest other woody plant. There were no shoots or tall grass 
at the base of the tree. Eighty marked fifth instars that would continue 
to feed for several days were released and observed as described in Expt. 
We 

All 80 larvae left the tree. However, 73 of them (91.3%) eventually 
returned to spin on its branches. Six others climbed trees about 46 m 
away and spun on their branches. The remaining larva crawled about 37 
m to the street and was lost. 

They crawled in a more or less straight line away from the trunk of the 
host tree, occasionally stopping to wave the head and thorax from side to 
side as if they were trying to get a visual fix on some object. They 
crawled from 1-25 m (x = 9.5 m) before returning to the tree. They 
always took a somewhat different path on the way back, indicating that 
they did not follow a silk or an odor trail, and suggesting that they used 
visual cues to orient to the tree. 


VoLUME 31, NUMBER 3 163 


TasBLE 4. Response of wandering Hyalophora cecropia larvae to black or white 
models presented either alone or simultaneously (Expt. VII). 


Larvae moving to: 


Black model White model 
No. larvae SE a re 

Models presented tested No. % No. % 

Black alone 40 36 90.0 = = 
White alone 40 — — 29 (ONS 
Black & white simultaneously 40 34 85.0 5 OAS 


Expt. VII. Orientation of migrating larvae to models. 


We tested the possibility that migrating larvae visually orient to tree 
trunks or shrubs, determining their responses to flat cardboard models 
which approximated the silhouette of a low juniper shrub. They were 
0.9 x 1.5 m rectangles painted either white or flat black. We tested 120 
larvae which had been placed in an isolated tree on an otherwise bare 
lawn a few days before the end of the feeding phase. They were tested 
only after they left the tree spontaneously. After a larva had crawled 
about 3 m from the tree, a model or a pair of models side by side, was 
placed a few meters ahead of the larva, parallel to, and about 1 m to the 
side of its anticipated path. The experiments were done on sunny days 
with the models placed so that their shadows were cast away from the 
larvae. 

The response of the larvae (Table 4) leaves no doubt that visual cues 
are an important component in finding spinning sites. There was a strong 
response to either the black or white model presented alone, although 
fewer larvae went to the white model. When black and white models 
were presented simultaneously, 97.5% of the larvae responded, 85% going 
to the black model. 

Positive responses usually consisted of abrupt changes in direction, the 
larva usually moving to the edge rather than the center of the model. 
If a model was moved while a larva was approaching, the larva stopped, 
raised the forward portion of its body, waved from side to side, and then 
moved toward the new position of the model. All larvae followed the 
model if it was moved, but some made gradual rather than abrupt turns 
toward its new position. 


DISCUSSION 


Our data and the data of Waldbauer & Sternburg (1967a) and Scar- 
brough et al. (1974) show that, generally speaking, overwintering 


164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


cecropia cocoons occur in two sorts of situations: 1) exposed to view on 
the branches of deciduous trees or shrubs, or 2) hidden near ground 
level among shoots or the stems of a shrub. The behavior involved in 
the selection of these sites is especially interesting because exposed co- 
coons are much more likely to be destroyed by woodpeckers than are 
hidden cocoons (Waldbauer & Sternburg, 1967b; Waldbauer et al., 
OO): 

The selection of the spinning site is significantly affected by the im- 
mediate environment of the feeding larva. Since feeding larvae are 
sedentary, we can assume that their location was ultimately determined 
by the ovipositing female. Larvae which feed on shrubs almost always 
spin the cocoon close to the ground on the same or an adjoining shrub. 
Larvae which feed in trees may spin on a branch of the tree, or they may 
leave and spin near ground level. They rarely return to a tree or climb 
another tree to spin unless other spinning sites are absent. In suburban, 
residential plantings, an average of 65%-71% of the larvae in trees 
ultimately left the tree to spin, either at its base or a more distant site. 

The onset of site selection is obviously controlled by a circadian 
rhythm. The cocoon site is always selected in daylight. Larvae usually 
begin to wander in the morning and usually begin to spin before night- 
fall. If not, they rest during the night, and resume wandering the next 
morning. Wandering in daylight rather than at night probably increases 
the risk of predation, but has the advantage of allowing the larvae to use 
visual cues in selecting a spinning site. 

Although we have not proved the point, our data suggest that the 
suitability of a potential cocoon site is determined by the larva’s negative 
phototaxis and positive thigmotaxis. This tends to hide the cocoon. We 
suggest that the larva is led to the spinning site by positive geotaxis and, 
sometimes, by its ability to orient to objects visually. An endogenous 
factor lengthens the wandering phase, decreasing the probability that 
larvae in trees will spin cocoons before reaching the ground. This is 
adaptive because a cocoon that is hidden among the leaves of a tree in 
summer, will probably be exposed to view in winter. 

The exposure of the larva to predation as it wanders in search of a 
spinning site seems to argue against the adaptive value of this behavior 
if one is not aware that most pupae in cocoons on trees are destroyed by 
woodpeckers during the winter (Waldbauer & Sternburg, 1967b; Wald- 
bauer et al., 1970). The wandering phase lasts an average of 3.5 hours, 
but the larva is in the open for only a brief portion of this time. We do 
not doubt that this relatively brief exposure is more than offset by the 
advantage of spinning in a site where the cocoon is hidden from view 
during winter. 


VoLUME 31, NUMBER 3 165 


SUMMARY 


The location of the overwintering cocoons of Hyalophora cecropia 
affects the probability that they will be attacked by vertebrate predators 
in winter. In this light we examined the selection of the cocoon site by 
the larvae, and the effect on this behavior of environmental factors in the 
suburban residential plantings in which cecropia is most common in cen- 
tral Illinois. Feeding larvae are sedentary and do not normally leave the 
host plants, shrubs and trees of many species. Between 3 and 9 hours 
(x = 5.7) after sunrise, mature fifth instars spend a mean of 38 minutes 
emptying the gut, and then begin to wander. Wandering usually lasts 
from less than an hour to 8 hours (X = 3.5), cocoon spinning beginning 
between 4 and 14 hours (x = 8.7) after sunrise. On cool days the larvae 
may rest and resume wandering the next morning. The cocoon site is 
always selected in daylight. Larvae which feed in shrubs almost always 
spin close to the ground on the same or a contiguous shrub. Larvae 
which feed on trees may spin on one of its branches, where the cocoon is 
exposed to view in winter, or they may migrate to spin near ground level 
in shoots at the base of the tree or in a nearby shrub where they are likely 
to be hidden in winter. In suburban plantings between 65% and 71% of 
the larvae in trees spin near ground level. Larvae locate distant shrubs 
by their ability to orient to silhouettes from a distance. The larva is ex- 
posed to predation as it wanders from a tree, but this is offset by the 
advantage of spinning the cocoon where it will be hidden from wood- 
peckers in winter. 


ACKNOWLEDGMENTS 


We are grateful to: Dr. George Sprugel, Jr., Chief of the Illinois 
Natural History Survey, and Dr. William H. Luckmann, Head of the 
Survey's Section of Economic Entomology, who gave us the use of a 
grove of trees for rearing larvae; Dr. J. E. Appleby of the Section of 
Economic Entomology who allowed us to use trees on some of his plots. 


LITERATURE CITED 


Frrcuson, D. C. 1972. Bombycoidea (in part), p. 155-275. In R. B. Dominick 
et al. (eds.), The moths of America north of Mexico. E. W. Classey, London. 
Marsu, F. L. 1937. Ecological observations upon the enemies of cecropia, with 
particular reference to hymenopterous parsites. Ecology 18: 106-112. 
ScarsBroucH, A. G. 1970. The occurrence of Hyalophora cecropia (L.) as related 
to urbanization. Ph.D. thesis. University of Illinois, Urbana, 217 p. 
ScarsroucH, A. G., G. P. WaLpBaurrR, & J. G. SteRNBuRG. 1970. <A method for 
associating cocoons and marked last-stage larvae. Ann. Ent. Soc. Amer. 63: 1481. 
1972. Response to cecropia cocoons of Mus musculus and two species of 
Peromyscus. Oecologia (Berl.) 10: 137-144. 


166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1974. Feeding and survival of cecropia (Saturniidae) larvae on various 
plant species. J. Lepid. Soc. 28: 212-219. 

VAN DER Kxioot, W. G. & C. M. Wituiams. 1953a. Cocoon construction by the 
cecropia silkworm. I. The role of the external environment. Behaviour 5: 141- 
SG 


1953b. Cocoon construction by the cecropia silkworm. II. The role of the 
internal environment. Behaviour 5: 157-174. 

WaALpBAvER, G. P. & J. G. SrerNBuRG. 1967a. Host plants and the locations of 
the baggy and compact cocoons of MHyalophora cecropia (Lepidoptera: 
Saturniidae ). Ann. Ent. Soc. Amer. 60: 97-101. 

1967b. Differential predation on cocoons of Hyalophora cecropia 

(Lepidoptera: Saturniidae) spun on shrubs and trees. Ecology 48: 312-315. 

1973. Polymorphic termination of diapause by cecropia: genetic and geo- 
graphical aspects. Biol. Bull. 145: 627-641. 

WALDBAUER, G. P., J. G. STERNBURG, W. G. GrEorcE, & A. G. ScArsroucH. 1970. 
Hairy and downy woodpecker attacks on cocoons of urban Hyalophora cecropia 
and other saturniids (Lepidoptera). Ann. Ent. Soc. Amer. 63: 1366-1369. 


ANTHERAEA POLYPHEMUS (SATURNIIDAE) AND BIBLIS HYPERIA 
(NYMPHALIDAE) IN TEXAS 


The statement was made (Ferguson 1972, Moths of North America, Fasc. 20.2B, 
E. W. Classey Ltd., London, p. 200) that although Antheraea polyphemus (Cramer) 
is the most widely distributed of North American Saturniidae it has not been reported 
from counties of the southern coastal bend of Texas. On 18 September 1976, I took 
eight males of this species in black light traps at the Risken Ranch (27°9’N, 97°41’ 
W) in Kenedy Co., definitely a southern coastal bend county. Three were kept as 
specimens, and one of the three was deposited in the U.S. National Museum. I had 
previously reported other, more northerly coastal bend records (Kleberg and San 
Patricio Co.) to J. F. Gates Clarke and at his suggestion sent voucher specimens to 
the National Museum. 

Biblis hyperia (Cramer) (Nymphalidae) is reported by Howe (1975, The butter- 
flies of North America, Doubleday, Garden City, NY, p. 125) to occasionally stray 
into the Brownsville region of Texas. In the Texas A & I University—Kingsville 
collection are three specimens (one female, two males) taken by members of my 
entomology class and myself at the R & B Welder Refuge near Sinton, San Patricio 
Co., Texas, on 30 November 1973. Some years previous to this date, Carroll Williams 
of the Corpus Christi (Texas) Museum mentioned to me that he had taken the 
species at the Welder Refuge. Good evidence exists that there is, or has been, a 
resident population of this species in the United States. Williams’ specimens were 
taken at the “Hackberry Motte,” mine ca. 6 mi distant at the “Bolsa,” both areas of 
dense wood along the Aransas River. 


James E. Ginuaspy, Biology Department, Texas A & I University, Kingsville, 
Texas 78363. 


VoLUME 31, NUMBER 3 167 


A NEW SUBSPECIES OF PIERIS SISYMBRII (PIERIDAE) 
FROM WESTERN GREAT PLAINS RELICT FORESTS 


Kurt JOHNSON 


Department of Biology, City University of New York, City College, 
Convent Avenue and 138th Street, New York, New York 10931 


Pieris sisymbrii Boisduval that occur in scarp woodlands on the western 
Great Plains have formerly been recognized as distinct from their Rocky 
Mountain counterparts (Johnson & Nixon, 1967; F. M. Brown, pers. 
comm.), but their extremely low density required several years for 
sufficient sampling. They represent a new, allopatric subspecies (de- 
scribed below )—part of a larger array of montane insects that occur 
northward from western Nebraska and originated in the vast conifer for- 
ests which once covered the area after the Wisconsin glaciation. They 
have been isolated in disjunct relict populations by drier climate and 
range fire. The present description is part of a larger study of speciation 
in this former montane area (Johnson, 1976, 1977; Johnson & Balogh, 
1976). 


Pieris sisymbrii nordini Johnson, new subspecies 
Bigs. is52 


Diagnosis. This subspecies can be distinguished from P. s. elivata Barnes & 
Benjamin by the following traits: Males, upper surface of the forewings: marking 
between veins Cu: and Mz reduced or nearly absent; extreme proximity of veins 
Ri and Re allowing only narrow costal margin, black coloration from dark bar at 
apex of discal cell extending costad and to base of wing (P. s. elivata: Ri and Re 
broadly parallel, allowing wide costal margin, white ground color invading between 
it and the marking in the cell). Both sexes, undersurface of the hindwings: vein 
RS bending broadly along costal margin locating distal RS marking, mounted speci- 
mens, quite distad and below the tornus [P. s. elivata: RS highly arched, very 
limited in extension along costal margin (making distal RS marking, mounted speci- 
mens, appear well beneath inner margin of forewings)]. Male genitalia (Fig. 2): 
valvae longer and more broad, area between articulations broadly concave, caudal 
end exceeding tip of uncus, uncus more toothed than tapered [P. s. elivata: shorter, 
caudally tapered valvae, barely exceeding uncus; area between the articulation not 
smoothly concave, uncus gradually tapered; P. s. sisymbrii, P. s. flavitincta Comstock 
Cie) 2). 

Male. Upper surface of wings: white; forewings with moderately dark sub- 
apical markings deeply incised between stem of veins R; + 4/Rs and M:; also less 
distinct marginal markings. Marking between veins CU: and Ms reduced or nearly 
absent. Dark coloration from marking, apex of discal cell, extending costad to base 
of wing. Hindwings above, vaguely dark along veins. 

Undersurface of wings: forewing markings as on upper surface but more greenish 
with yellow-gold veins. Hindwings, veins outlined uniformly and thickly with brownish 
green, veins yellow gold; postmedian “arrowhead”-shape markings not concise, 
more bulbous than tapered; no extremely lightened patch basad along the veins. 

Length of forewing: 19.0 mm (holotype), 17.0-19.0 mm (paratypes). 


168 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Pieris sisymbrii nordini, new subspecies: (A) holotype, male, upper 
surface; (B) same, undersurface; (C) allotype, female (yellow), upper surface; 
(D) same, undersurface. 


Female. Two morphs, white and yellow, the latter in high frequency [0.67 
(N=6)] compared with P. s. elivata, [0.22(N = 27)]. 

Upper surface of wings: as in male, with marginal and apical-subapical markings 
darker and more extensive; postmedian marking between veins CU: and Mz dark 
and extensive; also a dark marking between veins CUz and 2aA. Hindwings heavily 
powdered along veins, especially above ventral postmedian “arrowhead” markings 
and at apex of cell. 

Undersurface of wings: as in males, with dark postmedian markings between 
veins CU: and Ms, and CUz and 2aA. 

Length of forewing: 19.0 mm (allotype), 17.0-19.5 mm (paratypes). 

Male genitalia. Typical of the species, differing mainly as follows: valvae 
longer and more broad, caudal end exceeding length of uncus, area between artic- 
ulations broadly concave; uncus caudally more toothed than gradually tapered. 

Female genitalia. Typical of the species: sclerotized abdominal segments with 
apophyses; a large, toothed and bulbous-ended cylindrical signum in the wall of the 
oval-shape corpus bursa. 

Early stages. Unknown. 

Foodplant. Probably various Cruciferae (Brown et al., 1957; Ferris, 1971). 

Types. Holotype, male, Monroe Canyon, Sioux Co., Nebraska, 24 May 1964 
(K. Johnson), in the American Museum of Natural History (AMNH), genitalia in 


VoLUME 31, NUMBER 3 169 


P. S. NORDINI 


P. S. FLAVITINCTA (4 


AFDEAGUS 


We AWA 


UNCUS SACCUS 


Fig. 2. Genitalia of Nearctic Pieris sisymbrii subspecies: (1) P. s. elivata, (2) 
P. s. nordini, (3) P. s. sisymbrii, (4) P. s. flavitincta. Upper: lateral view, uncus, 
vinculum, saccus, valva (valvae only, sisymbrii, flavitincta). Middle: lateral view, 
aedeagus. Lower: left, dorsal view, uncus; right, dorsal view, saccus. 


vial KJ #2, same institution. Allotype, female, (representing morph of highest fre- 
quency, yellow), same locality, 24 May 1968 (K. Johnson), genitalia in vial KJ 
#19, same institution. Paratypes, males (battered): same locality, 20 May 1966 
(K. Johnson), genitalia in vial KJ #17, same institution; same locality, 26 May 
1987 (R. A. Leussler), genitalia in vial KJ #25, in collection of Ohio State Uni- 
versity, Columbus, Ohio; Hell Canyon, S of Hwy. 16, Custer Co., South Dakota, 31 
May 1968 (J. S. Nordin), in his collection: females (Y = yellow; W = white): 
(W), 5 mi. W of Pringle, Custer Co., South Dakota, 28 May 1966 (J. S. Nordin), 
in AMNH; (Y), Chadron, Dawes Co., Nebraska, 6 May 1962 (K. Johnson), geni- 
talia in vial KJ #20, in the Los Angeles County Museum; (W), Monroe Canyon, 
Sioux Co., Nebraska, 24 May 1968 (K. Johnson), genitalia in vial KJ #18, in the 
Allyn Museum of Entomology; (Y), Canon Region, N of Harrison, Sioux Co., 
Nebraska, 5 June 1914 (R. A. Leussler), genitalia in vial KJ #5, in collection of 
Ohio State University; (Y), Guffis Canyon Fire Road, Custer Co., South Dakota, 1 
June 1968 (J. S. Nordin), in his collection. 

Distribution. Black Hills of South Dakota and relict pine woodlands of Ne- 
braska and perhaps North Dakota (Fig. 3). 

Flight period. Mid-April (sight records, this author) to early June. 

Remarks. These specimens were compared with 53 specimens of P. s. elivata 
from Wyoming and Colorado, of which 10 genitalia were studied. Specimens were 


170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Missouri NORTH DAKOTA 
a 


KILLDEER 


“Usp 
RIEL ABIL 


MISSOURI R. 
BADLANDS 


Y VR 


SOUTH DAKOTA 


1/, HILLS % 
bi 


G 
Wp 


BEACK Himes 


Yi 


WILDCAT 
HILLS 


NEBRASKA 


Fig. 3. Known distribution of P. s. nordini, new subspecies, superimposed on map 
of present-day montane area of eastern Rocky Mountains and western Great Plains 
(black areas). Hatching indicates approximate distribution of post-Pleistocene cli- 
max conifer forest. Open pine-juniper woodland extended around it, especially 
into central Nebraska and westward to the Rocky Mountain forests. Open wood- 
lands, and then the forests, disappeared as conifer environments retreated upland 
leaving the present-day montane relicts. 


VOLUME 31, NUMBER 3 7k 


also examined of P. s. sisymbrii from California and Utah, of which 6 genitalia 
were examined (3¢,3@). 

Etymology. I am very pleased to name this subspecies after Dr. John S. Nordin 
(Warrington, Pennsylvania), whose collections and research concerning South 
Dakota butterflies comprise the largest single contribution to the knowledge of 
South Dakota’s butterfly fauna in recent years. 


DISCUSSION 


Butterfly subspecies endemic to the relict woodlands of the western 
Great Plains usually are distributed disjunctly from the Wildcat Hills 
(Scotts Bluff and Banner Cos., Nebraska) and the canyons of Goshen 
Co., Wyoming [these being relicts of the former “Cheyenne Ridge” 
(Clements, 1949)] northward to the Pine Ridge of northwestern Nebraska, 
the Black Hills of South Dakota, and the scarps along the Little Missouri 
River in western North Dakota. Puckering & Post (1960) mention a 
“possible” P. sisymbrii from just below the North Dakota border in 
Harding Co., South Dakota. It is not specifically stated whether the 
specimen figured is the specimen from Harding Co.; the figured speci- 
men does not match the description of P. s. nordini in major traits. This 
and the indistinctness of a marking between veins CU2 and 2,A (under- 
surface only), which if the specimen is a female should be much more 
distinct, suggest that their specimen may represent early spring P. 
protodice occidentalis Reakirt, as is also stated by Dr. A. B. Klots in 
Puckering & Post (1960). 


ACNOWLEDGMENTS 


I am very grateful to Dr. John S. Nordin, Dr. Clifford D. Ferris ( Uni- 
versity of Wyoming, Laramie), Dr. C. A. Triplehorn (Ohio State Uni- 
versity, Columbus), and John C. Downey (University of Northern 
Iowa, Cedar Falls) for loans of specimens or photographs and especially 
to Dr. Frederick H. Rindge, Curator of Lepidoptera, American Museum 
of Natural History, for providing facilities and advice during this study 
and editing the manuscript. 


LITERATURE CITED 


Brown, F. M., D. Err, & B. Rorcer. 1957. Colorado butterflies. Denver Mu- 
seum of Natural History, Denver. 368 p. 

Ciements, F. E. 1949. The dynamics of vegetation. H. W. Wilson, N.Y. xiii + 
296 p. 

Ferris, C. D. 1971. An annotated checklist of the Rhopalocera of Wyoming. 
Univ. Wyoming Agric. Exp. Stat. Sci. Monogr. 23, 75 p. 

Jounson, K. 1976. Concerning the name Anthocaris coloradensis Hy. Edwards 
with designation of a new subspecies (Pieridae). J. Lepid. Soc. 30: 252-259. 


172 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1977. Post-pleistocene environments and montane butterfly relicts of the 
western Great Plains. J. Res. Lep. 15: (in press). 

Jounson, K. & G. Batocu. 1976. A revision and taxonomic analysis of the 
Nearctic Lycaena rubidus complex. Bull. Allyn Mus. (in press). 

Jounson, K. & E. S. Nixon. 1967. The Rhopalocera of northwestern Nebraska. 
Am. Mid. Nat. 78: 508-528. 

Pucxerinc, D. L. & R. L. Posr. 1960. Butterflies of North Dakota. Dept. Agric. 
Ent., North Dakota Agric. College Publ. 1, 32 p. 


OVIPOSITIONAL MISTAKE BY A HACKBERRY BUTTERFLY 
(NYMPHALIDAE ) 


Ovipositional mistakes by female butterflies have been previously reported (Rem- 
ington, 1952, Psyche 59: 61-70; Dethier, 1959, Can. Ent. 91: 554-561; Neck, 1973, 
J. Lepid. Soc. 27: 22-33). Such mistakes may occur on toxic, exotic species which 
have palatable native congeners (Straatman, 1962, J. Lepid. Soc. 16: 99-103; 
Kendall, 1964, J. Lepid. Soc. 18: 129-157). 

Herein is described the behavior of a single female Asterocampa clyton texana 
(Skinner) (Nymphalidae) which was observed at 1810 hrs. CST on 7 August 1970 
in a residential area of Austin, Travis County, Texas. The adult was observed flying 
around the yard mosily at ground level. The individual landed momentarily on 
several ornamental plants: Chinese privet, Ligustrwm sinense Lour. (Oleaceae); 
heavenly bamboo, Nandina domestica Thumb. (Berberidaceae) and corona vine, 
Antigonon leptopus Hook. & Arn. (Polygonaceae). Subsequently, it landed on a 
blade of St. Augustine grass, Stenotaphrum secundatum Kuntze (Gramineae), upon 
which it quickly laid three eggs. It then landed on several dead branches of an 
adjacent Texas sugarberry, Celtis laevigata Willd. (Ulmaceae), its major larval food- 
plant in central Texas. Extrusion of the ovipositor was observed several times but 
no oviposition occurred. After flying out of the immediate area, the same individual 
returned at 1825 CST. It landed on the trunk of the privet plant and extruded 
its ovipositor but laid no eggs. It then flew out of sight and was not seen again. 

Asterocampa normally oviposit small to large egg masses on the underside of hack- 
berry leaves (Comstock, 1953, Bull. So. Cal. Acad. Sci. 52: 127-136; Langlois & 
Langlois, 1964, Ohio J. Sci. 64: 1-11). These observations involved a female A. 
clyton texana which had mated and was apparently intensely stimulated to oviposit 
although unable to locate new-growth leaves of C. laevigata. Such leaves were non- 
existent at the time of observation; the last significant rainfall had occurred on 28 
May (0.97 in.). 

These observations may provide insight into future research possibilities on 
strategies of oviposition in Asterocampa in semi-arid habitats. Several days previous 
to time of observation, weather conditions were overcast with high humidity as a 
result of fringe weather associated with Hurricane Celia; barometric pressure was 
slightly below normal levels. No substantial precipitation was recorded (0.22 in. on 
4 August). Prolonged periods of slightly depressed barometric pressure and elevated 
relative humidity would normally be a reliable predictor of sufficient rainfall to pro- 
duce new growth on Celtis which provides requisite oviposition sites. 


RayMonpd W. Neck, Pesquezo Museum of Natural History, 6803 Esther, Austin, 
Texas 78752. 


VOLUME 31, NUMBER 3 L738 


STUDIES ON RESTINGA BUTTERFLIES. I. 
LIFE CYCLE AND IMMATURE BIOLOGY OF MENANDER 
FELSINA (RIODINIDAE), A MYRMECOPHILOUS METALMARK 


Curtis J. CALLAGHAN 
IBM do Brasil, P.O. Box 1830, Rio de Janeiro, Brazil 


This paper is the first of a series that deals with various aspects of 
the ecology of butterflies which inhabit the sandy coastal plains near 
the sea in southern Brazil. Called the “restinga” in Portuguese, it is an 
unique habitat that contains a large number of endemic plants and ani- 
mals. Unfortunately, it is valuable land for real estate development 
and at present is being altered to make way for new residential and 
resort areas. Also, the new coastal highway from Rio de Janeiro to 
Santos, now under construction, is altering great expanses of restinga 
vegetation and in the process is opening up previously isolated areas to 
commercial exploitation. All this underlines the urgency with which 
the Brazilian coastal flora and fauna must be studied before their 
demise in the wake of “civilization.” 

The purpose of this paper is to describe the habitat and larval biology 
of Menander felsina (Hew.), an endemic restinga butterfly. The dis- 
cussion will include descriptions of oviposition, larval behavior, and re- 
lationships with ants. A future paper will deal with other aspects of 
adult behavior and population dynamics. 

Published information on the larval biology of the neotropical rio- 
dinids is very scanty, especially considering that there are over 1500 
recognized species. The majority of references deal with foodplant 
records or a description of larvae and pupae (Zikan, 1920, 1953; Stichel, 
1920; Beutelspacher, 1972; Monte, 1934; Raymundo, 1907; Santos, 1931; 
Gomes, 1940; Biezanko, 1949; Bertels, 1954; Mabilde, 1896; Ronna, 
1933; Mariconi, 1961; Brandao, 1942). To date, I know of only four 
studies on the relationship of riodinid larvae with ants, these being Ross 
(1966), Guppy (1904), Bruch (1926), and Bourquin (1953), which 
cover a total of six species. In contrast, there are 245 known myrmec- 
ophilous lycaenid larvae (Downey, 1961). In the present paper, the 
riodinid list is increased to seven species, a number which I suspect 
will grow in time with additional observations. 

The range of M. felsina is coastal Brazil from Joinville, Santa Catarina 
to Rio de Janeiro, the latter being the type locality (Hewitson, 1863). 
Seitz (1916) gives the distribution as being southern Brazil, and Stau- 
dinger (1888) mentions a specimen from Santa Catarina. The male 


174 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


butterfly described by Butler (1877) is undoubtedly something else 
since it is nothing like those from southern Brazil. Certainly the con- 
fusion in this case resulted from Hewitson (1863-1878) illustrating 
only the female, leaving the male to the imagination. Material ex- 
amined in the Museu Nacional and the Instituto Oswaldo Cruz col- 
lections indicates that the species is confined to the narrow coastal strip 
from Rio to Joinville, there being in addition to these localities speci- 
mens from Angra dos Reis between Rio and Santos. 


Study Area and Methods 


The study location is in a small woods at the base of a large rock 
(Fig. 1) some 20 m high called Pedra de Itauna at km 11 of the Rio- 
Santos highway. Here, the highway runs along a sand bar or “restinga” 
ca. 20 km long and 2 km wide that separates some small brackish water 
lakes from the open coast. An annual rainfall of ca. 1.5 m and an average 
temperature of 23°C place the area within the subtropical-humid classifi- 
cation of Holdridge (1967). The vegetation is not what one would expect 
in a subtropical-humid zone, however, since the sandy soils quickly 
drain moisture away, giving the landscape a rather desert-like ap- 
pearance. The flats on the seaward side are covered with meter-high 
scrub that consists of bromeliads, cactus (Cerus sp.), and low 
deciduous bushes intermixed with grasses and other annuals such as 
Sporobohas virginicus and Panicum racemesum and with patches of low 
woods ca. 5 m high in protected localities. The vegetation in these latter 
areas is very dense, making penetration difficult except by man-made 
paths and in occasional sandy clearings, 5-10 m in diameter, which are 
encountered sporadically in the woods. 

Typical genera of trees and bushes are Myrsine, Ocotea, Myrcia 
Tabebuia, Clusia, Eugenia, and Psidium. The forest floor is packed 
knee deep in bromeliads, and the trees and bushes are festooned with 
mosses, lichens, and other epiphytic plants that take advantage of the 
frequent fogs which roll in from the sea. Bromeliads are represented 
by the genera Nidularium, Billbergia, Aechmea, Vriesia, and Tillandsia; 
Araceas by Philodendron and Anthurium; the Pteridophytes by Ly- 
copodium, Polypodium, Polystichum, and Elaphoglosum; and the Gesner- 
aceae by Codonanthe and Hypocyrta among others (Joly, 1970). Gallery 
forests with trees up to 10 m high are found in the better protected 
parts on the leeward side of the restinga. These support a flora and 
fauna similar to those farther inland, with Heliconia plants and other 
shade-loving varieties replacing the bromeliads on the forest floor. 

The habitat of M. felsina is the area of low woods just inside the 
seaward edge. Here, the adults can be found feeding on flowers and, 


VOLUME 31, NUMBER 3 175 


Figs. 1-9. 1, Habitat of Menander felsina showing woods at base of rock; 2, 
Norantea brasiliensis, foodplant of M. felsina larvae; 3, M. felsina, female on leaf 
of foodplant; M. felsina, third instar larva; 5, M. felsina, fifth and sixth instar larvae; 
6, ant (Camponotus crassus ssp.) “drumming” on larva of M. felsina; 7, ant 
(Camponotus crassus ssp.) feeding at honey glands of M. felsina; 8, M. felsina, 
prepupal larva, note honeydew on upper part of larva; 9, M. felsina, pupa. 


in the case of males, patrolling along paths and in the sandy clearings. 
The M. felsina will fly outside the woods on the flats for oviposition and 
feeding. 

The observations that form the basis of this study were made over a 


176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Ws I 


Figs. 10-12. Menander felsina: 10, egg; 11, first instar larva; 12, final instar 
larva, side and top views. 


period of four years and some 46 visits. Ova were obtained after ob- 
serving Ovipositing females in the field and immatures were aquired 
by rearing them from eggs or collecting them on the foodplant, Noran- 
tea brasiliensis Choisy (Marcgraviaceae) (Figs. 2, 3). These were 
brought into the lab with the foodplant and raised in fruit jars. Attempts 
to induce females to oviposit on foodplant in the lab were unsuccessful; 
the insects remained in the corner of the cage nearest the light until 
dying three or four days after being caged. Of the 10 larvae brought in 
from the field, six pupated and four emerged as adults. Causes of 
death for larvae appeared to be injuries due to handling. The reasons 
for the pupae not emerging are unknown. In no case, however, were 
parasites found on larvae or pupae. Of eight eggs collected, five 
hatched and two larvae were raised to the third instar. Some larvae 
died in the first instar because we had no foodplant for them. 


VOLUME 31, NUMBER 3 LTT. 


Observations on ants were made both in the field and in the lab, and, 
in the latter case, on ants that were kept with the larvae. 


Description of Immature Stages 


Egg (Fig. 10). About 0.4 mm in diameter, flattened above and below, giving 
it the appearance of a rounded tire, covered with network of lines. Micropyle a small 
circle with a dot in the middle. Color brown, matching stem on which egg 
is laid. Duration: 10-11 days. 

First instar larva (Fig. 11). Newly hatched larva 1.3 mm in length; light green 
in color except for head and prothorax, which are dark brown. Forward edge of 
prothorax with 10 long, flexible setae extending over head. Rest of thoracic and 
abdominal segments have scaly lateral protrusions, three to a segment, the middle 
one being the largest, and a double row of black spots on dorsum. A transparent 
carapace attached to anal segment and extending past the rear of the larva has 
six long, flexible setae along its edge similar to those on prothorax. Very small, in- 
conspicuous spiracles located along sides of abdominal segments 1-8 and on supra- 
pedal lobe of prothorax. Duration about 6 days; head capsule width 0.2 mm. 

Second instar larva. Coloring as in first instar but with scattered green specks 
on body. Setae of the prothorax same as in first instar only with two having de- 
veloped into flattened lobes reaching the length of head. The scaly lateral pro- 
trusions present but folded under body. Plate at end of abdomen has setae as in 
the first instar, though proportionally reduced in length. Spiracles as in first instar. 
Duration 4 days; total body length 3 mm; head capsule width 0.3 mm. 

Third instar larva (Fig. 4). Coloring as in second instar. Head brown with 
white specks on face. Setae of prothorax reduced to two lobes with one long seta 
coming from base of each and with two shorter ones on the sides. Tail plate setae 
reduced to small projections, and protrusions on sides of body missing. Honey 
glands on eighth abdominal segment well-developed with spiracles immediately 
above them. Other spiracles as in first instar. A small slit found on upperside of 
metathorax, corresponding to the tubercles mentioned by Ross (1966). Duration 5 
days; total body length 5 mm; head capsule width 0.5 mm. 

Fourth instar larva. Color as in third instar. Setae of prothorax consist of 
two groups of three, flat, elongated lobes, one larger than the other two. Two large 
setae found on sides of prothorax. Legs bear tufts of setae, otherwise similar to 
third instar. Duration 4 days; total body length 7 mm; head capsule width 0.9 mm. 

Fifth instar larva (Fig. 5). Similar to fourth instar. Duration 5 days; total body 
length ca. 16 mm; head capsule width 1.2 mm. 

Sixth instar larva (Figs. 5, 12). Similar to fourth and fifth instars only larger. 
Four days after molting larva enters into prepupal stage, the body turning mottled 
brown, except for patch of original green color dorsally. Prepupa (Fig. 8) lasts for 
5 days, after which the pupa is formed. Length of prepupal larva 22 mm; head 
capsule width 1.8 mm. 

Pupa (Fig. 9). Light to dark brown mottled with greenish shading dorsally, 
attached by thin girdle passing over middle of body and by silk pad under last 
segment. Wing cases extend halfway along sides. Duration 22 days; total length 
21.5 mm; width at widest part of body 5 mm. 


DISCUSSION 


The foodplant (Norantea brasiliensis) of M. felsina at the study lo- 
cation is woody with rounded succulent leaves growing alternately 
around stems that spread out in all directions from the base of the 
plant to a maximum height of 1.5 m. The flowers appear from late 


178 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


September-January in the form of small red balls with nectar caps 
hanging below them growing off an extension of the stem. Individual 
plants are widely but sparingly distributed around the restinga, gen- 
erally in the protection of clumps of larger bushes. All plants dis- 
covered in the study area showed signs of larval damage. 

Oviposition occurs from 1230-1400. The female alights near the end 
of a stem of N. brasiliensis and walks down it in a spiraling motion ex- 
ploring the surface with her abdomen. A single egg is quickly deposited 
on the main stem near the base of a leaf. She then flies a short distance 
away to rest on the upper surface of a leaf, wings outspread. Within 
two minutes she returns to the food plant, depositing another egg on a 
different stem of the same plant or on a second plant if there is one in 
the same vicinity. One female was observed placing four eggs within 
12 min. A total of five ovipositions were observed, and in no case were 
more than four eggs deposited by any one female in any given session. 

The tiny larvae hatch 10-11 days after oviposition and immediately 
move up to the leaf buds of the plant, where they begin to feed. They 
spend their time hidden inside the curled portion of the bud, betraying 
their presence only by the frass that they leave outside the opening. 
After the second molt they move out and start feeding on the older 
leaves as well as the buds, always eating the edges and ends of the 
leaves. Feeding is during the early morning, late afternoon, and evening 
hours and on cloudy days. During the hot part of the day the larvae 
move to cooler spots under leaves or near the stem of the plant and re- 
main motionless, their flat profiles and green color allowing them to 
blend in perfectly with the leaves on which they rest. To determine 
whether they were active at night, two visits were made to the food- 
plant after dark. Using a flashlight with a piece of red plastic over the 
lens, two larvae were located resting on the plant. They showed no 
interest in feeding or any other activity during the 2 hours they were 
watched. This inactivity during the night was noted with larvae raised 
in the lab. However, molting, entering into the prepupal stage, and pupa- 
tion all took place during the night in the lab. Larvae of about the same 
size will remain together for feeding and pupating. However, it was 
more common to find solitary larvae, one to a stem of foodplant. When 
larvae of different instars were raised together, they showed no in- 
terest in each other and always fed separately on different leaves. No 
cannibalism was observed. When moving onto a leaf the first time, they 
always laid down a trail that consisted of strands of silk with a weaving 
motion of the head. This trail was used to maintain a grip on the sur- 
face of the leaf. If knocked off, the larvae would invariably remain 
dangling by a thread of silk. 


VoLUME 31, NUMBER 3 179 


Starting with the third instar, the M. felsina larvae are myrmecophilous, 
being attended by ants that are a subspecies of Camponotus crassus. 
The early instars are completely ignored by ants. Upon encountering 
a larva, the ants will caress it with their antennae, rapidly alternating 
the strokes, first with one antenna and then the other, much as one 
would play a drum (Fig. 6). The larva responds to this “drumming” 
by extending two small glands located on the eighth abdominal segment 
from which is emitted a small quantity of clear liquid that the ants 
eat (Fig. 7). The ants remain in the vicinity of the larva for some time, 
returning now and then for more “drumming” and liquid. In the lab, I 
tickled a larva with a hair, ant fashion, and managed to get it to extend 
the glands. But these were quickly withdrawn and the larva started to 
move off, showing signs of being disturbed. Undoubtedly the larva was 
able to distinguish between ants and other stimuli. 

In fact, an ant need not actually caress the larva; its mere presence 
nearby is enough to cause the larva to protrude the organs, an action 
that may serve as an attractor for the ants. The means by which the 
larvae know when an ant is near could not be determined during the 
study. 

The honey glands located on the eighth abdominal segment were the 
only active myrmecophilous structures observed. On the metathorax 
are two slits high on the back corresponding to the lateral tubercles of 
Ross (1966), but prolonged observation of these under magnification 
failed to show any physiological change in them, even with ants in 
attendance. This is in contrast to the protrusion of the lateral tubercles 
noted in various other riodinid larvae such as Lemonias rossi (Ross, 
1966) and Audre sp. (Callaghan, in prep.); these tubercles possibly 
serve to attract ants or frighten predators (Ross, 1966). Likewise, the 
“horns” of the prothorax in M. felsina appeared completely passive in 
relation to ants, unlike the vibrating pupilla of L. rossi. When disturbed, 
the M. felsina larvae nod the head rapidly up and down, creating vibra- 
tions inaudible to the human ear, but this seemed not to be directly re- 
lated to the relationship with ants. The function of the “horn” struc- 
tures in M. felsina remains a mystery, though, since ants were observed 
stroking the head and neck area, they could have an attractive or sen- 
sitive role. 

The ants frequent the N. brasiliensis plants, eating plant secretions. 
When a larva is located, the discovering ant will communicate this in- 
formation to nearby ants. An attendant ant was observed leaving a larva, 
walking over to another ant with whom it touched antennae, and being 
followed back to the larva, where both ants commenced “drumming” 
and partaking in the larva’s fluids. Soon the first ant left, and the second 


180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


remained in attendance. Similar observations were made by Ross (1966), 
though in this case one ant passed honeydew to the other to communicate 
the presence of larvae. 

It is apparent that the ants protect the larvae. In no case were larvae 
found parasitized by wasps, tachinid flies, or fungi, the constant attention 
of the ants undoubtedly serving to discourage this type of predation 
(Edwards, 1878). Norantea brasiliensis frequented by ants were largely 
free of other insect life, the exceptions being flying insects visiting the 
flowers and another lepidopterous larva found feeding on the leaves, 
but not attended by the ants. In contrast, other kinds of plants had numer- 
ous spiders and other small insects. To test the reaction of the ants to in- 
truding organisms, small spiders were released on the N. brasiliensis near 
ants attending larvae. Upon discovering the intruders, the ants took up 
a defensive position, typical of Camponotus, by rearing up on the hind 
legs with the abdomen tucked between them. The intruders always 
beat a hasty retreat under these circumstances. 

Observations of M. felsina indicate that the larvae remain on the 
foodplant 24 hours a day from hatching until pupal diapause, always 
with the ants nearby. The only time when larvae were discovered with- 
out ants was on one particular plant during a rainstorm. 

Shortly after the fifth instar, the larva moves off the leaf where it 
had fed and crawls down the stem to the lower part of the plant. Here, 
it takes a head-down position on the stem or among the leaf litter nearby 
and assumes the mottled brown-green color of the prepupa. Observa- 
tions of specimens in the lab indicate that this takes place at night. If 
disturbed, the larva will move off to find a more suitable spot. During 
this stage, the honey glands often secrete large quantities of honeydew, 
thus attracting the ants, which can always be found around the prepupal 
larvae. This undoubtedly provides protection during a very vulnerable 
period in the larva’s life. After 5 days, the larval skin is shed and the 
chrysalid case hardens, again taking place during the night-time hours. 
The ants at this point lose interest and abandon the pupa. The imago 
emerges 21 days later. 


ACKNOWLEDGMENTS 


I wish to thank Drs. Woodruff Benson and Keith Brown of the 
Universidade Estadual de Campinas for reading and making helpful 
suggestions on the manuscript, Drs. Luiz Emilio de Mello Filho and 
Alfredo Rei de Rego Barros of the Museum Nacional for determination 
of the foodplant and use of the museum facilities, and Dr. Cincinnato 
R. Gongalves of the Universidade Federal Rural do Rio de Janeiro for 
determining the ant species. 


VOLUME 31, NUMBER 3 181 


LITERATURE CITED 


BerTEts, A. 1954. Trabalhos entomoldégicos no Instituto Agronémico do Sul. Bol. 
Tec. Ind. Agron. Sul. R. S., Maio 10: 1-68. 

BEUTELSPACHER, C. R. 1972. Some observations on the Lepidoptera of Brome- 
liads. J. Lepid. Soc. 26: 133-136. 

Brezanxo, C. M. 1949. Relac&o dos principais insetos prejudiciais observados nos 
arredores de Pelotas nas plantas cultivades e selvagens. Agros Pelotas, Setembro 
2(3): 15-313. 

Bourguin, F. 1953. Notas sobre la metamorfosis de Hamearis susanae Orfila 
1953 con oruga mirmecofila {Lep. Riod.). Rev. Soc. Ent. Argent. 16: 83. 
BRANDAO FinHo, J. S. 1942. As orquideas sdo também parasitadas por pragas e 

doencas. Sit. e faz. Sao Paulo, Jul. 7(7): 54-56. 

Brucu, C. 1926. Orugas mirmecofilas de Hameris (sic) epulus signatus Stich. 
Rev. Soc. Ent. Argent. 1: 1-9. 

Butier, A. 1877. Mr. A. G. Butler on the Lepidoptera of the Amazons. Trans. 
Ent. Soc. Lond. 1877: 126-143. 

Downey, J. C. 1961. Myrmecophily in the Lycaenidae (Lepidoptera). Proc. 
North-Central Branch Ent. Soc. Amer. 16: 14-15. 

Epwarps, W. H. 1878. Notes on Lycaena pseudargiolus and its life history. 
Can. Ent. 10: 1-44. 

Gomes, J. G. 1940. Chave de campo para determinacao das principais pragas 
dos citrus. Rev. Soc. Bras. Agron. Rio de Janeiro, Mar. 3(1): 58-108. 

Guppy, J. 1904. Notes on the habits and early stages of Trinidad butterflies. 
Trans. Ent. Soc. Lond. 1904: 225-228. 

Hewirson, W. C. 1863. Illustrations of new species of exotic butterflies. 5(3). 
John Van Voorst, London. 

Hoxuprince, L. R. 1967. Life zone ecology. Revised Edition. Tropical Science 
Center, San Jose. Costa Rica. 206 p. 

Jory, A. B. 1970. Conheca a vegetacio Brasileira. Ed. Univ. de Sao Paulo e 
Poligono. 165 p. 

Masitpz, A. P. 1896. Guia pratico para os principiantes colecionadores de in- 
setos. Porto Alegre. Grundlach & Schuldt. 238 p. 

Mariconi, F. 1961. Contribuicfo para o conhecimento de alguns insetos que 
depredam a goiabeira (Psidium guajava). Publ. Cientifica no. 2, Inst. Gen. 
Esc. Sup. Agric. “Luiz de Queiroz,” pp. 35-67. 

Monte, O. 1934. Borboletas que vivem em plantas cultivadas. Bol. Agric. 
Zootech. Vet., Minas Gerais. 7(3 & 4). 

Raymunpo, B. 1907. Contribuicdo para a hist. Nat. dos lep. do Brasil, III Con- 
greso Cintifico Latino-Americano, Rio de Janeiro, 1905. Tomo III, Livro 
B.—Imprensa Nac. 

Ronna, E. 1933. Catdlogo dos insetos até hoje nas plantas do Rio Grande do Sul, 
Egatéa P. Alegre, 18(1-2): 47-53. 

Ross, G. N. 1966. Studies on Mexican butterflies IV. The ecology and ethology 
of Anatole rossi, a myrmecophilous metalmark (Lep: Riodinidae). Ann. Ent. 
Soc. Amer. 59: 985-1004. 

Santos, E. 1931. Inimigos e doencas das fruteiras. Edicao Bibl. Agr. D’ “O 
Campo,” Rio de Janeiro. 

Serrz, A. (ed.) 1916. Macrolepidoptera of the World, Vol. 5, American Rhapalo- 
cera. Alfred Kernen Verlag, Stuttgart, 1139 p. 

STAUDINGER, O. 1888. Exotische schmetterlinge von Dr. O. Staudinger und Dr. 
E. Schatz. I. Theil. Fuerth G. Loewonsohn, 333 p. 

SticHEL, H. 1920. Biologische beitriigue zur schmetterlingsfauna Brasilians. z.d. 
Deut. Ver. f. wiss. u. Kunst in S40 Paulo 1: 145-157. 


182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ZIKAN, J. F. 1920. Biologische Beitrage zur Schmetterlingsfauna Brasilians. z.d. 
Deut. Ver. f. wiss. u. Kunst in S40 Paulo 1: 145-157. 

. 1953. Beitrage zur Biologie von 19 Riodinininen-Arten (Riodinidae- 

Lepidoptera). Dusenia 4(5, 6): 403-418. 


NOTES AND NEWS 


Letters to the Editor 
Dear Mr. Godfrey, 


Referring to the article of Mr. Hans Epstein in “Notes and News” in the Journal 
of the Lepidopterists’ Society, vol. 31, number 1, pp. 73-74, I have to protest strongly 
against Epstein’s remark that “F. Bryk and C. Eisner . . . are heavily responsible for 
the oversplitting; for instance, by now 200 odd subspecies of Palaearctic P. apollo 
L. have been described.” I regard it as a malicious remark. Objectively, many more 
apollo subspecies have been described by other authors than Bryk and myself. 


Sincerely yours, 
Curt EISNER 


Dear Dr. Godfrey: 


The repeated recent mentioning of Papilio xuthus occurring in Hawaii (vol. 30: 
149 and vol. 31: 75) and the conjecturing about from where it may have immigrated 
needs a clarification from local Hawaiian records. The first specimen was recorded 
during April 1971 at Salt Lake, Honolulu, near Hickam Air Force Base. The species 
spread rapidly in the following years over all the Islands: 1972 Kauai, 1974 (June) 
Maui and Hawaii, 1974 (August) Molokai, 1974 (September) Lanai. Hawaiian 
entomologists are convinced that it came here by means of military planes, probably 
from Guam where it previously had its easternmost distributional limit. It is not 
alone in its way of immigration. Over the last few years we got also from Guam the 
banana skipper (Erionota thrax), the first specimen of which was caught at Hickam 
Air Force Base in August 1973. Three recent newcomers among the Sphingidae were 
also first recorded around Hickam Air Force Base. Their easternmost distributional 
limit before they came here was Okinawa: Theretra nessus (August 1974), 
Deilephila nerii (September 1974), Macroglossum pyrrhostictum (July 1976). Very 
often live specimens are intercepted by the quarantine service of the Department of 
Agriculture. So was recently recorded a 2 of Psilogramma menephron on a military 
plane from the West Pacific (Guam?). Also commercial planes occasionally bring 
in interesting species: in a PAN AM plane from Pago Pago (Samoa) came a live ¢ 
of Deilephila torenia, a beautiful and rare, dark olive-green phenotype. One of the 
latest live interceptions on a military plane was a $ of Hippotion boerhaviae from 
Guam. Among the butterflies there suddenly appeared this year (January) large 
numbers of the gulf fritillary ( Agraulis vanillae), and nobody has an explanation how 
that may have happened. 

Sincerely, 
J. C. E. Riorre 


VoLUME 31, NUMBER 3 183 


BIONOMIC NOTES ON SOME BAGWORM MOTHS 
(PSYCHIDAE) OF TEXAS 


RayMonpD W. NEcK 
Pesquezo Museum of Natural History, 6803 Esther, Austin, Texas 78752 


Jones & Parks (1928) listed 13 species of bagworms that were known to 
occur in the state of Texas with an additional three species whose 
presence was “anticipated.” Four were later relegated to synonyms at the 
species level and two were apparent errors, leaving a total of seven species 
that actually had been definitely recorded from Texas (Davis, 1964). The 
first Texas record of one of the “anticipated” species has only recently 
been reported (Neck, 1976), bringing the total to eight. 

The four synonyms followed by their present status are as follows: 
(1) Thyridopteryx vernalis Jones = T. ephemeraeformis (Haworth); (2) 
Oiketicus bonniwelli Barnes & Benjamin = O. townsendi townsendi 
Townsend; (3) Oiketicus dendrokomos Jones = O. townsendi dendro- 
komos Jones; and (4) Platocoeticus jonesi Barnes & Benjamin = Crypto- 
thelea gloverii (Packard). The first error is an “Oiketicus sp.” which 
Jones & Parks (1928) state exists at high elevations in extreme western 
Texas and southeastern Arizona. Davis (1964) lists this form under 
“unidentified larval cases.” He speculates that this may be an extreme 
northern extension of Oiketicus kirbyi, but his records are restricted to 
Arizona. The other species, Prochalia pygmaea Barnes & McDunnough, 
is more of a mystery. Jones & Parks (1928) state that it occurs from 
Bexar Co. (San Antonio) southward, whereas Davis (1964) records only 
a questionable identification at Dickinson (between Houston and Gal- 
veston) and states that the range is “much of the Atlantic and Gulf 
Coastal Plain from South Carolina to perhaps as far west as Texas.” His 
westernmost confirmed locality is New Orleans. 

Five species of bagworm moths are discussed below. Thyridopteryx 
meadi H. Edwards and Oiketicus townsendi Townsend (with two sub- 
species ) are not discussed because of the author's unfamiliarity with these 
two species. No further discussion of Oiketicus toumeyi Jones is required 
because the few data accumulated for this species in Texas have been 
previously published (Neck, 1976). 


Thyridopteryx ephemeraeformis (Haworth) 
Thyridopteryx ephemeraeformis (Haworth), the evergreen bagworm, 
is quite common over most areas of the eastern United States, frequently 
becoming a local economic pest on various species of evergreens. Craig- 
head (1950) reported that this species is known to flourish in Texas on 


184 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


cedars (Juniperus spp.), willows (Salix spp.), and cypress (bald cypress, 
Taxodium distichum (1L.) Rich.). 

In early June 1969, the author received a collection of this species that 
had been feeding on Barbados cherry (Malphigiaceae: Malphigia glabra 
L.) although two specimens had switched to Japanese yew (Taxaceae: 
Taxus cuspidata Sieb. & Zucc.) and another two had switched to an orna- 
mental holly (Aquifolicaeae: Ilex sp.). These latter two plants were 
located within several feet of the Barbados cherry. Unopened blossoms 
of the Barbados cherry were utilized in construction of the larval bags. 

Bagworms are generally found on a large number of larval foodplants 
(see records in Davis, 1964) and change foodplants during their larval 
lifetime. Since this collection contained 52 individuals that had remained 
on Barbados cherry, a larval foodplant crossover experiment was devised. 
Rather than continuing to feed the larvae on Barbados cherry, they were 
fed pyracantha (Rosaceae: Pyracantha coccinea Roem.) for an initial 
4-week period. For a second 4-week period, the larvae were switched to 
gum-elastic (Sapotaceae: Bumelia lanuginosa ( Michx.) Pers.). At the end 
of the first four-week period a total of 17 live larvae and 11 pupae out of 
the original collection were still alive. A total of 24 larvae had died. By 
the end of the second phase of the experiment, all of the remaining larvae 
had died. Of the 11 pupae (104; one 2), only four emerged (32; one 
2). Although individual bagworm larvae have been reported to switch 
foodplants (as indeed, four of the collection had switched), this particu- 
lar pair of plants was found to be unacceptable to these larvae with few 
exceptions. No larvae were able to survive the second change of food- 
plant. 

The single female emerged in mid-July with male emergence occurring 
in late July and early August. The female pupal case measured 27 mm in 
length. The mean pupal case length and mean larval bag length for males 
were, respectively, 17.7 (SE = 0.19: N = 3) and 40.7 mm(Sie—e2.a- 
N = 3). The same two measures for the dead pupae (those which never 
emerged) were 15.9 (SE = 0.6; N =7) and 42.9 mm (SE = 1.1; N =7). 
Student's t-tests revealed that the difference between the respective 
means of the dead and emerged pupae was significant for pupal case 
length (t = 2.7; p < 0.05) but not for larval bag length (t = 1.5; p > 0.1). 
This difference in pupal length represents variations in the amount of leaf 
material consumed by the larvae. However, the fact that the larval bag 
length of the dead pupae was equal to (actually slightly longer than) 
that of the emerged pupae indicates that these dead pupae represented 
larvae which had shrunk below each individual larva’s maximum size 
because of limited or total refusal to feed on the foodplant material pro- 
vided in the laboratory. Emergence of an adult from a pupa is at least 


VoLUME 31, NUMBER 3 185 


partially dependent upon the amount and quality of resources ingested 
by a larva even though the ingestion level may well have been sufficient 
for pupation. 

A further collection of T. ephemeraeformis from Lubbock, Lubbock 
Co., was received in early October 1969. These individuals, which had 
been feeding on ornamental eastern red cedar (Cupressaceae: Juniperus 
virginiana L.), included two live male pupae, nine live female pupae, one 
empty female pupal case, two dead larvae, and two parasitized larvae. One 
male emerged on 15 October with five females emerging by 14 October. 
Parasites that had attacked one of the larvae had emerged through exit 
holes, leaving cocoons similar to those constructed by Iphiaulax (Hyme- 
noptera: Ichneumonidae). Wasps in the other bag were preparing to pu- 
pate when received. The adult wasps, which emerged on 14 October, were 
identified as a hyperparasite, Habrocytus thyridopterigis How. (Pteromali- 
dae), which has been reported from this bagworm species in West Virginia 
(Kulman, 1965). Judging from the size of the larval bag that contained 
these wasps, this hyperparasite attacks the parasites of rather young 
larvae. Dead specimens of Spilochalcis sp. (probably mariae (Riley) ) 
have been found in larval cases of this species. The mean larval bag and 
pupal case length for males were 38.5 (SE = 1.5; N = 2) and 15 mm 
(SE = 1.0; N = 2), respectively. The same measures for females were 
43.4 (SE = 1.1; N = 10). A t-test revealed significant differences be- 
tween males and females for both larval bag length (t = 4.1; p < 0.005) 
and pupal case length (t = 14.1; p < 0.001) despite the smal! sample 
sizes involved. 

Thyridopteryx ephemeraeformis has been found on numerous host 
plants in the Austin area. A large, extinct colony was discovered on 
Oriental arbor vitae (Pinaceae: Thuja orientalis L.). Specimens have 
also been found on this plant in San Marcos, Hays Co. In Austin, isolated 
larval cases have been found on peach (Rosaceae: Prunus persica 
Batsch), cedar elm (Ulmaceae: Ulmus crassifolia Nutt.) , plateau live oak 
(Fagaceae: Quercus fusiformis Small), turk’s cap (Malvaceae: Malva- 
viscus arboreus Cav. var. drummondii (T. & G.) Schery), Texas persim- 
mon (Ebenaceae: Diospyros texana Scheele), agarita (Berberidaceae: 
Berberis trifoliolata Moric.), pyracantha (Rosaceae: Pyracantha coccinea 
Roem. ), willow baccharis (Compositae: Baccharis salicina T. & G.), Mex- 
ican plum (Rosaceae: Prunus mexicana Wats.); bald cypress (Taxo- 
diaceae: Taxodium distichum (L.) Rich.), greenbriar (Liliaceae: Smilax 
bona-nox 1..), ashe juniper (Cupressaceae: Juniperus ashei Bucch.), 
Texas sugarberry (Ulmaceae: Celtis laevigata Willd.), slender bamboo 
(Gramineae: Bambusa sp.), and western soapberry (Sapindaceae: 
Sapindus saponaria L. var. drummondii (H. & A.) L. Benson). These 


186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


individual larvae apparently result from dispersal by first-instar larvae. 
These isolated larvae contained the two sexes in approximately equal 
numbers (eight ¢; six 2). 

In northeast Texas (Titus Co.), T. ephemeraeformis has been found 
feeding on the following plants: eastern red cedar (Cupressaceae: 
Juniperus virginiana L.), American elm (Ulmaceae: Ulmus americana 
L.), slippery elm (Ulmaceae: Ulmus rubra Muhl.), alder (Corylaceae: 
Alnus serrulata (Aiton) Willd.), and American hop hornbeam (Cory- 
laceae: Ostrya virginica (Miller) K. Koch.). 


Oiketicus abbotii (Grote) 


Oiketicus abbotii (Grote) is restricted to the Atlantic and Gulf Coast 
from Virginia to Texas (Davis, 1964). Although the possibility exists 
that O. abbotii is part of a polytypic species, O. kirbyi Guilding, which 
would include the two above species and O. townsendii, Davis (1964) 
reports that the larval bags of these three taxa are quite distinct. The 
larval bags of O. abbotii and T. ephemeraeformis are of similar size but 
are easily distinguished by a single character. Twigs on larval bags of 
O. abbotii are placed transversely so that a cross section of the bag 
reveals an angular polygon. Twigs on larval bags of T. ephemeraeformis 
are placed longitudinally so that a cross section of the bag is roughly 
circular. 

The author has found O. abbotii in Brownsville, Cameron Co., on a 
wide variety of host plants, e.g., tepeguaje or giant lead tree (Le- 
guminosae: Leucaena pulverulenta (Schlect.) Benth.), mesquite (Le- 
guminosae: Prosopis glandulosa Torr.), avocado (Lauraceae: Persea 
americana Mill.), Texas sugarberry (Ulmaceae: Celtis laevigata Willd.), 
Japanese honeysuckle (Caprifoliaceae: Lonicera japonica Thumb. ), arbor 
vitae (Pinaceae: Thuja orientalis L.), and common rose (Rosaceae: Rosa 
sp.). The only infestation which could be called a colony was found on a 
single pyracantha bush (Rosaceae: Pyracantha coccinea Roem. ). 

Collection of larvae from the aforementioned pyracantha bush oc- 
curred in mid-July 1967. Larvae were quite active at this time but be- 
came relatively inactive in the latter part of July although increased 
activity became apparent in early August (all larvae observed in labora- 
tory). Earliest adult emergence (?) occurred on 30 August, but the 
next one (2) did not emerge until 15 September. Final emergence ( ¢ ) 
occurred 4 October. Females tend to emerge earlier; five of the first six 
adults were female and the final three to emerge were males. 

Of 33 larval cases collected in late June and early July at various 
localities in Brownsville, seven (21.1%) showed signs of parasitism. In 
late June, 10 adults of Iphiaulax manteri Nett. (Hymenoptera: Braconi- 


VOLUME 31, NUMBER 3 187 


dae) emerged from one of the bags. The cocoons of the wasps were 
present in two layers of five each, with the individual cocoons being 
parallel to the long axis of the bag. Stephens (1962) reported Iphiaulax 
sp. [near I. sublucens (Blanch.) + I. diversus (Vierick)] as attacking 
O. kirbyi in Costa Rica. Larvae of this form act as ectoparasites that 
gradually make their way to the center of the host. 

In late July a mud wasp, Pachodynerus astraeus Cameron (Hyme- 
noptera: Vespidae) emerged from one larval case. Examination of the 
interior of the case revealed two additional wasps. Each of the three 
cells were separated by a 2 mm thick layer of dried mud. The entire 
lining of the bag was covered by a layer of dried mud of 1 mm thickness. 
The bottom part of the bag was closed by a plug that was sculptured by 
the mother wasp. Davis (1964) reports a similar occurrence with bags 
of O. toumeyi being used by Pachodynerus acuticarinatus (Cameron). 
Whether the wasps utilize only empty bags or kill the inhabitant larva 
is unknown. Several additional bags of O. abbotii containing similar wasp 
cells from which adult wasps had previously emerged were collected. 

Oiketicus abbotii also exhibits the common sexual dimorphism. Larval 
bag and pupal case mean lengths for males of the pyracantha colony 
were 50.5 (SE = 1.4; N = 6) and 18.7 mm (SE = 0.8; N = 6) respectively, 
whereas the two lengths for females were 61.0 (SE = 1.0; N = 5) and 
25.2 mm (SE = 0.9; N = 5). The means are significantly different be- 
tween the sexes for both larval bag length (t = 8.1; p < 0.001) and pupal 
Gasetlencth(t — 3-2: p < 0.001). 

Several additional infestations on pyracantha in Brownsville were 
discovered by or reported to the author. Explanation for this abundance 
on pyracantha at that time is unknown. Possibly this occurrence illus- 
trated a temporary host race as discussed by Jones & Parks (1928). A 
return to these infestation sites in the summer of 1969 revealed no popula- 
tions. If a host plant race had developed, its occurrence was temporary. 

At localities other than Brownsville, only isolated foodplant records are 
available. One larval bag was found on southwestern bernardia (Euphor- 
biaceae: Bernardia myricaefolia (Scheele) ) at Goliad State Park, Goliad 
Co. At Lake Corpus Christi State Park, San Patricio Co., larvae have 
been observed feeding on Texas persimmon (Ebenaceae: Diospyros 
texana Scheele), guayacan (Zygophyllaceae: Porlieria angustifolia 
(Engelm.) Gray), and little lead tree (Leguminosae: Leucaena leuco- 
cephala (Lam.) de Wit). 


Cryptothelea gloverii (Packard) 


Cryptothelea gloverii (Packard) occurs along the Atlantic and Gulf 
Coasts from South Carolina to Central America. The general form of the 


188 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


larval case in this species is similar to that of T. ephemeraeformis but 
only one-half to one-third the length. In Austin, C. gloverii has been 
found infesting retama (Leguminosae: Parkinsonia aculeata L.), western 
soapberry (Sapindaceae: Sapindus saponaria L. var. drummondii (H. & 
A.) L. Benson), Texas sugarberry (Ulmaceae: Celtis laevigata Willd.), 
and common yarrow (Compositae: Achillea millefolium L.). In Browns- 
ville, C. gloverii had been found on pyracantha (Rosaceae: Pyracantha 
coccinea Roem.) and lime prickly ash (Rutaceae: Zanthoxylem fagara 
(L.) Sarg.). The record on pyracantha was a fairly large colony on the 
same bush as the colony of O. abbotii. Small bits of leaves, bark and 
fruits were attached to the bags. The bags were found attached to al- 
most all parts of the plant, e.g., trunk, stems, leaves, and even other bag- 
worms. Larvae sought the security of the internal branches prior to 
pupation. 

Collections were made during the first half of July, at which time the 
bags varied from 5 to 16 mm in length, with the majority being from 13- 
15 mm. Adult emergence occurred during mid-July with male pupal case 
length (N = 5) being very constant at 6-7 mm. Numerous mites were 
noticed on many of the bags. These mites have been predaceous on eggs 
as reported by Stephens (1962) for O. kirbyi in Costa Rica. On 12 July, 
the hatching of a total of 245 larvae from one larval case within ca. 3 hr 
was observed, with most of the hatching occurring within the first 2 hr. 
Behavioral patterns, i.e., orientation of abdomen and construction on 
initial larval bag, of the larvae were similar to those reported by Kaufman 
(1968) for T. ephmeraeformis. 

A return trip was made in December 1969 to the Brownsville locality 
where O. abbotii and C. gloverii had been observed in the summer 
months of 1967. No trace of either species could be found. Food supply 
failure did not seem to be a logical reason for the local extermination 
because the pyracantha plant was in excellent condition and exhibited no 
evidence of previous extreme defoliation. Although there had been in- 
secticide use to the east and southeast (directions of prevailing winds), 
drift is thought to have been minimal because ground-level hand spray- 
ing was employed. Three bird nests, probably constructed by mocking- 
birds, Mimus polyglottus (L.) (Passeriformes: Mimidae), a voracious 
insectivorous bird, were present in the bush. Stephens (1962) reported 
damage to a substantial number of bags of O. kirbyi which he attributed 
to an unknown bird species. Davis (1964) believes that some bird pre- 
dation of bagworms (cases that exhibited holes) is the result of wood- 
peckers. 

Overzealous collecting can be ruled out because bags in the top of the 
bush were not collected. These upper bags were not on the bush at the 


VoLUME 31, NUMBER 3 189 


time of the re-check. This fact raises the possibility that Hurricane 
Beulah, the eye of which passed within 20 mi of the area in September 
1967, led to the decimation of this colony. 


Astala confederata (Grote & Robinson) 


Astala confederata (Grote & Robinson), the lawn bagworm, occurs in 
the eastern United States west to about the 100th meridian. Larvae have 
been found feeding on Texas spear grass, Stipa leucotricha Trin. & Rupr., 
and Johnson grass, Sorghum halepense (L.) Pers. (both Gramineae). 
The larval stage of this species is an inconspicuous ground-level feeder 
until it attaches to a tree trunk or building wall and thereupon pupates 
in April and May (Jones & Parks, 1928). A series of bags was collected 
at Austin in mid-May 1969. Most bags showed prior adult emergence, 
although one bag still contained a live larva. Adult emergence from those 
bags containing live pupae occurred for several days following collection, 
with several egg masses present in other bags hatching on 1 June. The 
adult male of A. confederata is somewhat larger than the previous species 
and much darker, being almost black. 


Astala edwardsi (Heylands ) 


Astala edwardsi (Heylands), the chalk-hills bagworm, is restricted to 
Texas and Oklahoma (Davis, 1964), preferring barren areas where thin 
herbage only partially covers the limestone-derived soils. Jones & Parks 
(1928) report that the larvae eat both dead and living vegetable matter. 
I have observed one specimen feeding on thatch on the soil surface. The 
odd pencil-shaped bags are normally found on posts or tree trunks in 
September as the larvae ascend from the ground level. I have observed 
this species from the following localities in Texas: Crutchfield Ranch, 
Burnet Co.; 3 km S of Seguin, Guadalupe Co.; Austin, Travis Co.; and 
Longfellow, Pecos Co. 


ACKNOWLEDGMENTS 


I thank G. Ajilvsgi, W. Bleier, J. R. Crutchfield, and B. Maguire, Jr. for 
supplying some of the specimens. 


LITERATURE CITED 


CRAIGHEAD, F. C. 1950. Insect enemies of eastern forests. U.S. Dept. Agr. Mise. 
Rube 6o7: 679 p. 

Davis, D. R. 1964. Bagworm moths of the westem hemisphere. Bull. U.S.N.M. 
ZAA- 233 pp: 

Jones, F. M. & H. B. Parxs. 1928. The bagworms of Texas. Tex. Agr. Exp. Sta. 
Bull. 382: 26 p. 


190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


KauFMAN, T. 1968. Observations on the biology and behavior of the evergreen 
bagworm moth, Thyridopteryx ephemeraeformis (Lepidoptera: Psychidae). Ann. 
Ent. Soc. Amer. 61: 38-44. 

KutmMan, H. M. 1965. Natural control of the bagworm and notes on its status as 
a forest pest. J. Eco. Ent. 58: 863-866. 

Neck, R. W. 1976. Oiketicus toumeyi Jones: A bagworm moth new to the Texas 
fauna. J. Lepid. Soc. 30: 218. 

STEPHENS, C. S. 1962. Oiketicus kirbyi (Lepidoptera: Psychidae). A pest of 
bananas in Costa Rica. J. Eco. Ent. 55: 381-386. 


BUTTERFLIES ASSOCIATED WITH AN ARMY ANT SWARM RAID 
IN HONDURAS: THE “FEEDING HYPOTHESIS” AS AN 
ALTERNATE EXPLANATION 


Drummond (1976, J. Lepid. Soc. 30: 237-238) reported that a few female 
Mechanitis isthmia Bates (Nymphalidae: Ithomiinae) and male Graphium philolaus 
(Boisduval) (Papilionidae: Papilioninae) were attracted to a swarm raid of army 
ants (Eciton burchelli (Westwood) ) in Honduras. As an explanation, Drummond 
suggested a “reproductive odor hypothesis” to account for the attraction of female 
Mechanitis to the swarm raid: the strong, unpleasant odor mimicked the courtship 
scent of male Mechanitis, thus causing females of this species to follow the swarm. 
As Drummond indicated, the attraction of male Graphium to the swarm raid was 
puzzling since, like ithomiines, males produce the courtship scent. Recent literature 
on the feeding behavior of selected butterflies suggests an alternative explanation 
to these interesting observations. 

The acquisition of nutrients by adult butterflies of both sexes may be of widespread 
importance (Gilbert, 1972, Proc. Natl. Acad. Sci., U.S.A. 69: 1403-1407; 1976, 
Biotropica 8: 282-283; Arms et al., 1974, Science 185: 372-374). Adult butterflies 
are attracted to nutrient sites having an odor of decay (Gilbert, 1972, in litt.; Young, 
1975a, Stud. Neotrop. Fauna 10: 19-56; 1975b, Rev. Biol. Trop. 23: 101-123; 
Young & Muyshondt, 1973, Carib. J. Sci. 13: 1-49). Ithomiine butterflies are at- 
tracted to fresh deposits of bird droppings splashed on leaves (pers. obs.). Drum- 
mond (in litt.) suggested that the male Graphium might be responding to an odor 
stimulis that elicits food searching behavior. These observations suggest that the 
attraction to nutrient sites by adult butterflies involves odors of decay, although this 
is only speculation at the present time. 

Assuming that odors of decay cause the attraction of adult butterflies to nutrient 
sites, I suggest a “feeding hypothesis” as an alternative explanation for Drummond’s 
findings: both the Mechanitis and Graphium butterflies were being “fooled” by the 
swarm raid odors. The odors of decay associated with the swarm raid triggered food 
searching behavior by these butterflies, causing them to follow the army ants. Such 
an explanation accounts for the attraction of both sexes to the ants. Under the 
feeding hypothesis, the attraction of butterflies to puddling sites, bird-droppings, 
manure heaps, etc. is aided by responses to characteristic odors associated with these 
sources of nutrients (amino acids, sodium, etc. ). 


ALLEN M. Younc, Invertebrate Division, Milwaukee Public Museum, Milwaukee, 
Wisconsin 53233. 


VoLUME 31, NUMBER 3 191 


TWO NEW SPECIES OF CLEARWING MOTHS (SESIIDAE) 
FROM EASTERN NORTH AMERICA CLARIFIED 
BY SEX PHEROMONES 


W. DonaLp DuckworTH! AND THOMAS D. EICHLIN2 


Recently, J. H. Tumlinson and colleagues (1974) and Yonce et al. 
(1974) identified chemical compounds from females of Sanninoidea 
exitiosa (Say) (peachtree borer) and Synanthedon pictipes (Grote & 
Robinson) (lesser peachtree borer), which may prove to be major com- 
ponents of the female sex pheromone systems of the Sesiidae as a whole. 
Males of the peachtree borer respond to a mixture containing mostly the 
Z,Z isomer of 3,13-octadecadien-l-ol acetate (Z,Z ODDA), while 
responses of the lesser peachtree borer males are inhibited by the 
presence of even small quantities of the above isomer (Karandinos et al., 
1977). The E,Z isomer of ODDA is the major sex attractant for males of 
the lesser peachtree borer. 

Since the initial isolation of the two pheromones, studies have been, 
and are continuing to be, conducted utilizing the geometric isomers of 
ODDA singly and in various combinations in traps in a variety of environ- 
ments, mostly in Florida, Georgia, South Carolina, Ohio, Wisconsin, 
Washington and Calitornia. Cross attraction of different species to indi- 
vidual isomers or combinations has been demonstrated (Nielson & Balder- 
ston, 1973; Nielson et al., 1977; Karandinos et al., 1976). Most of the 
accumulated data on cross attractancy is still unpublished, but in sum- 
mary, about 30% (42 species to date) of the known North American sesiid 
fauna north of Mexico, representing the full phylogenetic range of the 
family, have been captured in traps baited with the pheromones. In addi- 
tion, a number of sesiid species have been captured using the pheromones 
in such widely differing areas as Mexico, Brazil, Costa Rica, Japan and 
Portugal (specific data as yet unpublished). 

While the major impetus for research in sesiid pheromone identifica- 
tion and field screening has been the development of control measures 
for pest species, it is becoming increasingly apparent that a valuable new 
tool for improving our general understanding of sesiid biology and evolu- 
tion has been discovered. There are significant gaps in our knowledge 
of sesiids, particularly in regard to distribution of species, relative 
abundance, seasonal periodicity, species diversity, etc., primarily due to 


1 Department of Entomology, National Museum of Natural History, Smithsonian Institution, 
Washington, D.C. 20560 : SULA. 

2 Laboratory Services/Entomology, Division of Plant Industry, California Department of Food 
and Agriculture, Sacramento, California 95814. 


192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the diurnal flight period and fugitive behavior of the adults, coupled with 
the endophagous boring habit of the larvae. Through the use of phero- 
mones for field sampling of adult males it is now possible to more readily 
detect the presence of sesiids in a given habitat and greatly enhance the 
investigation of life cycles, behavior, and related phenomena. 

In addition to the obvious benefits afforded by the use of sesiid sex pher- 
omones for sampling purposes, it is important not to overlook the funda- 
mental nature of pheromone systems in the evolution of the group. Clear- 
wing moths are remarkably homogeneous structurally (Duckworth & 
Eichlin, 1974) and distinguishing closely related species is frequently diffi- 
cult. Also, as our knowledge of distributions improves, many species previ- 
ously thought to be allopatric are proving to be at least partially overlap- 
ping in their ranges. In instances where closely related species occur 
sympatrically, chemical compatibility may play a critical role in achieving 
reproductive isolation. For example, two very closely related sesiid species 
which attack Viburnum, Synanthedon viburni Engelhardt and S. fatifera 
Hodges, occur sympatrically in Wisconsin. Recent investigations 
(Karandinos, per. com.; Roelofs & Comeau, 1969) suggest that repro- 
ductive isolation is achieved, at least in part, by each species utilizing a 
different isomeric pheromone system. This phenomenon for isolation 
is probably also operative in areas of probable sympatry (ie., Georgia 
and South Carolina) of Synanthedon kathyae n. sp. and the closely 
related species S. alleri (Engelhardt) which are discussed later in this 
paper. While it is overly simplistic to assume that pheromone systems 
alone have determined reproductive isolation in sesiid populations, it 
seems reasonable to assume that they are of increased importance in 
sympatric species where habitat preference, circadian and seasonal cycles, 
geographic distribution and other isolating mechanisms are less effective. 

Biological studies on various sesiid species have been initiated as a 
result of information initially gained through field testing of sesiid sex 
pheromones. These studies are also contributing to our understanding of 
the systematics of sesiids. For example, Nielsen & Purrington (1975) 
present data on flight periods of Podosesia syringae (Harris) in Ohio 
which suggests the existence of a previously undescribed, sympatric 
species of Podosesia which is temporally isolated from P. syringae but 
virtually indistinguishable structurally. Similarly, pheromone studies in 
South Carolina by R. L. Holloway, Clemson University, have uncovered 
the presence of a previously unknown species and helped to clarify the 
status of another undescribed species, both of which are described in 
this paper. 

As was the case in previous publications by the authors (Duckworth 
& Eichlin, 1973 and 1976), the following descriptions result from con- 


VoLUME 31, NUMBER 3 193 


tinuing revisionary studies on the Western Hemisphere Sesiidae and 
preparations for a fascicle on the Sesiidae for publication in The Moths of 
America North of Mexico. 

We wish to acknowledge with our appreciation the following indi- 
viduals and institutions who have provided specimens used in the present 
study: J. G. Franclemont, Cornell University, Ithaca, N.Y.; F. H. Rindge, 
American Museum of Natural History, New York, N.Y.; B. Wright, Nova 
Scotia Museum, Halifax; J. L. Sharp, Insect Attractants and Basic 
Biology Laboratory, USDA, ARS-Southern Region, Gainesville, Florida; 
C. E. Yonce, USDA, ARS-Southeastern Fruit and Tree Nut Research Sta- 
tion, Byron, Georgia; and R. L. Holloway, Clemson University, Clemson, 
South Carolina. For technical assistance we want to thank Laura S. 
Keller, University of California, Davis, for the drawings; Charles S. 
Papp, Scientific Illustrator, Special Services; and Magda R. Papp, Bio- 
logical Technician, Laboratory Services, California Department of Food 
and Agriculture. For numerous beneficial suggestions on improving the 
manuscript we wish to thank R. C. Froeschner and W. D. Field, National 
Museum of Natural History, Smithsonian Institution, and J. A. Powell, 
University of California, Berkeley. 


Genus Synanthedon Hiibner 
Synanthedon kathyae Duckworth & Eichlin, n. sp. 


Male: Antenna blue-black, clavate, tufted with scales apically, ciliate ventrally. 
Proboscis well-developed. Labial palpus smooth, yellow. Head with vertex blue- 
black, front blue-black, white lateroventrally, occipital fringe yellow. Thorax blue- 
black, with subdorsal yellow stripes, and mostly yellow laterally beneath wings. Abdo- 
men blue-black, dorsally with segments four and five yellow, ventrally yellow except 
segments two and three blue-black, anal tuft elongate, blue-black. Prothoracic leg 
mostly yellow, some blue-black often medially on coxa; mesothoracic leg blue-black, 
tarsi yellow; metathoracic leg with femur blue-black, tibia yellow with blue-black at 
base and on apical one-third, tarsi yellow. Forewing mostly hyaline, with very narrow 
margins, veins and discal spot blue-black, lightly powdered yellow on costal and anal 
margins dorsally, ventrally more strongly powdered yellow on margins and between 
veins apically. Hindwing hyaline, with narrow blue-black margins, costa yellow, fringe 
blue-black, becoming yellow at wing base. Male genitalia as in Fig. 1, typical of 
species placed in the genus Thamnosphecia (= Synanthedon) Spuler by Engelhardt 
(1946). Wing length of both sexes, 8-11 mm. 

Female: Antenna as for male but lacking ventral cilia. Scale patterns like the 
male, with slightly broader apical margin on forewing and anal tuft brush-like. Fe- 
male genitalia as in Fig. 2. 

Host: Unknown. 

Distribution: Halifax, Nova Scotia; Lewisboro, Westchester Co. and Long 
Island, New York; and near Oconee State Park, Oconee Co., South Carolina. 

Types: Holotype: ¢, Halifax, Nova Scotia, summer 1965, J. A. Godbout, Genitalia 
Slide ¢, by T. D. Eichlin, USNM 76020, deposited in Nova Scotia Museum, Halifax. 
Paratypes 5: 1 @ with same data as holotype, Genitalia Slide 2, by T. D. Eichlin, 
USNM 76036, in Nova Scotia Museum, Halifax; 1 2, Babylon, L.L., N.Y., 17.VII.87, 


194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-2. Ventral view, genitalia of Synanthedon kathyae: (1, left) male (left 
valve removed); (2, right) female. 


F. S. Blanton, Cornell University; 1 9, N.Y., Lewisboro, Westchester Co., 24-VII- 
1971, M. & T. M. Favreau, Genitalia Slide By M. R. Papp, CDA 225, in AMNH; 
2 6, Oconee County, SC, ZZ ODDA Pheromone, Date: 25-VI-76, R. L. Holloway 
Coll., one specimen labeled, Genitalia Slide By M. R. Papp, CDA 221, in NMNH. 

Discussion: On the basis of similar male genitalia, this species is closely related 
to Thamnosphecia alleri Engelhardt. The latter is known from Georgia, Florida, 
Alabama and Mississippi and differs from Synanthedon kathyae by having the fore- 
wings mostly opaque and the yellow markings replaced with orange. 

R. L. Holloway captured two specimens of S. kathyae in South Carolina in traps 
baited with Z,Z ODDA. By contrast, both J. L. Sharp in Florida and C. E. Yonce in 
Georgia have been collecting Thamnosphecia alleri in nearly all months of the year 
(57 captures in 1975-76) using the E,Z isomer only. 

Nothing is known of the biology of kathyae or alleri; Engelhardt (1946) believed 
the habitat of alleri to be open woodlands bordering on swamps. 

This species is known only from the six specimens of the type series. It is named 
for Kathy Eichlin, who plays a continuing supportive role in all of the sesiid studies. 


VOLUME 31, NUMBER 3 195 


Fig. 3. Ventral view, male genitalia (left valve and tegumen-uncus complex re- 
moved) of Carmenta odda. . 


Genus Carmenta Edwards 
Carmenta odda Duckworth & Eichlin, n. sp. 


Male: Antenna clavate, tufted with scales apically, ciliate ventrally, brown-black 
and with yellow dorsally. Proboscis well-developed. Labial palpus yellow dorsally. 
Head with vertex brown-black, mixed with yellow anteriorly, front rubbed on type, 
probably brown-black with white laterally, occipital fringe yellow, rubbed dorsally. 
Thorax brown-black, yellow subdorsal stripes, yellow beneath wings and on meta- 
thorax dorsally. Abdomen brown-black, dorsally with all segments narrowly banded 
posteriorly with yellow except third, fourth with widest band, ventrally with segments 
one and two solid pale yellow, four banded yellow, others with some yellow on poste- 
rior margin, anal tuft rubbed off. Legs missing from this specimen except forecoxae 
which are brown-black with yellow on lateral one-half. Forewing mostly hyaline but 
opaque costal margin spreading apically to cover area to below M: at wing margin, 
basal one-half with wing powdered orange on veins and margins, outer one-half of 
discal spot yellow-orange, yellow-orange more extensive ventrally but apparently 
not powdered in apical area. Hindwing hyaline, very narrow margins, some yellow 
powdering on costa. Male genitalia as in Fig. 3. Wing length, 9 mm. 

Host: Unknown. 

Distribution: Trenton, Edgefield Co., South Carolina. 

Holotype: ¢, Edgefield Co., SC., VI-11-1975, Coll. R. L. Holloway, ZZ phero- 
mone, Genitalia Slide By T. D. Eichlin, CDA 179, (U.S.N.M. No. 73592); in NMNH. 

Discussion: This species is described from a male specimen taken in a phero- 
mone trap coated with a sticky adhesive which resulted in the poor condition. How- 
ever, this capture represents the first new species in the U.S. to be discovered by the 
use of the sesiid pheromone, Z,Z 3,13-octadecadien-l-ol acetate and is named after 
this useful survey tool. 

Carmenta odda superficially resembles Ramosia arizonae (Beutenmiiller), Synan- 
thedon arkansasensis Duckworth & Eichlin, and S. refulgens (Edwards) (see Duck- 


196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


worth & Eichlin, 1973: Engelhardt, 1946), but the structure of the saccular ridge on 
the valva of C. odda is unlike any known species of Sesiidae in North America. 


LITERATURE CITED 


Ducxwortu, W. D. & T. D. E1rcuuin. 1973. New speces of clearwing moths 
(Lepidoptera: Sesiidae) from North America. Proc. Ent. Soc. Washington. 75: 
150-159. 

1974. Clearwing moths of Australia and New Zealand (Lepidoptera: 

Sesiidae). Smithsonian Contributions to Zoology. 180: 1-45. 

1976. A new species of clearwing moth (Lepidoptera: Sesiidae) from 
northern Mexico and southeastern Arizona. Proc. Ent. Soc. Washington. 78: 
304-308. 

ENGELHARDT, G. P. 1946. The North American clear-wing moths of the family 
Aegeriidae. Bull. U.S. Natl. Mus. 190: 1-222. 

Karanpinos, M. G., J. H. Tumuinson, & T. D. Ercutin. 1977. Field evidence of 
synergism and inhibition in the Sesiidae sex pheromone system. J. Chem. Ecol. 
3: 5/-64. 

NiELsEn, D. G. & C. P. Batperston. 1973. Evidence for intergeneric sex attrac- 
tion among aegeriids. Ann. Ent. Soc. Amer. 66: 227-228. 

NIELSEN, D. G., F. F. Purrtncton, J. H. Tumuinson, R. E. DoonirrLe, & C. E. 
YoncE. 1975. Response of male clearwing moths to caged virgin females, 
female extracts, and synthetic sex attractants. Environ. Ent. 4: 451-454. 

RoELors, W. L. & A. Comeau. 1969. Sex pheromone specificity: Taxonomic and 
evolutionary aspects in Lepidoptera. Science. 165: 398-400. 

TUMLINSON, J. H., C. E. Yonce, R. E. DoonirtLe, R. R. HEATH, C. R. Gentry, & 
E. R. MircHeti. 1974. Sex pheromones and reproductive isolation of the 
lesser peach tree and the peach tree borer. Science. 185: 614-616. 

YonceE, C. E., J. H. Tumuinson, C. R. Gentry, & E. R. Mrrcuetnt. 1974. Extrac- 
tion and field bioassay of the sex pheromone of the lesser peach tree borer. 
Environ. Ent. 3: 569-570. 


VOLUME 31, NUMBER 3 197 


GENERAL NOTES 


CATOCALA (NOCTUIDAE) SPECIES TAKEN IN CLAY COUNTY, 
TENNESSEE 


While collecting Lepidoptera in southwestern Michigan and on seven collecting 
trips of one or two annually to Clay Co., Tennessee from 1970-1976, I became 
aware of the large Catocala fauna of the latter area as compared to Michigan where 
many of the same species are rare or absent. Subsequently I made a special effort to 
collect this genus in Tennessee during 1975-1976. The majority of records listed 
below were made during this period. 


MATERIALS AND METHODS 


Collecting was conducted from 20 June-22 August within a two-mile radius of 
Celina in north-central Tennessee. The general countryside, being a part of the 
Cumberland Mountains, is hilly and heavily forested. Common trees here include 
White and Black Oak (Quercus alba L. & Q. ellipsoidalis Hill), Mockernut and Shag- 
bark Hickory (Carya tomentosa Nutt. & C. ovata Mill.), Honey Locust (Gleditsia 
triacanthos L.), Black Walnut (Juglans nigra L.), American Beech (Fagus grandi- 
folia Ehrhart), Tulip Poplar ( Liriodendron tulipifera L.), Black Maple (Acer nigrum 
F. Michaux), and White Ash (Fraxinus americana L.). The forested collecting areas 
generally consisted of about 25% hickory, 35% oak, 20% beech and maple, and 20% 
miscellaneous species. Most night collecting was done during a new moon phase with 
night temperatures averaging 60°—70°F. 

Four basic collecting methods were used for collecting Catocala: bait, ultraviolet 
and mercury vapor light, tapping trees, and a 150-watt incandescent light. Baiting 
was done along a path in a oak-hickory woods. The bait, consisting of beer, sugar, 
molasses, and fermented fruit, was applied to about 40 trees at dusk and checked 
periodically until about 0200 hours. A single 15-watt unfiltered fluorescent black- 
light tube set up in front of a sheet hung alongside the place of residence and a 
150-watt incandescent porch light were checked at regular intervals throughout 
every night. In 1976 a 175-watt mercury vapor lamp was used along with the other 
light sources. Tapping trees with a wood mop handle to flush resting moths during 
the daytime was done primarily in the same oak-hickory woods that baiting was done. 
Data was recorded in a field notebook as soon as possible on species behavior and 
numbers. 

The Catocala species and forms were determined by referring to the works of 
Barnes & McDunnough (1918, Mem. Amer. Mus. Nat. Hist. 3(1), 47 p., 21 pl.) 
and Sargent (1976, Legion of night. Univ. Mass. Press, Amherst, 215 p., 8 pl.), as 
well as examining Catocala collections of M. C. Nielsen and Michigan State University 
at East Lansing. The lone C. gracilis specimen, although worn, was determined by 
the presence of the short basal forewing dash often used in separating gracilis from 
the very similar C. sordida. 

All specimens collected have been deposited in my personal collection in Kala- 
mazoo, Michigan, given to Mike Larkin also of Kalamazoo, or donated to Western 
Michigan University. 

7 RESULTS AND DiscussION 

As of August 1976 the Catocala species total for Clay Co., Tennessee is 41. A list 
of the species arranged according to McDunnough (1938, Mem. So. Calif. Acad. 
Sci., Vol. 1. 275 p.) that includes earliest and latest dates taken and other capture 
data is shown in Table 1. The dates given in Table 1 coincide only with my collect- 
ing trips and should not be construed as conclusive. Many of the species have longer 
flight periods than shown. However, species whose condition and/or numbers that 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


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JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


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VoLUME 31, NUMBER 3 201 


appeared to be affected by the restrictive collecting dates are subsequently discussed. 
Although additional coverage of Clay Co. will undoubtedly add more species and 
behavior data, certain interesting conclusions can be drawn from Table 1. 

The 5 July collecting date for C. piatrix is very early for Tennessee, as this species 
peak flight is normally early to mid August. This could account for the small number 
taken of a fairly common species. The C. robinsoni male taken on 6 August is like- 
wise a very early date for the species. I have seen fresh robinsoni in the M. C. 
Nielsen Collection from Missouri with mid-September dates. The 2 males of C. 
sappho, and to a lesser extent a female, captured in early August were very worn, 
suggesting a mid-July flight period in Tennessee. Specimens of C. ulalume and C. 
maestosa were fresh, indicating their capture dates probably fall within the species 
main flight period. The late June dates recorded for C. clintoni are quite advanced. 
All specimens were worn and this species is generally considered to be the earliest 
Catocala on the wing. Eggs were secured from one female. The flight period of the 
smaller species begins about 20 June in Tennessee as nearly all specimens taken at 
that time were fresh. In cases where the small number of species taken prohibited an 
accurate determination of the flight period, Forbes (1954, Cornell Univ. Agric. Expt. 
Stat., Mem. 329. 433p.), Sargent (1976), and various Catocala collections were 
consulted to arrive at more accurate flight times. 

More specimens were taken at bait than by other methods because of large num- 
bers of a few species, e.g., C. obscura, residua, retecta, vidua, cerogama, illia, and 
cara. These same species are also found commonly at bait in Michigan. Baiting 
success was greatest right after dusk and became progressively worse into the night. 
C. angusi was the only species recorded as a consistent late night feeder, never 
appearing till a few hours after dusk. Dry, warm, still nights were more productive 
than heavy, humid nights, such as after a rain. I believe high humidity may dilute 
the scent of the bait. Moon phase did not appear as a factor in baiting success. 

Light attracted a wider variety of species than any other method with small 
Catocala being especially susceptible. The most productive night of collecting at 
bait and light occurred when a distant electrical storm was brewing. 

Shagbark Hickory produced more Catocala than all other species of trees tapped. 
Hickories composed only about 25% of the total tree species, but yielded 70% of the 
underwings. Tapping was best during hot, dry afternoons. If any electrical discharge 
was in the air, such as an oncoming storm, the results were phenomenal. I have 
scared as many as 10-12 moths off Shagbarks about 0.67 m in diameter. During cool 
days, moths might rest higher in trees using more exposed positions for thermo- 
regulation, which could account for the very poor results during those times. I have 
flushed Catocala from under shale outcroppings in woods. I could not make any 
positive identifications, but the species (about 5) were all large with yellow hind- 
wings, perhaps being C. nebulosa. This species is reported to select unusual resting 
places (Sargent, 1976). 

In addition to the results recorded in Table 1, other interesting Catocala observa- 
tions were made. C. obscura and residua intergrade very closely in Clay Co. The two 
species ranged from very distinct to almost totally indistinguishable. This observation 
for this area agrees with that made by Sargent (1976). C. obscura alone exhibits 
much individual variation. Some specimens are large (70 mm), with greenish 
tinged, dark, well maculated forewings that are marked with a black dash extending 
from slightly below the apex to the outermost point of the pm line. Specimens at 
the other extreme are small (62 mm) with light, uniform dusky brown primaries that 
have very little maculation. These variations, with little blending, are found in both 
sexes. All C. obscura from Michigan are of the latter description. Clay Co. lies 
within the range of C. retecta luctuosa, and appears to be a blend zone with many 
intergrades between the nominate form and the subspecies taken here. Both 
nominate C. cara and C. c. carissima were very distinct with no intergradation. I 
found this interesting, but because of the small sample size, I hesitate to speculate 


202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


further on this observation. The single specimen of the typical C. relicta was a posi- 
tively identified sight record. Sargent (1976) mentions Kentucky as the southern 
limit of this species, however Mather (pers. comm.) informed me of a single record 
from Mississippi. All C. illecta (males) were captured between 0300 and 0400 hours 
and could suggest that this is an active late night species. The lone capture of C. 
miranda was made at 0400 hours. This species’ rarity might be due in part to its 
being a late flyer. Sargent (1976) gives some Atlantic Coast states, with Pennsyl- 
vania, as the range of C. miranda, but besides this specimen, I have found another 
one from Tennessee in the Michigan State University collection at East Lansing, 
Michigan. 

The only regional list of Catocala I could find for any area close to Clay Co. was 
from Mather (pers. comm.) for Mississippi. Mather did not include habilis, judith, 
flebilis, subnata, cerogama, miranda, or coccinata. Species with only one or two 
records include serena, obscura, residua, sappho, palaeogama, relicta, gracilis, clintoni, 
and minuta. According to the Catocala ranges given by Sargent (1976) and assess- 
ment of local foodplants, the following might be expected to occur in Clay Co., but 
have not been recorded: C. consors A. & S., agrippina Stkr., marmorata Edw., junctura 
WIk., sordida Grt., crataegi Saund., mira Grt., titania Dodge, and dulciola Grt. 

The diversity of Catocala species encountered in Clay Co. is remarkable for only 
two years of fairly concentrated collecting. I would greatly appreciate hearing from 
anyone who has collected this genus in the Tennessee or Kentucky area. 


ACKNOWLEDGMENTS 


I wish to thank Mogens C. Nielsen of Lansing, Michigan for the encouragement 
and. initial review of this paper. He generously allowed me examination of his 
personal collection and provided access to the Michigan State University collection. 
I am also grateful to an anonymous reviewer of an earlier version of this paper. 


WayneE A. Mitter, 1491 North Hills Drive, Kalamazoo, Michigan 49007. 


APPARENT LONG-DISTANCE DISPERSAL BY PIERIS OCCIDENTALIS 
(PIERIDAE) 


Colonizing ability and the turnover rate of local populations of animals depend 
to a considerable degree on vagility. Records of “stray” butterflies are common in 
the literature, although it is rarely possible to deduce the manner of dispersal or 
even a minimum distance traveled. Altitudinal displacements sometimes offer good 
opportunities of this sort (Shapiro, 1973, J. Res. Lep. 12: 231-235; 1974, Ibid. 138: 
157-161). The western white (Pieris occidentalis Reakirt) and the checkered white 
(P. protodice Boisduval & LeConte) are a pair of sibling species which generally 
replace each other, occurring at high and low altitudes respectively through most of 
western North America. Both are believed to be colonizing species, with vigorously 
dispersing adults (Shapiro, 1975, Amer. Midl. Nat. 93: 424-433). P. protodice seems 
unable to overwinter in Donner Pass (Placer/Nevada cos., California) at 7000 feet, 
but regularly colonizes the area in summer. Records of downslope dispersal by P. 
occidentalis, which is not known to breed in the Sierra below 6000 feet, are much 
rarer. On 13 August 1975 a worn female was taken at 5000 feet in the South Yuba 
River Canyon, Nevada Co., and I considered this a noteworthy “low elevation” record 
for California (Shapiro, unpublished). 

On 17 October 1976 I was collecting a sample of P. protodice at Rancho Cordova, 
Sacramento Co., California, elevation 65 feet, when I noticed a dark female vigorously 
rejecting a courting male. I collected the two, and was astonished to find that the 


VoLUME 31, NUMBER 3 203 


female was a worn P. occidentalis of the “calyce” phenotype. Her phenotype is quite 
unambiguous and easily told from the 24 female protodice collected the same after- 
noon, which represent a large range of variation from summer to “vernalis’ pheno- 
types. The habitat is annual grassland and foothill woodland developed on dredge 
tailings, and is a classic locality for large populations of P. protodice breeding on the 
weedy mustard Brassica geniculata. No other occidentalis were observed alive, and 
no occidentalis tendency appeared in a sample of 76 protodice collected for study in 
an hour. 

The nearest 6000-foot contours on the map are 50—70 miles from Rancho Cordova 
(USGS 1:250,000 “Sacramento” topographic sheet). P. occidentalis of similar pheno- 
type and in mixed condition were still flying in Donner Pass at 7000 feet on 15 
October 1976. Prevailing winds during the week prior to the record were very light 
northeasterly, i.e., downslope, and weather conditions were clear, warm, and dry. 
My trip was not pre-scheduled so there is no chance of a deliberate release having 
been made as a hoax by a student, and in the huge protodice population any 
deliberate release would probably not have been recognized. I see no alternative but 
to regard this as an extreme instance of individual long-range dispersal in a non- 
migratory species. The fact that it turned up in what is probably the best protodice 
habitat on the floor of the Sacramento Valley suggests habitat selection by the female 
once she did get down near sea level, since the ecologies of the two species are so 
similar. How many such individual dispersals actually occur? And what are the 
potential consequences, say for gene flow, of events which are so unlikely to come 
to our attention? Records indicating long-range dispersal, properly documented, are 
not trivial, and have considerably more scientific value than their traditional role as 
bizarre additions to local lists. 


Artuur M. Sapiro, Department of Zoology, University of California, Davis, Cali- 
fornia 95616. 


THE BEHAVIOUR OF AN INEBRIATED OPSIPHANES CASSIAE 
(BRASSOLIDAE ) 


A female of Opsiphanes cassiae L., known in Costa Rica as “mariposa pacayera”™ 
or palm butterfly, drifted at dusk from the forest into a room where it flew and 
perched in places where light was subdued, until it arrived at a table where a bottle 
of a sound red wine (Rubion, Paul Masson, California, alcohol 12%/vol.) had been 
recently uncorked and left to breathe. The insect circled the bottle a few times be- 
fore landing on the table surface and assuming a normal resting position. I then 
placed a drop of wine some 10 cm away from the butterfly which shyly retreated 
from my hand, remained motionless for a moment and then approached the drop, 
extended its proboscis and drank at its leisure. After some five minutes there were 
obvious changes in the behaviour of the insect. 

First, some very slow up and down flapping of wings, followed by forewings 
being lowered and directed forward with brisk movements several times, hindwings 
remaining upright. With forewings in this position, the hindwings also moved for- 
ward, without lowering, until these were propped far ahead of their normal resting 
position. The abdomen remained motionless. Antennae were lowered until they 
touched the table and then shot backwards. Movement of fore-, hindwings and 
antennae were repeated several times in the same order of events. 

After a brief period of inactivity, a hopping spastic side-walking took place 
alternating with wing and antennae motions as well as a tremulous and agitated 
moving of the legs. More wine was offered to the insect which sipped it directly 
from my fingertip after a somewhat hesitant uncoiling of proboscis. Another sequence 


204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of the behaviour described above was observed until all wings were placed flat on 
the table although they were not limp and flaccid. A few forward strokes of fore- 
wings followed by a very fast vibratory flapping preceded a period of inaction. A 
few minutes later the butterfly took flight in a close-spiralling pattern towards 
an incandescent light, hitting the hot bulb several times, alighting and again at- 
tempting flight to the light source close to which it finally perched. After a few hours 
it resumed normal behaviour and flew away the next day. 

As these butterflies, like other Brassolidae, feed, among other things, on fermenting 
juices of fruit, it would be interesting to determine the alcoholic content of some of 
their foods and the possible influence on butterfly behaviour and survival in nature. 


Luis D. Gomez, Departamento de Historia Natural, Museo Nacional de Costa 
Rica, San Jose, Costa Rica. 


VOLUME 31, NUMBER 3 205 


OBITUARY 
PROFESSOR PHILIP M. SHEPPARD, D PHIL, FRS (1921-1976) 


Philip Sheppard’s most enduring human achievement was to apply his interest in 
the workings of natural selection to the solution of a medical problem: by considering 
the way natural selection operates on blood groups through the interactions of mother 
and. foetus, he and Sir Cyril Clarke invented and then successfully developed the 
‘Liverpool jab’ which prevents rhesus haemolytic disease of the newborn, up to now 
a major cause of miscarriage, infantile death, and brain-damaged children. 

Professor Sheppard’s research in evolution was very much in the Darwinian tradi- 
tion: a blend of experimental science and natural history which greatly valued the 
accumulated wisdom of naturalists, both amateur and professional. He thought of 
Lepidoptera as extremely valuable research tools in this respect, and used them more 
than any other group: in contributing to evolutionary biology by using Lepidoptera 
he contributed immensely to the field of lepidopterology, in which his monumental 
achievement (again mainly in collaboration with Sir Cyril Clarke) has been the 
elucidation of the genetics and evolution of mimicry, chiefly in three of the ex- 
traordinary polymorphic Papilio, now rounded out by studies of Hypolimnas and 
(posthumously) of Heliconius. This work has been a brilliant confirmation and 
extension of the theories of mimicry propounded by A. J. Nicholson, R. A. Fisher, and 
E. B. Ford, with the last of whom Sheppard did his doctoral research in Oxford 
(taking a D Phil in 1951) after serving during the second world war first in the 
RAF (Volunteer Reserve) and then as a prisoner of the Third Reich (he used to say, 
when lecturing on warning coloration, that what a man regarded as palatable de- 
pended on how hungry he was). 

His first studies included a now justly famous demonstration (in collaboration with 
A. J. Cain) that characteristics which most biologists thought of as trivial (in this 
case the colour and striping of the shells of terrestrial snails) could be of considerable 
adaptive importance to the organism and could be subject to detectable natural 
selection. This aspect of his work has been the one most widely followed by other 
workers, Cain and he having founded in effect, a college for the study of micro- 
evolution in snails. His book, Natural selection and heredity, which integrates this 
work into that of the ‘English’ school of evolutionary genetics, has been widely used 
by students. 

Sheppard spent some time on fieldwork, research and teaching in South East 
Asia, the United States, Finland, Trinidad, Brasil and other parts of the neotropics; 
most of his work was carried out at the University of Liverpool, which he joined as 
a senior lecturer in 1956, becoming Professor of Genetics in 1963. There he embarked 
on his wonderfully fruitful collaborations with C. A. Clarke on the genetics of people 
and of butterflies. He was elected to a Fellowship of the Royal Society in 1965, and 
was awarded the Darwin Medal in 1974. 

During his long fight against leukaemia his abnormal energy and enthusiasm 
for scientific enquiry continued with no noticeable diminution. Much work remains 
to be published by colleagues: the appended bibliography (which is probably in- 
complete even for earlier works) shows an outline of his achievements. 


Publications of P. M. Sheppard 


1950. Cain, A. J. & P. M. Sheppard. Selection in the polymorphic land. snail 
Cepaea nemoralis. Heredity 4: 275-294. 
1951. Sheppard, P. M. Fluctuations in the selective value of certain phenotypes 
in the polymorphic land snail Cepaea nemoralis. Heredity 5: 125-134. 
Sheppard, P. M. A quantitative study of two populations of the moth 
Panaxia dominula. Heredity 5: 349-378. 


206 


1952. 


1953. 


1954. 


1955. 


1956. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Sheppard, P. M. Natural selection in two colonies of the polymorphic land 
snail Cepaea nemoralis. Heredity 6: 233-238. 

Sheppard, P. M. A note on non-random mating in the moth Panaxia 
dominula. Heredity 6: 239-241. 

Cain, A. J. & P. M. Sheppard. The effects of natural selection on body 
colour in the land snail Cepaea nemoralis. Heredity 6: 217-231. 

Sheppard, P. M. Polymorphism and population studies. Symp. Soc. exp. 
Biol. 7: 274-289. 

Sheppard, P. M. Polymorphism, linkage and the blood groups. Amer. Nat. 
87: 283-294. 

Sheppard, P. M. Cancer of stomach and ABO blood groups. Brit. med. J. 
Tie 2PX0), 

Clarke, C. A. & P. M. Sheppard. Further observations on hybrid swallow- 
tails. Entomol. Rec. 65 (Supp.): 1-12. 

Sheppard, P. M. Evolution in bisexually reproducing organisms. In: Evolu- 
tion as a process (J. Huxley, A. C. Hardy and E. B. Ford, eds.), pp. 201- 
218, London, Allen and Unwin. 

Cain, A. J. & P. M. Sheppard. The theory of adaptive polymorphism. Amer. 
Nat. 88: 321-326. 

Cain, A. J. & P. M. Sheppard. Natural selection in Cepaea. Genetics 39: 
89-116. 

Sheppard, P. M. Genetic variability and polymorphism: synthesis. Cold 
Spring Harbor Symp. Quant. Biol. 20: 271-275. 

Clarke, C. A. & P. M. Sheppard. The breeding in captivity of the hybrid 
Papilio rutulus female < Papilio glaucus male. Lepid. News 9: 46—48. 
Clarke, C. A. & P. M. Sheppard. A preliminary report on the genetics of the 

machaon group of swallowtail butterflies. Evolution 9: 182-201. 

Clarke, C. A. & P. M. Sheppard. The breeding in captivity of the hybrid 
swallowtail Papilio machaon gorganus Fruhstorfer female « Papilio hos- 
piton Gene male. Entomologist 88: 266-270. 

Clarke, C. A., W. K. Cowan, J. W. Edwards, A. W. Howel-Evans, R. B. 
McConnell, J. C. Woodrow, & P. M. Sheppard. The relationship of the 
ABO blood groups to duodenal and gastric ulceration. Brit. Med. J. ii: 
643. 

Sheppard, P. M. Ecology and its bearing on population genetics. Proc. Roy. 
Soc. B 145: 308-315. 

Cain, A. J. & P. M. Sheppard. Adaptive and selective value. Amer. Nat. 90: 
202-203. : 

Clarke, C. A. & P. M. Sheppard. A further report on the genetics of the 
machaon group of swallowtail butterflies. Evolution 10: 66-73. 

Clarke, C. A. & P. M. Sheppard. MHand-pairing of butterflies. Lepid. News 
10: 47-53. 

Clarke, C. A. & P. M. Sheppard. The genetics of some mimetic forms of 
Papilio dardanus Brown and Papilio glaucus Linn. Proc. Roy. Ent. Soc. 
ondm@n2 40-41 

Clarke, C. A., R. B. McConnell, & P. M. Sheppard. The ABO blood groups 
and secretor status in duodenal ulcer. Heredity 10: 126. 

Clarke, C. A., J. W. Edwards, D. R. W. Haddock, A. W. Howel-Evans, R. 
B. McConnell, & P. M. Sheppard. ABO blood groups and secretor charac- 
oan duodenal ulcer—population and sibship studies. Brit. Med. J. ii: 
Tas 

Clarke, C. A. & P. M. Sheppard. The ABO blood groups in duodenal ulcer— 
a study of sibships. Acta Genet. 6: 570-574. 

McConnell, R. B. & P. M. Sheppard. The secretor character and disease. 
Acta Genet. 6: 574-579. 


VoLuME 31, NUMBER 3 207 


1957. 


1958. 


1959. 


Sheppard, P. M. The Rh blood groups. Lancet 1957: 212. 

Cain, A. J. & P. M. Sheppard. Some breeding experiments with Cepaea 
nemoralis (L.). J. Genet. 55: 195-199. 

Clarke, C. A. & P. M. Sheppard. The breeding in captivity of the hybrid 
Papilio glaucus female x Papilio eurymedon male. Lepid. News 11: 
201-205. 

Clarke, C. A., R. B. McConnell, & P. M. Sheppard. Blood groups and dis- 
ease. Lancet 1957: 212. 

Clarke, C. A., R. B. McConnell & P. M. Sheppard. ABO bicod groups and 
duodenal ulcer. Brit. Med. J. i: 758-759. 

Sheppard, P. M. Evolution in bisexually reproducing organisms. In: Evolu- 
tion as a process, 2nd ed. London, Allen & Unwin. 

Sheppard, P. M. Natural selection and heredity. London, Hutchinson. 

Clarke, C. A. & P. M. Sheppard. Genetics and race-crosses in Papilio dar- 
danus Brown. Proc. Roy. Ent. Soc. Lond. C 23: 1-3. 

Clarke, C. A. & P. M. Sheppard. A genetic investigation of mimicry. Proc. 
X. Int. Cong. Genet. 2: 52-53. 

Clarke, C. A. & P. M. Sheppard. Disease in relation to human polymorphic 
systems. Proc. X. Int. Cong. Genet. 2: 52. 

Sheppard, P. M. & S. Walker. An experimental study on the dwarf broom 
Cytisus scoparius spp. prostratus. Ann. Mag. Field Studies “The Lizard” 
1: 12-13. 

Clarke, C. A., R. Finn, R. B. McConnell, & P. M. Sheppard. The protection 
afforded by ABO incompatibility against erythroblastosis due to Rhesus 
Anti-D. Int. Arch. Allergy N.Y. 13: 380. 

Howel-Evans, W., R. B. McConnell, C. A. Clarke, & P. M. Sheppard. 
Carcinoma of the oesophagus with keratosis palmaris et plantaris (tylo- 
sis). Quart. J. Med. 27: 413—429. 

Sheppard, P. M. Natural selection and some polymorphic characters in 
man. In: Natural selection in human populations. Symp. Soc. Study 
Human Biology 2. pp. 35-48. 

Sheppard, P. M. The evolution of mimicry: a problem in ecology and 
genetics. Cold Spring Harbor Symp. Quant. Biol. 24: 131-140. 

Sheppard, P. M. Blood groups and natural selection. Brit. Med. Bull. 15: 
134-139. 

Sheppard, P. M. Natural selection and heredity (2nd edition), London, 
Hutchinson. 

Clarke, C. A. & P. M. Sheppard. The breeding of Papilio dardanus Brown in 
England. Technique and some results. Entomologist 92: 89-95. 

Clarke, C. A. & P. M. Sheppard. The genetics of some mimetic forms of 
Papilio dardanus Brown, and Papilio glaucus Linn. J. Genet. 56; 236— 
260. 

Clarke, C. A. & P. M. Sheppard. The genetics of Papilio dardanus Brown. 
I. Race cenea from South Africa. Genetics 44: 1347-1358. 

Clarke, C. A., D. A. Price-Evans, R. B. McConnell, & P. M. Sheppard. Secre- 
tion of blood group antigens and peptic ulcer. Brit. Med. J. i: 603-607. 

Clarke, C. A., W. Howel-Evans, R. B. McConnell, & P. M. Sheppard. Car- 
cinoma of oesophagus in association with tylosis. Brit. Med. J. ii: 1100. 

Kitchin, F. D., W. Howel-Evans, C. A. Clarke, R. B. McConnell, & P. M. 
Sheppard. PTC taste response and thyroid disease. Brit. Med. J. i: 
1069-1074. 

Werrett, W. R., A. J. Candy, J. O. L. King, & P. M. Sheppard. Semi-albino: 
a third sex-linked allelomorph of silver and gold in the fowl. Nature 
184: 480. 


208 


1960. 


1961. 


1962. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Clarke, C. A. & P. M. Sheppard. The evolution of dominance under disrup- 
tive selection. Heredity 14: 73-87. 

Clarke, C. A. & P. M. Sheppard. The evolution of mimicry in the butterfly 
Papilio dardanus. Heredity 14: 163-173. 

Clarke, C. A. & P. M. Sheppard. Super-genes and mimicry. Heredity 14: 
175-185. 

Clarke, C. A. & P. M. Sheppard. The genetics of Papilio dardanus Brown. 
II. Races dardanus, polytrophus, meseres, and tibullus. Genetics 45: 
439-457. 

Clarke, C. A. & P. M. Sheppard. The genetics of Papilio dardanus Brown. 
III. Race antinorii from Abyssinia and race meriones from Madagascar. 
Genetics 45: 683-698. 

Clarke, C. A., R. B. McConnell, & P. M. Sheppard. ABO blood groups and 
secretor character in rheumatic carditis. Brit. Med. J. i: 21-23. 

Clarke, C. A., R. B. McConnell, & P. M. Sheppard. Blood groups in 
rheumatic carditis. Brit. Med. J. i: 566. 

Clarke, C. A., R. B. McConnell, & P. M. Sheppard. A genetical study of the 
variations in ABH secretion. Ann. Hum. Genet. Lond. 24: 295-307. 
Clarke, C. A., R. B. McConnell, R. Finn, F. D. Kitchin, J. C. Woodrow, 
D. A. Price-Evans, & P. M. Sheppard. The role of inheritance in com- 

mon diseases. Lancet 1960: 526-527. 

Clarke, C. A., W. T. A. Donohoe, R. Finn, R. B. McConnell, P. M. Sheppard, 
& D. S. H. Nicol. Data on linkage in man: ovalocytosis, sickling and 
the Rhesus blood group complex. Ann. Hum. Genet. 24: 283-287. 

Cain, A. J., J. M. B. King, & P. M. Sheppard. New data on the genetics of 
polymorphism in the snail Cepaea nemoralis L. Genetics 45: 393-411. 
Sheppard, P. M. Recent genetical work on polymorphic mimetic Papilios. 
In: Insect Polymorphism (J. S. Kennedy, ed.), pp. 20-29. London, Roy. 

Ent. Soc. 

Sheppard, P. M. Differential transmission of Down’s syndrome (Mongolism ). 
Lancet 1961: 1455-1456. 

Sheppard, P. M. Some contributions to population genetics resulting from 
the study of the Lepidoptera. Ady. Genetics 10: 165-216. 

Sheppard, P. M. Darwinism up to date. Impulse 18: 6—10. 

Cain, A. J. & P. M. Sheppard. Visual and physiological selection in Cepaea. 
Amer. Nat. 95: 61-64. 

Clarke, C. A., R. B. McConnell & P. M. Sheppard. Data on linkage in man. 
The nail-patella syndrome, family S. Ann. Hum. Genet. Lond. 25: 25-27. 

Finn, R., C. A. Clarke, W. T. A. Donohoe, R. B. McConnell, P. M. Sheppard, 
D. Lehane, & W. Kulke. Experimental studies on the prevention of Rh 
haemolytic disease. Brit. Med. J. i: 1486-1490. 

Turner, J. R. G., C. A. Clarke & P. M. Sheppard. Genetics of a difference 
in the male genitalia of east and west African stocks of Papilio dardanus 
(Lep.). Nature 191: 935-936. 

Sheppard, P. M. Some aspects of the geography, genetics and taxonomy of 
a butterfly. In: Taxonomy and geography, pp. 135-152. London, Syst. 
Assn. Publication no. 4. 

Clarke, C. A. & P. M. Sheppard. Offspring from double matings in swallow- 
tail butterflies. Entomologist 95: 199-203. 

Clarke, C. A. & P. M. Sheppard. The hybrids between Papilio machaon L. 
female and Papilio maackii male and between Papilio polyxenes asterias 
Fabr. female and Papilio maackii Men. male (Lepidoptera, Nymphalidae). 
Entomologist 95: 269-271. 


Clarke, C. A. & P. M. Sheppard. Disruptive selection and its effect on a 


VOLUME 31, NUMBER 3 209 


1963. 


1964. 


metrical character in the butterfly Papilio dardanus. Evolution 16: 214— 
226. 

Clarke, C. A. & P. M. Sheppard. Blood-groups and disease. Lancet 1962: 
967-968. 

Clarke, C. A. & P. M. Sheppard. The genetics of Papilio dardanus Brown. 
IV. Data on the race ochracea, race flavicornis, and further information 
on races polytrophus and dardanus. Genetics 47: 909-920. 

Clarke, C. A. & P. M. Sheppard. The genetics of the mimetic butterfly 
Papilio glaucus. Ecology 43: 159-161. 

Sheppard, P. M. & L. M. Cook. The manifold effects of the medionigra 
gene of the moth Panaxia dominula and the maintenance of a poly- 
morphism. Heredity 17: 415-426. 

Cohen, C., W. H. Allton jr., R. Finn, C. A. Clarke, R. B. McConnell, J. C. 
Woodrow, W. Kulke, D. Lehane, & P. M. Sheppard. Iso-immunization in 
the rabbit with antibody-coated erythrocytes. Nature 193: 990-992. 

Clarke, C. A.. W. T. A. Donohoe, R. B. McConnell, J. H. Martindale, & P. M. 
Sheppard. Blood groups and disease: previous transfusion as a potential 
source of error in blood typing. Brit. Med. J. i: 1734-1736. 

Sheppard, P. M. The genetics of mimicry. Proc. XVI Int. Cong. Zool. 4: 
150-156. 

Sheppard, P. M. Some genetic studies of Miillerian mimics in butterflies of 
the genus Heliconius. Zoologica 48: 145-154. 

Clarke, C. A. & P. M. Sheppard. Frequencies of the melanic forms of the 
moth Biston betularia (L.) on Deeside and in adjacent areas. Nature 
198: 1279-1282. 

Clarke, C. A. & P. M. Sheppard. The results of some experimental breeding 
in the Lepidoptera and their application to man. Proc. Roy. Ent. Soc. 
Lond. C 28:—33—34. 

Clarke, C. A. & P. M. Sheppard. Interactions between major genes and poly- 
genes in the determination of the mimetic patterns of Papilio dardanus. 
Evolution 17: 404-413. 

Duncan, C. J. & P. M. Sheppard. Continuous and quantal theories of sensory 
discrimination. Proc. Roy. Soc. B. 158: 343-363. 

Clarke, C. A., C. G. C. Dickson, & P. M. Sheppard. Larval colour pattern in 
Papilio demodocus. Evolution 17: 130-137. 

Clarke, C. A., R. B. McConnell & P. M. Sheppard. Transplacental foetal 
loss. Lancet 1963: 642. 

Clarke, C. A., W. T. A. Donohoe, R. B. McConnell, J. C. Woodrow, R. Finn, 
J. R. Krevans, W. Kulke, D. Lehane, & P. M. Sheppard. Further experi- 
mental studies on the prevention of Rh haemolytic disease. Brit. Med. J. 
i: 979-984. 

Sheppard, P. M. Natural selection and mimicry. Discovery 25: 35-39. 

Sheppard, P. M. Protective colouration in some British moths. Entomologist 
97: 209-216. 

Sheppard, P. M. Experimental studies on microevolution in populations of 
the Lepidoptera. Proc. XII Int. Cong. Ent. London: 268. 

Clarke, C. A. & P. M. Sheppard. Genetic control of the melanic form 
insularia of the moth Biston betularia (L.). Nature 202: 215-216. 
Clarke, C. A. & P. M. Sheppard. The hybrid between Papilio polyxenes 
asterias Fabr. female and Papilio maackii Men. male. Entomologist 97: 

131-133. 

Clarke, C. A., R. Finn, R. B. McConnell, J. C. Woodrow, D. Lehane, & P. M. 

Sheppard. Prevention of Rh haemolytic disease. Brit. Med. J. i: L110. 


210 


1965. 


1966. 


1967. 


1968. 


1969. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Davis, J. C., A. P. Wade, G. S. Wilkinson, R. W. Smithells, C. O. Carter, 
C. A. Clarke, & P. M. Sheppard. Steroid excretion of dehydroepiandro- 
sterone in young mothers of Mongols. Lancet 1964: 782-785. 

Sheppard, P. M. The monarch butterfly and mimicry. J. Lepid. Soc. 19: 
227-230. 

Sheppard, P. M. Sex-limited polymorphisms in microevolution. Can. Ent. 
97: 442-445. 

Sheppard, P. M. Mimicry and its ecological aspects. Genetics Today (Proc. 
XI Int. Cong. Genet., Hague) 3: 553-560. 

Clarke, C. A. & P. M. Sheppard. Prevention of Rhesus haemolytic disease. 
Lancet 1965: 348. 

Duncan, C. J. & P. M. Sheppard. Sensory discrimination and its role in the 
evolution of Batesian mimicry. Behaviour 24; 269-282. 

MacDonald, W. W. & P. M. Sheppard. Cross-over values in the sex chromo- 
somes of the mosquito Aedes aegypti and evidence of the presence of 
inversions. Ann. Trop. Med. Parasitol. 59: 74-87. 

Woodrow, J. C., C. A. Clarke, W. T. A. Donohoe, R. Finn, R. B. McConnell, 
P. M. Sheppard, D. Lehane, S. H. Russell, W. Kulke, & C. M. Durkin. 
Prevention of Rh haemolytic disease: a third report. Brit. Med. J. ii: 
279-283. 

Bourdillon, R. E., C. A. Clarke, A. P. Ridges, P. M. Sheppard, P. Harper, & 
S. A. Leslie. ‘Pink Spot’ in the urine of schizophrenics. Nature 208: 
453-455. 

Clarke, C. A. & P. M. Sheppard. A local survey of the distribution of 
industrial melanic forms in the moth Biston betularia and estimates of 
the selective values of these in an industrial environment. Proc. Roy. Soc. 
B 165: 424-439. 

Sheppard, P. M. & E. B. Ford. Natural selection and the evolution of 
dominance. Heredity 21: 139-147. 

Sheppard, P. M. Natural selection and heredity (8rd edition), London, 
Hutchinson. 

Clarke, C. A., P. M. Sheppard, & J. A. Bishop. Some studies of protective 
colouration in Lepidoptera. Proc. R. Ent. Soc. Lond. C 32: 25-26. 
Karunaratne, P. B., C. A. Clarke, & P. M. Sheppard. The genetics of 
Chilasa clytia lankeswara (the common mime). Entomologist 100: 27- 

28. 

Cain, A. J., P. M. Sheppard, & J. M. B. King. Studies on Cepaea. I. 
The genetics of some morphs and varieties of Cepaea nemoralis (L.). 
Phil. Trans. Roy. Soc. B 253: 383-396. 

Clarke, C. A., P. M. Sheppard, & I. W. B. Thornton. The genetics of the 
mimetic butterfly Papilio memnon L. Phil. Trans. Roy. Soc. B 254: 37— 
89. 

Clarke, C. A., F. M. M. Clarke, & P. M. Sheppard. Mimicry and Papilio 
memnon. Some breeding results from England. Malayan Nature J. 21: 
201-219. 

Sheppard, P. M. Evolutionary genetics of animal populations: the study of 
natural populations. Proc. XII Int. Cong. Genet. Tokyo 3: 261-279. 
Clarke, C. A. & P. M. Sheppard. Rhesus sensitization and abortion. Brit. 

Med. J. 4: 748. 

Ford, E. B. & P. M. Sheppard. The medionigra polymorphism of Panaxia 
dominula. Heredity 24: 561-569. 

Sheppard, P. M., W. W. MacDonald, R. J. Tonn, & B. Grab. The dynamics 
of an adult population of Aedes aegypti in relation to dengue haemorrha- 
gic fever in Bangkok. J. Anim. Ecol. 38: 661-702. 

Sheppard, P. M., W. W. MacDonald, & R. J. Tonn. A new method of 


VOLUME 31, NUMBER 3 211 


1970. 


1971. 


1972. 


1973. 


measuring the relative prevalence of Aedes aegypti. Bull. Wild. Hlth. Org. 
40: 467—468. 

Tonn, R. J., P. M. Sheppard, W. W. MacDonald, & Y. H. Bang. Replicate 
surveys of larval habitat of Aedes aegypti in relation to dengue haemor- 
rhagic fever in Bangkok, Thailand. Bull. Wld. Hlth. Org. 40: 819-829. 

Clarke, C. A. & P. M. Sheppard. Is Papilio gothica (Papilionidae) a good 
species? J. Lepid. Soc. 24: 229-233. 

Tonn, R. J.. W. W. MacDonald, P. M. Sheppard, & S. Iatanasen. The 
magnitude of seasonal changes in larval populations of Aedes aegypti in 
Bangkok, Thailand. Bull. Wld. Hlth. Org. 42: 943-950. 

Allen, W. R. & P. M. Sheppard. Copper tolerance in some Californian pop- 
ulations of the monkey flower, Mimulus guttatus. Proc. Roy. Soc. Lond. 
B 177: 177-196. 

Clarke, C. A. & P. M. Sheppard. Further studies on the genetics of the 
mimetic butterfly Papilio memnon L. Phil. Trans. Roy. Soc. B 263: 25- 
70. 

Clarke, C. A. & P. M. Sheppard. The genetics of the mimetic butterfly 
Papilio polytes L. Phil. Trans. Roy. Soc. B 263: 431-458. 

Clarke, C. A. & P. M. Sheppard. [Untitled communication on the genetics 
of Hypolimnas bolina.] Proc. Roy. Ent. Soc. Lond. 37: 25. 

Clarke, C. A. & P. M. Sheppard. Genetic and environmental factors in- 
fluencing pupal colour in the swallowtail butterflies Battus philenor (1. ) 
and Papilio polytes L. J. Ent. A 46: 123-133. 

Clarke, C. A., P. M. Sheppard, & A. Willig. The use of ecdysone to break a 
two and a half year pupal diapause in Papilio glaucus 2 x Papilio 
rutulus 6 hybrids. Entomologist 105: 137-138. 

Sheppard, P. M. (editor). Practical Genetics, London, Blackwell Scientific 
Publications. 

Sheppard, P. M. Population and ecological genetics. In: Practical Genetics 
(P. M. Sheppard, ed.), pp. 296-317. London, Blackwell Scientific Publ. 

Sheppard, P. M. & J. A. Bishop. The study of populations of Lepidoptera 
by capture-recapture methods J. Res. Lep. 12: 135-144. 

Bishop, J. A. & P. M. Sheppard. An evaluation of two capture-recapture 
models using the technique of computer simulation. In: The mathematical 
theory of the dynamics of biological populations (M. S. Bartlett and 
R. W. Hiorns, eds.), pp. 235-252. London, Academic Press. 

Clarke, C. A. & P. M. Sheppard. Genetics and mimicry in butterflies: some 
observations from breeding experiments. Proc. Roy. Ent. Soc. Lond. 37: 
35-36. 

Clarke, C. A. & P. M. Sheppard. [Discussion of genetics and mimicry in but- 
terflies: some observations from breeding experiments] Proc. Roy. Ent. 
Soc. Lond. 38: 3-4. 

Clarke, C. A. & P. M. Sheppard. The genetics of four new forms of the 
mimetic butterfly Papilio memnon L. Proc. Roy. Soc. Lond. B 184: 1-14. 

Clarke, C. A., O. M. McKendrick, & P. M. Sheppard. Potatoes and spina 
bifida. Brit. Med. J. ii 3: 251-254. 

Clarke, C. A., P. M. Sheppard, & A. G. Smith. The genetics of fore and 
hindwing colour in crosses between Danaus chrysippus from Australia 
and from Sierra Leone (Danaidae). J. Lepid. Soc. 27: 73-77. 

Clarke, C. A., O. M. McKendrick, & P. M. Sheppard. Potatoes and spina 
bifida. Quart. J. Med. 42: 821-822. 

Bailey, W. J., C. A. Clarke, P. M. Sheppard, & D. J. Whittle. The status of 
the industrial melanics of Biston betularia (L.) in the Isle of Man. 
Entomologist 106: 210-214. 


212 


1974. 


IS), 


1976: 


tie 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Brown, K. S. jr., P. M. Sheppard, & J. R. G. Turner. Quaternary refugia in 
tropical America: evidence from race formation in Heliconius butterflies. 
Proc. Roy. Soc. Lond. B 187: 369-378. 

Sheppard, P. M. Millerian mimicry and Amazonian forest refuges. Heredity 
Sorts: 

Clarke, C. A. & P. M. Sheppard. [Exhibit of set specimens of hybrids which 
they had bred, using the hand-pairing technique. Papilio.] Proc. Roy. 
Ent. Soc. Lond. 39: 39-40. 

Sheppard, P. M. Natural Selection and Heredity (4th edition). London, 
Hutchinson. 

Turner, J. R. G. & P. M. Sheppard. Absence of crossing-over in female 
butterflies (Heliconius). Heredity 34: 265-269. 

Clarke, C. A. & P. M. Sheppard. The genetics of the mimetic butterfly 
Hypolimnas bolina (l.) Phil. Trans. Roy. Soc. B 272: 229-265. 

Clarke, C. A., P. M. Sheppard, & V. Scali. All-female broods in the butterfly 
Hypolimnas bolina (.) Proc. Roy. Soc. Lond. B 189: 29-37. 

Clarke, C. A., D. Hobson, O. M. McKendrick, S. C. Rogers, & P. M. Sheppard. 
Spina bifida and anencephaly: miscarriage as possible cause. Brit. Med. 
J. 1975: 743=746. 

Woodrow, J. C., C. A. Clarke, W. T. A. Donohoe, R. Finn, R. B. McConnell, 
P. M. Sheppard, D. Lehane, F. M. Roberts & T. M. D. Gimlette. Mech- 
anism of Rh prophylaxis: an experimental study on specificity of immuno- 
suppression. Brit. Med. J. ii: 57-59. 

Clarke, C. A., D. Hobson, O. M. McKendrick, P. M. Sheppard, & S. C. 
Rogers. Aetiology of anencephaly and spina bifida. Brit. Med. J. 1976: 
455. 

Clarke, C. A., P. M. Sheppard, & U. Mittwoch. Heterochromatin poly- 
morphism and colour pattern in the tiger swallowtail butterfly Papilio 
glaucus L. Nature 263: 585-587. 

Whittle, P. D. j., C. A. Clarke, P. M. Sheppard, & J. A. Bishop. Further 
studies on the industrial melanic moth Biston betularia (.) in the north- 
west of the British Isles. Proc. Roy. Soc. Lond. B 194: 467-480. 

Clarke, C. A. & P. M. Sheppard. A new tailed female form of Papilio mem- 
non L. and its probable genetic control. Syst. Ent. 2: 17-19. 

Clarke, C. A. & P. M. Sheppard. Data suggesting absence of linkage between 
two loci in the mimetic butterfly Hypolimnas bolina (Nymphalidae). J. 
kepid), Soc) oie Igo 148, 

Sheppard, P. M. & J. R. G. Tumer. The existence of muellerian mimicry. 
Evolution (in press). 

Sheppard, P. M., J. R. G. Turner, K. S. Brown jr., W. W. Benson, & 
M. Singer. Genetics and the evolution of muellerian mimicry in Heliconius 
butterflies. Phil. Trans. Roy. Soc. B (in prep). 


ACKNOWLEDGMENTS 


Thanks to Denise O’Leary and Christie Helquist for much help in compiling the 
bibliography. 


J. R. G. Turner, Division of Biological Sciences, State University of New York 
at Stony Brook, Stony Brook, New York 11794. 


VOLUME 31, NUMBER 3 913 


OBITUARY 


PETER J. HERLAN (1909-1977) 


Peter J. Herlan, Curator of Natural History at the Nevada State Museum, three 
times host of meetings of the Lepidopterists’ Society, and highly respected member of 
the Pacific Slope Branch, died 4 April 1977, at age 67. 

Peter Herlan was born in Seattle, 14 December 1909. He majored in botany at 
the University of Washington, where he was a member of the swimming team and 
played football on a Rosebowl team. He served as a seaman in the Merchant 
Marine and as an Inspector for Boeing Aircraft. During the Second World War he 
enlisted in the Air Force, served as a gunner, and was wounded in action. 

He moved to Nevada in 1946 on the advice of an Air Force physician, for the 
benefit of the dry climate on his wounds. 

He first worked for the Nevada Museum as a driver of a mobile unit for museum 
exhibits that toured the state. He later became Curator of Natural History. He had 
extensive field experience with plants, birds, and mammals, but was best known for 
his work with insects, especially the butterflies. He led numerous field expeditions 
for the Nevada Museum. Upon the retirement of James W. Calhoun as Director of the 
Museum in 1973, Peter Herlan was Acting Director of the Museum for two months, 
until a permanent Director was chosen. 

Under Peter Herlan’s direction, the collections and exhibits of the Nevada State 
Museum were greatly enlarged and improved, and extensive cooperation between the 
various museums and with the University of Nevada was maintained, so that exhibits 
were set up for public viewing. He also was instrumental in backing conservation 
legislation at the state level. 

Peter Herlan described one butterfly subspecies, naming the very distinctive 
Limenitis archippus lahontani Herlan (1970 [1971]; J. Res. Lep. 9: 217-222). 
He has had one named in his honor, Phyciodes orseis herlani Bauer, and another 
has been proposed by Kurt Johnson (in press). 

Pete, as he was affectionately known to his many friends and colleagues, was the 
only truly active lepidopterist residing in Nevada. Under his leadership, the meetings 
of the Lepidopterists’ Society were held three times in Carson City with the Nevada 
State Museum as host. The Pacific Slope Meetings were held there in 1964 and in 
1976. The combined National and Pacific Slope Meetings were held there in 1970. 


214 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Pete’s knowledge of the remote corners of Nevada, and his and Barbara’s hos- 
pitality, made the Nevada Museum, and at times the Herlan’s home, the point of 
departure for many collectors visiting Nevada. Pete worked closely with many other 
students of butterflies, including David Bauer, the late O. E. Sette, and others; and 
the results of their studies have done much to help clarify the problems relating to 
Nevada Lepidoptera. 

Pete was a loyal friend, one of nature’s gentlemen, cheerful, outgoing, and always 
willing to help. He will be greatly missed by all who were fortunate enough to 
have known him. 

He is survived by his wife Barbara, of Carson City, and by three sons, several 
grandchildren, and two sisters. 


J. W. Tren, 125 Cedar Lane, San Jose, California 95127. 


VoLUME 31, NUMBER 3 


bo 
— 
Ot 


BOOK REVIEW 


A FreLtp GUIDE TO THE BUTTERFLIES OF THE WEsT INpDIEs, by Norman D. Riley. 
1975. Quadrangle/The New York Times Book Co., 10 East 53 St., New York, N.Y. 
10022, 224 p., illus. + 24 plates. Price $12.50 (U.S.). 


Although Riley's Butterflies of the West Indies is a welcome addition to the litera- 
ture of the butterflies of the area, it has several shortcomings that detract from the 
usefulness of the book. These shortcomings will be discussed section by section. 

Illustrations. In general the illustrations by Gordon Riley were found to be very 
good, but those by Brian Hargreaves, especially of the Satyridae and the Lycaenidae, 
left a lot to be desired. In some cases, it is doubtful that they will help identify the 
species involved. 

Introduction. The introduction is very good. The only criticism is that on Fig. 2 
the veins of the wings are identified in the usual European manner, i.e., using 
numbers. This might not create any complications, except that in the key to the 
families the veins are identified by their names. For the specialist this inconsistency 
will not pose any problem, but for the amateur it probably will. 

Glossary. The glossary was found to be satisfactory. 

Taxonomic discussions. In the taxonomic discussions many errors and omissions 
were found. Some of these might be because the author apparently has not done a 
great amount of collecting in the area and has worked mostly with preserved museum 
specimens and the available literature. This method of obtaining information about 
the biology of an area is not the best one, but in this case it is probably excusable 
because the West Indies are a politically and geographically heterogenous group of 
islands where travel to some of the smaller islands is very difficult, expensive, and 
time consuming. 

What is not excusable, are those errors and omissions that pertain to islands such 
as Cuba, Jamaica, and Puerto Rico which have rather well known lepidopterous 
faunas. The more obvious errors and omissions are the following: Danaus gilippus 
cleothera is not mentioned from Puerto Rico, although it has been reported before; 
Danaus cleophile is reported from Puerto Rico where it does not occur, the specimen 
reported in the literature as belonging to this species was described by Clark in 
1941 as Danaus plexippus portoricensis. 

Lycorea ceres cleobaea is reported from Jamaica. As far as I know, this is the first 
time this butterfly has been reported from the island, but no additional information 
is given about the new record. 

Prepona antimache insulicola is placed in synonymy of Prepona amphitoe Godart 
1823, this I find to be correct, but Puerto Rico is not included in the distribution of 
the species even though it has been reported many times from the island. 

Marpesia petreus is a common butterfly in Puerto Rico, not a rare one as is stated 
in the book. 

There is no subspecies of Eurema dina present in Puerto Rico. Therefore, the 
statement that the black border is present in the hindwing only in the Puerto Rican 
subspecies is erroneous. The species present in Puerto Rico is the closely related 
Eurema leuce sanjuanensis, which does not have a black border in the hindwing. 
Probably the author was confusing E. dina with Eurema portoricensis, the male of 
which has a black border in the hindwing. 

Although admittedly seasonal, Kricogonia lyside is a very common butterfly in 
Puerto Rico. The statement that the records of K. lyside of Cuba should probably 
refer to K. cabrerai is erroneous. Salvador de la Torre was one of the first lepidopter- 
ologists to consider castalia and lyside as conspecific. He published a paper on the 
genus Kricogonia in Cuba in which he clearly distinguished between lyside and 
cabrerai. 

Aphirissa statira cubana breeds in Puerto Rico, where it has been found to breed 


216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


on “quenepa’ (Melicocca bijuga L.) and in Cassia sp. Therefore, the statement that 
the specimens collected in Puerto Rico are migrants is incorrect. 

Battus polydamas is present in Puerto Rico as subspecies thyamus Rothschild & 
Jordan, not as subspecies policrates Hoffer. 

Papilio machaonides is reported for the first time from Puerto Rico, but again no 
additional information is given, even though the butterfly used in the illustration is a 
male apparently collected in Puerto Rico. Papilio androgeus epidaurus occurs in 
in Puerto Rico, but the island is not included in the distribution of the species. 

The statement that Epargyreus zestos is not present in any of the Greater Antilles 
is incorrect, it has been reported from Puerto Rico by Wolcott (The Insects of Puerto 
Rico), Comstock (The Insects of Puerto Rico and the Virgin Islands. Lepidoptera 
Rhopalocera), and Brown and Heineman (Jamaica and its Butterflies). Comstock, 
and Brown and Heineman also report it from Hispaniola. 

Ephyriades zephodes is reported as present in Puerto Rico although the only 
record that I know is a single specimen at the British Museum, however E. arcas 
philemon which is a rather common skipper in Puerto Rico is not reported as present 
in the island. 

Synapte malitiosa malitiosa is not reported from Jamaica, although it has been 
reported by Brown and Heineman to be rather widespread on the island. 

Euphyes singularis insolata is reported for the first time from Puerto Rico, but again 
the information is rather scanty. 

Panoquina panoquinoides panoquinoides has been collected in Puerto Rico a num- 
ber of times but it is not listed as present on the Island. 

Checklist. The checklist has also some errors and omissions. In some cases, spe- 
cies reported from an island in the discussion are not included under that island in 
the checklist (e.g., Danaus gilippus jamaicensis and Cyanophris crethona from 
Jamaica, Strymon acis mars and Euphyes singularis from Puerto Rico). On other 
occasions, species not reported from an island in the discussion are included in the 
checklist for the island (e.g., Cyanophris crethona from Hispaniola and Polites 
dyctinna from Puerto Rico). There is some inconsistency on what is considered 
doubtful or vagrant, e.g., Authochton neis, Colgia calchas, and Nisoniades bessus 
are considered vagrants in Jamaica, and Antigonus nearchus is considered as doubtful. 
But all of them are considered as doubtful by Brown and Heineman because they 
doubted the correctness of the locality labels of the specimens in the British Museum. 

Some misspellings were found throughout the checklist: Urbanus tellus instead of 
teleus, Anetia briaria instead of briarea, and Calisto herophilus instead of herophile. 

Bibliography. The bibliography is good, it includes the most important references 
for the area. 

In summary, I consider this a good book which could have been an excellent one 
if the author had paid more attention to certain details. 


Stuart J. Ramos, Department of Biology, University of Puerto Rico, College Sta- 
tion, Mayaguez, Puerto Rico 00708. 


EDITORIAL STAFF OF THE JOURNAL 
GrorcE L. Goprrey, Editor 


Illinois Natural History Survey, Natural Resources Building 
Urbana, Illinois 61801 U.S.A. 


| _ Wr11aM H. Aten, Associate Editor JAMEs G. STERNBURG, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study 
of Lepidoptera. Contributors should prepare manuscripts according to the following 
instructions. 

Text: Manuscripts should be submitted in duplicate, and must be typewritten, 
entirely double-spaced, employing wide margins, on one side only of white, 84% X 11 
inch paper. Titles should be explicit and descriptive of the article’s content, including 
the family name of the subject, but must be kept as short as possible. The first men- 


- tion of a plant or animal in the text should include the full scientific name, with 


authors of zoological names. Insect measurements should be given in metric units; 
times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). 
Underline only where italics are intended. References to footnotes should be num- 


% bered consecutively, and the footnotes typed on a separate sheet. 


Literature Cited: References in the text of articles should be given as, Sheppard 
- (1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically 
_ under the heading Lirerature Crrep, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, 
London. 209 p. 

196la. Some contributions to population genetics resulting from the 
study of the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard 
(1961, Ady. Genet. 10: 165-216) or (Sheppard 1961, Sym. Roy. Entomol. Soc. 
London 1: 23-30). 

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

Tables: Tables should be numbered consecutively in Arabic numerals. Headings 
for tables should not be capitalized. Tabular material should be kept to a minimum 
and must be typed on separate sheets, and placed following the main text, with the 
approximate desired position indicated in the text. Vertical rules should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer’s errors. Excessive author’s changes at this time will be charged 
to authors at the rate of 75¢ per line. A purchase order for reprints will accompany 
the proofs. 

Correspondence: Address all matters relating to the Journal to the editor. Short 
manuscripts such as new state records, current events, and notices should be sent to 
the editor of the News: Jo Brewer, 257 Common Street, Dedham, Massachusetts 
02026 U.S.A. 


ALLEN PRESS, INC. ee ee LAWRENCE, KANSAS 
usS~x 


CONTENTS 


SELECTION OF THE COCOON SPINNING SITE BY THE LARVAE OF HYALO- 
PHORA CECROPIA (SATURNUDAE). A. G. Scarbrough, J. G. Stern- 
burg, and G. P. Waldbauer eee 


A New SuBSPECIES OF PIERIS SISYMBRII ( PIERIDAE) FROM WESTERN 
GreaT Puains Reticr Forests. Kurt Johnson __..----— 


Stupies ON ReEsTINGA BuTTerFLIES. I. Lir—E CYCLE AND IMMATURE 
BioLocy OF MENANDER FELSINA (RIODINIDAE), A MyRMECOPH- 
mous MerarmMark. Curtis J. Callaghan ___-_ 


Bionomic NorEes ON SOME BAGworm Morus ( PsycHIDAE) OF TEXAS. 
Raymond W. Neck 0 


Two New Sprecirs oF CLEARWING Morus (SEsSIIDAE) FROM EASTERN 
NortH AMERICA CLARIFIED BY SEX PHEROMONES. W. Donald 
Duckworth and Thomas D. Eichlin 


GENERAL NOTES 


Antheraea polyphemus (Saturniidae) and Biblis hyperia (Nymphalidae) 
in Texas.. James E. Gillaspy 
Ovipositional mistake by a hackberry butterfly (Nymphalidae). Raymond 
Wii Neck hE Oe 
Butterflies associated with an army ant swarm raid in Honduras: the 
“feeding hypothesis” as an alternate explanation. Allen M. Young —— 
Catocala (Noctuidae) species taken in Clay County, Tennessee. Wayne 
A, “Miller 8 PO a 
Apparent long-distance dispersal by Pieris occidentalis (Pieridae). Arthur 
M. Shapiro, et Os 
The behavior of an inebriated Opsiphanes cassiae (Brassolidae). Luis 
D. Gomez 


153 


167 


173 


183 


19] 


_ Volume 31 1977 Number 4 


5 


t 


JOURNAL 


_ LeprpopTerists’ SociETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
Brin. > Publié par LA SOCIETE DES LEPIDOPTERISTES 

_ Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


27 December 1977 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


J. W. TrpEn, President KENELM W. Puiuip, Vice President 
I. F. B. Common, Ist Vice President Jutian P. DonaAnHuE, Secretary 
LionEL Hiccins, Vice President Joun M. Sniper, Treasurer 


Members at large: 


F. S. CHEw R. A. ARNOLD J. F. EMMEL 
D. F. Harpwick E. D. CasHatr R. R. GATRELLE 
J. B. ZrEGLER R. E. STANFORD AvP. Pra 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and 
formally constituted in December, 1950, is “to promote the science of lepidopterology 
in all its branches, . .. . to issue a periodical and other publications on Lepidoptera, 
to facilitate the exchange of specimens and ideas by both the professional worker and 
the amateur in the field; to secure cooperation in all measures” directed towards 
these aims. 

Membership in the Society is open to all persons interested in the study of 
Lepidoptera. All members receive the Journal and the News of the Lepidopterists’ 
Society. Institutions may subscribe to the Journal but may not become members. 
Prospective members should send to the Treasurer full dues for the current year, 
together with their full name, address, and special lepidopterological interests. In 
alternate years a list of members of the Society is issued, with addresses and special 
interests. There are four numbers in each volume of the Journal, scheduled for 
February, May, August and November, and six numbers of the News each year. 


Active members—annual dues $13.00 
Student members—annual dues $10.00 
Sustaining members—annual dues $20.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $18.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
John M. Snider, 3520 Mulldae Ave., San Pedro, Calif. 90732 U.S.A. 


Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA 


by Cyr F. pos Passos 


Price: Society members, $5.00 U.S.; non-members, $7.50 U.S. Paper covers, revisions 
of the Melitaeinae and Lycaenidae supplied separately. 


Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni- 
versity of Louisville, Louisville, KY 40208, U.S.A. 


The Lepidopterists’ Society is a non-profit, scientific organization. The known 
office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second 
class postage paid at Lawrence, Kansas, U.S.A. 66044. 


aor 
~ 


ee sete 


=; 


ee a 


J OURNAL OF 


Tue LEPIDOPTERISTS’ SOCIETY 


ohne 31 1977 a ey 


PRESIDENTIAL ADDRESS 1977: THIRTY YEARS 
AND COUNTING! 


So. NICOLAY 
1500 Wakefield Drive, Virginia Beach, Virginia 23455 


_ During the early summer of 1947, some 100 lepidopterists, amateur 
and professional alike, received a mimeographed, 12-page document, 
dated May 1947 and entitled, The Lepidopterists’ News. Across the 
top of the page, placed squarely beneath the masthead were the words 
in full capital letters—WELCOME TO CHARTER MEMBERSHIP 
Dee, GEPIDOPTERISTS. SOCIETY. 

For this document and the promise of more to follow, we had each 
paid the magnificent sum of $1.00 for a full year’s subscription! Volume 
1, number 1 contained 12 typewritten, mimeographed pages. The for- 
mat set the tone for the remainder of the year’s output. The first para- 
graph on page 1 presented a very clear picture of what was to follow: 

“Here is the first number of the periodical announced in the March 
letter. The first few numbers, like this one, will each contain twelve 
mimeographed pages, and in this first issue are several of the regular 
features which are planned for your interest and enlightenment. On 
page 3 is the review of Ford’s Butterflies, which will be followed regu- 
larly by reviews of other books of direct interest to lepidopterists. Pages 
4-7 carry monthly notices of papers on Lepidoptera in the current 
literature. Page 8 contains the first of a series of brief biographies of 
important American and foreign Lepidopterists. Page 9 introduces a 
series of discussions on life history studies of Lepidoptera, concerned 
more with methods of study than with published papers and_ books. 
Pages 10-12 are devoted to collecting trips planned by members, your 
list of exchange notices and other requests, and miscellaneous items. 
We urge you to contribute items for these pages. In the serial subjects, 


1Read at the Annual Meeting of the Lepidopterists’ Society in Boulder, Colorado on 24 July 
TELE 


218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


such as on pages 3, 8 and 9, we expect to be able to present articles by 
authorities on specific phases of the subjects. The final issue of the 
year will be accompanied by a list of the names, addresses, and special 
interests of all members.” 

By the end of the year, the initial 101 members of May had grown 
to 220, the News page total had reached 117 and contained an incred- 
ible wealth of information, most of it as timely and interesting today 
as it was then. To determine how well the original concept has held 
up, one need only read our present-day Journal and News. 

As most, if not all of you, are aware, the Lepidopterists’ Society 
was initiated by Dr. Charles L. Remington, then a graduate student 
at Harvard. During the first few months of its life, the entire staff of 
the Society, editorial and administrative, consisted of three people: 
Charles Remington and Harry Clench, editors and Mrs. Jeanne Reming- 
ton. In September of the first year, Harry Clench moved to Michigan 
and the entire work of publishing the News and conducting Society 
business was accomplished by the Remingtons. 

In 1948, dues went from the original $1.00 to $2.00 in order to, as 
Charles Remington so aptly put it, “provide urgently needed typo- 
graphical help for the enormous task of preparing the final copy of each 
News for the lithoprinter without reducing the size of the News volume.” 
Some additional editorial help was added in the personages of Peter 
Bellinger, Dr. Diaronoff, C. F. dos Passos and Takashi Shirozu who 
assisted in examining the large number of scientific journals, and ab- 
stracting all papers on Lepidoptera to be included in the News. Total 
membership in the Society nearly doubled by the end of 1948 to 397. 
Volume 2 contained a total of 124 pages. 

Thus did the Lepidopterists’ Society and its publication, the News 
continue in almost the same format under the guidance of its primary 
founder until 1950. It had, however, become apparent for some time 
that the task of publishing the News, in addition to performing all other 
administrative functions was more than two could handle; some issues 
in late 1949 and most numbers in 1950 were combined and even then 
some appeared late. Early in 1950, Cyril F. dos Passos prepared a draft 
constitution and by-laws, and the president pro-tem, Dr. J. H. McDun- 
nough appointed 3 temporary committees to prepare for the first annual 
meeting. At the meeting, held in New York on December 29-30, 1950, 
ballots previously mailed to all members were counted and _ the first 
full slate of officers in the Lepidopterists’ Society were duly elected 
and took office. Just a little footnote of sadness to the above—in 
looking over the names of the officers, I note that only our first secre- 


VoLUME 31, NuMBER 4 219 


tary, Dr. Fred Rindge and our first elected treasurer, Dr. Ben Ziegler 
are still living. 

In 1951, the News took on a different look with an attractive grey 
paper cover. The efforts of editors, Charles and Jeanne Remington, 
were assisted by associate editors, F. Martin Brown, Peter Bellinger and 
Eugene Munroe. Volume 5 contained 126 pages, plus a membership 
list of 15 pages. Total membership had reached nearly 500. 

The year 1952 brought the first radical change in appearance to the 
Lepidopterists News: volume 6 was, for the first time printed in type- 
set on a high grade, slick paper. in the 6 by 9 inch format we are all 
familiar with in today’s Journal. And, although it was not clearly stated, 
it had become obvious that the News was becoming a quarterly publica- 
tion with 2 and sometimes as many as 3 numbers combined and mailed 
together in a single issue. Presentation of material and format remained 
essentially the same for the next 5 years. Elections of officers were 
held each year to fill vacancies as they occurred in the various offices 
as well as in the Executive Council. In 1955, with volume 9, dues for 
regular members, which had been raised to $3.00 in 1952, were raised 
again to $4.00 per year. 

Volume 12 in 1958, although carrying the masthead of the Lepidop- 
terists News was the final issue to appear in the now familiar format 
and was, in fact, the precursor for our current Journal. Beginning with 
volume 10 in 1956, the membership list was already being mailed sep- 
arately as a mimeographed list of sizeable proportions. Volume 12, 
issued in 3 parts of 2 numbers each, contained a total of 240 pages and 
set the model for the Journal that followed. The News as we know it 
today began publication at this time with Dr. J. W. Tilden as editor. 
This new News, economically mimeographed or multilithed, provided 
the vehicle for material of a temporary nature but of immediate interest; 
the Journal was a continuation of the old News with no major change in 
format, content, size or editors. Volume 13 in 1959 appeared for the 
first time with the masthead of Journal of the Lepidopterists’ Society. 
Issued quarterly, it carried a total of 256 pages. 

From 1960, when dues rose to $5.00 per year for regular member- 
ship and then to $6.00 the following year, publication costs remained 
relatively stable for the next 10 years. The cost of a Sustaining Mem- 
bership, which, by the way, has always been one of the most beneficial 
ways of helping the Society financially, remained steady at $15.00 for 
the 10 years between 1961-1971. Total membership remained within 
the 500-600 bracket for about 6 years, between 1952 and 1959. From 


220 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


that time until the present, the Society has grown at the rate of a bit 
less than 10% per year. 

But, enough of figures and statistics. For those who really wish to 
go into the statistical history of the Society in complete detail, I recom- 
mend a copy of our Commemorative Issue. And, on this subject, I 
come to the main point of this presentation. The Society has, in recent 
years, experienced increasing difficulty in finding qualified members 
willing to give of their time and talents in order that the Society might 
continue to function smoothly and efficiently. The Commemorative 
Issue, originally conceived to celebrate our 25th anniversary, has been 
6 years in the works; we now hope to have it in the hands of individual 
members before the end of this, our 3lst year! This is not meant as an 
indictment of those who have expended so much time and effort to 
see it published; on the contrary, I would hate to try to estimate the 
incredible number of hours and days Roy Kendall has put into this 
project. And he is but one of a number who have had a hand in its 
creation. I use the example of the Commemorative Issue to amplify 
the following point. 

As President of the Society for the past year, I have, with the help 
of other council members, tried to find a successor for two of our most 
critically important officers, the Editor of the Journal and our Treas- 
urer. We have written letters, made personal contacts and appeals, 
and otherwise have attempted by those limited means within our reach 
to find suitable and qualified personnel to fill these most critical posi- 
tions. The 3-year terms of our Journal editor, Dr. George Godfrey, 
who has done such an outstanding job these past years and that of our 
Treasurer, Dr. John Snider, who has paid our bills and kept us solvent, 
will expire on 1 January 1978. At the time this was written, we had 
not been able to find interested and suitably qualified members willing 
to accept these positions. 

The Lepidopterists’ Society is now, and always has been, a non-profit 
organization. It has no salaried officers or employees, and all work is 
accomplished by members of the Society on a voluntary basis. My 
purpose in presenting the brief history of the growth of the Society 
was to illustrate that the needs of the Society also grew throughout the 
years, as did its size and the scope of its endeavors. These needs were 
not only monetary, as illustrated by the steady increase in dues from 
the original sum of $1.00 to the current $13.00; the Society, from its 
original 2-man operation has now an elected, functioning Executive 
Council or Staff, however you wish to call it, of 15 officers plus two 
separate editorial staffs, one for the Journal and one for the News. 


VoLUME 31, NUMBER 4 221 


As in most organizations, certain staff positions require more time and 
work than others, but all contribute their time and efforts without 
salary or remuneration of any sort. 

A number of entomological societies have flourished in years past 
but have eventually disappeared. I do not know the detailed history 
of any of them, but I would make an educated guess that the most 
common cause of their demise was a lack of interest . . . a lack of in- 
terest by the membership at large in seeing to it that the organization 
survived those crises, fiscal or organizational, that occasionally arise 
in every society from time to time. Payment of dues and the money de- 
rived therefrom is vitally important for survival of any society of 
this sort; but no less so is the active and continued support of its mem- 
bers who give of their time as well as their money. A careful reading 
of Dr. Snider's fiscal report for the past 3 years indicates that we paid 
our bills, that our income exceeded our expenses by a reasonable and 
fair amount and that we have a comfortable, if not excessive, balance 
to our credit. Dr. Snider has declined, and rightfully so, to continue 
beyond his 3-year term of office. 

It is with an audible sigh (Thank God) of relief, that I can now 
report to you that 2 days ago one of our members volunteered to have 
his name placed on the ballot for the coming year in the position of 
Treasurer, to serve for the next 3 years. 

Just a few days ago I received a letter from an individual requesting 
information on membership in the Society. It was a very short letter 
and I will quote it in part: 

“Dear Sir, I am writing to find if you are still associated with the 
Lepidopterists News/Journal. I would like to get membership but my 
letter to Sarasota, Florida was returned stamped unknown. I realize 
the issue I used is old, but I can’t believe this paper has quit.” 

Ladies and gentlemen, I too, find it hard to believe that this ‘paper 
is even near quitting. I find it hard to believe that out of a total mem- 
bership that must now be in the neighborhood of 1200 plus, there are 
not a number of individuals with talent, experience and, most of all, 
the desire to serve in order that the Lepidopterists’ Society will survive. 
Even if the current crises were solved today (and I believe that they 
will be) I can foresee many similar difficulties in the future unless more 
members, particularly the younger and/or newer members of the So- 
ciety find it also interesting and rewarding to serve with their time and 
talents in addition to just paying and receiving. 

This message is designed to reach each and every member of the 
Society. Those of you here in Boulder listening to this, are, for the most 


PAP) JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


part, those who have and/or are serving the Society in many ways. To 
those of you who will read this . . . when a member of the Executive 
Council contacts you with a request for your help in whatever capacity 
it might be, please reconsider that initial urge to say, “No.” Most of 
us find that 24 hours in one day, just aren't enough to accomplish all 
that we would like. So consider joining those who are in that same 
boat, for it is the busiest individual of all, who seems to find the time 
to add one more task to the growing list, and serve. 


NOTES AND NEWS 


Editorial Farewell 


Three years ago the Society asked me if I would take the editorship of the Journal. 
I received such an impressive letter about my qualifications for the position that I 
felt compelled either to accept or tell the individual who wrote to me that he was 
a liar. I finally agreed to take the responsibility, hoping that if I did a respectable 
job we would both save face. 

At the close of my term as your editor, I want to thank my associate editors, the 
reviewers, and other members of the Society plus several non-members who assisted 
and encouraged me during the past three years. Based on letters and comments 
that I have received, the Journal has remained the best publication solely devoted 
to Lepidoptera. The quality of the Journal is a reflection of those persons who 
have supported me. I especially want to single out William H. Allen for functioning 
as my technical editor in spite of his hectic schedule as a budding journalist. For 
the current volume I thank Paul H. Faber for providing the cover illustration of 
Abbott’s Sphinx, Sphecodina abbottii (Swainson). 

It is my pleasure to announce that Dr. Austin P. Platt will be the new editor ef- 
fective 1 January 1978. More information about Dr. Platt will appear in the NEWS. 
I wish him the same cooperation and assistance that I received. Please see his 
special notice to contributing authors on page 274 of this issue. 


George L. Godfrey 


VOLUME 31, NUMBER 4 223 


PHENOTYPIC WING PATTERN MODIFICATION BY VERY 
BRIEF PERIODS OF CHILLING OF PUPATING ARICIA 
ARTAXERXES VANDALICA (LYCAENIDAE). 
ARICIA STUDIES NO. 16 


Ove H@EcH-GULDBERG AND ARNE LINDEBO HANSEN 


Natural History Museum, Aarhus, Denmark 


Since Dorfmeister “in 1845” showed that divergent forms were devel- 
oped when butterfly pupae were kept at increased or decreased tempera- 
ture, many scientists have worked on this problem because the effect of 
temperature might explain some of the aberrant forms which now and then 
are found in natural populations. Some important papers on temperature- 
induced forms in Lepidoptera are referred to below that concern the age 
of the pupa at the time of cooling, the temperature used, and the duration 
of the treatment. 

Merrifield (1893) and Standfuss (1896) independently found that 
pupae chilled immediately after pupation died or produced crippled but 
normal-patterned imagines. To obtain altered imagines the pupae were 
chilled not earlier than 12 hours (h) after pupation, when the sensitive 
period begins. The chilling must continue for at least 14 days. Siiffert 
(1924), working on Araschnia levana L. and Aglais urticae L., found a 
sensitive period some days after pupation; there was no effect with 
earlier chilling. The cooling period was 10 days. Reinhardt (1969), who 
definitely solved the problems of Araschnia’s seasonal dimorphism, only 
conducted a few experiments with pupae as young as 0-6 h; the other 
experiments were from 6-36 h after pupation. Kiihn (1926) also found 
no changes in Vanessinae if cooling took place immediately after pupa- 
tion. The effect increased from 6-24 h after, but there was no effect 
after 72 h. The temperature used was low, from —3 to —-10°C, for two 
days. Very exact experiments were conducted by Kohler & Feldotto 
(1935), who, working on Vanessinae, found the heat-sensitive period to 
be from 0-48 h after pupation. Every pattern element was found to have 
its special sensitive period. In these experiments only increased tem- 
perature, for short periods, was used. In most of these experiments only 
Nymphalidae were examined, but Merrifield also investigated Geometri- 
dae and Kiihn also worked with certain moths. 

Krodel (1904) performed experiments with three species of Lycaeni- 
dae: Plebeius (Lycaena) argus L., Lysandra (Lycaena) coridon Poda, 
and Agrodiaetus (Lycaena) damon Schiff. The pupae were never less 
than 5 h old; in the course of 6 days and nights they were exposed to 12 


224 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


CLEP eS 


Fig. 1. Wild rare forms of Aricia artaxerxes vandalica: (left) 9, underside, 
Tversted Strand, Jutland, 16 July 1961, spot value 3—f. “caeca”; (right) ¢, upper- 
side, Tornby Strand, Jutland, 11 July 1960, f. “albicostalis” (= “albivenata” ). 


cold spells each of 6 h duration, during which the temperature went 
down to -14°C. In this way Krodel produced numerous cold forms, 
partly corresponding to forms taken in the open, and he was of the 
opinion that these forms could be explained as ancient characters which 
for many generations have remained latent, but which reappear under 
certain circumstances, such as cold treatment. Lorkovic (1938) found 
obsolescence of eyespots by cooling pupae of Polyommatus icarus Rott. 
Hgegh-Guldberg (1971) cooled pupae (1-24 h old) of the same species 
at +2 to +5°C for 4 weeks and observed many changed imagines. 
Lorkovic (1943) observed reduction of eyespots by cooling—for 3 weeks 
—“young” pupae of Everes argiades Pall. 

Another genus of Lycaenidae is Aricia, and the following deals entirely 
with A. artaxerxes F. and A. agestis Schiff. These are two closely related 
species widely distributed in Europe. In both species, but especially in 
some A. artaxerxes populations, aberrant forms occur quite often. Two rare 
forms are depicted in Fig. 1, to be compared with the normal appearance, 
Figs. 2A & D. It is shown in Fig. 2D that the blackish upperside has 
marginal lunules most distinct on the hindwings, and without the white 
nervures at the tip of forewings which are seen in Fig. 1. The normal 
underside, Fig. 2A, has a complex system of eyespots when compared 
with the aberrant form in Fig. 1. This eyespot pattern varies gradually 
from that extreme; the degree can be expressed as “spot value,” which 
registers the number of spots and centres on one forewing and hindwing, 
including all pupillations plus the sum of their centres. Maximum is 42: 


VOLUME 31, NuMBER 4 225 


Fig. 2. Wild average forms and chilled specimens of Aricia artaxerxes vandalica. 
Undersides: (A) ¢ from nature (61,60), Tornby Strand, Jutland, 4 June 1961, spot 
value 25, normal form; (B) Pl @ (73,61), ex Tornby Strand, pupa 1 min, chilled 
9 h, spot value 12; (C) Pl @ (73,125), ex Tomby Strand, pupa 15 min, chilled 2 
“nights,” spot value 3, f. “caeca.” Uppersides: (D) ¢ from nature (66,53), Tversted 
Strand, Jutland, 6 July 1966, no white elements, normal form; (E) Pl @ (73,59), 
ex Tornby Strand, pupa 1 min, chilled 2 “nights,” f. “snelleni,” f. “panalbisignata”; 
(F) Pl 2 (73,62), ex Tomby Strand, pupa 1 min, chilled 9 h, f. “albicostalis” 
(= “albivenata’” ). 


1+ 1,7+7 (counting the lowest double discal spot as 1); hindwing 
4+4,1+41, 8+ 8 (the second discal spot from below is counted as | 
even if double). It is very rare to find a spot number exceeding 35. The 
minimum is 3—only the two discoidal spots remain, the hind one without 


226 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


pupil (f. “panobsoleta” (=“caeca”) + f. “carteri”). In A. artaxerxes 
vandalica the normal spot value varies from 26 to 33 with nearly equal 
representation of each value and very few individuals with lower or 
higher counts. 

Jarvis and Hgegh-Guldberg have conducted numerous experiments by 
rearing large series of butterflies from eggs of both species. Young 
pupae have been cooled for various periods, but pupae less than | h old 
have been avoided because they are very fragile. Jarvis (1959) demon- 
strated that four conditions had to be fulfilled if changes are to be ob- 
tained: (1) temperature must be constant, (2) it must be +1 to +3°C, 
(3) the cooling period must be 20 days or more, (4) the age of the pupa 
must be 1-2 h (or up to 24 h), but most extreme results are obtained in 
the younger pupae. 

Hgegh-Guldberg (1968), after examining large numbers of bred sibs, 
confirmed this. He also had the same results at +2 to +5°C, and he 
added a fifth condition: that a positive result also depended on a par- 
ticular genotype in the individual and in the population. Jarvis also 
suggested that extreme forms, such as f. “caeca” (Fig. 1, left) found in 
the wild, might be due to accidental cooling on cold nights during a 
critical phase of pupation. This was a tempting theory, but in experi- 
ments such a phase of short duration had never produced these forms. 
Since pupation in nature in these species takes place in the early summer, 
where a long-lasting, very low and constant temperature is not found, it 
seemed impossible to explain aberrant wild forms as cold forms. Later, 
other experiments by Hgegh-Guldberg & Jarvis (1969), with conditions 
which might occasionally be found in nature, also produced no changes 
in the imaginal pattern. These experiments were with prepupae (24 h 
before pupation) and pupae (2-12 h after pupation), cooled at various 
temperatures for shorter periods. Thus, still no possibility existed for 
explaining natural aberrant forms as the result of cold. 

But a solution to the problem was found by starting cooling just 
around the time of pupation. At that time the sensitivity is so high that 
only a short cooling sometimes suffices to create changes in the imaginal 
pattern (Fig. 3). First the author (Hgegh-Guldberg, 1974a) had to 
establish the timetable for pupation of Aricia by careful observation of 
prepupae; thus it is possible to judge when the shedding of the larval 
skin will occur. 


EXPERIMENTAL PROCEDURE 


One hundred and thirty-four specimens of Aricia artaxerxes vandalica 
were reared in single vials indoors from eggs laid by two females from 
Tornby Strand and Skallerup Klit found on 17 June 1973. Thirty-three 


bo 
bo 
~| 


VOLUME 31, NuMBER 4 


mature larva 


EES prepupa, 180 min. before pupation 
- 2-3 d 
{_- prepupa, 120-30 min. before ee 


: 


- formation of prepupa 


ED Prepupa, 15-2 min. before 


pupation 


BOS pupa, 0-10 min. after 


| P4490 R 

i] ~ er 
| Zf, ) older 
I 

I 

i 


16 days 


| 
I 
I 
| 
| 
I 
| 
| 
I 
I 
I 
I 
| 
I 
| 
I 
I 
| 
| 
I 
J 
| 
| 


YS 


I 

! 

I 

I | 
I | 
| I 
! | 
| I 
I I 
I. | 
I | 
| I 
| I 
I | 
| I 
I | 
I I 


“emergence 


aay ny, 
no changes no changes 


| 


Normal habitus Some individuals exhibit changes: 


appearance of white elements 


reduction of spots f 
(upper sides) 


(under side) (upper side) 


no white (under sides) 

elements white in cell, white nervures 
and along border on forewing 
of hindwing and 
forewing 


The same forms (and many others) are created in most imagines, if 
1-24 h. old pupae are constantly chilled for 2-4 weeks, and every 
imago exhibits some changes if the chilling period is 6 weeks. 


y 


Fig. 3. Effect of chilling Aricia artaxerxes vandalica at +2 to +5°C for 1, 2 or 3 
“nights” and then returning to room temperature. 


individuals served as controls; they were kept at room temperature 
(+18-20°C) until emergence. Seventy mature larvae and prepupae, 
whose time for pupation was judged from the appearance, and 31 pupae 
in the first minutes (min) after pupation were cooled at +2 to +5°C for 
one, two, or three periods of 9-12 h, the latter two cases with room- 
temperature periods of 12 h in between. All pupae were kept at room 
temperature for the remaining 14-16 days of the pupal period. Normally 
the pupation takes place at any time, so the start of cooling could be at 


228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. The effect on “spot value” of cooling Aricia artaxerxes vandalica during 
critical phases of pupation. 


No. with 
} low spot 
Experimental treatments: No. Died Crippled value x? Pp 


Room temperature 
controls 33 3 
Cooling at +2-5°C 


Age in relation Cooling 


Stage of to time of periods 
development pupation “nights” 
Mature Y 5 1 
larva 3 8 0.1405 FOr, 0.7 
Prepupa —2 days oy} 4 1 
—1 day 2 4 0.1388 0.5 0.7 
—2 to 15h 1 3 
—3 to 2.0 h DA Z, 1 
—2h 3 4 1 % 0.2477 0.3 0.5 
—1 to 0.5 h 1 1 2 
Dh 11 lt 1 4 
3 15 i 3 1.1758 072 0.3 
—15 to 2 min IL o i 
Ds 3 1 3 
3 ] il 7.8526 0.01 
Pupa 0 to 10 min 1 16 5 
Y, in 1 3 
8) 8 il 3.1662 0.1 0.05 
x? is calculated from nC ade be|) — 14n)?2/(a ai b)(c + d)(a + c)(b + d) 
a = number of controls with normal spot value. 
b = number of controls with low spot value. 
c = number of experimental animals with normal spot value. 


= number of experimental animals with low spot value. 
P is the probability that low spot values occur with the same frequency as in the control group. 


any time of the day and night. These conditions are as close as possible 
to natural conditions occurring in early Danish summer, i.e., one, two, or 
three cold nights rising to 15-25°C in the daylight hours. 


RESULTS 


In some of the cooled imagines, differences were found partly in the 
presence of white elements of the upperside and partly in the “spot 
value.” It is seen from Table 1 that in the cooled groups there is a 
tendency toward low spot values; this tendency is statistically significant 
at the P < 0.05 level just around pupation. Of the individuals with low 
spot values, the specimen in Fig. 2C is a true f. caeca (panobsoleta), a 
very rarely found form in nature. It had been cooled 15 min, before 


VoLUME 31, NUMBER 4 237 


TaBLE 4. Mating status and free lipid content of a sample of 111 females col- 
lected at Site Alpha on 25-26 January 1977. 


No. of spermatophores in bursa Free lipids in abdomens 
0 1 2 8 None Moderate f Gonsiicrble 
85% 11% 2% 2% 24% 39% 37% 


tain extent can override short daylength, 6ogenesis is minimal to absent 
at daylengths of less than 11-12 h and temperatures of less than 20°C 
(Barker & Herman, 1976). Site Alpha is at approximately 20° North 
Latitude where at winter solstice the light period is 10 h 48 min (Finch 
& Trewartha, 1949). By 1 February, daylength would be critical, i.e., 
ca. 11.3 h (Beck, 1968), with the stage set for the massive panmictic 
mating ceremony to be triggered by rising temperatures. 

We obtained experimental evidence in support of this thermal trigger- 
ing hypothesis. Fifteen females and 14 males collected at Site Alpha 
on 27 January were taken to Amherst, Massachusetts and placed in 
nylon cages the following evening in a controlled environment room 
at 22-30°C and 75-81% relative humidity. By 31 January, with only 
two days of exposure to the 15 h light period, 9 of the 15 pairs had mated. 

These physiological findings are in remarkable agreement with the 
observations on spring and summer breeding dates and range of the 
monarch in eastern North America (Urquhart, 1960; Williams et al., 
1942). Since Gogenesis occurs slowly at low temperatures and does not 
peak until 28°C (Barker & Herman, 1976), the mated females are free 
to leave the colony and migrate northward initially unencumbered with 
large numbers of eggs, but with an increasing rate of egg maturation 
as both seasonal temperature and daylength increase. 


Robust Condition of the Butterflies 


The general appearance of the majority of monarchs at Site Alpha 
was of exceptionally high quality compared to butterflies from over- 
wintering sites in California (Tuskes & Brower, l.c.). Few butterflies 
were tattered and the abdomens of most appeared robust. The dissection 
sample indicated that three-fourths of the females had moderate to con- 
siderable amounts of free lipids (Table 4) and well developed fat bodies 
(Brower, Calvert & Hedrick, ms in prep.) as is characteristic of fall mi- 
grants (Beall, 1948; Cenedella, 1971; Brown & Chippendale, 1974). The 
butterflies also appeared to be in a favorable state of water balance. 
On the clear days we observed them flying down along the draw to the 


238 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Thermoregulatory struggle of more than 10,000 monarch butterflies which 
released their grip in response to smoke from a small fire (less than one meter in 
diameter ) which drifted laterally and upwards through the clusters in the trees. The 
ambient temperature in the forest was about 13°C at the time of the incident (ca 
1500), 2°C below the thoracic temperature at which monarchs are able to fly. 
Original 35 mm Kodachrome by George D. Lepp. 


creek and many thousands were drinking at moist spring sites and along 
the creek itself. 

Nectaring was observed on several flowering herbs and shrubs, in- 
cluding at least one species in the genus Bidens, two in Lupinus, four in 
Senecio, and two in Stevia. However, the numbers of butterflies so 
vastly exceeded the available flowers in the area that nectar during 
their overwintering period cannot be an important source of carbo- 
hydrate. 


Predation by Orioles 


We also observed attacks by birds, identified with the use of Peterson 
& Chalif (1973). At 1120 on 26 January, a small flock of Bullock’s 
Oriole (Icterus bullockii Swainson) flew into a fir tree which had no 


> 


Fig. 2. Because of the low ambient temperature, many of the smoked-down 
butterflies could not fly back to the clusters. This picture, taken the next morning, 


VOLUME 31, NuMBER 4 


shows them having crawled up into less vulnerable positions on projecting rocks 
and vegetation. By noon, the temperature had risen sufficiently and virtually all 
the butterflies had flown back up into the clusters. Original 35 mm Kodachrome by 
George D. Lepp. 


240 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


roosting monarchs on it. From this tree they flew singly to an adjacent 
fir laden with monarchs and upon landing displaced several butterflies, 
which either dropped to the ground or flew off into the forest. Only 
clustering butterflies were attacked and the birds remained in a small 
area feeding upon them for about 45 min. Ingestion per se was not ob- 
served, but one monarch wing floated down and beak snapping could 
be heard frequently among the clusters. A kiskadee-like flycatcher 
(probably Pitangus sulphuratus L.) was observed perching near a 
cluster, but was not actually seen feeding on the monarchs. Two other 
patterns of apparent bird predation occurred extensively along the log- 
ging cut bordering the eastern side of Site Alpha. Numerous individual 
wings littered the forest floor along with many maimed but still living 
butterflies, including ones with one or more wings missing and others 
lacking their abdomens. Comparable predation has been noted in Cali- 
fornia (Kammer, 1970). 


Mortality Caused by Man 


Overall, natural mortality seemed to be low at Site Alpha. However, 
upslope from the colony, dead butterflies littered the ground to an ex- 
tent that their odor of decomposition was strongly evident. Based on 
Calvert’s observaticn of the upper boundary of the site on 31 December, 
we believe that a substantial percentage of the colony may have been 
eaten and trampled by cattle led to Site Alpha by local ranchers. 


Likelihood of Many Overwintering Sites in Mexico 


Urquhart (1976) and Urquhart & Urquhart (1976) maintained that 
overwintering of the monarch butterfly in Mexico is geographically re- 
stricted, involving as few as four sites in one general area. We believe 
that, as is the case in California (Williams et al., 1942) numerous over- 
wintering sites will be found in Mexico at locations having ecological 
characteristics similar to those at Site Alpha. The principal reason for 
our contention is that roosting in a very limited area would make the 
butterflies highly vulnerable to fire. Not only does overwintering occur 
during the dry season, but this area of Mexico is in the Trans-Mexico 
Volcanic Belt (Anon., 1961). Past volcanic activity must have set many 
large fires which would have decimated whole colonies. 

The pressing need to mount a conservation effort to preserve Site 
Alpha has been described elsewhere (Brower, 1977). 


VoLUME 31, NuMBER 4 241 


SUMMARY 


More than 14.25 million monarch butterflies in prime condition over- 
winter in a 1.5 ha site located in a coniferous forest in the trans-volcanic 
belt of Mexico. The site is characterized by thermal stability, low wind 
velocity, and high humidity. The reproductive status of the butterflies 
and their thermoregulatory activities in the colony support conclusions 
from experimental physiology. Genetic implications of mass mating 
behavior prior to dispersal and northward migration appear great. 


ACKNOWLEDGMENTS 


We are grateful to the following people for help during the initial stages 
of our search for the location of Site Alpha: Natalie Drake, Elizabeth 
Mahan, Dr. Victoria Foe, Dr. Michael Dennis, Dr. Richard Lindley, Dr. 
Paul Tuskes, and Dr. Steven P. Lynch. We also thank Dr. Rudolf M. 
Schuster and H. E. Ahles for identifying the plants and Christine M. 
Moffitt and Helen S. Smith for critically reading the manuscript. The 
research was supported by a grant (DEB 7514265) from the National 
Science Foundation with L. P. Brower as Principal Investigator. 


LITERATURE CITED 


Anonymous. 1961. Tectonic map of Mexico. Geological Society of America. 

Barker, J. F. ann W. S. Herman. 1973. On the neuroendocrine control of 
ovarian development in the monarch butterfly. J. Exptl. Zool. 183: 1-9. 

1976. Effect of photoperiod and temperature on reproduction of the 
monarch butterfly, Danaus plexippus. J. Insect Physiol. 22: 1565-1568. 

Beatt, G. 1948. The fat content of a butterfly, Danaus plexippus Linn., as af- 
fected by migration. Ecology 29: 80-94. 

Beck, S. D. 1968. Insect photoperiodism. Academic Press, N.Y. and London. 
viii + 288 p. 

Brower, L. P. 1961. Studies on the migration of the monarch butterfly. 1. Breed- 
ing populations of Danaus plexippus and D. gilippus berenice in south central 
Florida. Ecology 42: 76-83. 

1962. Biology of the monarch butterfly. Ecology 43: 181-182. 

1977. Monarch migration. Natural History 86(6): 40-53. 

Brower, L. P. & J. C. HuserrH. 1977. Strategy for survival: behavioral ecology 
of the monarch butterfly. 16 mm, sound, color, 30 minute film. Harper and 
Row, Inc., 10 E. 53rd St., N.Y. 10022. 

Brown, J. J. & G. M. CurppENDALE. 1974. Migration of the monarch butterfly, 
Danaus plexippus: energy sources. J. Insect Physiol. 20: 1117-1130. 

Burns, J. M. 1968. Mating frequency in natural populations of skippers and 
butterflies as determined by spermatophore counts. Proc. Nat. Acad. Sci. 
USA 61: 852-859. 

CENEDELLA, R. J. 1971. The lipids of the female monarch butterfly, Danaus 
plexippus, during fall migration. Insect Biochem. 1: 244-247. 

Fincn, V. C. & G. T. TREwartuHa. 1949. Elements of geography, 3rd ed. McGraw- 
Hill Book Co., Inc. N.Y. xii + 711 p. 


242 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Herman, W. S. 1973. The endocrine basis of reproduction inactivity in monarch 
butterflies overwintering in central California. J. Insect Physiol. 19: 1883-1887. 

1975. Endocrine regulation of post-eclosion enlargement of male and 
female reproductive glands in monarch butterflies. Gen. Compar. Endocrin. 
26: 534-540. 

Hitt, F. H., Jn, A. M. WENNER, & P. H. Wetts. 1976. Reproductive behavior 
in an overwintering aggregation of monarch butterflies. Amer. Midl. Nat. 95: 
10-19. 

Kammer, A. E. 1970. Thoracic temperature, shivering, and flight in the monarch 
butterfly, Danaus plexippus (L.). Z. Vergl. Physiol. 68: 334-344. 

1971. Influence of acclimation temperature on the shivering behavior of 
the butterfly Danaus plexippus (L.). Z. Vergl. Physiol. 72: 364-369. 

Kammer, A. E. & J. Braccrt. 1973. Role of the wings in the absorption of radiant 
energy by a butterfly. Comp. Biochem. Physiol. 45A: 1057-1064. 

Pan, M. L. & G. R. Wyarr. 1976. Control of vitellogenin synthesis in the monarch 
butterfly by juvenile hormone. Developmental Biol. 54: 127-134. 

Pererson, R. T. & E. L. Cuauir. 1973. A field guide to Mexican birds. Houghton 
and Mifflin Co., Boston. xxii + 298 p. 

SANCHEZ, O. S. 1976. La flora del Valle de Mexico. 3rd ed. Printed by the 
author, Mexico, D.F. viii + 520 p. 

ScumMipT-Korenic, K. 1975. Migration and homing in animals. Springer-Verlag, 
Berlin. xii + 99 p. 

Tusxes, P. M. & L. P. Brower. 1978. Overwintering ecology of the monarch 
butterfly, Danaus plexippus, in California. Ecol. Ent. 3: (in press). 

Urounart, F. A. 1960. The monarch butterfly. Univ. of Toronto Press, 


xxiv + 361 p. 
1976. Found at last: the monarch’s winter home. Nat. Geogr. Mag. 150: 
160-173. 


Uroqunart, F. A. & N. R. UrquHArtT. 1976. The overwintering site of the eastern 
population of the monarch butterfly (Danaus p. plexippus; Danaidae) in 
southern Mexico. J. Lepid. Soc. 30: 153-158. 

Urqunart, F. A., N. R. Urqunart, & F. Muncer. 1970. A study of a con- 
tinuously breeding population of Danaus plexippus in southern California com- 
pared to a migratory population and its significance in the study of insect 
movement. J. Res. Lep. 7: 169-181. 

WasseRTHAL, L. T. 1975. The role of butterfly wings in regulation of body 
temperature. J. Insect Physiol. 21: 1921-1930. 

WixuiaMs, C. B. 1930. The migration of butterflies. Oliver and Boyd, Edinburgh. 
reese ANS) 40% 

Wituiams, C. B., G. F. Cocxpitt, M. E. Gress, & J. A. Downes. 1942. Studies in 
the migration of Lepidoptera. Trans. Roy. Entomol. Soc. Lond. 92 (Part 1): 
NOI ZS3; 


VOLUME 31, NUMBER 4 243 


REVISION OF NORTH AMERICAN BOLORIA SELENE 
(NYMPHALIDAE) WITH DESCRIPTION 
OF A NEW SUBSPECIES 


STEVE KOHLER 


Montana Department of Natural Resources and Conservation, Division of 
Forestry, 2705 Spurgin Road, Missoula, Montana 59801 


This study began as an effort to determine the validity of the sub- 
species name nebraskensis (Holland) as applied to Boloria selene (Denis 
& Schiffermiiller ) populations in a limited area of southeastern Nebraska. 
As the study progressed, it became evident that the eastern limit of the 
range of nebraskensis was not known and that the relationship of ne- 
braskensis to myrina (Cramer) was not well understood in the pub- 
lished literature. It also was apparent that considerable variation in 
both size and markings was present within the described subspecies of 
selene and that a considerable amount of intergradation occurred in 
some geographic areas between the populations traditionally considered 
as subspecies of selene. For these reasons, the study was expanded to 
include the entire range of B. selene in North America (Fig. 1). 

The North American subspecies of selene were formerly placed under 
the species myrina in the genus Brenthis Hiibner (McDunnough, 1938). 
Clark (1941) recognized that myrina and the European selene were 
conspecific and placed the North American subspecies under selene. 
Studies of male genitalia by dos Passos & Grey (1945) reinforced Clark's 
placement of myrina under selene. These studies showed that all the 
North American representatives of the genus Brenthis, including selene, 
should be placed in the genus Boloria Moore. 

The populations considered subspecies of selene actually represent 
aggregations of populations homogeneous in wing characters but on a 
larger scale representing the distinct clustering of characters amid broad 
clines. Subspecies names are valuable in characterizing these groups 
as to phenotypic appearance, habitats, and ecological-range affinities. 
Subspecies names are used in this paper for these reasons. 

A total of 1,264 specimens, representing all of the named subspecies, 
was obtained from museum and private collections for examination. 
This paper presents the results of the study of these specimens, inte- 
grated with previous knowledge of the species. Seven subspecies are 
recognized. 

Methods. Characters used to differentiate the subspecies of selene are 
given in Tables 1 and 2. The degrees of dorsal maculation, dark dorsal 


244 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TaBLE 1. Comparison of characters of the North American subspecies of Boloria 
selene. 


nebras- sabulo- tolland- atroco- terrae- 
myrina kensis collis ensis albequina _ stalis novae 
Number of typical 
specimens examined 
Males 130 232 51 88 U5) 159 14 
Females 63 166 8 53 6 80 il 
FW length, mm 
Males 20.98 24.78 Upyey 19.55 19.17 20.08 20.67 
Females 21.85 DEO DTD 20.46 2.0.08 20.57 21.50 
Degree of maculation 
(dorsal )* 2.29 1.99 ES 2, 1.50 Das bro) PAB) BaesBre) 


Degree of dark mar- 

ginal wing scaling 

(dorsal )* IW 52 1.14 1.80 122. 3.00 3.19 1.93 
Degree of dark basal 

wing scaling 

(dorsal )* 1.70 87 1.20 DA 4,33 2.10 2.47 
Percentage of speci- 

mens with basal spot 


in discal cell of dor- 
sal HW obscured 30 6 15 79 100 60 93 


Percentage of speci- 
mens with anvil- 
shaped silver spot in 
cell Cuz of ventral 
HW divided 12 4 4 30 33 6 7 


Yellow scaling invad- 
ing cinnamon color 
of basal and discal 
areas of ventral HW? ++ ++ t4++ 44444 444+ + + 


1 Values represent an average obtained by visually rating each specimen on a scale of 0-5. 
2 Average appearance of each subspecies based on a composite of all specimens examined. 


marginal wing scaling, and dark basal wing scaling dorsally, given for 
each subspecies, represent averages of numerical values obtained by 
visually rating each specimen on a scale of zero to five. The values for 
the amount of yellow scaling that invades the cinnamon color of basal 
and discal areas of the ventral HW (hindwing) represent the average 
appearance of each subspecies based on a composite of all specimens 
examined. These are also on a scale of zero to five. Division of the 
anvil-shaped silver spot in cell Cu, of the ventral HW is not intended 
to be a main character for separation of subspecies. It is used to point 


out the close relationship of albequina (Holland) to tollandensis (Barnes 
& Benjamin ). 


VOLUME 31, NUMBER 4 237 


TABLE 4. Mating status and free lipid content of a sample of 111 females col- 
lected at Site Alpha on 25-26 January 1977. 


No. of spermatophores in bursa Free lipids in abdomens 
0 1 2 3 None Moderate Considerable 
85% 11% 2% 2% 24% 39% 37% 


tain extent can override short daylength, 6ogenesis is minimal to absent 
at daylengths of less than 11-12 h and temperatures of less than 20°C 
(Barker & Herman, 1976). Site Alpha is at approximately 20° North 
Latitude where at winter solstice the light period is 10 h 48 min (Finch 
& Trewartha, 1949). By 1 February, daylength would be critical, i.e., 
ca. 11.3 h (Beck, 1968), with the stage set for the massive panmictic 
mating ceremony to be triggered by rising temperatures. 

We obtained experimental evidence in support of this thermal trigger- 
ing hypothesis. Fifteen females and 14 males collected at Site Alpha 
on 27 January were taken to Amherst, Massachusetts and placed in 
nylon cages the following evening in a controlled environment room 
at 22-30°C and 75-81% relative humidity. By 31 January, with only 
two days of exposure to the 15 h light period, 9 of the 15 pairs had mated. 

These physiological findings are in remarkable agreement with the 
observations on spring and summer breeding dates and range of the 
monarch in eastern North America (Urquhart, 1960; Williams et al., 
1942). Since 6ogenesis occurs slowly at low temperatures and does not 
peak until 28°C (Barker & Herman, 1976), the mated females are free 
to leave the colony and migrate northward initially unencumbered with 
large numbers of eggs, but with an increasing rate of egg maturation 
as both seasonal temperature and daylength increase. 


Robust Condition of the Butterflies 


The general appearance of the majority of monarchs at Site Alpha 
was of exceptionally high quality compared to butterflies from over- 
wintering sites in California (Tuskes & Brower, l.c.). Few butterflies 
were tattered and the abdomens of most appeared robust. The dissection 
sample indicated that three-fourths of the females had moderate to con- 
siderable amounts of free lipids (Table 4) and well developed fat bodies 
(Brower, Calvert & Hedrick, ms in prep.) as is characteristic of fall mi- 
grants (Beall, 1948; Cenedella, 1971; Brown & Chippendale, 1974). The 
butterflies also appeared to be in a favorable state of water balance. 
On the clear days we observed them flying down along the draw to the 


238 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Thermoregulatory struggle of more than 10,000 monarch butterflies which 
released their grip in response to smoke from a small fire (less than one meter in 
diameter ) which drifted laterally and upwards through the clusters in the trees. The 
ambient temperature in the forest was about 13°C at the time of the incident (ca 
1500), 2°C below the thoracic temperature at which monarchs are able to fly. 
Original 35 mm Kodachrome by George D. Lepp. 


creek and many thousands were drinking at moist spring sites and along 
the creek itself. 

Nectaring was observed on several flowering herbs and shrubs, in- 
cluding at least one species in the genus Bidens, two in Lupinus, four in 
Senecio, and two in Stevia. However, the numbers of butterflies so 
vastly exceeded the available flowers in the area that nectar during 
their overwintering period cannot be an important source of carbo- 
hydrate. 


Predation by Orioles 
We also observed attacks by birds, identified with the use of Peterson 


& Chalif (1973). At 1120 on 26 January, a small flock of Bullock’s 
Oriole (Icterus bullockii Swainson) flew into a fir tree which had no 


> 


Fig. 2. Because of the low ambient temperature, many of the smoked-down 
butterflies could not fly back to the clusters. This picture, taken the next morning, 


VoLuME 31, NuMBER 4 


S 


<a 3+ ee) mares 
——-—re 


‘ae OO , 


een eH a“ 


» 
foal 


J 


shows them having c1 


‘-awled up into less vulnerable positions on projecting rocks 
and vegetation. By noon, the temperature had risen sufficiently and _ virt 


ually all 
the butterflies had flown back up into the clusters. Original 35 mm Kodachrome by 
George D. Lepp. 


240 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


roosting monarchs on it. From this tree they flew singly to an adjacent 
fir laden with monarchs and upon landing displaced several butterflies, 
which either dropped to the ground or flew off into the forest. Only 
clustering butterflies were attacked and the birds remained in a small 
area feeding upon them for about 45 min. Ingestion per se was not ob- 
served, but one monarch wing floated down and beak snapping could 
be heard frequently among the clusters. A kiskadee-like flycatcher 
(probably Pitangus sulphuratus L.) was observed perching near a 
cluster, but was not actually seen feeding on the monarchs. Two other 
patterns of apparent bird predation occurred extensively along the log- 
ging cut bordering the eastern side of Site Alpha. Numerous individual 
wings littered the forest floor along with many maimed but still living 
butterflies, including ones with one or more wings missing and others 
lacking their abdomens. Comparable predation has been noted in Cali- 
fornia (Kammer, 1970). 


Mortality Caused by Man 


Overall, natural mortality seemed to be low at Site Alpha. However, 
upslope from the colony, dead butterflies littered the ground to an ex- 
tent that their odor of decomposition was strongly evident. Based on 
Calvert’s observation of the upper boundary of the site on 31 December, 
we believe that a substantial percentage of the colony may have been 
eaten and trampled by cattle led to Site Alpha by local ranchers. 


Likelihood of Many Overwintering Sites in Mexico 


Urquhart (1976) and Urquhart & Urquhart (1976) maintained that 
overwintering of the monarch butterfly in Mexico is geographically re- 
stricted, involving as few as four sites in one general area. We believe 
that, as is the case in California (Williams et al., 1942) numerous over- 
wintering sites will be found in Mexico at locations having ecological 
characteristics similar to those at Site Alpha. The principal reason for 
our contention is that roosting in a very limited area would make the 
butterflies highly vulnerable to fire. Not only does overwintering occur 
during the dry season, but this area of Mexico is in the Trans-Mexico 
Volcanic Belt (Anon., 1961). Past volcanic activity must have set many 
large fires which would have decimated whole colonies. 

The pressing need to mount a conservation effort to preserve Site 
Alpha has been described elsewhere (Brower, 1977). 


VoLUME 31, NuMBER 4 241 


SUMMARY 


More than 14.25 million monarch butterflies in prime condition over- 
winter in a 1.5 ha site located in a coniferous forest in the trans-volcanic 
belt of Mexico. The site is characterized by thermal stability, low wind 
velocity, and high humidity. The reproductive status of the butterflies 
and their thermoregulatory activities in the colony support conclusions 
from experimental physiology. Genetic implications of mass mating 
behavior prior to dispersal and northward migration appear great. 


ACKNOWLEDGMENTS 


We are grateful to the following people for help during the initial stages 
of our search for the location of Site Alpha: Natalie Drake, Elizabeth 
Mahan, Dr. Victoria Foe, Dr. Michael Dennis, Dr. Richard Lindley, Dr. 
Paul Tuskes, and Dr. Steven P. Lynch. We also thank Dr. Rudolf M. 
Schuster and H. E. Ahles for identifying the plants and Christine M. 
Moffitt and Helen S. Smith for critically reading the manuscript. The 
research was supported by a grant (DEB 7514265) from the National 
Science Foundation with L. P. Brower as Principal Investigator. 


LITERATURE CITED 


Anonymous. 1961. Tectonic map of Mexico. Geological Society of America. 

Barker, J. F. anp W. S. Herman. 1973. On the neuroendocrine control of 
ovarian development in the monarch butterfly. J. Exptl. Zool. 183: 1-9. 

1976. Effect of photoperiod and temperature on reproduction of the 
monarch butterfly, Danaus plexippus. J. Insect Physiol. 22: 1565-1568. 

BEatt, G. 1948. The fat content of a butterfly, Danaus plexippus Linn., as af- 
fected by migration. Ecology 29: 80-94. 

Beck, S. D. 1968. Insect photoperiodism. Academic Press, N.Y. and London. 
viii + 288 p. 

Brower, L. P. 1961. Studies on the migration of the monarch butterfly. 1. Breed- 
ing populations of Danaus plexippus and D. gilippus berenice in south central 
Florida. Ecology 42: 76-83. 

1962. Biology of the monarch butterfly. Ecology 43: 181-182. 

1977. Monarch migration. Natural History 86(6): 40-53. 

Brower, L. P. & J. C. Huserru. 1977. Strategy for survival: behavioral ecology 
of the monarch butterfly. 16 mm, sound, color, 30 minute film. Harper and 
Row, Inc., 10 E. 53rd St., N.Y. 10022. 

Brown, J. J. & G. M. CurprpENDALE. 1974. Migration of the monarch butterfly, 
Danaus plexippus: energy sources. J. Insect Physiol. 20: 1117-1130. 

Burns, J. M. 1968. Mating frequency in natural populations of skippers and 
butterflies as determined by spermatophore counts. Proc. Nat. Acad. Sci. 
USA 61: 852-859. 

CENEDELLA, R. J. 1971. The lipids of the female monarch butterfly, Danaus 
plexippus, during fall migration. Insect Biochem. 1: 244-247. 

Fincn, V. C. & G. T. TREwarTHa. 1949. Elements of geography, 3rd ed. McGraw- 
Enll Book Co., Inc’ N.Y. xii + 711 p. 


242 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Herman, W. S. 1973. The endocrine basis of reproduction inactivity in monarch 
butterflies overwintering in central California. J. Insect Physiol. 19: 1883-1887. 

1975. Endocrine regulation of post-eclosion enlargement of male and 
female reproductive glands in monarch butterflies. Gen. Compar. Endocrin. 
26: 534-540. 

Hitt, F. H., Jn, A. M. WENNER, & P. H. Wetus. 1976. Reproductive behavior 
in an overwintering aggregation of monarch butterflies. Amer. Midl. Nat. 95: 
10-19. 

Kammer, A. E. 1970. Thoracic temperature, shivering, and flight in the monarch 
butterfly, Danaus plexippus (L.). Z. Vergl. Physiol. 68: 334-344. 

1971. Influence of acclimation temperature on the shivering behavior of 
the butterfly Danaus plexippus (L.). Z. Vergl. Physiol. 72: 364-369. 

Kammer, A. E. & J. Braccr. 1973. Role of the wings in the absorption of radiant 
energy by a butterfly. Comp. Biochem. Physiol. 45A: 1057-1064. 

Pan, M. L. & G. R. Wyarr. 1976. Control of vitellogenin synthesis in the monarch 
butterfly by juvenile hormone. Developmental Biol. 54: 127-134. 

Pererson, R. T. & E. L. Coauir. 1973. A field guide to Mexican birds. Houghton 
and Mifflin Co., Boston. xxii + 298 p. 
SancHEZ, O. S. 1976. La flora del Valle de Mexico. 3rd ed. Printed by the 

author, Mexico, D.F. viii + 520 p. 

Scumipr-Korenic, K. 1975. Migration and homing in animals. Springer-Verlag, 
Berlin. xii + 99 p. 

Tusxes, P. M. & L. P. Brower. 1978. Overwintering ecology of the monarch 
butterfly, Danaus plexippus, in California. Ecol. Ent. 3: (in press). 

Urounart, F. A. 1960. The monarch butterfly. Univ. of Toronto Press, 


xxiv + 361 p. 
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160-173. 


Urquuaert, F. A. & N. R. UrQquHaArt. 1976. The overwintering site of the eastern 
population of the monarch butterfly (Danaus p. plexippus; Danaidae) in 
southern Mexico. J. Lepid. Soc. 30: 153-158. 

UrquHart, F. A., N. R. Urngunart, & F. Muncer. 1970. A study of a con- 
tinuously breeding population of Danaus plexippus in southern California com- 
pared to a migratory population and its significance in the study of insect 
movement. J. Res. Lep. 7: 169-181. 

WassEeRTHAL, L. T. 1975. The role of butterfly wings in regulation of body 
temperature. J. Insect Physiol. 21: 1921-1930. 

WiuuiaMs, C. B. 1930. The migration of butterflies. Oliver and Boyd, Edinburgh. 
xii + 473 p. 

WituiaMs, C. B., G. F. Cocxpiii, M. E. Gress, & J. A. Downes. 1942. Studies in 
the migration of Lepidoptera. Trans. Roy. Entomol. Soc. Lond. 92 (Part 1): 
101-283. 


VoLUME 31, NuMBER 4 243 


REVISION OF NORTH AMERICAN BOLORIA SELENE 
(NYMPHALIDAE) WITH DESCRIPTION 
OF A NEW SUBSPECIES 


STEVE KOHLER 


Montana Department of Natural Resources and Conservation, Division of 
Forestry, 2705 Spurgin Road, Missoula, Montana 59801 


This study began as an effort to determine the validity of the sub- 
species name nebraskensis (Holland) as applied to Boloria selene (Denis 
& Schiffermiiller ) populations in a limited area of southeastern Nebraska. 
As the study progressed, it became evident that the eastern limit of the 
range of nebraskensis was not known and that the relationship of ne- 
braskensis to myrina (Cramer) was not well understood in the pub- 
lished literature. It also was apparent that considerable variation in 
both size and markings was present within the described subspecies of 
selene and that a considerable amount of intergradation occurred in 
some geographic areas between the populations traditionally considered 
as subspecies of selene. For these reasons, the study was expanded to 
include the entire range of B. selene in North America (Fig. 1). 

The North American subspecies of selene were formerly placed under 
the species myrina in the genus Brenthis Hiibner (McDunnough, 1938). 
Clark (1941) recognized that myrina and the European selene were 
conspecific and placed the North American subspecies under selene. 
Studies of male genitalia by dos Passos & Grey (1945) reinforced Clark's 
placement of myrina under selene. These studies showed that all the 
North American representatives of the genus Brenthis, including selene, 
should be placed in the genus Boloria Moore. 

The populations considered subspecies of selene actually represent 
ageregations of populations homogeneous in wing characters but on a 
larger scale representing the distinct clustering of characters amid broad 
clines. Subspecies names are valuable in characterizing these groups 
as to phenotypic appearance, habitats, and ecological-range affinities. 
Subspecies names are used in this paper for these reasons. 

A total of 1,264 specimens, representing all of the named subspecies, 
was obtained from museum and private collections for examination. 
This paper presents the results of the study of these specimens, inte- 
grated with previous knowledge of the species. Seven subspecies are 
recognized. 

Methods. Characters used to differentiate the subspecies of selene are 
given in Tables 1 and 2. The degrees of dorsal maculation, dark dorsal 


244 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Comparison of characters of the North American subspecies of Boloria 
selene. 


nebras- sabulo- tolland- atroco- terrae- 
myrina kensis collis ensis albequina _ stalis nowvae 
Number of typical 
specimens examined 
Males 130 232 Dit 88 15 159 14 
Females 63 166 8 53 6 80 il 
FW length, mm 
Males 20.98 24.78 22,.82 19.55 LES PET) 20.08 20.67 
Females Dl leretsy PY MG, Pe, 20.46 20.08 20.57 2.1.50 
Degree of maculation 
(dorsal )* 2.29 1.99 1.82 1.50 Dae ofo) GAD) Diao 


Degree of dark mar- 
ginal wing scaling 
(dorsal )’ Ley? 1.14 1.80 1-22 3.00 3.19 1.93 


Degree of dark basal 
wing scaling 
(dorsal )* KO 87 1.20 2.14 4.33 2.10 DAF 


Percentage of speci- 
mens with basal spot 


in discal cell of dor- 
sal HW obscured 30 6 15 79 100 60 93 


Percentage of speci- 
mens with anvil- 
shaped silver spot in 
cell Cuz of ventral 
HW divided Te, 4 4 30 33 6 7 


Yellow scaling invad- 
ing cinnamon color 
of basal and discal 
areas of ventral HW” ++ Shale “pase ahi sind et ate “F “h 


? Values represent an average obtained by visually rating each specimen on a scale of 0-5. 
2 Average appearance of each subspecies based on a composite of all specimens examined. 


marginal wing scaling, and dark basal wing scaling dorsally, given for 
each subspecies, represent averages of numerical values obtained by 
visually rating each specimen on a scale of zero to five. The values for 
the amount of yellow scaling that invades the cinnamon color of basal 
and discal areas of the ventral HW (hindwing) represent the average 
appearance of each subspecies based on a composite of all specimens 
examined. These are also on a scale of zero to five. Division of the 
anvil-shaped silver spot in cell Cu, of the ventral HW is not intended 
to be a main character for separation of subspecies. It is used to point 
out the close relationship of albequina (Holland) to tollandensis (Barnes 
& Benjamin). 


VOLUME 31, NuMBER 4 24 


Ot 


TaBLE 2. Comparison of width to height ratios! of median black lunules in cells 
Rs, Mi, Me, Ms, and CU; of the ventral FW of some North American Boloria selene 
subspecies. 


myrina nebraskensis sabulocollis tollandensis atrocostalis 

Number of 

specimens 

examined 116 OF 59 44 66 
Cell Rs 1.34 1.37 1.38 1.20 oe 
Cell Mi P27 1.38 1.25 L222, 1.17 
Cell M2 1.58 LT) 1.49 Loi 1.51 
Cell M; 1.12 AD oe) .90 1.00 
Cell Cu: 1.21 1.28 1.04 .95 1.13 


1 Values obtained by dividing width of the lunule by height. Values greater than 1 indicate 
oblate lunules. 


One of the more consistent characters used in the study was the shape 
of the median black lunules of the ventral FW (forewing) in cells M,, 
M;, and Cu;, but especially in M; and Cu,. The values given in Table 2 
are averages obtained by taking measurements of each spot on every 
specimen using a binocular dissecting microscope with a micrometer 
disc grid in the eye-piece. These measurements of width and height of 
each individual spot were then adjusted to a ratio number by dividing 
width by height. Lunules that were oblate had numbers greater than 
one. Numbers near one or less indicated the lunules were not flattened 
and elongated. 

Terminology used for describing veins, cells, and wing areas follows 
that given for butterflies in general and specifically for Boloria by Ehr- 
lich & Ehrlich (1961). 

No consistent or reliable characters which could serve to differentiate 
subspecies were found in the genitalia. 


Boloria selene myrina (Cramer ) 
(Figs, 2.6) 


Papilio myrina Cramer, 1777, vol. 2: 141, pl. 189, figs. B., C. 

Papilio myrinus: Herbst, 1798. 

Argynnis myrissa Godart, 1819, p. 253. 

Argynnis myrina: Scudder, 1863, p. 166; 1868, p. 379. Edwards, 1876, p. 161. 

Brenthis myrina nubes Scudder, 1889, p. 595. 

Brenthis myrina ab. nubes: McDunnough, 1938, p. 16. 

Boloria selene myrina ab. nubes: dos Passos, 1964, p. 89. 

Brenthis myrina: Dyar, 1900, p. 487. Cary, 1901, p. 307; 1906, p. 439. Elrod, 1906, 
p. 69. Franzen, 1914, p. 366. McDunnough, 1928, p. 274. Holland, 1931, p. 105. 
Clark, 1932, p. 102. dos Passos & Grey, 1934, p. 190. McDunnough, 1938, p. 15. 
Davenport & Dethier, 1938, p. 158. Leussler, 1938, p. 216. Rysgaard, 1939, p. 
193. Comstock, 1940, p. 53. Brooks, 1942, p. 34. 


246 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Brenthis selene myrina:; Clark, 1941, p. 384. 

Brenthis selene marilandica Clark, 1941, p. 384. 

Boloria myrina: dos Passos & Grey, 1945, p. 4. 

Boloria selene marilandica: Klots, 1951, p. 89. dos Passos, 1964, p. 89. 

Boloria selene myrina: Klots, 1951, p. 89. dos Passos, 1964, p. 89. Shepard (in 
Howe ), 1975, p. 246. 


Diagnosis. This subspecies is most closely allied to nebraskensis in coloration and 
pattern of markings. Series of the two are readily separated by size, since FW length 
of nebraskensis averages ca. 4 mm greater than myrina in both males and females. 

The pattern of dorsal dark markings on the orange ground color of myrina is 
heavier than on nebraskensis and is comparable to atrocostalis (Huard). In the 
basal portion of the wings dorsally, myrina also has more dark scaling than nebras- 
kensis, and the dark basal spot in the discal cell is frequently obscured with dark 
scales (ca. 30% of specimens). This spot is obscured in only 6% of nebraskensis 
specimens (Table 1). The black marginal band dorsally is a little heavier on myrina 
than nebraskensis. 

In both subspecies, the black median lunules in cells Ms, Ms, and Cu; of the 
ventral FW tend to be elongated or oblate (Table 2). This tendency is less pro- 
nounced in myrina than in nebraskensis. 

Male. Dorsal surface: ground color bright orange, with pattern of dark markings 
sharply demarcated; black wing margins of FW and HW not heavy, and enclosed 
row of orange marginal spots distinct and visable along entire margin; basal areas 
of both wings moderately dusted with dark scales, often obscuring basal spot in 
discal cell of HW. 

Ventral surface: apical patch of FW solid cinnamon-brown, not infused with 
yellow scales; orange of FW ground color usually extending distally to cinnamon- 
brown apical patch without fading to yellow; black median lunules in cells M,, Ma, 
and Cu, of FW oblate, elongated on long axis of wing; cinnamon-brown ground 
color of HW mostly solid, especially in basal area, with little dusting of yellow 
scales; postmedian row of light spots of HW all prominent and well silvered; anvil- 
shaped silver submedian spot in cell Cu, of HW seldom divided. 

Length of forewing: 20.98 mm average. 

Female. Similar to male but orange ground color of dorsal surface slightly more 
pale. 

Length of forewing: 21.85 mm average. 

Type locality. The type locality given by Cramer in his original description of 
myrina was New York. Cramer’s specimen most likely came from southeastern New 
York, rather than northern New York, which was probably quite inaccessible at that 
time. 

Holland (1928) felt that specimens in his possession from Hunter and Saratoga, 
New York, Pennsylvania, and other localities in New England agreed with the some- 
what crude figure given by Cramer and accepted the Hunter and Saratoga speci- 
mens as being topotypical. Holland pointed out that the discal and basal markings 
of the wings on the upper side of these specimens were not as heavy as in Cramer's 
original drawing and that they more nearly represented the “general run” of speci- 
mens in New England and the Middle States. He also mentioned that he had 
specimens from Hunter, New York and the Allegheny Mountains near Cresson, 
Pennsylvania that were as heavily marked as the specimen shown on Cramer's 
plate. Specimens examined for this study from some localities, including Schuylkill 
Haven, Pennsylvania (Fig. 2) were similarly marked. 

Distribution. Southern portions of New York, Vermont, and New Hampshire 
south to central Virginia and West Virginia, including Massachusetts, Rhode Island, 
Connecticut, New Jersey, Pennsylvania, Maryland, and Delaware, west through 
southern Michigan, extreme northeast Illinois, the southern portions of Wisconsin 
and Minnesota to eastern South Dakota (Fig. 1). 


VOLUME 31, NuMBER 4 247 


myrina 


nebraskensi 


sabulocollis NX YU eeee00 
tollandensis WYYyyy - —— 


_albequina 
atrocostalis 
terraenovae 


intergradation 


2) 
@ 
©) 
® 
© 
© 
@ 


Fig. 1. Distribution of the subspecies of Boloria selene in North America. 


Remarks. The subspecies myrina has been referred to by some authors ( Klots, 
1951; Shepard in Howe, 1975) as being average for the species in both size and 
markings. Because of this, many specimens representing intergrades between atro- 
costalis and myrina and between myrina and nebraskensis, as well as specimens of 
nebraskensis over much of its range, have been lumped together under myrina. 

Specimens from central Ohio, northern Indiana, northern Illinois (except the 
extreme northeast corner), and Iowa previously considered to be myrina have been 
found to be nebraskensis. There is intergradation with atrocostalis along the entire 
northern limit of the range of myrina, and some evidence of intergrading with 
nebraskensis to the south in Illinois and northeastern Nebraska. 

Two aberrations of myrina have been given names. Ab. nubes (Scudder) was de- 
scribed from a male collected in Sanbornton, New Hampshire, in which the median 
spots were enlarged and somewhat suffused. The type is now in the Museum of 
Comparative Zoology at Harvard. The name nivea (Gunder) was given to an albino 
male collected May 28, 1900, on Staten Island, New York. The ground color, both 
dorsal and ventral, is white, with no maculation change. The type is in the Ameri- 
can Museum of Natural History. 

The type locality of marilandica (Clark) is Beltsville, Maryland, “From the boggy 
pasture on the south side of the road from the Beltsville, Md. railway station to the 
Dept. of Agriculture experiment farm.” It is illustrated in Clark (1932, plate 3, 
figs. 5 & 6). The distinctive features listed in the original description were larger 
size, richer color, and heavier dark markings. There are no real consistent or sig- 
nificant differences between this local form (probably now extinct) and myrina, 
and it is here considered a synonym of myrina. The type is in the U.S. National 
Museum. 

Records. The following records represent 193 typical specimens (130 8, 63 9° ) 
of myrina and additional intergrades examined: 


248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


myrina 


Connecticut: Fairfield, Hartford, and Windham cos. DELAWARE: Water Gap. 
MARYLAND: Baltimore and Garrett cos. MassacuusETts: Hampshire and Norfolk 
cos. MicuicaAN: Indian Lake; Empire; Ramona; Cass, Delta, Jackson, Oakland, 
Ottawa, Van Buren, and Washtenaw cos. NrEw Jersey: Jamestown; Bergen, 
Hunterdon, Middlesex, Monmouth, Passaic, and Union cos. New York: Feurabush; 
E. Berne; Flushing; Ravenwood; McLean; New York City; Big Island; Somers; Fulton 
Co. PENNSYLVANIA: Lackawanna, Perry, and Schuylkill cos. Ruopr Istanp: Oak 
Lawn; Providence Co. Sour Daxora: Brown and Day cos. Wisconsin: Dous- 
man; Milwaukee; Brown, Milwaukee, Ozaukee, Walworth, Washington, and 
Waukesha cos. 


nr. myrina 


Inntinois: McHenry Co. Massacnuserrs: Mt. Greylock; Readville; Berkshire Co. 
MicuicaNn: Presque Isle Co. VERMONT: Bennington Co. Wisconsin: Waupaca Co. 


Boloria selene nebraskensis (Holland) 
(Figs. 2, 6) 


Brenthis myrina var. nebraskensis Holland, 1928, p. 36; 1931, p. 105, pl. LV, fig. 8 
(male type). 

Brenthis myrina race nebraskensis: Leussler, 1938, p. 216. 

Brenthis myrina nebraskensis: McDunnough, 1938, p. 16. 

Brenthis selene nebraskensis: Clark, 1941, p. 384. 

Boloria selene nebraskensis: Klots, 1951, p. 89. Brown, 1957, p. 64. dos Passos, 1964, 
p. 89. Shepard (in Howe), 1975, p. 246. 

Boloria selene nebrascensis: Forbes, 1960, p. 160. 


Diagnosis. Series of nebraskensis can be separated quite easily from the other 
subspecies of selene on the basis of large size alone, although other distinct features 
are present (Tables 1 & 2). 

The dorsal wing margins of nebraskensis have the least amount of black of any 
of the subspecies, and the basal areas of both wings dorsally also have the least 
suffusion of dark scales. The basal spot in the discal cell of the dorsal HW is 
seldom completely obscured by dark scales (only 6% of the 398 specimens exam- 
ined). The black median lunules in cells M2, Ms, and Cu: of the ventral FW are 
oblate and elongated as in myrina, but in nebraskensis this tendency is more 
pronounced. 

Male. Dorsal surface: ground color bright orange; normal pattern of dark mark- 
ings very sharply demarcated; black margins of both wings reduced, and enclosed 
row of orange marginal spots very distinct, large, and visible along entire margin; 
very little suffusion of black scales at wing bases; basal spot in discal cell of HW 
almost never obscured by dark scales; postbasal spot in discal cell of HW relatively 
small, leaving space equal to at least one-half diameter of spot between spot and 
veins Rs and M; above and below cell. 

Ventral surface: ground color of FW very bright orange and usually extending 
completely to apical patch; apical patch bright cinnamon-brown, usually with little 
infusion of yellow scales; black median lunules in cells M2, Ms, and Cu, of FW 
strongly oblate; slight dusting of yellow scales on cinnamon-brown ground color of 
HW in basal and limbal areas (postmedian band); postmedian row of light spots 
of HW all prominent and well silvered; anvil-shaped silver submedian spot in cell 
Cu. of HW almost never divided. 

Length of forewing: 24.78 mm average. 


VOLUME 31, NuMBER 4 249 


Fig. 2. Adults of B. selene: (a) myrina male, Schuylkill Haven, Schuylkill Co., 
Pa.; (b) same, ventral; (c) myrina male, New York City, N.Y.; (d) same, ventral; 
(e) nebraskensis male, Valley, Douglas Co., Neb.; (£) same, ventral; (g) nebrasken- 
sis male, Goose Lake Prairie State Park, Grundy Co., Ill.; (h) same, ventral. All 
photos natural size. 


250 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Female. Similar to male except orange of dorsal ground color slightly more pale. 

Length of forewing: 25.73 mm average. 

Type locality. The type series consisted of four male specimens collected by 
Mr. E. A. Dodge. These specimens are presently at the Carnegie Museum in Pitts- 
burgh. The type locality is Dodge Co., Nebraska. 

Distribution. The range of nebraskensis is much more extensive than previous 
authors have reported. Typical specimens occur from east central Nebraska east 
through northern Iowa, northern Illinois, and northern Indiana to central Ohio 
(Fig. 1). It intergrades with myrina in some areas of the northern part of its 
range. 

Remarks. B. s. nebraskensis was described by Holland in 1928 as a new variety 
of B. myrina. The sole basis for the name was the exceptional size of four male 
specimens collected by Mr. E. A. Dodge in Dodge Co., Nebraska. No description 
was given, except to call attention to size. Holland (1931) later discussed briefly 
nebraskensis in the revised edition of The Butterfly Book and illustrated a male 
type on Plate LV. Again, the only distinguishing character mentioned in the text 
was the large size. 

My interest in nebraskensis was stimulated by discussion with Mr. Orville D. 
Spencer of Lincoln, Nebraska in 1968. Of particular interest was the fact that 
although good series of large typical specimens from Nebraska were present in 
some museums, none had been collected in recent years (Johnson, 1972). 

I began actively seeking colonies of nebraskensis in Nebraska in the summer of 
1968. That year colonies were located at Dead Timber State Recreation Area, 
Dodge Co., and Valley, Douglas Co. In 1969, several females taken from the Valley 
colony in July were kept alive and ova obtained. From these ova, 231 adult speci- 
mens (115 ¢, 116 2) were reared. Unfortunately, data on the immature stages 
were not kept, nor were photographs taken of the stages, nor,were the stages 
studied in detail. The primary purpose of rearing was to obtain adult specimens. 

In Nebraska, nebraskensis appears to have three broods per year; the first at the 
beginning of June from overwintering larvae, the second in mid-July, and the third 
in mid-August. Habitat where recent nebraskensis specimens were collected at 
Valley was wet meadow associated with river bottom and willow. Illinois habitats 
are similar (Conway, in litt.). 

Records. Typical specimens of nebraskensis examined, including the reared Valley 
series, numbered 232 ¢ and 166 2. These records and additional intergrades are 
as follows: 


nebraskensis 


ILtinois: Chicago; Cook, Grundy, LaSalle, Mercer, and Stephenson cos. INDIANA: 
La Porte Co. Iowa: Winneshiek Co. NepraAska: Douglas Co. Onno: Franklin, 
Medina, and Richland cos. Sour Daxora: Brookings Co. 


myrina/nebraskensis 
NEBRASKA: Dodge and Madison cos. 


In the western panhandle of Nebraska, extreme western South Dakota, south- 
western North Dakota, and the northeastern prairie region of Colorado, specimens 
of selene exhibit characters which are different from nebraskensis. This area is iso- 
lated geographically and includes the major remaining conifer forest and montane 
areas of the Great Plains. Because of this geographic isolation and the differences 
observed in specimens from this area, the following new subspecies of B. selene is 
described. 


VOLUME 31, NumBeErR 4 


bo 
Ol 
— 


g 


Fig. 3. Adults of B. selene: (a) sabulocollis male, holotype, Smith Lake, Sheri- 
dan Co., Neb.; (b) same, ventral; (c) sabulocoilis male, paratype, Timnath, Larimer 
Co., Colo.; (d) same, ventral; (e) sabulocollis male, paratype, Mineral Springs, 
Slope Co., N.Dak.; (f) same, ventral; (g) tollandensis male, Tolland, Gilpin Co., 
Colo.; (h) same, ventral. All photos natural size. 


Dbz, JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Boloria selene sabulocollis Kohler new subspecies 
(Figs.33, 6.7) 


Diagnosis. This subspecies is intermediate in size between myrina and nebras- 
kensis. The main characters separating it from nebraskensis and myrina, which it is 
quite similar to dorsally, are on the ventral wing surfaces. The median black lunules 
in cells Ms and Cu; of the ventral FW are “square-shaped,” not oblate and elongated 
as in myrina and nebraskensis. There is more dusting of yellow scales on the 
cinnamon-brown ground color of the ventral HW of sabulocollis. 

Male. Dorsal surface: normal pattern of dark markings sharply demarcated on 
orange ground color; row of orange marginal spots enclosed in black marginal band 
distinct and visible along entire margin of both wings, some suffusion of dark scales 
at wing bases, but basal spot in discal cell of HW seldom obscured; postbasal spot 
in discal cell of HW relatively small, leaving space equal to at least one-half diam- 
eter of spot between edge of spot and veins Rs above and Mz; below discal cell. 

Ventral surface: ground color of FW dull orange, fading to yellow-orange before 
reaching apical patch; cinnamon-brown of apical patch dusted with yellow scales; 
black median lunules in cells Ms; and Cui of FW “square-shaped,” not oblate and 
elongated; considerable dusting and infusion of yellow scales in cinnamon-brown 
ground color of HW, with row of black spots in postmedian cinnamon-brown band 
surrounded by halos of yellow scales, and infusion of yellow scales in basal area of 
all cells; anvil-shaped silver submedian spot in cell Cus of HW almost never divided. 

Length of forewing: 22.82 mm average; holotype 22 mm. 

Female. Similar to male, but with orange of dorsal ground color slightly more 
pale. 

Length of forewing: 23.75 mm average; allotype 25 mm. 

Types. Holotype: male, Smith Lake, Sheridan Co., Nebraska, 1.viii.68, S. Kohler; 
placed in the collection of the American Museum of Natural History. 

Allotype: female, same data as holotype; placed in the collection of the American 
Museum of Natural History. 

Paratypes (57): NersraskA: Smith Lake, Sheridan Co., 1.viii.68, S. Kohler, 22 
6, 1 9; Dewey Lake, Cherry Co., 15.vii.69, S. Kohler, 1 ¢; Clear Lake, Cherry 
Co., 13.vi.29, 2 &; North Platte, Lincoln Co., 7.vii.29, 4 ¢, 292. Sours Daxora: 
S. of Lead, Hwy. 229, 5800’, Lawrence Co., 10.vii.66, J. S. Nordin, 1 ¢; Ditch Cr., 
S. of Deerfield Res., Pennington Co., 19.vii.70, J. S. Nordin, 2 ¢; Custer State Park, 
5000’, Custer Co., 31.vii.64, J. S. Nordin, 2 ¢. Norra Daxora: Mineral Springs, 
Slope Co., 20.vi.61, 23.vi.60, 10.vii.60, J. Oberfoell, 12 4, 1 9. Cotorapo: Timnath, 
Larimer Co., 18.vii.72, R. E. Stanford, 2 ¢, 1 2; nr. Timnath, 4865’, Larimer Co., 
7.vi.73, R. E. Stanford, 2 %; nr. Timnath, Larimer Co., ex. ova 18.vii.72, em. 3- 
S.ixs 2) we Stantordse dag saan Oy. 

Paratypes will be placed in the collections of the American Museum of Natural 
History, the Los Angeles County Museum, the National Museum of Natural History, 
the Allyn Museum of Entomology, J. S. Nordin, J. Oberfoell, J. R. Heitzman, R. E. 
Stanford, M. S. Fisher, and S. Kohler. 

Distribution. The range of sabulocollis is limited to western Nebraska, the 
prairie region of northeastern Colorado, western South Dakota, and extreme south- 
western North Dakota (Fig. 1). Specimens from Albany Co., Wyoming show 
intergradation between sabulocollis and tollandensis. 

Remarks. A knowledge of the paleobotany of the western Great Plains is im- 
portant in understanding how sabulocollis may have become isolated. Johnson (1975, 
1976) points out that the present coniferous forests of the western Great Plains 
(Black Hills, Pine Ridge, Cheyenne Ridge, etc.) are the remains of extensive forest 
areas that once covered much of the region. The subspecies sabulocollis probably 
represents a relict of more extensive populations that were associated with the 
former coniferous forest. 


VoLuME 31, NuMBER 4 253 


Colonies in western Nebraska are situated at the marshy edges of lakes and 
adjacent wet meadows in the sandhills region, an area characterized by thousands 
of square miles of large, stabilized sand dunes covered with range grasses. The 
western part of this region, once covered by open pine-juniper forest, has abundant 
ground water, with many small lakes. Further east, the region becomes drier, The 
sandhills probably are an effective barrier to intergradation with nebraskensis to 
the east, although this possibility exists along the Platte and Elkhom rivers. The 
colonies in southwestern North Dakota and western South Dakota are isolated from 
the east by extensive arid areas lacking suitable habitat. 

The name sabulocollis is a combination of the Latin words fer sand (sabulo) 
and hill (collis), which is descriptive of the area of the type locality. 


Boloria selene tollandensis (Barnes & Benjamin) 
(igiss ou 4h) 


Brenthis myrina tollandensis Barnes & Benjamin, 1925, p. 44. Holland, 1928, p. 37. 
Klots, 1937, p. 328. McDunnough, 1938, p. 16. 

Brenthis myrina var. tollandensis: Holland, 1931, p. 106. 

Brenthis selene tollandensis: Clark, 1941, p. 384. 

Boloria selene tollandensis: Remington, 1952, p. 67. Brown, 1957, p. 64. Albright, 
1960, p. 158. Newcomer & Rogers, 1963, p. 171. dos Passos, 1964, p. 89. Shepard 
(in Howe), 1975, p. 246. 

Boloria myrina (=selene) tollandensis: Brown, 1954, p. 64. 


Diagnosis. This is the most lightly marked of the subspecies, with the pattern 
of dark maculation reduced dorsally, especially on the HW. Suffusion of dark scales 
in the basal areas of the wings dorsally is comparable to atrocostalis, and the basal 
spot in the discal cell of the dorsal HW is usually obscured by dark scaling. The 
postbasal spot in the discal cell of the dorsal HW is characteristically very large in 
relation to the cell, often filling almost the entire width of the cell. The size of 
this spot is accentuated by the otherwise fine maculation of the HW. Ventrally, 
tollandensis exhibits extensive infusion of yellow into the cinnamon-brown basal and 
discal areas of the HW. 

Male. Dorsal surface: Ground color orange to yellow-orange; black marginal band 
reduced, and enclosed orange marginal spots very distinct and visible along entire 
margin of both wings; normal pattern of black markings reduced and narrow on 
both FW and HW, with exception of postbasal spot in discal cell of HW, which is 
enlarged and often fills almost entire width of cell, accentuated by otherwise fine 
markings of HW; considerable suffusion of dark scales at wing bases, especially on 
HW where dark scales form rather abrupt line across middle of discal cell, usually 
obscuring basal spot in cell. 

Ventral surface: ground color of FW _ yellow-orange; apical cinnamon-brown 
patch of FW much reduced and powdery, with brown areas largely replaced by 
clear yellow-buff; black median lunules in cells M; and Cu; of FW “square shaped,” 
not oblate; extensive yellow scaling in HW, with cinnamon-brown of postmedian 
band between postmedian and submarginal rows of silver spots reduced to powdery 
patches between veins Rs and Ms and between veins Ms and Cu, and remaining 
brown of basal area of HW powdered with yellow scales; anvil-shaped silver sub- 
median spot in cell Cur of HW frequently divided (30% of specimens). 

Length of forewing: 19.55 mm average. 

Female. Similar to male but with orange of dorsal ground color noticeably more 
pale, especially on outer half of both FW and HW. 

Length of forewing: 20.46 mm average. 

Type locality. The type locality of tollandensis is Tolland, Gilpin Co. (not 
Moffat), Colorado. The original description (Barnes & Benjamin, 1925) lists Tol- 


254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


g 


Fig. 4. Adults of B. selene: (a) tollandensis male, Avon, Cache Co., Ut.; (b) 
same, ventral; (c) tollandensis/albequina male, Moxee Bog, Yakima Co., Wash.; (d) 
same, ventral; (e) tollandensis/albequina male, Atlin, B.C.; (£) same, ventral; (g) 
albequina male, Homer, Alaska; (h) same, ventral. All photos natural size. 


land as being in Moffat Co. This must be an error. The type series consisted of 


the holotype ( 4 ), allotype (2), and 12 paratypes (5 6,7 @). All of these were 
in the Barnes collection, now at the U.S. National Museum. 


Distribution. The range of typical tollandensis extends from north-central New 


VoLUME 31, NuMBER 4 


bo 
Ul 
Ol 


Mexico north through central Colorado and Wyoming, west through northeastern 
Utah, southern Idaho, and extreme southwestern Montana, and north through most 
of the Idaho panhandle. 

Remarks. Nearly typical specimens have been examined from extreme northern 
Idaho. In eastern Oregon and Washington, and most of British Columbia, tollan- 
densis intergrades with albequina (Holland) (Figs. 4 and 8). Intergrading with 
the eastern atrocostalis occurs over a wide band including most of Alberta and 
Montana, the eastern Idaho panhandle, and the northwest corner of Wyoming. 
Intergrading with sabulocollis occurs in eastern Wyoming. 

Brown (1957) states that tollandensis in Colorado is single-breoded. Utah tol- 
landensis appears to be double-brooded, and there is evidence that iniergrades from 
Montana have two broods per year, since fresh adults have been collected from the 
same locality in both mid-June and mid-August. 

Habitat listed by both Brown (1957) and Ferris (1971) for tollandensis in Colo- 
rado and Wyoming is willow bogs at rather high altitude, 9,000 to 10,000’ for 
Colorado. The Utah specimens illustrated in Fig. 4 were taken at ca. 5500’ eleva- 
tion in an open boggy pasture near the mouth of a canyon. Montana specimens 
have been taken as low as 3300’ in wet open meadows and river bottom near 
Missoula and at higher elevations in Glacier National Park and elsewhere in the 
state. 

The aberration “serratimarginata” (Gunder) was described in 1926 from Vernon, 
British Columbia. The specimen, a male, was taken 12 August, 1904. The sub- 
marginal row of black markings of the dorsal HW are fused with the black mar- 
ginal band, giving the margin a serrated appearance. The type is in the American 
Museum of Natural History (dos Passos, 1938). 

Records. One hundred forty-one typical specimens (88 4, 53 2 ) of tollandensis 
and additional intergrades were examined. Records of these specimens are as 
follows: 


tollandensis 


Cotorapo: Rocky Mtn. Nat. Park; nr. Northgate; Boulder, Conejos, Gilpin, Grand, 
Lake, Routt, and Summit cos. Ipano: Blaine, Fremont, and Latah cos. MONTANA: 
Beaverhead Co. New Mexico: Rio Arriba and Sandoval cos. Urau: Vineyard; 
Cache, and Utah cos. Wyominc: NE of Cora; Fremont, Johnson, and Sublette cos. 


nr. tollandensis 
Ipauo: Bonner and Custer cos. OrEcon: Crook Co. Wyominc: Big Hom Mts.; 
Albany, Carbon, and Teton cos. 


albequina/tollandensis 


Brirish Cotumpia: Beaverfoot Range; Smithers; Atlin; Fernie; Lillooet; Robson; 
Cranbrook; and Laird Hot Springs. WasHincron: Yakima Co. 


atrocostalis/tollandensis 


Auserta: Didsbury; Banff; Carbon; Pine Creek; Bilby; and Slave Lake. Monrana: 
Cascade, Flathead, Gallatin, Granite, Jefferson, Lake, Missoula, Ravalli, and Sweet 
Grass cos. Wyominc: Yellowstone Nat. Park; Park and Teton cos. 


Boloria selene albequina (Holland) 
(Fig. 4) 
Brenthis albequina Holland, 1928, p. 40, figs. 4, 5, and 6; 1931, p. 107, pl. LY, figs. 
19, 20, and 21 (types). McDunnough, 1932, p. 269; 1938, p. 16 (as synonym ot 
atrocostalis ). 


256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Brenthis albequina var. baxteri Holland, 1928, p. 42. 

Brenthis myrina atrocostalis ab. baxteri: McDunnough, 1938, p. 16. 
Boloria selene albequina: dos Passos, 1964, p. 89. 

Boloria selene albequina ab. baxteri: dos Passos, 1964, p. 89. 


Diagnosis. Dorsally, albequina is similar to atrocostalis. Both have heavy dark 
wing margins, but albequina differs from atrocostalis in having a greater suffusion 
of dark scales in the basal areas of the wings (Table 1). There is also a tendency 
for the pattern of dark spots which are not obscured by the dark basal suffusion to 
appear “blurred” or “smudged,” not sharply demarcated as in atrocostalis. 

The ventral surface of the HW of albequina also differs from that of atrocostalis 
in having considerable infusion of yellow scales into the cinnamon-brown of the 
basal and discal portions, as does tollandensis. In atrocostalis, these areas are largely 
solid cinnamon-brown. 

Male. Dorsal surface: ground color of both wings dull orange; black marginal 
band heavy, and enclosed orange marginal spots obliterated on FW from apex 
posterior at least to cell M:, and remaining spots on FW and HW reduced to small 
orange points; normal pattern of black markings heavy and usually appearing slightly 
“blurred” or “smudged,” not sharply demarcated; extensive suffusion of black scales 
into basal portions of wings, always obscuring basal spot in discal cell of HW and 
often postbasal spot, with suffusion forming more or less solid black patch on HW. 

Ventral surface: ground color of FW dull yellow-orange; apical cinnamon-brown 
patch of FW much reduced and powdered with yellow scales, surrounded by dull 
yellow-buff; black median lunules in cells M; and Cui of FW “square-shaped,” not 
oblate; considerable yellow scaling on HW, with cinnamon-brown of postmedian 
band between postmedian and submarginal rows of silver spots reduced and pow- 
dery; silver spot of postmedian band in cell M2 of HW lacking, and spot in cell Ms; 
often indistinct; anvil-shaped silver submedian spot in cell Cus of HW frequently 
divided (39% of specimens). 

Length of forewing: 19.17 mm average. 

Female. Similar to male. 

Length of forewing: 290.08 mm average. 

Type locality. This subspecies was described from seven males collected at White 
Horse Pass, Yukon Territory, by W. F. O. Baxter. The types are in the Carnegie 
Museum. 

Distribution. The range of typical albequina is limited to southwest Alaska and 
adjacent portions of the Yukon Territory east into the far west portion of the North- 
west Territories (Fig. 1). 

Remarks. In northern British Columbia, albequina intergrades with tollandensis. 
Many British Columbia specimens have been incorrectly referred to as albequina. 
Specimens from Alberta, Montana, and northwestern Wyoming, many of which 
were also previously referred to as albequina, represent a wide band of intergrada- 
tion between tollandensis and atrocostalis. These specimens exhibit dark wing mar- 
gins dorsally, as in atrocostalis, but the pattern of markings is reduced as in 
tollandensis, especially on the HW. In addition, the postbasal spot in the discal cell 
is often enlarged, covering nearly the width of the cell in many specimens, as it 
does in tollandensis. Ventrally, the HW of these specimens exhibits characteristics 
of both tollandensis and atrocostalis (Figs. 5 and 8). 

It has been suggested by some authors, as early as 1932 (McDunnough) and 
as recently as 1975 (Shepard, in Howe), that the name albequina be reduced to 
the synonymy of atrocostalis. It is my opinion that this decision is based primarily 
on the study of specimens from zones of intergradation with atrocostalis and tol- 
landensis and that the name albequina should be used for the Alaska, Yukon, and 
Northwest Territories populations. 

In the original description of albequina, Holland (1928) placed considerable 
emphasis on the anvil-shaped silver submedian spot in cell Cuz of the ventral HW. 


VoLUME 31, NuMBER 4 957 


Fig. 5. Adults of B. selene: (a) atrocostalis male, Lake Jacques Cartier, Lauren- 
tides Park, Que.; (b) same, ventral; (c) atrocostalis/tollandensis male, Florence, 
Ravalli Co., Mont.; (d) same, ventral; (e) terraenovae male, Doyle’s Station, New- 
foundland; (f) same, ventral; (g) terraenovae male, Doyle’s Station, Newfoundland; 
(h) same, ventral. All photos natural size. 


258 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


He pointed out the tendency of this spot to be divided in albequina and the inner 
half reduced in size. McDunnough (1932) felt that many of the other differences 
mentioned by Holland were due to the rubbed condition of the specimens and that 
the stress Holland placed on the division of the silver spot was unwarranted, since 
this condition was also present in specimens of atrocostalis from the north shore of 
the Gulf of St. Lawrence. Of 239 typical specimens of atrocostalis examined in this 
study, only 6% showed division of the submedian silver spot in cell Cun of the 
ventral HW. This compares with the division of this spot being displayed in 39% 
of the specimens of albequina examined. Of 141 typical specimens of tollandensis 
examined, 30% had this spot divided. The tendency for the spot to be divided in 
both albequina and tollandensis and the extensive yellow scaling of the HW of both 
show that albequina is more closely allied to tollandensis than atrocostalis. 

The name baxteri was given by Holland (1928) to an aberrant female of the 
same series as the types of albequina from White Horse Pass, Yukon Territory. 
The dark basal and median markings of the dorsal wing surfaces are enlarged and 
connected on this aberration. 

Records. A total of 21 typical specimens (15 ¢, 6 2) of albequina were 
examined. This included a series that was examined for me by Mr. Kenelm Philip, 
Alaska Institute of Arctic Biology. The records are as follows: 


albequina 


AuasKA: Kenai Peninsula; Kodiak; Goldstream Valley, nr. Fairbanks; Chatanika 
River, NE of Fairbanks; nr. Talkeetna; and mile 154.7, new Anchorage-Fairbanks 
Hwy. 


nr. albequina 


ALBERTA: Laggan (Lake Louise). 


Boloria selene atrocostalis (Huard) 
(Figs. 5, 8) 


Argynnis atrocostalis (Provancher and) Huard, 1926 (1927), p. 131. 

Brenthis myrina atrocostalis: McDunnough, 1930, p. 107; 1938, p. 16. dos Passos & 
Grey, 1934, p. 190. 

Brenthis myrina ab. jenningsae Holland, 1928, p. 36. 

Brenthis myrina var jenningsae: Holland, 1931, p. 105, pl. LVI, fig. 9 (male type). 

Brenthis myrina atrocostalis ab. jenningsae: McDunnough, 1932, p. 269; 1938, p. 16. 

Boloria selene atrocostalis ab. jenningsae: dos Passos, 1964, p. 89. 

Brenthis selene atrocostalis: Clark, 1941, p. 384. 

Boloria selene atrocostalis: Klots, 1951, p. 89. dos Passos, 1964, p. 89. Shepard (in 
Howe), 1975, p. 246. 


Diagnosis. The subspecies atrocostalis is separated from myrina by the heavy 
dark marginal wing scaling dorsally, which often almost completely eliminates the 
normal row of orange marginal spots. It also has more suffusion of dark scales in 
the basal portions of the wings dorsally than does myrina, but not as much as 
albequina. Ventrally, the cinnamon-brown ground color of the HW of atrocostalis 
is not reduced or infused with yellow scales, as in albequina. The ventral FW 
median black lunules in cells M; and Cu: of atrocostalis are more “square-shaped,” 
instead of oblate as in myrina. 

Male. Dorsal surface: ground color bright orange; black marginal band heavy 
and broad, with enclosed orange marginal row of spots largely obscured by black, 
especially on FW, or completely obliterated by black on both FW and HW of 


VoLUME 31, NuMBER 4 259 


Fig. 6. Adults of B. selene: (a) myrina female, New York City, N.Y.; (b) 
same, ventral; (c) nebraskensis female, Valley, Douglas Co., Neb.; (d) same, ven- 
tral; (e) nebraskensis female, Goose Lake Prairie State Park, Grundy Co., Ill; (£) 
same, ventral; (g) sabulocollis female allotype, Smith Lake, Sheridan Co., Neb.: 
(h) same, ventral. All photos natural size. 


260 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


some specimens; normal pattern of black markings heavier, but sharply outlined on 
orange ground color; some suffusion of dark scales at bases of both wings, extending 
distally near inner margin of FW to median row of spots, and on HW filling ca. 
one-half of discal cell; basal spot in discal cell of HW usually obscured by dark 
scales. 

Ventral surface: ground color of FW dull orange, extending to apical patch with 
no fading; apical patch of FW large, and solid cinnamon-brown in color, with little 
or no powdering of yellow scales; black median lunules in cells M; and Cu, of FW 
“square-shaped,” not oblate; cinnamon-brown ground color of HW in basal portion 
and in postmedian band between postmedian and submarginal rows of silver spots 
solid, with little or no infusion of yellow scales; all postmedian light spots of HW 
prominent and well silvered. 

Length of forewing: 20.08 mm average. 

Female. Similar to male except orange ground color of dorsal surface slightly 
more pale and black dorsal wing margins less broad and heavy, with orange marginal 
row of spots more conspicuous. 

Length of forewing: 20.57 mm average. 

Type locality. The type locality of atrocostalis is Chicoutimi, Quebec. 

Distribution. The range of atrocostalis extends from Nova Scotia west through 
Canada to western Saskatchewan and as far north as Churchill and Lynn Lake, 
Manitoba. In the United States, atrocostalis is found in Maine, most of Vermont 
and New Hampshire, the northern portions of New York, Michigan, Wisconsin, 
Minnesota, and in eastern North Dakota (Fig. 1). 

Remarks. Huard (in Provancher & Huard, 1926) used the name atrocostalis at 
the species rank, but shortly after, McDunnough (1928) examined the type and 
found that it represented what he considered to be the eastern Canadian form of 
myrina. McDunnough then applied the name at subspecies rank to eastern Canada 
specimens. 

The subspecies atrocostalis intergrades with tollandensis in Alberta, Montana, and 
northwestern Wyoming, with albequina in the northwestern part of its range and 
myrina along much of the southern limit of its range. 

The name jenningsae (Holland) was applied in 1928 to an aberrant male speci- 
men taken by Mrs. O. E. Jennings at Jellicoe, on Thunder Bay, Ontario. The 
transverse markings of the wings are fused and enlarged. The type is in the 
Carnegie Museum. 

Records. Two hundred thirty-nine typical specimens (159 ¢, 80 9) and addi- 
tional intergrades were examined: 


atrocostalis 


Maine: Rangeley; E. Haepswell; Gilead; Seboomook Lake; Androscoggin, Penobscot, 
and Piscataquis cos. Manirospa: Bereton Lake; Kelwood; Red Rock Lake; Broken- 
head; Churchill; Betula Lake; Rennie; McCreary; Telford; and Riding Mts. Micut- 
GAN: Cheboygan, Chippewa, Emmet, Ostego, and Schoolcraft cos. MINNESOTA: 
Arago; Aitkin, Anoka, Cass, Cook, Crow Wing, Dakota, Lake, Otter Tail, St. Louis, 
Sherburne, and Rice cos. New Brunswick: Edmundston; St. Leonard; and Dor- 
chester. New Hampsuire: Randolph; Jefferson; Belknap, Carrol, Coos, and Grafton 
cos. Ontario: Nakina; Hymers; Savanne; Nipegon; Nipegon Lake; One Sided 
Lake; Upsala; Poland; Geraldton; Sudbury; Wabigoon; Guelph; Goodwood; Toronto; 
Charlton; and Blind River. QuEesEc: Montreal; Casapedia River; Lytton; Lanoriae; 
Laval; Lomininque; Bois de Filion; Laurentides Park; St. Calixte de Kilkenny; 
Metapedia; Ste. Flavien; and St. Agapit. SaskKATCHEWAN: Rivercourse and Lloyd- 
minster. VERMONT: Caledonia and Windsor cos. Wisconsin: Chippewa, Dunn, 
Forest, Lincoln, Marathon, Oneida, Price, and Sawyer cos. 


VOLUME 31, NuMBER 4 261 


Fig. 7. Adults of B. selene: (a) sabulocollis female, paratype, Timnath, Larimer 
Co., Colo.; (b) same, ventral; (c) sabulocollis female paratype, Mineral Springs, 
Slope Co., N.Dak.; (d) same, ventral; (e) tollandensis female, Tolland, Gilpin Co., 
Colo.; (£) same, ventral; (g) tollandensis female, Avon, Cache Co., Ut.; (h) same, 
ventral. All photos natural size. 


262 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Boloria selene terraenovae (Holland) 
(Fig. 5) 


Brenthis myrina terrae-novae Holland, 1928, p. 36. dos Passos, 1935, p. 87. McDun- 
nough, 1938, p. 16. 

Brenthis myrina var. terrae-novae: Holland, 1931, p. 105, pl. LV, fig. 13 (male 
type). 

Brenthis selene terrae-novae: Clark, 1941, p. 384. 

Boloria selene terrae-novae: Klots, 1951, p. 89. 

Boloria selene terraenovae: dos Passos, 1964, p. 89. Shepard (in. Howe), 1975, p. 
264. 


Diagnosis. The dorsal wing surfaces of the majority of terraenovae specimens are 
much like myrina. The wing margins are not as heavily marked with black scales 
as are those of atrocostalis. Basal suffusion of dark scales is a bit heavier than in 
myrina, but few specimens approach the degree of suffusion in Holland’s types. 

The ventral surfaces of the HW are a different shade of brown, being con- 
sistently a deep, bright mahogany rather than the cinnamon-brown of the other 
subspecies. Occasionally, specimens of atrocostalis will approach this mahogany 
color of the HW, but have the heavier black wing margins characteristic of that 
subspecies. 

Male. Dorsal surface: ground color bright red-orange; black marginal band not 
heavy, and enclosed orange marginal row of spots visible along entire margin of 
both wings in most specimens; normal pattern of black markings sharply demarcated 
on ground color; suffusion of dark scales in basal areas of both wings, in HW 
filling ca. one-half of discal cell; basal spot in discal cell of HW almost always 
obscured by dark scales. 

Ventral surface: ground color of FW deep orange, usually extending to apical 
patch with little or no change in color; apical patch of FW not reduced, and solid 
bright mahogany-brown, with no powdering of yellow scales; ground color of HW 
uniform deep, bright mahogany, and solid, with little or no infusion of yellow 
scales; yellow areas of HW more or less restricted to cells Rs, Mi, Ms, and Cu; in 
area between submedian and postmedian rows of silver spots and to normal patch 
in cells M. and Ms; just basad of submarginal row of silver spots; row of black limbal 
spots on HW overlaid with mahogany scales, with mahogany color masking black 
so that spots appear dark brown; all postmedian silver spots prominent, except spot 
in cell Mz, which is reduced in most specimens. 

Length of forewing: 20.67 mm average. 

Female. Similar to male, but orange of dorsal ground color more pale. 

Length of forewing: 21.50 mm average. 

Type locality. The type locality of terraenovae is listed only as Newfoundland. 
Mr. C. F. dos Passos (1935) examined the holotype and two paratypes and com- 
mented that, although the labels did not show from what area of Newfoundland 
they came, it was probably the eastern part of the island. The types are in the 
Carnegie Museum. 

Distribution. The range of terraenovae is limited to the Island of Newfoundland. 

Remarks. In Nova Scotia and possibly New Brunswick, terraenovae intergrades 
to some extent with atrocostalis. Boloria selene have been recorded from Labrador, 
and these may represent neither terraenovae nor atrocostalis, but too few specimens 
are available to make sound judgments. 

In the original description by Holland (1928), the only distinguishing factor 
mentioned was that the inner halves of both wings were suffused dorsally by dark 
scales, obscuring the wing markings in these areas. Such specimens are not repre- 
sentative of the normal run of specimens from Newfoundland, as evidenced by the 
specimens examined for this study. In 1935, dos Passos reached similar conclusions 


VoLUME 31, NuMBER 4 263 


Fig. 8. Adults of B. selene: (a) tollandensis/albequina female, Moxee Bog, 
Yakima Co., Wash.; (b) same, ventral; (c) atrocostalis female, Lytton, Que.; (d) 
same, ventral; (e) atrocostalis/tollandensis female, Florence, Ravalli Co., Mont.; (f) 
same, ventral. All photos natural size. 


when he examined a series of 32 males and 11 females from Newfoundland and 
compared them with the types. He found that some specimens were as heavily 
marked with dark scales in the basal wing areas but that most of the specimens were 
not. 

Records. Fifteen typical specimens (14 ¢, 1 2) of terraenovae were studied. 
These records and additional intergrades are as follows: 


terraenovdae 


NEWFOUNDLAND: Doyle’s Station; Codroy; and Spruce Brook. 


nr. terraenovae 


Nova Scotia: Mt. Uniacke; Port Shoreham; and Passboro. 


264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


CONCLUSIONS 


Although Boloria selene is quite variable in both size and markings, 
several recognizable and consistent populations exist within its range in 
North America. There is considerable intergradation among these pop- 
ulations at the limits of their ranges. Over the course of this study, the 
ranges of these recognizable populations have been more precisely de- 
fined, as have the zones of intergradation among populations. 

In general, selene in North America exhibits a gradual intergradation 
of characters throughout its range, more evident on north-south lines, 
but also present on east-west lines. Names are available to apply at 
subspecific rank to populations sufficiently differentiated on these clines, 
and are useful tools when so applied. It is felt that the existence of 
these clines has not been obscured by the recognition of too many 
subspecies names and that those names recognized in this paper best 
portray the population changes on the clines. 

In the west, typical tollandensis from the mountains of central Colo- 
rado and New Mexico form the southern pole of a cline and gradually 
intergrade into albequina in southwestern Alaska at the northern pole 
of the cline. A quite variable intermediate population occurs in a broad 
band including eastern Oregon and Washington, most of British Co- 
lumbia, Alberta, northern Idaho and Montana, and northwestern Wy- 
oming. This intermediate population is influenced by intergrading from 
the east, by atrocostalis on the north and sabulocollis to the south. 
Specimens from much of this intermediate population may show char- 
acteristics predominantly of tollandensis, albequina, or atrocostalis, or 
characteristics that are intermediate. 

In the western Great Plains, a relict population (sabulocollis), which 
probably became isolated as coniferous forest began its demise in this 
area, has been newly described. 

In the east, the north-south intergradation is more stepped and closely 
parallels environmental factors, chiefly habitat. The northern popula- 
tion (atrocostalis) of the coniferous boreal forest zone intergrades into 
the central population (myrina), associated with the deciduous eastern 
forest zone, and then into the southern population (nebraskensis) of 
the prairie zone, which has never supported forests. 

On the east coast, the population named marilandica is a local pheno- 
typic expression influenced by habitat similar to that of nebraskensis. 
The marilandica population is restricted to a prairie habitat of the lower 
coastal plain in Maryland, while myrina is present in the piedmont and 
deciduous forest zones. Because it is a local phenotypic population 


Y 


VoLUME 31, NuMBER 4 265 


differing only slightly from myrina, marilandica has been relegated to 
the synonymy of myrina. 

A weaker east-west cline is formed by terraenovae and atrocostalis in 
the northeastern portion of the range of selene. The name terraenovae 
is retained to apply to the sufficiently differentiated eastern pole of this 
cline. 

Through application of this gradual intergradation concept, ranges of 
some selene populations have been significantly expanded (nebraskensis 
and tollandensis), while others have been reduced (albequina and 
myrina ). 

As a result of this study, the following treatment of Boloria selene in 
North America is proposed: 


Boloria selene (Denis & Schiffermiiller), 1775 


a. Ss. myrina (Cramer), 1777 
myrinus ( Herbst), 1798 
myrissa (Godart ), 1819 
ab. “nubes” (Scudder), 1889 
ab. “nivea” (Gunder), 1928 
marilandica (Clark), 1941 
b. s. nebraskensis (Holland), 1928 
s. sabulocollis Kohler, 1977 
. s. tollandensis (Barnes & Benjamin), 1925 
ab. “serratimarginata” (Gunder), 1926 
e. s. albequina (Holland), 1928 
ab. “baxteri’” ( Holland), 1928 
. s. atrocostalis (Huard), 1927 
ab. “jenningsae” (Holland), 1928 
g. s. terraenovae (Holland), 1928 


ao 


lum] 


ACKNOWLEDGMENTS 


Sincerest appreciation is given to the following for their contributions 
to this study. Without their efforts in supplying specimens and col- 
lection records, reviewing and offering suggestions to the manuscript, 
and furnishing valuable information and assistance, this work could not 
have been completed: Norman L. Anderson (Dept. Zoology and En- 
tomology, Montana State Univ., Bozeman), Edward U. Balsbaugh, Jr. 
(Entomology-Zoology Dept., South Dakota State Univ., Brookings), A. 
E. Brower, F. Martin Brown, Harry K. Clench (Section of Insects, 
Carnegie Museum, Pittsburgh), Patrick J. Conway, Julian P. Donahue 
(Asst. Curator, Dept. of Entomology, Natural History Museum of Los 
Angeles County), Ernst J. Dornfeld (Dept. of Zoology, Oregon State 
Univ., Corvallis), J. Gordon Edwards (Dept. of Biological Sciences, 
San Jose State Univ., San Jose), J. Donald Eff, Scott L. Ellis, John H. 
Fales, Clifford D. Ferris (Bioengineering, Univ. of Wyoming, Laramie), 


266 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


William D. Field (Curator of Lepidoptera, National Museum of Natural 
History, Washington, D.C.), Mike Fisher, L. Paul Grey, J. Richard 
Heitzman, John Hinchliff, Ronald R. Hooper, Ronald L. Huber, Kurt 
Johnson (Dept. of Biology, City Univ. of New York), Alexander B. 
Klots, James H. Lowe (School of Forestry, Univ. of Montana, Missoula), 
David V. McCorkle (Dept. of Biology, Oregon College of Education, 
Monmouth), L. Joseph Melton, II, Lee D. Miller (Curator, Allyn 
Museum of Entomology, Sarasota), John S. Nordin, James Oberfoell, 
Kenelm W. Philip (Inst. of Arctic Biology, Univ. of Alaska, College), 
Frederick H. Rindge (Curator of Lepidoptera, The American Museum 
of Natural History, New York), James C. Schmulbach (Dept. of Biology, 
Univ. of South Dakota, Vermillion), Orville D. Spencer, and Charles A. 
Triplehorn (Dept. of Zoology and Entomology, The Ohio State Univ., 
Columbus ). 


LITERATURE CITED 


ALBRIGHT, R. 1960. A record of Boloria selene in Oregon. J. Lepid. Soc. 14: 158. 

Barnes, W. & F. H. Benjamin. 1925. A new subspecies of Brenthis myrina 
Cram. (Lepid.: Nymphalidae). Ent. News 36: 44. 

Brooks, G. S. 1942. A revised check list of the butterflies of Manitoba. Can. Ent. 
74; 31-36. 

Brown, F. M. 1954. Some notes on Boloria in central Colorado (Nymphalidae). 
Lepid. News 8: 64-66. 

1957. Colorado butterflies. Denver Museum of Natural History, Denver, 
368 p. 

Cary, M. 1901. Notes on the butterflies of Sioux County, Nebraska. Can. Ent. 
33: 305-311. 

1906. On the dinurnal Lepidoptera of the Athabaska and Mackenzie 
region, British America. U.S. Nat. Mus. Proc. 31: 425-457. 

Ciark, A. H. 1932. The butterflies of the District of Columbia and vicinity. U-S. 
Nat. Mus. Bull. 157, 337 p. 

1941. Notes on the American representatives of the butterfly genus 
Argynnis. J. Wash. Acad. Sci. 31; 381-384. 

Comstock, W. P. 1940. Butterflies of New Jersey. J. N. Y. Ent. Soc. 48: 47-84. 

Cramer, P. “1779”(1775)—1782. Papillons exotiques des trois parties du monde, 
lAsie, Afrique et l’Amerique. S. J. Baalde, Amsterdam, Barthelemy Wild, 
Utrecht, 4 vol. 

Davenport, D. & V. G. Deruer. 1938. Bibliography of the described life his- 
tories of the Rhopalocera of America north of Mexico 1889-1937. Ent. Amer. 
17: 155-196. 

pos Passos, C. F. 1935. Some butterflies of southern Newfoundland with de- 
scriptions of new subspecies (Lepid. Rhopal.). Can. Ent. 67: 82-88. 

. 1938. The types of Lepidoptera described by J. D. Gunder. Amer. Mus. 

Nov. 999, 16 p. 

1964. A synonymic list of the Nearctic Rhopalocera. Lepid. Soc. Mem. 1, 
YA5 p. 

pos Passos & L. P. Grey. 1934. A list of the butterflies of Maine with notes con- 
cerning some of them (Lepid., Rhopalocera). Can. Ent. 66: 188-192. 

1945. <A genitalic survey of Argynninae (Lepidoptera, Nymphalidae). 

Amer. Mus. Nov. 1296, 21 p. 


VoLUME 31, NuMBER 4 267 


Dyar, H. G. 1900. Papers from the Harriman Alaskan Expedition, XII, ento- 
mological results (6): Lepidoptera. Proc. Wash. Acad. Sci. 2:487-501. 

Epwarps, W. H. 1876. Farther notes upon Argynnis myrina. Can. Ent. 8: 161-— 
163. 

Eure|uicu, P. R. & A. H. Exruicu. 1961. How to know the butterflies. Wm. C. 
Brown Co., Dubuque, Iowa, 262 p. 

Exrop, M. J. 1906. The butterflies of Montana. Univ. Mont. Bull. 10, 174 p. 

Ferris, C. D. 1971. An annotated checklist of the Rhopalocera [butterflies] of 
Wyoming. Sci. Monogr. 23, Agric. Exp. Stat., Univ. of Wyoming, Laramie, 75 


Dp. 

Forses, W. T. M. 1960. Lepidoptera of New York and neighboring states. Part 
IV. Cornell Univ. Agric. Exp. Stat. Mem. 371, 188 p. 

FRANZEN, J. W. 1914. Minnesota butterflies (Lepid.). Ent. News 25: 363-371. 

Gopart, J. B. 1819. Encyclopedie methodique. Histoire naturelle entomologie, ou 
histoire naturelle des Crustaces, des Arachnides et des Insectes. Agass, Paris, 
vol. 9. 

GunbER, J. D. 1926. Several new aberrant Lepidoptera (Rhopalocera). Ent. News 
37: 1-9. 

. 1928. Additional transition forms (Lepid., Rhopalocera). Can. Ent. 60: 
162-168. 

Herest, J. F. W. 1798 (1785-1806). Natursystem aller bekannten in-und Aus- 
landischen Insecten; nach dem system des Carl Von Linne’ bearbeitet (Von C. 
G. Jablonsky) fortgestzt Von J. F. W. Herbst. Berlin, Pauli, 723 p. 

Hoiuanp, W. J. 1928. III. Notes upon some North American species and varieties 
of the genus Brenthis. Ann. Carnegie Mus. 19: 35-45. 

1931. The butterfly book. Rev. Ed., Doubleday & Co., Inc., New York, 
A424 p. 

Jounson, K. 1972 (1973). The butterflies of Nebraska. J. Res. Lepid. 11: 1-64. 

1975 (1976). Post-Pleistocene environments and montane butterfly relicts 

on the western Great Plains. J. Res. Lepid. 14: 216-232. 
1976. Concerning the name Anthocaris coloradensis Hy. Edwards with 
designation of a new subspecies (Pieridae). J. Lepid. Soc. 30: 252-259. 
Kiors, A. B. 1937. Some notes on Colias and Brenthis (Lepidoptera, Pieridae and 
Nymphalidae). J. N. Y. Ent. Soc. 45: 311-333. 

1951. <A field guide to the butterflies. Houghton Mifflin Co., Boston, 
349 p. 

LeussLer, R. A. 1938. An annotated list of the butterflies of Nebraska with the 
description of a new species (Lepid.: Rhopalocera). Ent. News 49: 3-9, 76— 
80, 213-218, 275-280. 

McDunnoucu, J. 1928. Notes on Canadian diumal Lepidoptera. Can. Ent. 60: 
266-275. 

. 1930. The Lepidoptera of the north shore of the Gulf of St. Lawrence. 

Can. Ent. 62: 107-108. 

1932. Notes on some diurnal Lepidoptera figured in Holland’s revised 

butterfly book. Can. Ent. 64: 267-270. 

1938. Check list of the Lepidoptera of Canada and the United States of 
America. Part I, Macrolepidoptera. Mem. S. Calif. Acad. Sci. 1, 275 p. 

Newcomer, E. J. & W. H. Rocrers. 1963. Notes on Boloria selene (Nymphalidae ) 
in the Pacific Northwest. J. Lepid. Soc. 17: 171-172. 

PROVANCHER, L. & V. A. Huarp. 1926. Faune entomologique de la province 
Quebec, sixieme ordre, les lepidopteres. Nat. Can. 54: 129-143. 

REMINGTON, C. L. 1952. The biology of nearctic Lepidoptera. I. Foodplants and 
life histories of Colorado Papilionoidea. Psyche 59: 61-70. 

Ryscaarpb, G. N. 1939. <A preliminary study of the superfamily Papilionoidea in 


268 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the northern portion of Pine County, Minnesota (Lepidoptera). Ent. News 50: 
191-196. 

ScuppER, S. H. 1863. A list of the butterflies of New England. Proc. Essex Inst. 
3: 161-179. 

1868. Supplement to a list of the butterflies of New England. Proc. 

Boston Soc. Nat. Hist. 11: 375-384. 

1889. Butterflies of the eastern United States and Canada, with special 
reference to New England. Vols. 1-3, Published by the Author, Cambridge, 
Mass. 

SHEPARD, J. H. 1975. Genus Boloria Reuss. Pp. 243-252 in Howe, W. H. (ed.) 
The butterflies of North America. Doubleday & Co., Inc., Garden City, New 
York. 


VoLUME 31, NuMBER 4 261 


Fig. 7. Adults of B. selene: (a) sabulocollis female, paratype, Timnath, Larimer 
Co., Colo.; (b) same, ventral; (c) sabulocollis female paratype, Mineral Springs, 
Slope Co., N.Dak.; (d) same, ventral; (e) tollandensis female, Tolland, Gilpin Co.., 
Colo.; (£) same, ventral; (g) tollandensis female, Avon, Cache Co., Ut.; (h) same, 
ventral. All photos natural size. 


262 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Boloria selene terraenovae (Holland) 
(Fig. 5) 


Brenthis myrina terrae-novae Holland, 1928, p. 36. dos Passos, 1935, p. 87. McDun- 
nough, 1938, p. 16. 

Brenthis myrina var. terrae-novae: Holland, 1931, p. 105, pl. LV, fig. 13 (male 
type). 

Brenthis selene terrae-novae: Clark, 1941, p. 384. 

Boloria selene terrae-novae: Klots, 1951, p. 89. 

Boloria selene terraenovae: dos Passos, 1964, p. 89. Shepard (in Howe), 1975, p. 
264. 


Diagnosis. The dorsal wing surfaces of the majority of terraenovae specimens are 
much like myrina. The wing margins are not as heavily marked with black scales 
as are those of atrocostalis. Basal suffusion of dark scales is a bit heavier than in 
myrina, but few specimens approach the degree of suffusion in Holland’s types. 

The ventral surfaces of the HW are a different shade of brown, being con- 
sistently a deep, bright mahogany rather than the cinnamon-brown of the other 
subspecies. Occasionally, specimens of atrocostalis will approach this mahogany 
color of the HW, but have the heavier black wing margins characteristic of that 
subspecies. 

Male. Dorsal surface: ground color bright red-orange; black marginal band not 
heavy, and enclosed orange marginal row of spots visible along entire margin of 
both wings in most specimens; normal pattern of black markings sharply demarcated 
on ground color; suffusion of dark scales in basal areas of both wings, in HW 
filling ca. one-half of discal cell; basal spot in discal cell of HW almost always 
obscured by dark scales. 

Ventral surface: ground color of FW deep orange, usually extending to apical 
patch with little or no change in color; apical patch of FW not reduced, and solid 
bright mahogany-brown, with no powdering of yellow scales; ground color of HW 
uniform deep, bright mahogany, and solid, with little or no infusion of yellow 
scales; yellow areas of HW more or less restricted to cells Rs, M:, Ms, and Cu; in 
area between submedian and postmedian rows of silver spots and to normal patch 
in cells M. and M; just basad of submarginal row of silver spots; row of black limbal 
spots on HW overlaid with mahogany scales, with mahogany color masking black 
so that spots appear dark brown; all postmedian silver spots prominent, except spot 
in cell M2, which is reduced in most specimens. 

Length of forewing: 20.67 mm average. 

Female. Similar to male, but orange of dorsal ground color more pale. 

Length of forewing: 21.50 mm average. 

Type locality. The type locality of terraenovae is listed only as Newfoundland. 
Mr. C. F. dos Passos (1935) examined the holotype and two paratypes and com- 
mented that, although the labels did not show from what area of Newfoundland 
they came, it was probably the eastern part of the island. The types are in the 
Carnegie Museum. 

Distribution. The range of terraenovae is limited to the Island of Newfoundland. 

Remarks. In Nova Scotia and possibly New Brunswick, terraenovae intergrades 
to some extent with atrocostalis. Boloria selene have been recorded from Labrador, 
and these may represent neither terraenovae nor atrocostalis, but too few specimens 
are available to make sound judgments. 

In the original description by Holland (1928), the only distinguishing factor 
mentioned was that the inner halves of both wings were suffused dorsally by dark 
scales, obscuring the wing markings in these areas. Such specimens are not repre- 
sentative of the normal run of specimens from Newfoundland, as evidenced by the 
specimens examined for this study. In 1935, dos Passos reached similar conclusions 


VOLUME 31, NUMBER 4 263 


Fig. 8. Adults of B. selene: (a) tollandensis/albequina female, Moxee Bog, 
Yakima Co., Wash.; (b) same, ventral; (c) atrocostalis female, Lytton, Que.; (d) 
same, ventral; (e) atrocostalis/tollandensis female, Florence, Ravalli Co., Mont.; (£) 
same, ventral. All photos natural size. 


when he examined a series of 32 males and 11 females from Newfoundland and 
compared them with the types. He found that some specimens were as_ heavily 
marked with dark scales in the basal wing areas but that most of the specimens were 
not. 

Records. Fifteen typical specimens (14 &, 1 2) of terraenovae were studied. 
These records and additional intergrades are as follows: 


terraenovae 


NEWFOUNDLAND: Doyle’s Station; Codroy; and Spruce Brook. 


nr. terraenovae 


Nova Scotia: Mt. Uniacke; Port Shoreham; and Passboro. 


264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


CONCLUSIONS 


Although Boloria selene is quite variable in both size and markings, 
several recognizable and consistent populations exist within its range in 
North America. There is considerable intergradation among these pop- 
ulations at the limits of their ranges. Over the course of this study, the 
ranges of these recognizable populations have been more precisely de- 
fined, as have the zones of intergradation among populations. 

In general, selene in North America exhibits a gradual intergradation 
of characters throughout its range, more evident on north-south lines, 
but also present on east-west lines. Names are available to apply at 
subspecific rank to populations sufficiently differentiated on these clines, 
and are useful tools when so applied. It is felt that the existence of 
these clines has not been obscured by the recognition of too many 
subspecies names and that those names recognized in this paper best 
portray the population changes on the clines. 

In the west, typical tollandensis from the mountains of central Colo- 
rado and New Mexico form the southern pole of a cline and gradually 
intergrade into albequina in southwestern Alaska at the northern pole 
of the cline. A quite variable intermediate population occurs in a broad 
band including eastern Oregon and Washington, most of British Co- 
lumbia, Alberta, northern Idaho and Montana, and northwestern Wy- 
oming. This intermediate population is influenced by intergrading from 
the east, by atrocostalis on the north and sabulocollis to the south. 
Specimens from much of this intermediate population may show char- 
acteristics predominantly of tollandensis, albequina, or atrocostalis, or 
characteristics that are intermediate. 

In the western Great Plains, a relict population (sabulocollis), which 
probably became isolated as coniferous forest began its demise in this 
area, has been newly described. 

In the east, the north-south intergradation is more stepped and closely 
parallels environmental factors, chiefly habitat. The northem popula- 
tion (atrocostalis) of the coniferous boreal forest zone intergrades into 
the central population (myrina), associated with the deciduous eastern 
forest zone, and then into the southern population (nebraskensis) of 
the prairie zone, which has never supported forests. 

On the east coast, the population named marilandica is a local pheno- 
typic expression influenced by habitat similar to that of nebraskensis. 
The marilandica population is restricted to a prairie habitat of the lower 
coastal plain in Maryland, while myrina is present in the piedmont and 
deciduous forest zones. Because it is a local phenotypic population 


VoLUME 31, NuMBER 4 265 


differing only slightly from myrina, marilandica has been relegated to 
the synonymy of myrina. 

A weaker east-west cline is formed by terraenovae and atrocostalis in 
the northeastern portion of the range of selene. The name terraenovae 
is retained to apply to the sufficiently differentiated eastern pole of this 
cline. 

Through application of this gradual intergradation concept, ranges of 
some selene populations have been significantly expanded (nebraskensis 
and tollandensis), while others have been reduced (albequina and 
myrina ). 

As a result of this study, the following treatment of Boloria selene in 
North America is proposed: 


Boloria selene (Denis & Schiffermiiller), 1775 


a. s. myrina (Cramer), 1777 
myrinus (Herbst), 1798 
myrissa (Godart), 1819 
ab. “nubes” (Scudder), 1889 
ab. “nivea” (Gunder ), 1928 
marilandica (Clark), 1941 
b. s. nebraskensis (Holland ), 1928 
c. s. sabulocollis Kohler, 1977 
d. s. tollandensis (Bames & Benjamin), 1925 
ab. “serratimarginata” (Gunder), 1926 
e. s. albequina (Holland), 1928 
ab. “baxteri” (Holland), 1928 
f. s. atrocostalis (Huard), 1927 
ab. “jenningsae” (Holland), 1928 
g. s. terraenovae (Holland), 1928 


ACKNOWLEDGMENTS 


Sincerest appreciation is given to the following for their contributions 
to this study. Without their efforts in supplying specimens and col- 
lection records, reviewing and offering suggestions to the manuscript, 
and furnishing valuable information and assistance, this work could not 
have been completed: Norman L. Anderson (Dept. Zoology and En- 
tomology, Montana State Univ., Bozeman), Edward U. Balsbaugh, Jr. 
(Entomology-Zoology Dept., South Dakota State Univ., Brookings), A. 
E. Brower, F. Martin Brown, Harry K. Clench (Section of Insects, 
Carnegie Museum, Pittsburgh), Patrick J. Conway, Julian P. Donahue 
(Asst. Curator, Dept. of Entomology, Natural History Museum of Los 
Angeles County), Ernst J. Dornfeld (Dept. of Zoology, Oregon State 
Univ., Corvallis), J. Gordon Edwards (Dept. of Biological Sciences, 
San Jose State Univ., San Jose), J. Donald Eff, Scott L. Ellis, John H. 
Fales, Clifford D. Ferris (Bioengineering, Univ. of Wyoming, Laramie), 


266 | JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


William D. Field (Curator of Lepidoptera, National Museum of Natural 
History, Washington, D.C.), Mike Fisher, L. Paul Grey, J. Richard 
Heitzman, John Hinchliff, Ronald R. Hooper, Ronald L. Huber, Kurt 
Johnson (Dept. of Biology, City Univ. of New York), Alexander B. 
Klots, James H. Lowe (School of Forestry, Univ. of Montana, Missoula), 
David V. McCorkle (Dept. of Biology, Oregon College of Education, 
Monmouth), L. Joseph Melton, III, Lee D. Miller (Curator, Allyn 
Museum of Entomology, Sarasota), John S. Nordin, James Oberfoell, 
Kenelm W. Philip (Inst. of Arctic Biology, Univ. of Alaska, College), 
Frederick H. Rindge (Curator of Lepidoptera, The American Museum 
of Natural History, New York), James C. Schmulbach (Dept. of Biology, 
Univ. of South Dakota, Vermillion), Orville D. Spencer, and Charles A. 
Triplehorn (Dept. of Zoology and Entomology, The Ohio State Univ., 
Columbus ). 


LITERATURE CITED 


ALBRIGHT, R. 1960. A record of Boloria selene in Oregon. J. Lepid. Soc. 14; 158. 

Barnes, W. & F. H. BENyAmin. 1925. A new subspecies of Brenthis myrina 
Cram. (Lepid.: Nymphalidae). Ent. News 36: 44. 

Brooks, G. S. 1942. A revised check list of the butterflies of Manitoba. Can. Ent. 
7A: 31-36. 

Brown, F. M. 1954. Some notes on Boloria in central Colorado (Nymphalidae). 
Lepid. News 8: 64—66. 

1957. Colorado butterflies. Denver Museum of Natural History, Denver, 


368 p. 
Cary, M. 1901. Notes on the butterflies of Sioux County, Nebraska. Can. Ent. 
se Se sBIUIL. 


. 1906. On the dinurnal Lepidoptera of the Athabaska and Mackenzie 
region, British America. U.S. Nat. Mus. Proc. 31: 425—457. 

Criark, A. H. 1932. The butterflies of the District of Columbia and vicinity. U.S. 
Nat. Mus. Bull. 157, 337 p. 

1941. Notes on the American representatives of the butterfly genus 
Argynnis. J. Wash. Acad. Sci. 31: 381-384. 

Comstock, W. P. 1940. Butterflies of New Jersey. J. N. Y. Ent. Soc. 48: 47-84. 

CraMeEer, P. “1779’(1775)—-1782. Papillons exotiques des trois parties du monde, 
lAsie, Afrique et !Amerique. S. J. Baalde, Amsterdam, Barthelemy Wild, 
Utrecht, 4 vol. 

Davenport, D. & V. G. Dernier. 1938. Bibliography of the described life his- 
tories of the Rhopalocera of America north of Mexico 1889-1937. Ent. Amer. 
Mila 106; 

pos Passos, C. F. 1935. Some butterflies of southern Newfoundland with de- 
scriptions of new subspecies (Lepid. Rhopal.). Can. Ent. 67: 82-88. 

. 1938. The types of Lepidoptera described by J. D. Gunder. Amer. Mus. 

Nov. 999, 16 p. 

1964. A synonymic list of the Nearctic Rhopalocera. Lepid. Soc. Mem. 1, 
145 p. 

pos Passos & L. P. Grey. 1934. A list of the butterflies of Maine with notes con- 
cerning some of them (Lepid., Rhopalocera). Can. Ent. 66: 188-192. 

1945. A genitalic survey of Argynninae (Lepidoptera, Nymphalidae). 

Amer. Mus. Nov. 1296, 21 p. 


VoLUME 31, NUMBER 4 267 


Dyar, H. G. 1900. Papers from the Harriman Alaskan Expedition, XII, ento- 
mological results (6): Lepidoptera. Proc. Wash. Acad. Sci. 2:487-501. 

Epwarps, W. H. 1876. Farther notes upon Argynnis myrina. Can. Ent. 8: 161-— 
163. 

Eueticu, P. R. & A. H. EnruicH. 1961. How to know the butterflies. Wm. C. 
Brown Co., Dubuque, Iowa, 262 p. 

Exrop, M. J. 1906. The butterflies of Montana. Univ. Mont. Bull. 10, 174 p. 

Ferris, C. D. 1971. An annotated checklist of the Rhopalocera [butterflies] of 
Wyoming. Sci. Monogr. 23, Agric. Exp. Stat., Univ. of Wyoming, Laramie, 75 


Dp. 

Forses, W. T. M. 1960. Lepidoptera of New York and neighboring states. Part 
IV. Cornell Univ. Agric. Exp. Stat. Mem. 371, 188 p. 

FRANZEN, J. W. 1914. Minnesota butterflies (Lepid.). Ent. News 25: 363-371. 

Gopart, J. B. 1819. Encyclopedie methodique. Histoire naturelle entomologie, ou 
histoire naturelle des Crustaces, des Arachnides et des Insectes. Agass, Paris, 
vol. 9. 

Gunoper, J. D. 1926. Several new aberrant Lepidoptera (Rhopalocera). Ent. News 
37: 1-9. 

. 1928. Additional transition forms (Lepid., Rhopalocera). Can. Ent. 60: 
162-168. 

Hersst, J. F. W. 1798 (1785-1806). Natursystem aller bekannten in-und Aus- 
landischen Insecten; nach dem system des Carl Von Linne’ bearbeitet (Von C. 
G. Jablonsky) fortgestzt Von J. F. W. Herbst. Berlin, Pauli, 723 p. 

Hoiuanp, W. J. 1928. III. Notes upon some North American species and varieties 
of the genus Brenthis. Ann. Carnegie Mus. 19: 35-45. 

1931. The butterfly book. Rev. Ed., Doubleday & Co., Inc., New York, 
424 p. 

Jounson, K. 1972 (1973). The butterflies of Nebraska. J. Res. Lepid. 11: 1-64. 

1975 (1976). Post-Pleistocene environments and montane butterfly relicts 

on the western Great Plains. J. Res. Lepid. 14: 216-232. 
1976. Concerning the name Anthocaris coloradensis Hy. Edwards with 
designation of a new subspecies (Pieridae). J. Lepid. Soc. 30: 252-259. 
Kiors, A. B. 1937. Some notes on Colias and Brenthis (Lepidoptera, Pieridae and 
Nymphalidae). J. N. Y. Ent. Soc. 45: 311-333. 

1951. A field guide to the butterflies. Houghton Mifflin Co., Boston, 
349 p. 

LeEussLeR, R. A. 1938. An annotated list of the butterflies of Nebraska with the 
description of a new species (Lepid.: Rhopalocera). Ent. News 49: 3-9, 76- 
80, 213-218, 275-280. 

McDwunnoucu, J. 1928. Notes on Canadian diumal Lepidoptera. Can. Ent. 60: 
266-275. 

. 1930. The Lepidoptera of the north shore of the Gulf of St. Lawrence. 

Can. Ent. 62: 107-108. 

1932. Notes on some diurnal Lepidoptera figured in Holland’s revised 

butterfly book. Can. Ent. 64: 267-270. 

1938. Check list of the Lepidoptera of Canada and the United States of 
America. Part I, Macrolepidoptera. Mem. S. Calif. Acad. Sci. 1, 275 p. 

Newcomer, E. J. & W. H. Rocers. 1963. Notes on Boloria selene (Nymphalidae ) 
in the Pacific Northwest. J. Lepid. Soc. 17: 171-172. 

PRovANCHER, L. & V. A. Huarp. 1926. Faune entomologique de la province 
Quebec, sixieme ordre, les lepidopteres. Nat. Can. 54: 129-143. 

REMINGTON, C. L. 1952. The biology of nearctic Lepidoptera. I. Foodplants and 
life histories of Colorado Papilionoidea. Psyche 59: 61-70. 

Ryscaarp, G. N. 1939. A preliminary study of the superfamily Papilionoidea in 


268 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the northern portion of Pine County, Minnesota (Lepidoptera). Ent. News 50: 
191-196. 

ScuppER, S. H. 1863. A list of the butterflies of New England. Proc. Essex Inst. 
3: 161-179. 

1868. Supplement to a list of the butterflies of New England. Proc. 

Boston Soc. Nat. Hist. 11: 375-384. 

1889. Butterflies of the eastern United States and Canada, with special 
reference to New England. Vols. 1-3, Published by the Author, Cambridge, 
Mass. 

SHEPARD, J. H. 1975. Genus Boloria Reuss. Pp. 243-252 in Howe, W. H. (ed.) 
The butterflies of North America. Doubleday & Co., Inc., Garden City, New 
York. 


VoLUME 31, NuMBER 4 277 


entrance to the bait. Fermenting bananas served as bait every day except one when 
pineapple was placed with the banana. The bait was replaced every 2-3 days. The 
trap was suspended from the NW corner of the author’s garage. The base of the 
trap was 1.3 m above ground level. 

Cuts were made in the trunk of a gmelina (Gmelina arborea) to attract other 
Charaxes. 

All Charaxes collected were caught on bright sunny days. Specimens were identi- 
fied to species and those in good condition were stored in paper triangles. Sex of 
the specimens was not determined. 


RESULTS 


The following species and numbers were trapped using banana as bait. Charaxes 
estesipe (Godart): 3 September 1973, 15 November 1973, 17 November 1973, 23 
April 1974; Charaxes tiridates (Cramer): 23 December 1973, 28 December 1973; 
Charaxes boueti Feisthamel: 11 December 1973, 9 January 1974, 21 January 1974, 
24 April 1974; Charaxes jasius Reiche: 16 December 1973, 18 December 1973, 29 
December 1973, 9 January 1974, 19 April 1974; Charaxes viola Butler: 16 December 
1973, 23 December 1973; Charaxes castor Cramer: 23 October 1973, 11 November 
1973, 18 December 1973; Charaxes brutus Cramer: 6 August 1973, 30 September 
1973; Charaxes catachrous Staudinger: 21 December 1973; Charaxes achaemenes 
Felder: 15 December 1973, 21 December 1973; Charaxes lucretius (Cramer): 25 
December 1973, 6 January 1974; Charaxes lactitinctus Karsch: 24 April 1974. 

Only one Charaxes eupale (Drury) was collected, on the day when the trap was 
baited with banana and pineapple. One C. castor was caught feeding on the sap 
from the G. arborea, and one C. lactitinctus was caught feeding on feces and urine 
(animal species unknown) on the Egede-Affa road. 


DISCUSSION 


The collection of C. ewpale on the banana-pineapple combination is interesting but 
not signficant. Owen & Chanter (1972, J. Ent. (A) 46: 135-143) reported numerous 
C. eupale attracted to banana bait in Sierra Leone. Boorman (1965, Nigerian butter- 
flies. Part III: Nymphalidae-Section I, Ibadan Univ. Press, Ibadan, 16 p.) noted 
that C. achaemenes records are few from Nigeria, mainly from savannah areas north 
of Ibadan. The collection of C. achaemenes in Enugu will add to the recorded 
distribution of the species. C. lactitinctus is considered rare by Boorman (Ibid.). 
Nigeran records are mainly from the Enugu area. Collection of this species from a 
derived savannah and urban garden situation probably indicates a lack of intensive 
collection in similar areas of Nigeria. 


ACKNOWLEDGMENTS 


I wish to thank Dr. E. U. Balsbaugh, Jr., Curator of the North Dakota State 
University Insect Collection, for encouragement during the preparation of this paper. 
I am grateful to Mr. J. C. Okafor, Botanist, Forest Department, Enugu, for identifica- 
tion of the plants. Special thanks go to my wife, Francisca, and to my collecting 
companion, Sean Lewis. 


R. D. Pererson II, Entomology Department, North Dakota State University, Fargo, 
North Dakota 58102. 


1 Published with the approval of the Director, North Dakota Agricultural Experiment Station as 
Journal Article No. 769. 


278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


SUPPLEMENT TO “A CHECKLIST OF THE BUTTERFLIES OF 
GRANT COUNTY, NEW MEXICO AND VICINITY” 


Although drought conditions prevailed in the Southwest during much of the 
summer of 1976, some localities in the mountains north of Silver City, New Mexico 
received a moderate amount of moisture. The Burro Mountains and the high plateau 
areas such as L-S Mesa were very dry, but both Cherry Creek Canyon and the 
Mimbres River Valley supported fairly lush vegetation in July. On the east slope of 
the Black Range below Emory Pass, there was an especially good crop of wild- 
flowers along Highway 90. Butterflies were abundant about these flowers for the 
first time in a number of years. Many species were taken that had not been seen in 
Grant and Sierra counties for at least five years. 

In April 1976, the author published a checklist of the butterflies of the Grant Co., 
New Mexico region (J. Lepid. Soc. 30: 38-49). This paper reported the results of 10 
years of collecting. Collecting in July 1976 produced three species new to the area 
and a number of range extensions. The purpose of this note is to update the checklist 
published earlier. The list for this region now stands at 160 confirmed species with 
several additional sight records. Additional locality data were obtained for 48 species. 

The locality alphabetic codes are those used in the original checklist and shown in 
Figures 1 and 2 of that paper. One additional code, MRV, has been introduced. This 
applies to the Mimbres River Valley from San Lorenzo to just south of Sherman. 
The three new species entries are preceded by an asterisk. 

As in the past, the author’s cousin, Ralph A. Fisher, Jr. assisted him in the field 
and he is responsible for many of the Sierra Co. records. 


HESPERIDAE-PYRGINAE 


Epargyreus clarus huachuca (Dixon). Grant Co.: SP. 

* Erynnis lucilius afranius (Lintner). Grant Co.: BM, CCC, LR, SP. Sierra Co.: 
EP. July. This species, which had not been seen prior to 1976, was quite 
common. Perhaps it was overlooked or mistaken in the field for E. persius 
fredericki H. A. Freeman, which is also common in the region. This species was 
confirmed by genitalic examination. 

E. meridanus meridanus Bell. Grant Co.: BM, LS, T. 

E. pacuvius pacuvius (Lintner). Grant Co.: July. 

E. tristus tatius (Edwards). Grant Co.: CCC, GR, MRV, SL, T. Sierra Co.: EP. 

* Pholisora alpheus (Edwards). Grant Co.: MRV. One male on 3 July 1976. 

P. catullus (Fabricius). Grant Co.: GR, MRV, SL. 

Pyrgus communis complex. Grant Co.: FH, MRV. 

Staphylos ceos (Edwards). Grant Co.: SL. 

Thorybes pylades (Scudder). Grant Co.; GC, IC, MRV, SP. 


HESPERIIDAE-HESPERINAE 


Amblyscirtes aenus ssp. Sierra Co.: EP. 

A. cassus Edwards. Grant Co.: GC. 

A. exoteria (Herrich-Schaffer ). Grant Co.: CCC, SP. Sierra Co.: EP. 
A. oslari (Skinner). Grant Co.: CCC. July. 

A. simius Edwards. Grant Co.: T. 

* A. texanae Bell. Grant Co.: MRV. One male on 3 July 1976. 
Atalopedes campestris (Boisduval). Grant Co.: MRV. July. 
Copaeodes aurantiaca (Hewitson). Grant Co.: MRYV, SL. 

Oarisma edwardsii (Barnes). Grant Co.: CCC, LGC. July. 

Piruna polingii (Barnes). Sierra Co.: EP. 


VoLUME 31, NUMBER 4 279 


PAPILIONIDAE 


Papilio multicaudata (Peale MS) Kirby. Grant Co.: GR. 
P. polyxenes asterius Stoll. Grant Co.: FH. July. 
P. rutulus arizonensis Edwards. Sierra Co.: EP. July. 


PIERIDAE 


Colias (Zerene) cesonia (Stoll). Grant Co.: Bayard. Sierra Co.: EP. July. 
Eurema mexicana (Boisduval). Grant Co.: GR. 

E. nicippe (Cramer). Grant Co.: GR, MRV. 

Phoebis sennae eubule (Linnaeus). Grant Co.: T. Sierra Co.: EP. July. 
Pieris protodice protodice (Boisduval & LeConte). Grant Co.: MRV. 


RIODINIDAE 
Apodemia nais (Edwards). Grant Co.: SP. 


LYCAENIDAE 


Atlides halesus halesus (Cramer). Grant Co.: SC, MRV. July. 

Callophrys (Callophrys) apama apama (Edwards). Grant Co.: SP. 

Callophrys (Mitoura) siva siva (Edwards). Grant Co.: MRV, SL. Sierra Co.: EP. 

Erora quaderna sanfordi dos Passos. Sierra Co.: EP. July. A series was taken in 
mid-July 1976. The specimens were absolutely fresh so that it appears this 
species is bi-voltine. It is normally collected from February—May. 

Hemiargus isola alce (Edwards). Grant Co.: GR, IC, MRV. 

Leptotes marina (Reakirt). Grant Co.: GR, MRV. 

Phaeostrymon alcestis oslari (Dyar). Grant Co.: GR, MRV. 

Strymon melinus franki Field. Grant Co.: LS, MRV, SP. Sierra Co.: EP. 


LIBYTHEIDAE 


Libytheana bachmanii ssp. Grant Co.: MRV. July. 


NYMPHALIDAE 


Asterocampa celtis montis (Edwards). Grant Co.: MRV. July. 
Chlosyne lacinia crocale (Edwards). Grant Co.: MRV, SL. July. 
Danaus gilippus strigosus (Bates). Grant Co.: MRV. 

Euptoieta claudia (Cramer). Grant Co.: GR, MRV. 

Limenitis weidemeyerii angustifascia Perkins & Perkins. Grant Co.: SP. 
Phyciodes campestris camillus Edwards. Grant Co.: MRV, SL. July. 

P. mylitta arizonensis Bauer. Grant Co.: IC. 

Phyciodes tharos (Drury). Grant Co.: MRV. July. 

Poladryas minuta arachne (Edwards). Sierra Co.: EP. 

Precis lavinia coenia (Hubner). Grant Co.: MRV, SL. 

Thessalia theona thekla (Edwards). Grant Co.: MRV, SL. Sierra Co.: EP. July. 


SATYRIDAE 


Cyllopsis pertipeda dorothea (Nabokov). Grant Co.: IC. 
Megisto rubricata cheneyorum (R. L. Chermock). Grant Co.: MRV. 


CuirForD D. Ferris,* College of Engineering, University of Wyoming, Laramie, 
Wyoming 82071. 


1 Published with the pagerove of the Director, Wyoming Agricultural Experiment Station as 
Journal Article no. JA 894. 

2 Research Associate, Allyn Museum of Entomology, Sarasota, Florida; Museum Associate, Los 
Angeles County Museum of Natural History, Los Angeles, Calif. 


280 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


A RECORD OF ANAEA AIDEA (NYMPHALIDAE) FROM 
SOUTHERN ILLINOIS 


Two worn females of Anaea aidea Guerin-Meneville (Nymphalidae) were taken 
on the morming of 3 September 1972 at Brookport, Massac Co., Illinois. They were 
captured while on the wing in a weedy, abandoned railroad yard east of town 
during approximately two hours of collecting by the author. Both specimens were 
of the summer form, i.e., their forewings had a blunt apex. No others were seen in 
the area, nor have I observed them there since. This appears to be the first record 
of the species in Illinois, as previously, the furthest east it had been reported was 
in Scott Co., Kansas and Jeff Davis Co., Texas by Field (1938, Bull. Univ. Kansas, 
Lawrence, Kansas, biol. ser. 39: 108). This is mainly a tropical and subtropical 
insect that occasionally strays north, which undoubtedly accounts for this record. 
The specimens are currently retained in the author’s collection. 


MicHaEL R. Jerrorps, Department of Entomology, 320 Morrill Hall, University 
of Illinois, Urbana, Illinois 61801. 


BUTTERFLIES AS PREY FOR CRAB SPIDERS (THOMISIDAE) 


Although crab spiders (Thomisidae) have long been known to capture various 
insect prey while concealed in flowers, there are surprisingly few published obser- 
vations where butterflies are the prey. Hobby (1930), Proc. Ent. Soc. London. 5: 
107-110) cited a 1902 record by E. B. Poulton when a Satyrid butterfly, Melanargia 
lachesis Hiibner, was captured by a crab spider in Spain. In another early record, 
Bell (1921, Bull. Brooklyn Ent. Soc. 16: 96-97) mentioned that Epargyreus tityrus 
Fabricius was captured by a crab spider, Misumena vatia (Clerck). Some years 
later, Voss (1953, Lepid. News. 7: 54) recorded that Ancyloxpha numitor (Fabri- 
cius) was similarly taken in Michigan, and Libythea bachmanii Kirtland and Polites 
mystic (Scudder) in Pennsylvania. None of the crab spiders were identified. 
Conway (1954, Lepid. News. 8: 28) reported the seizure of a Papilio glaucus 
Linnaeus by a crab spider in Illinois. Nielson (1958, Lepid. News. 12: 202) stated 
that a Lycaena phlaeas americana (Harris) escaped after being grasped temporarily 
by a crab spider in Michigan. In a more recent account, Jennings & Toliver (1976, 
J. Lepid. Soc. 30: 236-237) discussed capture of the pine butterfly, Neophasia 
menapia (Felder & Felder) in South Dakota by an immature crab spider, Misu- 
menops sp. 

It is of interest to record additional observations of butterflies which were at- 
tacked and captured by crab spiders. The senior author has been interested in this 
behavior, and since 1960 has made numerous observations in which crab spiders 
captured butterflies on various flowers in Maryland. In each case the prey and 
predator were collected and preserved, and the associated plant identified. The 
spiders were identified by the junior author. 

In Table 1 are given the butterflies captured by two species of crab spiders, 
Misumenoides formosipes (Walckenaer) or Misumenops celer (Hentz), along with 
collection information. The data indicate that the former species was found with 
the butterfly prey more often than the latter. Most of the butterflies preyed on 
were Hesperiidae, although much larger Papilionidae were also taken. Single species 
of Lycaenidae and Nymphalidae were captured. A total of 11 different species of 
butterflies were prey for these crab spiders; and 9 additional species of butterflies 
are recorded in this note. The captured butterflies varied from badly worn to nearly 
fresh condition. The senior author witnessed the actual attack on a male Wallen- 


(AInIq ) sospy} so1vyy sapoiahiyd 


sen "TT wnuqouups wnuhoody (OATeD ) JUIOg un[g gL ‘sny TE 
N AVGITVHAWAN 
TT DuDLUdU 1140911.) (q1OATeD ) JUIOg AzZoo1g 09 ‘sny CZ al omdeys apiupziz 40J1a sauvo0g 
AVGIMaAdS ap 
(zyUd}{ ) 1aj7a9 sdouawnsiyy Aq poinjderd 
‘TT DJOLDI snonvdqg (JIOATeD ) UMOJSUTVUN FT €2 349g 6I A (ArnIq, ) so“py} so1pyy saporohyg 
AVGITVHAWAN 
(‘]) vawisua pzapadsaT (sopteyD ) pIpousg 9g) ‘3dag 8 A (sliey]) ymuny snuyau uowhys 
AVGINAVOA'T 
(‘T]) asuaawp wmisi1D (sopreyD ) AD JUoxnjzeg QL 149g FZ A snevuuryT snonpjs snonnjs oyidog 
("]) asuaaip wnisiiD (sopreyD ) AWD JUoxnzeg 9) 3deg F W snevuury snonpjs snonnjs oyrdog 
AVGINOITIdV 
("[) asuaaiw wis.) (soprey) AD JUoxnzeg gL ‘31dag 9 A (toureID ) snw79 sninja snaihsindy 
‘TT wnuijzsajao9 wniwoyodny (sopreyD ) 910}g JUING gL ‘ideas 9 W (tourely ) snipj9 sninjo snaihsindy 
(]) asuaaip wnisi1D (soprey) AWD JUoxnzeg gL ‘3d9Sg F W (19urIeID ) snipj9 sninja snaihdindy 
TT wnyofidosshy wnitojodny (q19ATeD ) JUIOG wm] Gy ‘sny OF A (1oWIeID ) snwj9 snunja snaihaivdy 
(‘T) vyofmmunis osppyos (qroaTeD ) yuIOg win{ G) “3ny OT | (19uIvID ) snij9 sninja snashaindy 
‘TT DUDLUDU DLLOWY OD (qOATVD ) JUIOg Azoo1g 99 3d9S [TT A (1oureITD ) snivja sninja snashaivdy 
T wnaindind wmuioj,vdny (sojteyD ) UMoyURAIG eg ‘sny [TE — (19WeID ) snwjd snupj9 snaihaivdy 
‘ds pajppng (AoATeD ) JUIOgG Azoo1g coi) Maile Key (a) W ( y{IUIS ) sn100D DUWaLa'T 
TT wnoavupajhisuad wnuoshjog (}OATeD ) JULIO umnyg 6) ‘8s0ny QZ WW (snioniqey ) sozuunu pydhixojhouy 
(TT) asuaawp wns) (sopreyD ) yoIpeueg 9), ‘1d9Sg 2 W (a][ferye'T ) sajaojstwmay? saqyog 
‘TT SLIDS)NA D]JaUNI (sos10e4 90ULIg ) O[[IAS}og 69 ‘SV 6I W (1owleBIDn ) sp109 saqyog 
(7) vyofuuupis osppyos (SAIBJI “3S ) S9OKTUT “75 gL snY 7 W ( YILUUIS ) OY7,0 DIUaIsUAaT]D MA 
<H TT wnyofidosshy wnitojodny (sArey ‘3S ) FLOYoo'T zULOg 7L‘s0V 16 W (YIUIS ) OY70 DIUaLBUAT]D MM 
wc "TT wnaindind wmuojodn |] (jepunry ouuy ) UsARTy Arey gL ‘39S T W ([eAnpsiog ) sujsaduipo sapado)pjiy 
A ‘ds sajadv (}10ATeD ) JuIOg win[g cay) alley Gap TIN ([eAnpsiog ) siysadupo sapadojpiy 
2 ‘TT wnutysaja09 wni..ojodn y (jepunsy cuuy ) WoARFy Arey G) 309g ST al ([eAnpsiog ) s14sadup9 sapado)piy 
y "TT asuaqyoid wnyofis T, (WoATeD ) yUIOg win] g PL 3desg OT A ([eAnpsiog ) sijsadup9 sapado)pjiVy 
‘TT asuazoid wnyosis J, (WoATeD ) yUIOg winyg PL ies OL | ({eAnpsiog ) sijsadwpo sapadojpiy 
mal "TT wnaindind wmiozodn 7 (jJepuniy ouuy ) WoARFT Arey ZL ‘3ydag 9 W ([BAnpsiog ) sijsadwupo sapado)piy 
AVCIIMaAdS AP] 
: (rovuayoTeAA ) Sadisowsof{ saprouawnsipy Aq poinjder 
a SOFT TOMO (AyunOF) UMOT, 97eq x9S se1oedg 
ie ‘purpAieyy ul stopids qeto Aq poinjdeo soljieyng ‘[ ATV, 


282 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


grenia otho (Smith) on 21 August 74 which happened in an instant. The Wallen- 
grenia otho dated 22, August 76 was collected by Jonathan P. Haliscak. 


Joun H. Fass, 2809 Ridge Road, Neeld Estate, Huntingtown, Maryland 20639. 
DANIEL T. JENNINGS, 214 Main Street, Orono, Maine 04473. 


OVIPOSITION BEHAVIOR OF COLONIZED CALLOSAMIA PROMETHEA 
(SATURNIIDAE ) 


In maintaining colonies of Callosamia promethea (Drury) we use paper bags to 
collect eggs from fertile females. Just prior to the hatching of eggs, we insert food 
plant cuttings into the bags and leave them there until the newly-hatched larvae 
have migrated and started to feed (Miller & Cooper, 1977, J. Lepid. Soc. 31: 146— 
147). This procedure involves only minimal handling of eggs and larvae and can 
be used with maximum effectiveness when a large number of eggs is deposited in 
a single bag over a short period of time. This minimizes the time food plants must 
be kept in the bags and provides groups of larvae of the same approximate age. To 
determine the optimum period for collecting eggs, we conducted a detailed study 
of the oviposition behavior of C. promethea females from our colony. This paper 
reports the oviposition profile we have elucidated for this species. 

The adult moths used in the study were second generation individuals from a 
colony maintained in Frederick Co., Maryland. Pupae of both sexes were held in 4 
large indoor emergence cages. As the adults emerged, the females either mated in 
the emergence cages, or were transferred to portable outdoor cages (Miller & 
Cooper, 1976, J. Lepid. Soc., 30: 95-104) to attract males. Only females that 
mated on the day of emergence or the following day were used. The mated moths 
were observed closely so that the females could be transferred to oviposition bags 
as soon as the pairs separated. Each female moth was held in a bag overnight and 
transferred to a new bag each morning until death. After a period of time sufficient 
to allow all eggs to hatch, each bag was opened to record the total number of eggs 
and the total number of larvae hatched. 


Taste 1. Summary of oviposition data obtained from colonized Callosamia pro- 
methea females. 


Night Eggs Deposited 

After Number a 

Mating Females Number Cumulative % % Hatch 
] 28 1695 35.5 94.1 
2 28 1577 68.5 94.4 
oS 28 689 83.0 88.8 
4 28 Sie) 90.8 75.0 
5 28 27, 95.2 83.9 
6 Ue) 153 98.4 Wall 
o 16 60 Sou fillies) 
8 7 7 99.8 30.0 
9 4 6) 100 0 
10 1 0 100 0 


VoLuME 31, NUMBER 4 283 


60 


[] eecs vePositep 


PER FEMALE 


| LARVAE PRODUCED 


40 


20 


MEAN NUMBER OF EGGS OR LARVAE 


NIGHT AFTER MATING 


Fig. 1. Oviposition profile for colonized Callosamia promethea females. 


Table 1 summarizes the data we obtained for 28 C. promethea females. These 
individuals deposited a total of 4,771 eggs over a 10-day period. All females sur- 
vived for at least 5 days after mating; one individual lived for 10 days. The average 
longevity after mating was 6.8 days. The maximum number of eggs deposited by 
a single female that lived for 6 days was 262; the minimum was 80 eggs for a 
female that lived for 5 days. The average number of eggs deposited per female 
was 170.4. Data on percent hatch were variable, but the general trend was one of 
decreasing hatchability with increasing time after mating of the female. Percent 
hatch did not decrease markedly until after the 7th night. 

To determine an optimum period for collecting eggs, we considered our findings 
in terms of the average number of eggs (or larvae) produced by a female moth 
(Fig. 1). Collecting eggs beyond the 3rd night after mating would be of decreasing 
efficiency for the effort involved because, although the percent hatch remains rela- 
tively high through the 7th night, the actual number of eggs deposited beyond the 
3rd night is small. We have concluded from this study that the optimum period 
for collecting eggs under the conditions of our colony is during the first 3 nights 
after mating. 


Tuomas A. Minter & WiLLIAM J. Cooper, U.S. Army Medical Bioengineering 
Research and Development Laboratory, Fort Detrick, Frederick, Maryland 21701. 
(This research was not supported by government funds; the opinions contained 
herein are those of the authors and should not be construed as official or reflecting 
the views of the Department of the Army.) 


284 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


BOOK REVIEWS 


Lipro Rojo DE Los LEPIDOPTEROS IBERICOS, by M. G. de Viedma and M. R. Gomez 
Bustillo. 1976. Instituto Nacional Para la Conservacion de la Naturaleza, Madrid, 
Spain, 120 p., 82 figures (color photographs), 50 maps. Paper. Price $5.50 (U.S.). 
Available from SHILAP, Apartado no. 331, Madrid, Spain. 


This small paperback represents an entomological “first.” Patterned after the 
International Union for the Conservation of Nature’s (I.U.C.N.) Red Data Books 
on World endangered species, the authors present status summaries for 50 species of 
Iberian Lepidoptera. Each species is treated as Endangered (5 species), Vulnerable 
(4 species), Rare (20 species), Endemic (16 species), or Migratory (5 species). 
Butterflies receive most coverage, but 13 families are covered as follows: Hesperiidae 
(5 species), Lycaenidae (14 species), Satyridae (9 species), Nymphalidae (1 spe- 
cies), Papilionidae (3 species), Epipyropidae (1 species), Zygaenidae (4 species), 
Saturniidae (3 species), Endromididae (1 species), and Sphingidae (2 species). 
With the exception of a summary page in English, the book is entirely in Spanish, 
as one would expect. 

The treatment for each species includes sections on distribution and flight period, 
habitat and life cycle, status and population, and recommendations. 

The intent of the book is to be a preliminary assessment of Iberian Lepidoptera in 
need of some conservation effort or in need of continuing scrutiny. 

Apparently, the status of Iberian Lepidoptera lags behind that of the remainder of 
Western Europe, since only five somewhat unsuitable species are cited as En- 
dangered: Coenonympha oedippus (Satyridae) is thought to be extirpated in Iberia, 
but is widespread in other portions of Europe; Samia cynthia and Antheraea pernyi 
(both Saturniidae) are both Asiatic species brought to Europe by man in the 1800's 
for their possible silk-producing qualities. Epipyrops schawerdae (Epipyropidae) 
may be endemic, but is known only from a single individual, and represents an 
esoteric family not regularly sampled by collectors; Procris predotae (Zygaenidae), 
a Spanish endemic, is the most logical candidate of the five, but still remains little 
investigated. 

Documentation or even mention of threats to the habitats of the fifty species is 
largely lacking in the book. Under provisions of the United States’ Endangered 
Species Act of 1973 most of these species could not be listed because of the lack of 
appropriate documentation of threats to their habitat. 

It is heartwarming to see that the Spanish have begun to assess the status of their 
endemic Lepidoptera with an eye fixed toward future conservation efforts. Let us 
hope that the future efforts include further field study of potential candidates, and 
more importantly, acquisition, protection and/or management of the habitats upon 
which the more unique taxa depend. 

I recommend this book to all members with conservation interests and an aptitude 
for reading Spanish. 


Paut A. Oper, Office of Endangered Species, U.S. Fish and Wildlife Service, 
Washington, D.C. 20240. 


THE CLASSIFICATION OF EUROPEAN BUTTERFLIES, by L. G. Higgins. 1975. 320 p., 
402 line drawings. Wm. Collins and Sons, Ltd., Glasgow. Price: £7.50 net. 


Within the pages of this book are the justifications for a radically different classi- 
fication of the European butterflies, one that was long overdue. The data that are 
included in the present book were originally to have been incorporated into the 


VOLUME 31, NuMBER 4 285 


Higgins and Riley Field Guide to the Butterflies of Britain and Europe, but 
were not. In some respects it is unfortunate that this information was not placed 
in the Field Guide; the amateur would have been exposed to it and had the 
opportunity to make use of it or not as he saw fit. The present volume is more 
likely to have a limited audience and will most likely be purchased by the profes- 
sional lepidopterist and the advanced amateur only. The idea of placing a great 
deal of technical detail in basically an identification book is not new—Godman and 
Salvin interwove beautifully colored plates with illustrations of structural details 
very successfully in the Rhopalocera section of the Biologia Centrali-Americana, a 
work that has remained a standard reference for the last three-quarters of a century. 
The combination of these details in the Field Guide do the same thing for the 
European species, and the serious student cannot afford to be without it. 

The illustrations, none of which is colored, are highly informative, but not over- 
done. The reader searching for pretty pictures will be disappointed, unless he is 
impressed by pictures of structural details. These details are clearly illustrated and 
provide much of the basis for the “new systematics” of the European butterflies. 
With the illustrations at hand it would be difficult for a worker to seriously question 
the conclusions reached. 

The errors are few and far between. Most of the literature citations given are 
correct (I know I learned a great deal about who first proposed what higher cate- 
gory), and the consistency with which the oldest name is used for family-groups is 
laudable. The only exceptions to this are the use of Riodinidae instead of the 
older Nemeobiidae (there is an appeal pending before the International Commission 
on Zoological Nomenclature for the validation of Riodinidae) and the Pierinae is 
credited to Swainson in 1840 while the Pieridae is attributed to Duponchel, 1832. 
The “Code” is specific in its assertion that when one family-group name is proposed, 
all other family-group names on the same stem are proposed by implication. I 
suspect that the Pieridae-Pierinae situation was merely an oversight. Other errors of 
omission or commission largely may be ascribed to Dr. Higgins not being familiar 
with some of the work done on this side of the Atlantic, but American authors have 
been far less cognizant of what has been done in Europe, so Higgins’ omissions 
may be forgiven in this regard. 

On balance, this book is a magnificent piece of workmanship and of scholarship, 
perhaps the best that has been done on Palearctic butterflies in recent years. No 
thopalocerist who is more than a hobbyist can afford to be without it. Now the 
gauntlet has been flung—we Americans need to do a similar work on the Nearctic 
butterfly fauna! 


Lee D. Mitter, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, 
Florida 33580. 


286 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


INDEX TO VOLUME 31 


(New names in boldface ) 


aberrations, 99, 223 
Adams, J. W., 150 
Anaea aidea, 280 
Antheraea polyphemus, 166 
Apamea inebriata, 59 
Aricia artaxerxes vandalica, 223 
Asterocampa clyton texana, 172 
leilia, 111 
Austin, G. T., 111 
behavior, 35, 100, 111, 153, 172, 190, 
VS, QA. DS, BAW 
Biblis hyperia, 166 
biology, 17, 81, 100, 173, 232 
bionomics, 183 
Boloria selene, 243 
s. albequina, 255 
. atrocostalis, 258 
. myrina, 245 
nebraskensis, 248 
. sabulocollis, 252 
. terranovae, 262 
s. tollandensis, 253 
book reviews, 78, 79, 151, 215, 284, 
285 
Brassolidae, 203 
Brower, L. P., 232 
Bungalotis milleri, 97 
Callaghan, C. J., 173 
Callosamia, 23 
promethea, 146, 148, 282 
Calvert, W. H., 232 
Carmenta odda, 195 
Catocala, 1, 197 
Charaxes, 276 
Chlosyne lacinia, 35 
Choristoneura, 119 
Christian, J., 232 
Clarke, C. A., 139 
Cocytius duponchel, 34 
Codatractus yuecatanus, 96 
Colias cesonia, 22 
collecting, 197, 276, 278 
Cooper, W. J., 146, 147, 283 
Cudonigeria, 119 
houstonana, 119 
Danaidae, 232 
Danaus plexippus, 232 
dispersion, 202 
distribution, 16, 17, 34, 41, 61, 67, 68, 
10, 1385)9 1388) 143s GG Gre elon 
197, 248 269 27 Ge oo 


NDNnnnNDnDNHA NR 


Dominick, R. B., 109 
Duckworth, W. D., 191 
Eichlin, T. D., 191 
Fish same leeemla, 
Eoparagyractis plevie, 81 
Epargyreus deleoni, 91 
Erynnis tristus tatius, 99 
Eulithis molliculata, 269 
evolution, 124 
Fales, J. H., 282 
Ferguson, D. C., 57 
Ferris, C. D., 99, 279 
Fiance, S. B., 81 
Freeman, H. A., 89 
Garland, J. A., 71 
genetics, 139 
genitalia, 31, 45, 46) 48; 50} aI 52, 53; 
54, 60, 65, 94, 118, 120, 136, 144, 
169, 194, 195 
Geometridae, 71, 269 
Gillaspy, J. E., 166 
Glyphipterigidae, 124 
Godfrey, G. L., 79 
Gomez, L. D., 204 
Hansen, Ay i 225 
Hedrick, L. E., 232 
Heppner, J. B., 124 
Heran, Rojas 
Hesperiidae, 17, 70, 89, 99, 138, 278, 
281 
hibernation, 71 
Highfill, J. W., 146 
Hodges, R. W., 110 
Hgegh-Guldberg, O., 223 
Hyalophora cecropia, 153 
hybridization, 23 
Hypolimnas bolina, 139 
Immidae, 129 
Itame abruptata, 269 
Jeffords, M. R., 280 
Jennings, D. T., 282 
Johnson, K., 167 
Khalaf, K. T., 148 
Kohler, S., 243 
larval foodplants, 17, 35, 41, 81, 100, 
isisy, Wye lisse, WAGE) 
Libytheidae, 279 
life histories, 81, 100, 173 
Lycaenidae, 223, 279, 281 
McGuffin, W. C., 269 
Megalopyge opercularis, 147 


RL, PD mine ap 


VoLUME 31, NUMBER 4 


Megalopygidae, 147 
Melitaea saxatilis, 16 
Menander felsina, 173 
migration, 232 

Miljanowski, E. S., 76 

Miller, L. D., 285 

Milles ©. A. 146, 147, 283 
Miller, W. A., 202 

Miller, W. E., 118, 135 
Moeller, R. E., 81 

Munroe, E., 63 

Napier, A. H., 150 

Necker. We. 22,68, 172,-183 
Nekrutenko, Y. P., 77 

new family, 129 

new genus, 119 


new species, 47, 49, 51, 55, 58, 89, 91, 
95.96: 97, 135, 136, 193, 195 


new subspecies, 167, 252 
Nicolay, S. S., 217 
Nielsen, M. C., 149, 275 
Noctuidae, 1, 57 


notes and news, 73, 182, 222, 274 
Nymphalidae, 16, 35, 111, 139, 166, 


172, 276, 279, 280, 281 


obituaries, 76, 109, 150, 205, 213 


Obraztsov, N. S., 119 
Ochlodes yuma, 17 
Opler, P. A., 41, 284 
Opsiphanes cassiae, 203 
oviposition, 172, 282 
Papilio androgeus, 143 
Papilionidae, 100, 143, 279, 281 
parasites, 147, 183 
Parides iphidamas, 100 
Patterson, A. J., 143 
Peigler, R. S., 23 
Peterson, R. D., II, 277 
Petrova khasiensis, 135 
salweenensis, 136 
phenology, 1, 197 
Pieridae, 22, 68, 167, 202, 279 
Pieris occidentalis, 202 
napi oleracea, 68 
sisymbrii nordini, 167 
Polites sabuleti sabuleti, 70 
Polythrix guatemalaensis, 95 
Powell, J. A., 119 
predation, 22, 148, 183, 280 


287 


presidential address, 217 
Psychidae, 183 
Pyralidae, 41, 63, 81 
Pyrrhopyge hoffmanni, 89 
Ramos, S. J., 216 
Rhodophaea caliginella, 44 

cruza, 47 

durata, 49 

fria, 49 

kofa, 47 

neva, 51 

suavella, 55 

yuba, 55 
Rickard, M. A., 138 
Riodinidae, 173, 279 
Riotte, J. C. E., 152 
Sarsent ta Dae 
Saturniidae, 23, 144, 146, 148, 153, 166, 

282 
Satyridae, 279 
Scarbrough, A. G., 153 
Schweitzer DL Fo 72 
Scoparia huachucalis, 63 
Scott, JavA., 47 
Sesiidae, 191 
Shapiro, A. M., 203 
Sheppard, P. M., 139, 205 
Shields, O., 17 
Shull, E. M., 70 
Sieker, W. E., 80 
Sphingidae, 34, 275 
Sphinx luscitiosa, 275 
Stamp, N., 35 
Sternburg, J. G., 153 
Synanthedon kathyae, 193 
Systematics, 41 57, 63;"89> T19) 135, 
WG. MN Bas 

atreshiy fo tin 1G 
techniques, 144, 146 
Tilden, J. W., 214 
Tortricidae, 135 
Turner, J. RK: G., 212 
Auittles |. Ps 134. 
Typhedanus salas, 95 
Urbanus simplicius, 138 
Waldbauer, G. P., 153 
weather, 67 
Young, A. M., 100, 190 


288 


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use 


CONTENTS 


PRESIDENTIAL ApprREss 1977: Tuirry YEARS AND COUNTING. 
S.'S.. Nicolay 200 Ee a Ee a 


PHENOTYPIC WING PATTERN MODIFICATION BY VERY BRIEF PERIODS 
oF CHILLING OF PUPATING ARICIA ARTAXERXES VANDALICA (Ly- 
CAENIDAE). ARICIA Stupies No. 16. Ove Hgegh-Guldberg and 
Arne Lindebo Hansen 0) 


BIOLOGICAL OBSERVATIONS ON AN OVERWINTERING COLONY OF MON- 
ARCH BUTTERFLIES (DANAUS PLEXIPPUS, DANAIDAE) IN MEXICO. 
Lincoln P. Brower, William H. Calvert, Lee E. Hedrick, and 
John: Christian 2 


Revision OF NortH AMERICAN BOLORIA SELENE (NYMPHALIDAE) 
Wirn Description oF A New Susspecies. Steve Kohler 


A Stupy OF THE DISTRIBUTIONS OF PHYSOCARPUS OPULIFOLIUS AND 
Two GEOMETRIDS FEEDING ON IT. W. C. McGuffin —.-____ 


GENERAL NOTES 


Sphinx luscitiosa (Sphingidae ) feeding on decayed fish. M. C. Nielsen 
Charaxes species (Nymphalidae) from Engu, Anambra State, Nigeria. 
RD: Peterson Li oo Ne 
Supplement to “A checklist of the butterflies of Grant County, New Mex- 
ico and vicinity.” Clifford D) Ferris 0.) eae 
A record of Anaea aidea (Nymphalidae) from southern Illinois. Michael 
Reifeffords) (0g Ge 8 y's on RI Oe TUN SS 
Butterflies as prey for crab spiders (Thomisidae ). John H. Fales and 
Daniel 'T. Jennings 8 oo 
Oviposition behavior of colonized Callosamia promethea (Saturniidae). . 
Thomas A. Miller and William J. Cooper 


NOTES) AND) NEWS (C2 OUI B RS NU aii 222) 
BOOK: REVIEWS 00 Pie onl ei Ns EN, 284, 


InvExX TO VOLUME! 31) oo) s00> 05000 ie OS go 


223 


232 


243 


269 


275 
276 
278 
280 
280 


282 
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