

ys teeny ai had yt ea ee 
t a) sha 48 : uae, CH RVR AEE eee ’ : “fe 
; a 4 pad ded eae 8 a a aah HAM sa Cy eae i t atta Fs 
’ j 4 : a he Aaa sb Pa? wai Ppt 1 By tae “i. 
yd daa tiaens hate ee glade prahes ties gh athet yp Mal d beh Otg: bea Rar aa ier 
Pauli “eae has Yet Pie eed} Sema AU 
‘ " ‘8 Sha anity', eae : ahh 
‘ge. ‘ . 
: wy 
A f 
aa +e" 
eee ed 
a 4 
‘ 44 ah 
a sa 2 
, . : 4 
La . 
ede bee ae 
“ea ee 
> we : 
1 4 7 
. ‘ . 
x be es SE feast 
ae ae 
Pos EERE VA De BR 
at ats Se ee Oe Be 
‘ { ARS ie tt 
; a! * , 
: ' ‘ 3: 
2 va 
vu 
se } y 
, . Sst eerie fee ke 
' , ee 
, Te) 
A en 
, : rie tH oe 
ew Wa ! 
> 
ai 
4 , 
: e 
oie t ¢ : 4 
4 Gans 


: te sates 


bo wpetyts4- 2-53 
Zpe koi K BBs hs: 


* 


: 


i : 
bey 
eat 


ok atte ip 
gee A 


Raw neg 
bate pias 
, Se Ser 


eve Sey me £94 
ti aye 


Ri) pee) Ai, re es ee OD 
eR) SM, iat ‘ 


Volume 26 Vi AG~A7 1979 Number 1 
Ent, 


: JOURNAL, 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
| Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 


21 March 1972 


THE LEPIDOPTERISTS’ SOCIETY 
EXECUTIVE COUNCIL 


Lioyp M. Martin (Prescott, Ariz.) President 

J. F. Gates CLarke (Washington, D.C.) President-elect 
S. A. Ag (Nagoya, Japan) Ist Vice President 

Kerry S. Brown (Rio de Janeiro, Brasil) Vice President 
H. A. FREEMAN (Garland, Texas) Vice President 

S. S. Nico.ay (Virginia Beach, Va.) Treasurer 

LEE D. Miter (Sarasota, Fla.) Secretary 


Members at large (three year term): R. B. Dominick (McClellanville, S.C.) 

B. MATHER (Clinton, Miss.) 1972 1973 

M. Ocara (Osaka, Japan) 1972 J. P. Donanve (Los Angeles, Calif.) 1973 

E. C. Wexuinc (Merida, Mexico ) 1972 J. M. Burns (Cambridge, Mass.) 1974 

ANDRE BLANCHARD (Houston, Texas ) R. H. Carcasson ( Vancouver, B.C.) 1974 
1973 M. C. Nretson (Lansing, Mich.) 1974 


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 eight numbers of the News each year. 


Active members—annual dues $10.00 
Student members—annual dues $5.00 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $15.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia 23455. 


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


by Cyai F. pos Passos 


Price, postpaid: Society members—$5.00, others—$7.50; uncut, unbound signatures 
available for interleaving and private binding, same prices; hard cover bound, mem- 
bers—$8.00, others—$10.00. Revised lists of the Melitaeinae and Lycaenidae will be 


distributed to purchasers free (separately with paper covered copies and unbound 
signatures, bound in with hard covered copies) 


The Lepidopterists’ Society is a non-profit, scientific organization. The office of 
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520. 
Second class postage paid at Lawrence, Kansas, U.S.A. 66044. 


JOURNAL OF 


Tue LerrporprTreRIstTs’ SOCIETY 


Volume 26 1972 Number 1 


NOTES ON THE BALANOTES (MEYRICK) GROUP OF 
OIDAEMATOPHORUS WALLENGREN WITH DESCRIPTION 
OF A NEW SPECIES (PTEROPHORIDAE) 


Everetr D. CAsHATT 
Illinois State Museum, Springfield, Illinois 


The separation of Oidaematophorus balanotes (Meyrick), O. grandis 
(Fish), O. lacteodactylus (Chambers) and O. kellicottii (Fish) has been 
difficult. Barnes and Lindsey (1921) used alar expanse and maculation, 
length of palpus and certain genitalic characters to distinguish these 
species. The accumulation of additional material, however, reveals that 
all of the above characteristics are subject to great variation within species. 
To more clearly define the taxa of this complex, an intensive study of the 
genitalia of both sexes was made. For males consistent differences 
between species were found in a secondary structure (hereinafter referred 
to as the “clasper”’) on the inner surface of the left valva. Differences in 
the anterior margins of the eighth tergites and in the configurations of 
the anterior apophyses aid in separating the females. 

Non-genitalic characters were unreliable. Because it was difficult to 
associate males with females, the sexes were associated chiefly by lo- 
calities. Further studies will include rearing larvae to confirm the rela- 
tionships of the sexes. 


Oidaematophorus balanotes (Meyrick) 
(Figures 1, 6, 9; Map 1) 


Pterophorus balanotes Meyrick 1908: 503. 
Pterophorus aquila Meyrick 1908: 503. 
Oidaematophorus balanotes, Barnes and Lindsey 1921: 429. 


Alar expanse: 31—42 mm. 

Head: Scales brown, brownish white between antennal bases. Antenna brownish 
white. Labial palpus brownish white with brown tips, slender and erect, extending 
beyond antennal base. 

Thorax: Scales brownish white. Forewing scales brownish white, an indistinct 
brown dash extending from base to near cleft; usually one or two small dark brown 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


= AM 


a \! \ \ \\ 
\N\ 


‘\ NW AT 


VoLuME 26, NuMBER 1 3 


spots proximad to cleft; tips of some or all veins dark brown (photograph in Barnes 
and Lindsey 1921: Pl. XLIV, Fig. 12). Hindwing brownish white to pale brown. 
Legs brownish white; foreleg and midleg brown mediad, midtibia more heavily scaled 
than other tibiae. 

Abdomen: Scales brownish white with indistinct brown longitudinal lines on tergum, 
pleuron and sternum. 

Male genitalia (Fig. 1): Tegumen triangulate. Uncus attenuate, curved ventrad. 
Vinculum narrow, arched ventrad. Valvae asymmetrical with hair-pencil tufts on 
outer surfaces; left valva longer and wider, apex pointed, inner surface with a long, 
curved, finely drawn clasper. Juxta well developed, directed toward right side, right 
arm extending beyond left. Aedeagus slightly curved, coecum well developed. 

Female genitalia (Figs. 6, 9): Pouch for sex pheromone gland prominent (everted 
in drawing). Posterior apophysis long and slender. Anterior apophysis short, curved 
posteriad and ventrad. Tergite VIII with lateral margin excavate posteriad to anterior 
apophysis. Ostium bursae opening to left posterior margin of sternite VIII. Ductus 
bursae and corpus bursae membranous. Inception of ductus seminalis at anterior of 
corpus bursae, appendix bursae well developed. 

Type data: Holotype, male, Titusville, Florida, August, genitalia slide 10183, in 
collection of British Museum (Natural History); P. aquila Meyrick, holotype, female, 
Texas, in collection of British Museum (Natural History ). 

Food plant: Specimens in the U.S. National Museum Collection were reared from 
larvae which were boring in the stems of Baccharis sp. and Myrica sp. 

Specimens examined (40¢ ¢, 382 9; Map 1): Unitrep States, Arizona: Babo- 
quivari Mtns., Pima Co., 12, Aug. 1-15, 1933, O. C. Poling (USNM); same locality, 
4° 9, Aug. 15-30, 1923, O. C. Poling (USNM); Madera Canyon, Santa Rita Mtns., 
4880 ft., 26 ¢, Aug. 1-4, 1959, R. W. Hodges (USNM); Mohave Co., 2¢ 6, Aug. 
8-15 (USNM); Santa Catalina Mtns., Pinal Co., 26 ¢ Aug. 1-7 (USNM); same 
locality, 1¢, no date (USNM). Florida: Archbold Biological Station, Lake Placid, 
1°, May 1-7, 1964, R. W. Hodges (USNM); Lauderdale, 12, Feb. 11, 1923, D. M. 
Bates (USNM); St. Petersburg, 16, Jan. 31, 1951, R. Ludwig (USNM); same 
locality, 2¢ 6, April 24, 1914, R. Ludwig (USNM), same locality 1¢, Dec. 30, 1914, 
R. Ludwig (USNM); same locality, 12, Feb. 8-15 (USNM); same locality, 19, 
March 1-7 (USNM); same locality, 12, Oct. (USNM); same locality, 34 ¢, 32 9, 
no date (USNM); Siesta Key, Sarasota Co., 5 ¢ ¢, Jan. 17-20, 1951, C. P. Kimball 
(CPK); same locality, 12, June 6, 1957, C. P. Kimball (CPK); same locality, 19, 
Feb. 14, 1956, C. P. Kimball (CPK); Titusville, 1¢, Aug. (BMNH); Vero Beach, 
14, Feb., 1914, J. R. Malloch (USNM). Maryland: Highway 50 at South River, 1 ¢, 
Aug. 19, 1939, stem borer ex. Baccharis sp., J. F. G. Clarke (USNM); Lloyds, Dor- 
chester Co., 16, 19, July 10, 1907, H. S. Barber (USNM). Mississippi: Landon, 
19, Aug. 7, 1921, Larva in Myrica sp., L. E. Miles (USNM). South Carolina: Bluff- 
ton 19, Oct. 14, 1887 (USNM); Charleston, 192, July 9, 1898 (USNM). Texas: 
Brownsville, 12, June, F. H. Snow (USNM); same locality, 29 2, July 11, G. Dorner 
(USNM); same locality, 1¢, 29 2, ex. marsh willow, lot #44-27928; same locality, 
19, Nov. 14, 1928, F. H. Benjamin (USNM); same locality, 13, Nov. 18, 1927, F. H. 
Benjamin (USNM); same locality, 1¢, 329, Nov. 27, 1928, F. H. Benjamin 
(USNM); same locality, 6¢ 6, 42 2, no date (USNM); San Benito, 3¢ 6, 19, 
July 16-23 (USNM); same locality, 1¢, 22 9, Aug. (USNM); same locality, 2¢ ¢, 
Sept. 8-15 (USNM). 


< 


Figs. 1-2. 1, Male genitalia of Oidaematophorus balanotes (Meyrick), ventral 
view, aedeagus removed; 2, male genitalia of Oidaematophorus grandis (Fish), 
ventral view, aedeagus removed. (Scales = 1 mm.) 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


a Wwods 


[ SZ JANLIDNON 193M 08 se 06 ool sol oll Sil 
e N ‘ > l 1 IS z ; 
po Se =. b; cy * 
r ¢ st WT NOILDarOYd SYasIVv N ‘ : 
Se £ ae Saldads Mau ‘luua|d OQ @ 
2 SY83L3NO1IM 009 oor 002 ° ~ LS * 
: a 
SZ saw oor Ooe 002 *OO01 ° 


(siaquieyg) SnjAjoepoajoe| ‘0 X 


(YS!4) Iy}09I1]34 0 ¥ 
(ysi4) Sipueis Q oO 


VoLUME 26, NuMBER 1 5 


Because of its larger size, this species is not often confused with other 
related species in the southeastern United States. A West Coast species, 
O. grandis (Fish), is nearly as large, but slight genitalic differences and 
its apparent restriction to western California are sufficient to separate the 
two species. 

Wing maculation and the clasper of the male genitalia are extremely 
variable. Specimens from the same locality and collected on the same 
date may have all, a few, or none of the forewing vein tips marked with 
dark spots. The forewing cleft is typically marked with one or two dark 
spots, but the spots are absent on some specimens. The clasper of the left 
valva is usually more curved at the base than it is at the middle, but the 
curvature of the clasper is more variable in O. balanotes than in the other 
species studied. The clasper of the holotype is acutely curved near the 
middle but seems to be within the range of the species. The tip of the 
right valva is more acutely pointed than in the other related species 
studied. 


Oidaematophorus grandis ( Fish) 
(Figures 2, 10; Map 1) 


Lioptilus grandis Fish 1881: 141. 

Alucita grandis, Fernald, in Smith 1891: 87. 

Pterophorus grandis, Fernald 1898: 50. 

Pterophorus baccharides Grinnell 1908: 317. 
Oidaematophorus grandis, Barnes and Lindsey 1921: 430. 


Alar expanse: 30-34 mm. 

Head: Scale coloration and palpus as in O. balanotes. 

Thorax: Scales brownish white. Forewing brownish white with faded dark spots at 
tips of veins; cleft spot pale or absent (photograph in Barnes and Lindsey 1921: PI. 
XVII, Fig. 7). Hindwing pale brownish white to grayish white. Legs identical to 
those of O. balanotes. 

Abdomen: Scale coloration as in O. balanotes. 

Male genitalia (Fig. 2): Much like O. balanotes except clasper on left valva more 
acutely curved near base and right valva with a slight lobe near middle of hind margin. 

Female genitalia (Fig. 10): Identical to those of O. balanotes except anterior 
apophysis shorter and slightly thicker. 

Type data: I hereby designate a male in the collection of the Museum of Compara- 
tive Zoology as lectotype, labeled: “California, 1782,” genitalia slide EDC 258. Two 
paralectotypes (without abdomens), one male and one broken specimen, from Cali- 
fornia are in the collection of the U.S. National Museum. Grinnell’s two syntypes 
of P. baccharides were not examined. 

Food plant: The larvae bore in stems of Baccharides pilularis DC. Additional notes 
on the immature stages were recorded by F. X. Williams (in Barnes and Lindsey 
1921: 431-432). 


< 


Map 1. Distribution records for Oidaematophorus spp. It is important to note 
that these records perhaps indicate the distribution of collectors of Oidaematophorus 
rather than the actual distribution of the species. 


6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ro 


: 


1 Az 

if). YY y 

Huff 
yf oy 

4 Me SY 
Z 


Figs. 3-5. 3, Left valva of Oidaematophorus kellicottii ( Fish ) 
male genitalia of Oidaematophorus lacteodactylus (Chambers ) 
removed; 5, male genitalia of Oidaematophorus glenni Cash 
aedeagus removed. (Scales = 1 mm.) 


, ventral view; 4, 
, ventral view, aedeagus 
att, n. sp. ventral view, 


VoLUME 26, NUMBER 1 7 


Specimens examined (10¢ 6,222, Map 1): Unirep States, California: Berke- 
ley, 16, June 10, 1931, Don Meadows; Carmen, 2¢ ¢, June, A. A. Vachell; Los 
Angeles Co., 1¢, May, C. V. Riley; Mills College, Alameda Co., 14, 12, June 27-30, 
1908, G. R. Pilate; Monterey Co., 2¢ ¢, June 10, 1908, F. X. Williams; San Felipe 
Valley, San Diego Co., 19, Sept. 5, 1937, Don Meadows; Santa Clara, 2¢ ¢, no date; 
Wheeler Hot Springs, 1 ¢, July 29, 1943, (all USNM). 


Barnes and Lindsey (1921: 432) noted that a specimen (female) of O. 
grandis labeled “Plummer’s Island, Maryland, May” was probably an 
error either in their record or on the label. After dissecting the genitalia 
of this specimen, I find it to be O. glenni described below. 

Notes on the genitalia will aid in separating this western species from 
O. glenni in California. 


Oidaematophorus kellicottii (Fish) 
(Figures 3, 11; Map 1) 


Lioptilus kellicottii Fish 1881: 141. 

Alucita kellicottii, Fernald, in Smith 1891: 87. 

Pterophorus kellicottii, Fernald 1898: 49 

Pterophorus kellicotti (sic), Meyrick 1910: 17. 
Oidaematophorus kellicottii, Barnes and Lindsey 1921: 427. 


Alar expanse: 20-28 mm. 

Head: Scales light brownish white. Labial palpus slender and erect, but not reach- 
ing antennal base. 

Thorax: Scales light brownish white. Forewing scales brownish white with an indis- 
tinct brownish dash extending from base and fading out toward cleft; usually a dark 
spot at base of cleft; tips of some or all veins marked with a small dark brown spot 
(photograph in Barnes and Lindsey 1921: Pl. XLVII, Fig. 5). Hindwing brownish 
white to pale brown. Legs identical to O. balanotes. 

Abdomen: Scale coloration as in O. balanotes. 

Male genitalia (Fig. 3): Similar to those of O. balanotes except smaller; tips of 
valvae more rounded; clasper slightly shorter and directed laterad or mediad, distal 
half straighter, curvature more acute near base and at tip. 

Female genitalia (Fig. 11): Similar to those of O. balanotes except tergite VIII 
with anterior margin rounded; anterior apophysis straight and directed laterad. 

Type data: Lectotype, male, without abdomen, and one female paralectotype, 
Buffalo, New York, no date, in U.S. National Museum. 

Food plant: The larva bores into the stems of Solidago. The food habits were 
studied by Kellicott (in Barnes and Lindsey 1921: 429). 

Specimens examined (106 6, 72 2; Map 1): Unrrep States, Colorado: Glen- 
wood Springs, 1¢, June 8, 1893, W. Barnes (USNM). Florida: Archbold Biological 
Station, Lake Placid, 1¢, June 1-8, 1964, R. W. Hodges (USNM); St. Petersburg, 
19, May (USNM); Siesta Key, Sarasota Co., 16, Feb. 18, 1956, C. P. Kimball 
(USNM); same locality, 12, May 18, 1963, C. P. Kimball (CPK). Illinois: Putnam 
Co., 16, 22 9, July 7, 1961, M. O. Glenn (MOG); same locality, 1¢, July 16, 1960, 
M. O. Glenn (MOG). Iowa: Sioux City, 19, July 15, 1917, A. Lindsey (USNM). 
Massachusetts: Martha’s Vineyard, 12, July 19, F. M. Jones (USNM). Mississippi: 
Agriculture College, 1¢, July 23, 1920, F. H. Benjamin (USNM). New Jersey: 
Anglesca, 16, May 28, 1905, W. D. Kearfott (USNM); Essex Co. Park, 1¢, Aug. 2, 
1906, W. D. Kearfott (USNM); no data, 16, D. S. Kellicott (USNM). New York: 
no locality, 19, Oct. 16, 1880, Fernald Coll., labeled “type” (USNM). North Caro- 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


—=---- 
=< 


VoLUME 26, NuMBER 1 9 


lina: Tryon, 1¢@, June 2, 1904, Fiske (USNM); same locality, 1¢, Aug. 4, 1904, Fiske 
(USNM). 


The genitalia of this species are nearer those of O. balanotes than those 
of the other species; the differences are described above. Specimens of O. 
kellicottti are generally smaller and have a single distinct spot at the base 
of the cleft. 


Oidaematophorus lacteodactylus (Chambers ) 
(Figures 4, 7, 12; Map 1) 


Pterophorus lacteodactylus Chambers 1873: 72. 

Alucita subochracea Femald, in Smith 1891: 87 (in partim). 
Pterophorus subochraceus, Fernald 1898: 43 (in partim). 
Oidaematophorus lacteodactylus, Barnes and Lindsey 1921: 426. 


Alar expanse: 27-29 mm. 

Head: Scale coloration and palpus as in O. balanotes. 

‘Thorax: Scales brownish white. Forewing brownish white with diffuse light brown 
dash extending from base and fading near cleft; one or two indistinct brown spots at 
base of cleft; tips of some or all veins marked with dark brown (photograph in 
Barnes and Lindsey 1921: Pl. XLVII, Fig. 6). Hindwing brownish white. Legs 
identical to those of O. balanotes. 

Abdomen: Scale coloration as in O. balanotes. 

Male genitalia (Fig. 4): Similar to those of O. kellicottii except distal one-third of 
clasper more broadly curved and not as finely drawn, tip of right valva not sharply 
pointed. 

Female genitalia (Figs. 7, 12): Similar to those of O. kellicottii except anterior 
apophysis shorter, anterior margin of tergite VIII only slightly rounded. 

Type data: Holotype, male, Kentucky, no abdomen, in the collection of the Museum 
of Comparative Zoology. 

Food plant: Dr. J. F. G. Clarke reared specimens which were boring in stems of 
Solidago sp. in Washington. 

Specimens examined (1036 6, 492 2; Map 1): Uwnirep States, Illinois: Putnam 
Co., 22 2, July 8-11, M. O. Glenn (MOG), Kentucky: no further data, 1¢ (MCZ). 
Maryland: Plummer’s Island, 1¢, July 1, 1903, A. Busck (USNM). Washington: 
Almota, 8¢ 6,29 2, May 2-26, reared from Solidago sp., J. F. G. Clarke (USNM); 
Pullman, 1¢, May 15, 1935, J. F. G. Clarke (USNM). 


The identity of this species has been confusing. Barnes and Lindsey’s 
illustration (1921: Pl. LIV, Fig. 2) of the male genitalia matches a slide 
labeled “O. lacteodactylus, 67551” in the U.S. National Museum Collec- 
tion. Since the abdomen of the type specimen is lost, I am unable to 
compare the type’s genitalia with those of the study material. Most speci- 
mens that I have examined have a slight outward bend near the middle of 
the clasper which is not shown in Barnes and Lindsey’s figure. The length 


< 


Figs. 6-7. 6, Female genitalia of Oidaematophorus balanotes (Meyrick), ventral 
view; 7, female genitalia of Oidaematophorus lacteodactylus (Chambers), ventral 
view. (Scale = 1 mm.) 


10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


3 


Figs. 8-13. 8, Female genitalia of Oidaematophorus glenni Cashatt, n. sp., ventral 
view; 9-13, eighth abdominal tergite of female, dorsal view: 9, Oidaematophorus 
halanotes (Meyrick); 10, Oidaematophorus grandis (Fish); 11, Oidaematophorus 
kellicottii (Fish); 12, Oidaematophorus lacteodactylus (Chambers); 13, Oidae- 
matophorus glenni Cashatt, n. sp. (Scale = 1 mm.) 


and width of the valvae are highly variable, but the general shape and 
width of the clasper seem fairly constant. 

McDunnough (1927) was unable to find Lindsey’s slides or any speci- 
mens which resembled his illustration of the male genitalia. Therefore, 
he offered a new figure of the left valva (including the clasper) of a 


VoLUME 26, NUMBER 1 IU 


species that he considered to be O. lacteodactylus. I have examined 
specimens whose genitalia are identical to McDunnough’s figure, and I 
consider them to be new and distinct from O. lacteodactylus. This new 
species, O. glenni, is described below. 


Oidaematophorus glenni Cashatt, new species 
(Figures 5, 8, 13; Map 1) 


Oidaematophorus lacteodactylus: auctorum nec Chambers. 


Alar expanse: 24-34 mm. 

Head: Scale coloration identical to that of O. balanotes. 

Thorax: Forewing brownish white with dark spot at base of cleft, or spot may be 
absent; ends of veins usually marked with dark or faded brown spots. Hindwing 
brownish white. Legs with scale coloration as in O. balanotes. 

Abdomen: Scale coloration brownish white with indistinct pale brown longitudinal 
lines as in O. balanotes. 

- Male genitalia (Fig. 5): Similar to those of O. balanotes except clasper on left 
valva broadly curved, not attenuate, shorter and with tip flattened before terminating 
in a short, curved point. 

Female genitalia (Figs. 8, 13): Similar to those of O. balanotes except anterior 
margin of tergite VIII with a narrow dorsal fold; anterior apophysis short and pointed, 
reinforced by lateral ends of dorsal fold. 

Type data: Holotype, male, Putnam Co., IIl., June 12, 1953, M. O. Glenn, genitalia 
slide EDC 892; in the collection of the Illinois Natural History Survey. Paratypes 
(298 6,219 9; Map 1): California: Big Trees, Santa Cruz Co., 1¢, July 19, 1921, 
Don Meadows, genitalia slide EDC 187 (USNM); Half Moon Bay, 12, June 14, 1937, 
W. H. Lange (USNM). Colorado: no locality, 22 9, Bruce, genitalia slides EDC 
198, 272 (USNM). Illinois: Decatur, 1 ¢, no date, genitalia slide EDC 277 (USNM). 
All from Putnam Co.: 2¢ ¢, May 18, 1965, M. O. Glenn, genitalia slides EDC 894, 
895 (MOG); 146, May 25, 1964, genitalia slide EDC 898 (MOG); 14, June 5, 1956, 
reared from larva in roots of Solidago canadensis, 11675 (USNM); 14, June 7, 1956, 
reared from larva in roots of Solidago canadensis, 11757, genitalia slide 190 (USNM); 
14, June 9, 1955, larva reared on goldenrod (root borer), 11557, genitalia slide 191 
(USNM); 14, June 9, 1956, larva reared from roots of Solidago canadensis, 11457, 
genitalia slide EDC 891 (MOG), 19, June 9, 1963, genitalia slide EDC 897 (MOG); 
1g, Jume 14, 1956 larva in roots of Solidago canadensis, genitalia slide EDC 900 
(MOG); 19, June 14, 1967 (MOG); 14, June 16, 1956, larva in roots of Solidago 
canadensis, genitalia slide EDC 893 (MOG); 19, June 19, 1958, reared from larva 
boring in roots of goldenrod (MOG). Iowa: Homestead, 1 ¢, May 30, genitalia slide 
72-448 (USNM). Maryland: All from Plummer’s Island: 19, May 1, 1906, Aug. 
Busck, genitalia slide EDC 195; 14, June 5, W. V. Warner, genitalia slide EDC 204; 
1@, no date, Aug. Busck, genitalia slide EDC 197 (All USNM). Massachusetts: 
Barnestable, 1¢, June 20, 1954, C. P. Kimball, genitalia slide EDC 292 (CPK). 
New Hampshire: Hampton, 1¢, June 3, 1906, S. A. Shaw, genitalia slide 265 
(USNM). New Jersey: Essex Co. Park, 1¢, June 24, trap, W. D. Kearfott, geni- 
talia slide 72-447 (USNM). New York: Monroe Co., 1¢, June 10, 1947, C. P. 
Kimball, genitalia slide EDC 289 (CPK); same locality and collector, 1¢, June 27, 
1948, genitalia slide EDC 291 (CPK); Newfield, 16, May 27, 1960, R. W. Hodges, 
genitalia slide EDC 906 (USNM); Six Mile Creek, Ithaca, 2¢ ¢, May 29, 1959, R. W. 
Hodges, genitalia slides EDC 905, 909 (USNM). Pennsylvania: Pittsburgh, 1 ¢, May 
30, 1905, Henry Engel, genitalia slide EDC 203 (USNM). Virginia: Montgomery 
Co., 19, May 27, 1898, genitalia slide EDC 207 (USNM). Washington: All from 
Almota, reared from Solidago stalks by J. F. G. Clarke: 19, Jan. 3, 1935, genitalia 


12 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


slide JFGC 396; 12, March 18, 1934, genitalia slide EDC 209; 1¢, March 21, 1934, 
genitalia slide EDC 178; 14, April 9, 1934, genitalia slide JFGC 397; 192, April 10, 
1934, genitalia slide JFGC 394; 19, April 23, 1934, genitalia slide EDC 183; 14, 
May 3, 1935, genitalia slide EDC 181; 19, May 5, 1935, genitalia slide 391; 19, May 
9, 1934, genitalia slide EDC 202; 19, May 13, 1935, genitalia slide JFGC 393; Wawa- 
wai, 2¢ 6, 39 2, Jan. 9, 1935 reared from Solidago stalks, J. F. G. Clarke, genitalia 
slides EDC 179, 180, 182, 184 JFGC 395; same data 1¢@, Jan. 10, 1935, genitalia slide 
200 (All USNM). Wisconsin: Milwaukee Co., 2¢ 6, March 3, 1923, F. P. Breakey, 
genitalia slides EDC 267, W. H. Lange 520. 

Food plant: Dr. J. F. G. Clarke reared larvae which were boring in the stalks of 
Solidago sp. in Washington. Mr. Murray O. Glenn of Illinois collected and reared 
larvae from the roots of Solidago canadensis L. 

I take great pleasure in naming this species after Mr. Murray O. Glenn, 
Henry, Illinois, who through many years of collecting has contributed 
much toward our knowledge of Lepidoptera in Illinois. 

This species was considered to be O. lacteodactylus by McDunnough 
(1927). After examining genitalia slides which match the illustration of 
that species (Barnes and Lindsey 1921), I am convinced this species is 
distinct and separate from O. lacteodactylus and O. kellicottii. The clasper 
of the male genitalia of O. glenni has a flattened tip with a small oblique 
barb whereas the claspers of O. lacteodactylus and O. kellicottii have a 
finely drawn tip. 

In addition to the localities listed in the type series, McDunnough 
(1927) reports the species to be widely distributed in Canada (Alberta, 
Manitoba, Ontario, Saskatchewan ). 


ACKNOWLEDGMENTS 


I wish to thank Dr. R. W. Hodges of the Systematic Entomology Lab- 
oratory, U.S. Department of Agriculture, and Dr. W. D. Duckworth, U.S. 
National Museum (USNM), for their guidance; Dr. L. D. Miller, Allyn 
Museum of Entomology, for reviewing the manuscript; Dr. P. J. Darling- 
ton, Museum of Comparative Zoology (MCZ), and Mr. Paul Whalley, 
British Museum (Natural History) (BMNH) for their cooperation con- 
cerning type specimens, Messrs. M. O. Glenn (MOG), Henry, Illinois, 
and C. P. Kimball (CPK), Sarasota, Florida, for loaning their specimens 
(The letters in parentheses are used in the text to refer to the location of 
specimens examined ). 

This research was supported in part by the Biology Department, Catho- 
lic University of America, Washington, D.C. and the Division of Lepi- 
doptera and Diptera, Entomology Department, Smithsonian Institution, 
United States National Museum, Washington, D.C. 


LITERATURE CITED 


Barnes, W. anp A. W. Linpsey. 1921. The Pterophoridae of America, north of 
Mexico. Contrib. Nat. Hist. Lep. North America. 4: 281-483. 


VoLUME 26, NUMBER 1] 13) 


CHAMBERS, V.T. 1873. Micro-Lepidoptera. Can. Ent. 5: 72-75. 

FERNALD, C. H., in J. B. SMrrH. 1891. List of the Lepidoptera of Boreal America. 
Entomological Society, Philadelphia. 124 pp. 

1898. Pterophoridae of North America. Spec. Bull. Mass. Agri. Coll. 
64 pp. 

FisH, C. 1881. Pterophoridae. Can. Ent. 13: 140-143. 

GRINNELL, F. 1908. Notes on the Pterophoridae or plume-moths of southern 
California with descriptions of new species. Can. Ent. 40: 313-321. 

McDunnoucu, J. 1927. Contribution toward a knowledge of our Canadian plume- 
moths (Lepidoptera). Trans. Roy. Soc. Can. Series 3, Sect. 5, 21: 175-190. 

Meyrick, E. 1908. Notes and descriptions of Pterophoridae and Omeodidae. 
Trans. Ent. Soc. London. (Part 4) 1907: 471-511. 

1910. Lepidoptera Heterocera Family Pterophoridae. In Genera Insec- 

torum. 100: 1-23. 


ANNOTATED LIST OF THE BUTTERFLIES OF 
INDIANA, 1971 


ERNEST M. SHULL 
North Manchester, Indiana 


and 


F. SINEY BADGER 
Woodland Hills, California 


The entomologists at Purdue University, while having adequate funds 
available, necessarily concentrate their efforts and research on the insects 
considered agricultural pests. They are, however, building a world-wide 
collection of butterflies. At Indiana University funding for entomology 
is scarce and spent largely on teaching efforts. Thus the field of Rhopa- 
locera has been wide open and challenging for the serious collector in 
Indiana, as was appreciated by the authors who received much encourage- 
ment and help from members of the Lepidopterists’ Society. Although 
Indiana has had two major annotated state lists of Rhopalocera (Blatch- 
ley, 1891 and Montgomery, 1931), and more recently a list of the butter- 
flies of Perry County (Masters and Masters, 1969), nearly forty years 
have passed without the publication of a comprehensive state-wide list 
of the butterflies (Papilionoidea ) and skippers ( Hesperioidea ). 

The State of Indiana, customarily considered a flat agricultural area, 
in reality combines a number of different and interesting zoogeographic 
regions with natural habitats for a diversified flora and fauna. In short 


14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


these are the northeastern lakes and bogs, the northeastern and central 
plains, the southeastern lowlands and flats, the south central hills, the 
Ohio-Wabash river lowlands, the intrusion of the prairie and the Kankakee 
sand area from Illinois, and the northwestern snow belt extending south 
and west of Lake Michigan and including the typical lake dune country. 
Thus Indiana provides a rich and varied collecting area for entomologists. 

The authors, unknown to each other until 1968 and then only by corre- 
spondence, carried out intensive collecting in the north and central 
counties of Indiana at different periods which did not overlap. Both were 
challenged by the lack of up-to-date records and possibilities. They 
independently projected an annotated list of the Rhopalocera of the state 
with equal emphasis on the Papilionoidea and the more difficult Hesperioi- 
dea neglected either entirely or in part by Blatchley (1891), Montgomery 
(1931), and Masters and Masters (1969). This annotated list covers the 
officially recorded species. It combines the authors’ individual efforts 
and draws freely on previously published papers, as well as information 
from collectors who have penetrated one or two counties deep from ad- 
joining states. It also lists possible species not yet officially recorded for 
which the authors have diligently searched. 

Shull collected sporadically in Indiana from 1932 to 1946, and regularly 
from 1964 to 1971. He collected intensively in Wabash and Kosciusko and 
frequently in Allen, Whitley, Huntington, Miami, Steuben, and LaGrange 
counties. His collection of more than 15,000 specimens contains 107 
species. Badger collected from 1953 to 1962 intensively in Howard and 
Carroll counties and frequently in Fulton, Marshall, Pulaski, Marion, 
Brown and Lake counties. He collected 101 species with a total in excess 
of 10,000 specimens. The combined total of species collected is 118. Many 
other counties, particularly in the northern half of the state, were visited 
for desired species. 

Portions of the state felt to have been inadequately covered are the 
central western prairie areas for the skippers surviving from the western 
prairies, the south central areas from Spring Mill State Park south in- 
cluding French Lick and the southwestern counties adjoining Kentucky 
and Ohio. The authors have only spot-checked these areas and intensive 
collecting in them should be rewarding. The southwestern counties have 
proved unrewarding, but on occasion yield southern species wandering 
northwards when conditions are favorable. 


It is hoped that the following list will prove helpful to Indiana collectors 
and to others visiting and collecting in the state. A more thorough study 
of the indigenous and visiting species in many counties, especially in the 
southern parts, is greatly needed. 


VoLUME 26, NuMBER 1 LE 


HESPERIIDAE 


Panoquina ocola (Edwards) Ocola Skipper. Uncommon locally in northeastern coun- 
ties. Collected in August in Wabash and Kosciusko counties. 

Lerodea eufala (Edwards) Eufala Skipper. Scattered late summer records in Perry 
County (Masters and Masters, 1969). Recorded in September, Kosciusko County. 

Amblyscirtes vialis (Edwards ) Roadside Skipper. Common in the southern half of the 
state, but rare or absent in the northern counties. From mid-May to early Sep- 
tember. 

Atrytonopsis hianna (Scudder) Dusted Skipper. Rare in Lake and LaGrange counties 
in June. Not taken in southern parts. 

Euphyes dion dion (Edwards) Dion Skipper. Uncommon in the bogs of LaGrange, 
Wabash, Kosciusko, Fulton, and Marshall counties in late June and July. 

Euphyes dukesi (Lindsey) Dukes’ Skipper. Collected by Homer F. Price, July 24, 
1962, Steuben County and by Shull, July 27, 1970, Wabash County. 

Euphyes conspicua (Edwards) Black Dash. Uncommon to common in LaGrange, 
Steuben, Wabash, Marshall, Carroll, and Kosciusko counties in July. 

Euphyes bimacula (Grote & Robinson) Two Spotted Skipper. Uncommon in boggy or 
marshy meadows, from late June to early September, in LaGrange, Wabash, Kosci- 
usko, and Fuiton counties. 

Euphyes vestris metacomet (Harris) Dun Skipper. Common throughout the state, 
June to September. 

Poanes massasoit (Scudder) Mulberry Wing. Uncommon to common in July in the 
northeastern bogs of LaGrange, Steuben, Wabash, and Kosciusko counties. Also 
found in Fulton and Marshall counties. 

Poanes hobomok (Harris) Hobomok Skipper. Common in the northern half of the 
state from May to late August or early September. Diamorphic forms of the 
female pocahontas were found in Marshall, Howard, Cass, Marion, Fulton, and 
Kosciusko counties. 

Poanes zabulon (Boisduval & LeConte) Zabulon Skipper. Apparently more common 
in the southern half of the state; however, it has been collected in Marshall, How- 
ard, Cass, Marion, Fulton, and Kosciusko counties. May to September. 

Poanes viator (Edwards) Broad Winged Skipper. Uncommon in the northeastern 
counties. July in Steuben and Wabash counties. 

Atryone delaware delaware (Edwards) Delaware Skipper. Common throughout the 
state from late June to September. 

Atalopedes campestris ( Boisduval) Sachem. Common from late July to September in 
the northeastern counties. Scarce in late summer in Perry County, an extreme 
southern county (Masters and Masters, 1969). 

Pompeius verna verna (Edwards ) Little Glassy Wing. Common throughout the north- 
erm areas in June and July. September for Perry County. 

Wallengrenia otho egeremet (Scudder) Broken Dash. Very common everywhere from 
June to mid-September. 

Polites coras (Cramer) Peck’s Skipper. Abundant over most of the state. May 24 to 
early October. 

Polites themistocles (Latreille) Tawny Edged Skipper. Very common throughout the 
state. May 21 to October. 

Polites origines origines (Fabricius) Cross Line Skipper. Fairly common some years 
in Kosciusko, Wabash, Howard, and Marion counties. From May 25 to early 
October. 

Politas mystic (Scudder) Long Dash. Uncommon in LaGrange, Kosciusko, Wabash, 
Marshall, and Fulton counties in June. 

Hesperia metea (Scudder) Cobweb Skipper. Found sparingly in southern Indiana in 
May. 


16 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Hesperia sassacus (Harris ) Indian Skipper. Fairly common some years from late April 
to mid-June in Wabash and Kosciusko counties. 

Hesperia leonardus (Harris) Leonardus Skipper. Usually uncommon in the northern 
half of the state, from July to September. Fewer records from southern counties. 

Hylephila phyleus (Drury) Fiery Skipper. Uncommon throughout the state. From 
August 7 to September 21 in Wabash and Howard counties. 

Thymelicus lineola (Ochsenheimer ) European Skipper. Uncommon in June and July 
in Fulton, Wabash, and Kosciusko counties (Shull, 1968; Badger, unpublished, 
1962). In recent years its numbers have increased and it should be looked for in 
other counties. 

Oarisma powesheik (Parker) Powesheik Skipper. Uncommon locally in mid-June and 
July in the northwestern portions. Recorded in Lake County. 

Ancyloxypha numitor (Fabricius) Least Skipper. Common from late May to early 
October in the northern half of the state. Sparse in southern counties. 

Lerema accius (Abbott & Smith) Clouded Skipper. Status in the state is rather uncer- 
tain. Blatchley took it in Monroe County on July 3, 1886. New authentic records 
are needed to keep this species on the Indiana check list. 

Nastra Vherminier (Latreille) Swarthy Skipper. Uncommon and local throughout most 
of the state. Masters and Masters (1969) found it not uncommon in Perry County 
in certain fields. Sometimes common in Brown County, but rare in Kosciusko 
County. 

Pholisora catullus (Fabricius) Common Sooty Wing. Common throughout the state, 
from May 24 to September. 

Pyrgus communis communis (Grote) Checkered Skipper. Common everywhere from 
August to early November. 

Erynnis icelus (Scudder & Burgess) Dreamy Dusky Wing. Common in the northern 
half of the state in May and June. Not found in the southern counties. ? 

Erynnis brizo brizo (Boisduval & LeConte) Sleepy Dusky Wing. Common from late 
April to June in the northern half; uncommon southward. 

Erynnis persius (Scudder) Persius Dusky Wing. Collected by Badger, early May in 
the Dunes State Park, Lake County. Identified by Dr. J. M. Burns. 

Erynnis lucilius (Scudder & Burgess) Columbine Dusky Wing. Not uncommon from 
late May to mid-June in LaGrange County. Blatchley (1891) found it in Putman 
and Lake counties. 

Erynnis baptisiae (Forbes) Wild Indigo Dusky Wing. Shull collected a single speci- 
men 2 August 27, 1970 in Kosciusko County. Identified by Mr. H. A. Freeman. 

Erynnis zarucco (Lucas) Zarucco Dusky Wing. One record from Crawfordsville by 
Fred T. Hall (1936). 

Erynnis martialis (Scudder) Mottled Dusky Wing. Found locally throughout the state 
in June and July. 

Erynnis juvenalis juvenalis (Fabricius) Juvenal’s Dusky Wing. Common throughout 
the state from early May to June 5. 

Staphylus mazans hayhurstii (Edwards) Southern Sooty Wing. Uncommon in Allen, 
Kosciusko, Wabash, Marshall, Fulton, Howard, and Marion counties. Found in 
May, June, and July. 

Thorybes bathyllus (Smith) Southern Cloudy Wing. Fairly common throughout the 
state from mid-May to July. 

Thorybes pylades (Scudder) Northern Cloudy Wing. Common throughout the state 
from mid-May to July. 

Achalarus lyciades (Geyer) Hoary Edge. Uncommon in the northern counties in June 
and July. More common in Brown County and in the southern half of the state. 

Autochton cellus (Boisduval & LeConte) Golden Banded Skipper. Single specimen 
collected by Shull, June 7, 1971, Brown County State Park, 

Epargyreus clarus clarus (Cramer) Silver Spotted Skipper. Common in the northern 
half of the state from May to September. Uncommon in the southern counties. 


VoLUME 26, NuMBER 1 17 


PAPILIONIDAE 


Battus philenor philenor (Linnaeus ) Pipe Vine Swallowtail. Common throughout the 
state. May 1 to October 13 (late date). 

Papilio polyxenes asterius (Stoll) Black Swallowtail. Most common swallowtail 
throughout the state. April 27 to October 20. 

Papilio cresphontes (Cramer) Giant Swallowtail. Fairly common some years in the 
northern half of the state, usually in August. Found in June rarely and August in 
Kosciusko and Wabash counties. Masters and Masters (1969) refers to the Perry 
County subspecies as pennsylvanicus Chermock & Chermock (?). 

Papilio glaucus glaucus (Linnaeus) Tiger Swallowtail. Common throughout the state 
from April 2 to October 2. In northern Indiana the yellow females are less numer- 
ous than the black females. 

Papilio troilus troilus (Linnaeus) Spicebush Swallowtail. Common throughout the 
state. May 1 to October 13. 

Graphium marcellus (Cramer) Zebra Swallowtail. Fairly common throughout the state 
from April to October 22. 


PIERIDAE 


Pieris protodice protodice ( Boisduval & LeConte ) Checkered White. Widespread but 
not abundant throughout the state. April to October. 

Pieris napi oleracoa (Harris) Mustard White. Blatchley (1891) found it in Kosciusko 
County during the summer of 1890. On July 12, 1971 one specimen was collected 
in Mongo, LaGrange County by Shull. 

Pieris rapae (Linnaeus) European Cabbage Butterfly. Very common throughout the 
state from late March to early November. In 1969 it had five broods in Kosciusko 
County (Shull’s records). On June 17, 1969 a rare yellow female rapae was col- 
lected in Kosciusko County. 

Colias eurytheme eurytheme (Boisduval) Orange Sulphur; Alfalfa Butterfly. Common 
throughout the state. May to November. On September 5, 1969 a typical orange 
2 eurytheme was in copula with a typical yellow ¢ philodice in an alfalfa field 
in Kosciusko County. This species and the next have many varied summer and 
winter forms. __ 

Colias philodice philodice (Godart) Common or Clouded Sulphur. Very common in 
the whole state from mid-April to early November. 

Colias (Zerene) cesonia (Stoll) Dog Face. Rare in the northern half of the state in 
September. Occasional records from the southern parts in September and October. 

Phoebis sennae eubule (Linnaeus) Cloudless Sulphur. An uncommon local migrant 
into the southern counties. Rare in the north in the fall. Collected in Montgomery, 
Carroll, and Howard counties. 

Phoebis philea (Johansson) Orange Barred Sulphur. Blatchley (1891) recorded a 
single specimen from Jefferson County collected by Mr. G. C. Hubbard. The only 
other record is by Cooper (1938) who raised an imago from a larva found in 
Shelby County. Klots (1951) includes Indiana in its range. 

Eurema lisa ( Boisduval & LeConte) Little Sulphur. Usually common throughout the 
state from June to October; however, some years it may be absent or uncommon. 

Eurema nicippe (Cramer) Sleepy Orange. Uncommon in the northern half of the 
state from August to November. More common in late summer in the southern 
counties. 

Nathalis iole (Boisduval) Dainty Sulphur. Locally common in many areas from late 
summer to the end of November. 

Anthocaris midea ( Hiibner) Faleate Orange Tip. Not uncommon in central and south- 
ern Indiana from late April to early May. Some years it is rare or absent. 

Euchloe olympia olympia (Edwards) Olympia. Not uncommon some years in the 
northern half of the state from mid-April to mid-May. Badger found it in Lake 
and Pulaski counties. 


18 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


RIODINIDAE 


Calephelis borealis (Grote & Robinson) Northern Metalmark. Locally abundant in 
wooded areas during July in Montgomery, Brown, Howard, Marion, and Kosciusko 
counties. 

Calephelis muticum (McAlpine) Swamp Metalmark. Usually uncommon in the north- 
ern bogs in July and August. Occasionally common in the tamarack bog at Mongo, 
LaGrange County. 

Harkenclenus titus (Fabricius ) Coral Hairstreak. Uncommon in the northern half of 
the state in June and July. Masters and Masters (1969) found the subspecies 
mopsus (Hiibner) in Perry County in late June. 

Satyrium liparops strigosa (Harris) Striped Hairstreak. Common in northeastern Indi- 
ana, and probably throughout the state, from late June to July 10. 

Satyrium calanus falacer (Godart ) Banded Hairstreak. Common throughout the state 
from late June to mid-July. Former Indiana records by the name calanus (Florida 
Hairstreak ) belong to this classification. 

Satyrium caryaevorus (McDunnough) Hickory Hairstreak. Uncommon in June and 
July in Kosciusko and Wabash counties. 

Satyrium edwardsii (Saunders) Edward’s Hairstreak. Uncommon in the northern half 
of the state in June and July. 

Satyrium acadica acadica (Edwards) Acadian Hairstreak. Uncommon to locally com- 
mon in LaGrange, Kosciusko, Wabash, Marshall, Fulton, and Lake counties in late 
June and early July. 

Calycopis cecrops (Fabricius) Red Banded Hairstreak. Although both Blatchley 
(1891) and Klots (1951) list this species for Indiana, it must be rare indeed as 
we have not found it. Masters and Masters (1969) found it scarce in late May 
and July in Perry County. 

Callophrys (Incisalia) polios (Cook & Watson) Hoary Elfin. Found in the sand dunes 
of Lake County in May. Uncommon in April in some southern counties. 

Callophrys (Incisalia) irus (Godart) Frosted Elfin. Taken on lupine in May in Pulaski 
County. According to Ehrlich and Ehrlich (1961), it is found in northwestern 
Indiana. 

Callophrys (Incisalia) henrici turneri (Clench) Henry’s Elfin. Uncommon in April and 
May in the southern half of the state. Collected on redbud in Howard County. 

Callophrys (Mitoura) gryneus gryneus (Hubner) Olive Hairstreak. Rare and local 
near Red Cedar during May in Carroll County. This species is probably wide- 
spread in southern Indiana. There are records from Lake and Parke counties. 

Atlides halesus (Cramer) Great Purple Hairstreak. According to Ehrlich and Ehrlich 
(1961), strays have been taken in northern Indiana. The presence of mistletoe in 
Perry County makes it a likely area. We have not collected it in Indiana, but have 
taken it elsewhere. 

Euristrymon ontario (Edwards) Northern Hairstreak. Some years it is uncommon 
from mid-June to early July in Wabash and Kosciusko counties. Dogbane is its 
favored food flower in northern Indiana. 

Panthiades m-album (Boisduval & LeConte) White M Hairstreak. Strays have been 
found in southern Wisconsin, Ohio, and northern Indiana (Lake County). Mas- 
ters and Masters (1969) report that Lois Ann Winter found a male of this species 
in Perry County between May 5 and 14, 1962. Badger found a stray in Carroll 
County, May 1960. 

Strymon melinus melinus (Hiibner) Gray Hairstreak. Some years it is common and 
other years uncommon in the northern half of the state from June 29 to October 3. 
Common in southern parts from April to September. 

Feniseca tarquinius tarquinius (Fabricius ) Harvester. Uncommon to rare in the south- 


ern half of the state. Found between mid-May and mid-September in Howard 
County. 


VoLUME 26, NuMBER 1 I) 


Lycaena thoe (Guerin-Meneville) Bronze Copper. Fairly common in central and 
northern Indiana from June to early October. Uncommon in southern counties. 

Lycaena helloides (Boisduval) Purplish Copper. Uncommon in July and August in 
Wabash and Kosciusko counties. Occasionally found in northwestern Indiana 
(Lake County ). 

Lycaena dorcas (Kirby) Dorcas Copper. Found rarely in the bogs of LaGrange 
County. We have not yet found it. 

Lycaena epixanthe (Boisduval & LeConte) Bog Copper. Uncommon in July in 
Wabash County. Klots (1951) also records it in northern Indiana. Blatchley 
(1891) says it is rare in Lake County. 

Lycaena phlaeas americana (Harris) American Copper. Common in the northern 
counties from May 15 to September 24. Rare in extreme southern portions. 
Badger found the form fasciata in Howard County. 

Lycaeides melissa samuelis (Nabokov ) Karner Blue. Uncommon to rare in LaGrange 
and Wabash counties; more common in Lake County. The northeastern popula- 
tions formerly known as scudderi (Edwards) now belong to the races of melissa. 
Found from June to August. 

Everas comyntas comyntas (Godart) Eastern Tailed Blue. Common throughout the 
state from April to early October. 

Glaucopsyche lygdamus (Doubleday) Silvery Blue. Recorded from Wabash and 

LaGrange counties. Usually rare in May and June. 

Celastrina argiolus (Linnaeus) Spring Azure. Abundant in northeastern Indiana, 
where most of the specimens belong to the subspecies pseudargiolus (Boisduval 
& LeConte) and its forms. The subspecies lucia (Kirby) does not occur in the 
northern portions of the state. Argiolus occurs throughout the state from mid-April 
to mid-September. Klots (1951) gives a fine description of this group. 


LIBYTHEIDAE 


Libytheana bachmanii (Kirtland) Snout Butterfly. Usually common throughout the 
state from early April to September. 


NYMPHALIDAE 


Anaea andria andria (Scudder) Goatweed Butterfly. Scattered records from the cen- 
tral and southern parts of the state in April, July, and August. Found in July in 
Orange County and in August in Wabash County. 

Asterocampa celtis celtis (Boisduval & LeConte) Hackberry Butterfly. Common 
throughout the state, June 12 to September 4. 

Asterocampa clyton clyton ( Boisduval & LeConte) Tawny Emperor. Common in the 
entire state, but less so than the above species. From mid-June to September 14. 

Limenitis arthemis astyanax (Fabricius) Red Spotted Purple. According to Platt and 
Brower (1968) arthemis and astyanax are not distinct species. Therefore the 
present authors have combined them. Two specimens, having indistinct white 
bands, were collected June 21, 1969 in Wabash County. Rather prominent white- 
banded forms have been taken in a few northern counties. From June 5 to October 
7 throughout the state, with the astyanax form greatly outnumbering the arthemis 
form. 

Limenitis archippus archippus (Cramer) Viceroy or Mimic. Common throughout the 
state. Early June to October 13. 

Vanessa atalanta (Linnaeus) Red Admiral. Common throughout the state from late 
March to September 20. One very late date is October 26. 

Vanessa virginiensis (Drury) American Painted Lady. Usually common throughout 
the state; however, some years it is uncommon or absent. From late April to 
September 25. 

Vanessa cardui (Linnaeus) Painted Lady. Some years it is common to abundant 


20 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


throughout the state; from March to September 20. In 1969, it was strangely 
absent in northern Indiana. It is a highly migratory species. 

Junonia coenia coenia (Hiibner) Buckeye. Usually common or fairly common and 
quite variable throughout the state; from May to October 13, and rarely to early 
November. 

Nymphalis vau-album j-album (Boisduval & LeConte ) Compton Tortoise Shell. Found 
in Porter and Lake counties. Two specimens were collected in August, 1934 in 
Wabash County. Rare. 

Nymphalis milberti milberti (Godart) Milbert’s Tortoise Shell. Very common in 
northern Indiana, but uncommon in the southern counties. From March to No- 
vember 7. It may even hibernate as one was found on a log on December 29, 
1953 in Madison County. 

Nymphalis antiopa antiopa (Linnaeus ) Mourning Cloak. Common throughout most of 
the state; less common in the south. From February to October 25. 

Polygonia interrogationis (Fabricius ) Question Mark. Common from March to Novem- 
ber in the entire state. 

Poiygonia comma (Harris) Comma or Hop Merchant. Common throughout the state 
from March 4 to November 22; however, it may be rarely found much later on 
warm winter days. 

Polygonia satyrus (Edwards) Satyr Angle Wing. Recorded from Kosciusko and Wa- 
bash counties from May 30 to late July. Uncommon, rare, or absent some years. 

Polygonia progne (Cramer) Gray Comma. Common, uncommon, or absent some years 
in Wabash and Kosciusko counties. From June 20 to early November. Apparently 
uncommon in midsummer in several other counties. 

Chlosyne nycteis nycteis (Doubleday) Silvery Checkerspot. Fairly common through- 
out the state from May 17 to mid-October. 

Chlosyne gorgone carlota (Hiibner ) Gorgone Checkerspot. The only records are from 
Perry County by J. F. and Wilma L. Masters (1969). It should be looked for 
elsewhere. 

Chlosyne harrissi (Scudder) Harris’ Checkerspot. Our 1934 Wabash County records 
may be incorrect as they were not checked by an expert, but they are rarely taken 
from the Mongo tamarack bog in LaGrange County. Klots (1951) includes Indi- 
ana in its range. 

Phyciodes tharos tharos (Drury) Pearl Crescent. Very common throughout the state 
from April or mid-May to October. 

Phyciodes batessi ( Reakirt) Tawny Crescent. Perhaps there are a few authentic rec- 
ords for Indiana from late May to early June, but upon close examination our 
batesii turned out to be aberrant tharos. As Masters and Masters (1969) suggest, 
this species may be widespread from Brown County northward. 

Euphydryas phaeton (Drury) Baltimore. The nominate subspecies is uncommon in the 
northern half of the state in June and July. Specimens from Brown and other 
south-central counties, according to Masters and Masters (1969), may belong to 
the subspecies ozarkae, but ours from Brown County appear to be identical to 
the northern specimens. 

Boloria selene myrina (Cramer) Silver Bordered Fritillary. Common in the far north- 
ern counties from June 15 to September. Absent in the remainder of the state. 

Boloria toddi ammiralis (Hemming) Meadow Fritillary. Very common throughout the 
northern portions of the state from April 26 to October 1. 

Speyeria idalia (Drury) Regal Fritillary. Scattered records from widely separated 
counties during July and August. Uncommon to rare in Wabash, Kosciusko, Mar- 
shall, and Fulton counties. 

Speyeria atlantis (Edwards ) Atlantis Fritillary, Blatchley (1891) and Montgomery 
(1931) both reported it from Vanderburgh and Lake counties. No authentic, 


recent records from any portion of the state have come to our attention. It should 
be found along the Indiana-Michigan border counties. 


VOLUME 26, NUMBER 1 Dak 


Speyeria diana (Cramer) Diana. Old records are from Vanderburgh County ( Blatch- 
ley, 1891). According to Masters and Masters (1969), a male of this species was 
collected on July 15, 1962 just north of Troy in Perry County. 

Speyeria cybele cybele (Fabricius ) Great Spangled Fritillary. Common throughout the 
state. May 20 to September. 

Speyeria aphrodite (Fabricius) Aphrodite. In northeastern Indiana most specimens 
belong to the subspecies alcestis (Edwards). Some years it is common from June 
25 to late August. A few nominate subspecies have been collected in the extreme 
northern bogs. Also the aphrodite was found in Marshall, Fulton, and Howard 
counties. 

Euptoieta claudia claudia (Cramer) Variegated Fritillary. Uncommon throughout the 
state, but more found in the northeastern counties of LaGrange, Kosciusko, and 
Wabash. From June 22 (early) to October 15. 

Agraulis vanillae (Linnaeus) Gulf Fritillary. Blatchley (1891) reported it from Van- 
derburgh County. Lawrence James collected one in Clay County in July. 


DANAIDAE 


Danaus plexippus plexippus (Linnaeus) Monarch. Some years common throughout 
the state; normally from late April to early November, but more common in Sep- 
tember, when occasional southward migrations take place. 


SATYRIDAE 


Lethe portlandia anthedon (Clark) Pearly Eye. Uncommon throughout the state. 
Occasional records from Howard and Carroll counties. June to August. 

Lethe creola (Skinner) Creole Pearly Eye. Extremely rare in northeastern Indiana 
near the Michigan border. Masters and Masters (1969) report that Dr. E. M. 
Brackney took one on June 29, 1965 in Orange County. 

Lethe eurydice appalachia (R. L. Chermock) Eyed Brown. Uncommon to common in 
the northeastern counties from June to August. Found in LaGrange, Steuben, 
Wabash, and Marshall counties. Less common in the central portions of the state. 

Euptychia gemma gemma (Hubner) Gemmed Satyr. Recorded only in Perry County 
from late April to early October by J. H. and Wilma L. Masters. 

Euptychia mitchellii (French) Mitchell’s Satyr. Uncommon in July in our extreme 
northeastern county bogs. Collected in LaGrange County many years by Homer 
F’. Price. 

Euptychia cymela cymela (Cramer) Little Wood Satyr. Very common throughout the 
state from late May through July. 

Cercyonis pegala (Fabricius) Wood Nymph, Grayling. Common in the northern half 
of the state, but absent in the southern parts. From mid-May to October 1. The 
subspecies nephele (Kirby) is occasionally found in July from Kosciusko County, 
but C. p. alope (Fabricius ) is the common form in Wabash and Kosciusko coun- 
ties. Other subspecies seem to be present, but they may be only aberrant forms. 
This species, with its many overlapping or integrated forms, needs far more study 
in Indiana. 


HYPoTHETICAL List FOR INDIANA 


Collectors should look for the following species in Indiana: 


Hesperiidae 


Amblyscirtas samoset (Scudder). Mississippi Valley. 
Atrytone arogos (Boisduval & LeConte). Florida and Gulf States, north to New Jersey, 
Minnesota, Iowa, and Nebraska. 


bo 
bo 


JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Polites vibex (Geyer). Tropics north to Connecticut, Arkansas, and a dubious Wis- 
consin record. 

Hesperia uncas (Edwards). A western prairie species which sometimes enters the 
easter area. 

Hesperia attalus (Edwards ). Ohio and Wisconsin. 

Pyrgus centaureae (Rambur). New York to Colorado. 

Erynnis horatius (Scudder & Burgess ). Ohio. 

Thorybes confusis (Bell). Missouri, Maryland. Probable 9 June 17, 1962, Marshall 
County—Badger. 


Pieridae 


Ascia monuste (Linnaeus). Mississippi Valley to Kansas. 

Colias interior (Scudder). Michigan. 

Phoebis agarithe ( Boisduval ). Strays to Illinois. 

Eurema mexicana (Boisduval). Rarely to Michigan and Wisconsin. 
Kricogenia lyside (Godart). Strays to Illinois. 


Lycaenidae 


Callophrys (Incisalia) augustinus (Westwood). Illinois and Michigan. 

Callophrys (Incisalia) niphon (Hiibner). Michigan. 

Satyrium kingi (Klots & Clench). Harry K. Clench has written that it seems to occupy 
the same habitat as Lethe creola. 


Lycaena xanthoides (Boisduval). Upper Mississippi Valley, Kansas north through 
Nebraska, Minnesota, and Illinois. 


Plebejus saepiolus ( Boisduval). Michigan. 


Riodinidae 


Lephelisca virginiensis (Guérin-Méneville ). Ohio. 


Nymphalidae 


Polygonia faunus (Edwards). Canada to South Carolina, Iowa, and Michigan. 
Boloria eunomia (Esper). Michigan and Wisconsin. 


ACKNOWLEDGMENTS 


In this annotated list of Indiana butterflies, the authors have, in the 
main, followed the classifatory arrangement of Dr. Cyril F. dos Passos, 
A Synonymic List of the Nearctic Rhopalocera (1964), and his revisions 
of the Nearctic Melitaeinae (1969) and the Nearctic Lycaenidae (1970). 

A Guide to the Butterflies (1951) by Dr. Alexander Klots, The Butterfly 
Book (1940) by Dr. J. W. Holland, and How to Know the Butterflies 
(1961) by Dr. and Mrs. Paul R. Ehrlich constitute the primary sources 
used for identification; however, a few of the more difficult species and 
subspecies were either confirmed or identified by the following experts: 
Dr. J. M. Burns of Cambridge, Mass., Mr. Harry K. Clench of the Carnegie 
Museum, Pittsburgh, Pa., Mr. H. A. Freeman of Garland, Texas, and Drs. 
Frederick H. Rindge and Alexander B. Klots of the American Museum of 


VoLUME 26, NuMBER 1 23 


Natural History, New York. To these people the authors are greatly in- 
debted. The authors, of course, assume full responsibility for any in- 
accuracies which may have been reported in this study. 

Correspondence with the following collectors has been very helpful in 
determining the distribution of some species: Mr. Ray W. Bracher of 
Granger, Ind., Dr. J. W. Burns of Cambridge, Mass., Mr. Harry K. Clench 
of Pittsburgh, Pa., Mr. Julian P. Donahue and Mr. M. C. Nielsen of East 
Lansing, Mich., Dr. Richard Heitzman of Independence, Mo., Dr. R. R. 
Irwin of Chicago, Ill., Dr. Wilbur S. McAlpine of Union Lake, Mich., 
Mr. Homer F. Price of Payne, Ohio, Dr. Charles Remington of New 
Haven, Conn., Dr. P. Sheldon Remington of Greenwich, Conn., and Mr. 
James E. Shields of Indianapolis, Ind. 

Also appreciation is expressed to the coordinators and reporters of the 
Annual Summary reports, published by the Lepidopterists’ Society, for- 
merly in the Journal and recently in the News. 


LITERATURE CITED 


Bapcer, F. S. 1958. Euptychia mitchelli (Satyridae) in Michigan and Indiana 
tamarack bogs. Lepid. News 12: 41-46. 

Barnes, W. B. 1952. Zoogeographic regions of Indiana. Amer Midland Nat. 48: 
694-699. 

BLATCHLEY, W. S. 1886. Some southern Indiana butterflies. Hoosier Nat., Nov. 
& Dec. 

1891. Catalogue of the butterflies known to occur in Indiana. Ann. Rep. 
Indiana State Geol. 17: 365—408. 

Cooper, R. H. 1938. A breeding record for the red-barred sulphur (Callidryas 
philea Linn.) from Indiana. Ent. News 49: 261. 

pos Passos, C. F. 1964. A synonymic list of the nearctic Rhopalocera. Mem. Lepid. 
Soc. 1: VI & 145 pp. 

1969. A revised synonymic list of the Nearctic Melitaeinae with taxonomic 

notes (Nymphalidae). J. Lepid. Soc. 23: 115-125. 

1970. A revised synonymic catalogue with taxonomic notes on some 
Nearctic Lycaenidae. J. Lepid. Soc. 24: 26-38. 

EuruicH, P. R. & ANNE H. 1961. How to Know the Butterflies. Wm. C. Brown, 
Dubuque, Iowa. 262 p. 

Hatt, F. T. 1936. The occurrence of unusual Rhopalocera in Indiana. Proc. 
Indiana Acad. Sc. 45: 273-274. 

Hoiianp, W. J. 1940. The Butterfly Book. Doubleday, New York, 442 pp. + 77 
pls. 

Kiots, A. B. 1951. A Field Guide to the Butterflies. Houghton Mifflin, Boston, 
349 p. 

Linpsry, A. A., ed. 1966. Natural Features of Indiana. Indiana Acad. Sc. 597 pp. 

Masters, J. H. & WimMa L. 1969. An annotated list of the butterflies of Perry 
County and a contribution to the knowledge of lepidoptera in Indiana. Assoc. 
Minnesota Ent. 6; 1—25. 

MontcoMery, R. W. 1931. Preliminary list of the butterflies of Indiana. Proc. 
Indiana Acad. Sc. 40: 357-359. 

Puatr, A. P. AND L. P. Brower. 1968. Mimetic versus disruptive coloration in 


24 JouRNAL OF THE LEPIDOPTERISTS’ SOCIETY 


integrating populations of Limenitis arthemis and astyanax butterflies. Evolution 
22: 699-718. 

Prick, H. F. anp E. M. Suuxxi. 1969. Uncommon butterflies of northeastern 
Indiana. J. Lepid. Soc. 23: 186-188. 

SHuLt, E. M. 1968. Thymelicus lineola (Hesperiidae) in Indiana. J. Lepid. Soc. 
DAZ PA), 


THE LIFE HISTORY OF SCHINIA INTRABILIS 
(NOCTUIDAE ) 


D. F. Harpwick 


Entomology Research Institute, Canada Department 
of Agriculture, Ottawa, Ontario 


Schinia intrabilis Smith (1893, p. 331) feeds in the larval stage on the 
blossoms of the Arrowweed, Pluchea sericea (Nutt.) Coy. The arrow- 
weed is a willow-like composite that occurs abundantly around seeps and 
along river banks in the deserts of southern California. During the early 
blossoming period of its food plant, Schinia intrabilis is not a rare insect 
in appropriate desert habitats. 

According to Munz (1963), the Arrowweed is distributed from southern 
California eastward to Texas, but I have examined intrabilis only from 
as far east as Yuma and Ehrenberg, Arizona. The species is univoltine and 
the period of adult activity is co-ordinated in any area with the single 
annual blossoming period of the Arrowweed. Specimens in the Canadian 
National Collection from the deserts of southern California were taken on 
dates between the middle of March and the end of April. 


Behaviour 


Schinia intrabilis is evidently a species of exclusively or preponderantly 
nocturnal habits. In moist areas in which eggs and young larvae could be 
recovered without difficulty from Arrowweed heads, no adult activity was 
noted during daylight hours. 

Eggs are deposited in the Arrowweed head at a stage when the sepals 
of the bud have drawn apart at the top sufficiently to expose the florets. 
Females do not and probably cannot oviposit in the tough, leathery, un- 
opened buds. The egg is inserted among the florets from the top of the 
head. 

Three captive females deposited a mean of 83 eggs, and the maximum 


VoLUME 26, NUMBER 1 25 


eo > if 


Figs. 14. Schinia intrabilis Smith, its habitat and food plant. 1, Adult, Twenty- 
nine Palms, Calif. 2, Willis Palms, near Indio, Calif. 3, 4, Pluchea sericea ( Nutt.) 
Cov. 


deposited by a single female was 132. Eggs maintained at room tempera- 
ture hatched on the third and fourth days after deposition. 

The newly hatched larva bores into an adjacent floret and tunnels down 
through it toward the receptacle. The second and subsequent instars feed 
preponderantly on the developing seeds. Usually by the third stadium, 
the larva quits the first head and enters a second. Both third-and fourth- 
stadium larvae attack the heads from the top, and the half grown larvae 
must curl up within the individual head to remain concealed while feed- 
ing. Toward the end of the fourth stadium and during the fifth, the larva 
feeds on the heads from a position on the stem; it usually attacks the head 
by boring a hole through the side just above the heavy sepals that sur- 
round the base. At the cessation of feeding the larva makes its way to 
the surface of the ground and tunnels into the soil to pupate. 


Descriptions of Stages 


The following descriptions of immature stages were based on the prog- 
eny of seven females taken in the Indio area of southern California. The 


26 JouRNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Yi iy ty Ly yy 7 


Figs. 5-8. Schinia intrabilis Smith, ultimate-stadium larvae. 5, 6, Left lateral; 7, 
8, dorsal. 


durations of stadia listed are those obtained from rearings maintained at 
room temperature. Rearing techniques employed were the same as those 
outlined by Hardwick (1958). The estimate of variation, following the 
means for various values, is the standard deviation. 


Adult (Fig. 1). Head and thorax light olivaceous fawn. Abdomen paler, cream or 
creamy-grey. Forewing fawn or fawn-grey, often with an olivaceous suffusion. Trans- 
verse anterior line white, angling outward from trailing margin, then curving sharply 
inward and terminating not on costal margin but at base of wing. Basal space fawn 
or fawn-grey. Transverse posterior line white, angling inward from costa near apex 
to a point below reniform spot then curving outwardly to meet trailing margin near 
outer angle. Median space variably suffused with white or pale grey and thus usually 
paler than basal and subterminal spaces. Orbicular and claviform spots absent; reni- 
form evident as a dark, ill-defined shade. Subterminal line cream, regular, close to 
outer margin of wing and parallel to it. Fringe and narrow terminal space concolorous 
with subterminal space. Hind wing white with a brown outer-marginal band and a 
large brown discal spot; a pale median streak in outer-marginal band. Inner margin 
of wing often suffused with brown. Fringe white with a yellowish or brownish basal 
line. Underside of forewing shining cream with a large dark-brown reniform spot 


and a light-brown submarginal band. Hind wing white, with or without brown spots 
on disc and near anal angle. 


Expanse: 23.5 +1.3 mm (47 specimens ). 

Egg. Very pale yellow when deposited, and exhibiting little colour change until a 
few hours before hatching when head capsule becomes visible through chorion. 

Dimensions of egg: length, 0.872 + 0.025 mm; diameter, 0.510 + 0,042 mm (25 
eggs). 

Incubation period: 3.5 + 0.5 days (104 eggs). 

First-Stadium Larva. Head medium to dark smoky-brown. Prothoracic and suranal 


VoLUME 26, NUMBER 1 Dif, 


Figs. 9, 10. Schinia intrabilis Smith, apical abdominal segments of pupa. 9, 
Ventral; 10, right lateral. 


shields concolorous with head or somewhat paler. Trunk white, cream, or pale grey. 
Spiracles with light- to medium-brown rims. Legs medium smoky-brown. 

Head width: 0.262 + 0.009 mm (15 larvae). 

Duration of stadium: 3.7 + 0.9 days (33 larvae). 

Second-Stadium Larva. Head light orange-brown, mottled dorsally with light to 
medium smoky-brown. Prothoracic shield fawn to light orange-brown, variably 
mottled and emarginated with chocolate-brown. Suranal shield approximating pro- 
thoracic shield in colour and similarly marked with brown. Trunk light greyish-cream 
to pale greenish-grey, usually with two pairs of dorsal and a pair of lateral white 
lines. Spiracles with dark-brown rims. Thoracic legs fawn to light orange-brown, 
variably suffused with smoky-brown. 

Head width: 0.446 + 0.034 mm (25 larvae). 

Duration of stadium: 3.0 + 0.8 days (33 larvae). 

Third-Stadium Larva. Head warm cream mottled dorsally with light orange. Pro- 
thoracic shield fawn, mottled with white and brown and with a white median line. 
Suranal shield white, lightly mottled with pale brown. Trunk green with numerous 
longitudinal white lines. Mid-dorsal band light greyish-green, often with a white 
median line. Subdorsal area consisting of white marginal lines and a median greyish- 
green band; median band often with a diffuse white median line. Supraspiracular 
area greyish-green with a white median line. Spiracular band white. Ventral region 
light grey. Rims of spiracles and bases of setae dark brown or black. Thoracic legs 
pale fawn suffused with light smoky-brown. 

Head width: 0.70 + 0.03 mm (18 larvae). 

Duration of stadium: 3.3 + 0.7 days (33 larvae). 

Fourth-Stadium Larva. Head cream, mottled dorsally with very pale fawn. Pro- 
thoracic shield light green with three longitudinal white lines. Suranal shield light 
green or light fawn variably marked with cream. Trunk greenish-grey at beginning 
of stadium, becoming leaf-green after feeding; trunk with numerous longitudinal lines 
of cream or pale grey. Mid-dorsal band green with a median longitudinal shade of 
cream or pale grey. Subdorsal area cream or pale grey with a pair of median longi- 
tudinal green lines. Supraspiracular area cream or pale grey margined by green lines 
and with a pair of green median lines. Spiracular band white with a discontinuous 
green median shade. Suprapodal area green, mottled with cream and lightly marked 


28 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


with fawn. Mid-ventral area greyish-green. Spiracles with black rims. Thoracic legs 
straw-coloured, often tinged with green proximally, and occasionally lightly suffused 
with fawn. 

Head width: 1.10 + 0.03 mm (25 larvae). 

Duration of stadium: 4.0 + 0.6 days (33 larvae). 

Fifth-Stadium Larva (Figs. 5-8). Larvae occurring in two colour phases, pale fawn 
and green. Head straw-coloured, faintly mottled with orange, and suffused with 
green in green specimens. Prothoracic shield pale orange or pale green; three narrow 
white longitudinal lines usually evident; shield marked with black around bases of 
setae. Suranal shield pale fawn or green, marked with white and with black setal 
bases. Trunk pale fawn or light green. Mid-dorsal band somewhat darker than re- 
mainder of dorsum. Subdorsal and supraspiracular areas undistinguished, mauve-grey 
with six or seven longitudinal lines of pale fawn or bright green; longitudinal lines 
often irregular and discontinuous. Spiracular band white with a median shade of 
fawn or green. Suprapodal area pale fawn or green, lightly marked with mauve-grey. 
Mid-ventral area pallid fawn or pale green. Thoracic legs pale fawn, suffused with 
green in green specimens. 

Head width: 1.75 + 0.08 mm (7 larvae). 

Duration of stadium: 5.4 + 1.1 days (33 larvae). 

Pupa (Figs. 9, 10). Medium orange-brown, the appendages often suffused with 
green. Spiracles on a level with general surface of cuticle; spiracular sclerites weakly 
projecting. Anterior marginal areas of abdominal segments 5, 6 and 7, each with a 
rather wide band of prominent pitting. Proboscis terminating well anterior to apexes 
of wings. Apex of tenth abdominal segment broadly rounded and bearing two short, 
fine setae. 

Length from anterior end to posterior margin of fourth abdominal segment: 8.1 + 
0.4 mm (23 pupae). 


ACKNOWLEDGMENTS 


I appreciate the help of my associate, Mr. Eric Rockburne, who mea- 
sured the immature stages and drew the cremaster area of the pupa. Mr. 
John E. H. Martin of the Entomology Research Institute took the photo- 
graphs accompanying this paper. 


LITERATURE CITED 


Harpwick, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed 
species of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae. 
Can. Ent. Suppl. 6. 

Munz, P. A. 1963. A California flora. University of California Press, Berkeley. 

SmirH, J. B. 1893. Descriptions of Noctuidae from the Death Valley. I] t is 
5: 328-334, y. Insect Life 


NATURAL INTER-BREEDING OF CLOSE NYMPHALID GROUPS 


On June 12, 1971 on the hilltops of Mother Cabrini Shrine in Jefferson County 
Colorado, a freshly emerged female of Melitaea pola arachne Edwards was found an 
copulation with a male Chlosyne gorgone carlota Reakirt. The male was hanging and 


the female was flying. The time was 0920 and the legend was Joel Jablonski 
RAYMOND J, JAE, 1286 South Umatilla St., Denver, Colorado 80223. 


VOLUME 26, NUMBER 1 29 


THE LIFE HISTORY OF SCHINIA PALLICINCTA 
(NOCTUIDAE) 


D. F. Harpwick 


Entomology Research Institute, Canada Department 
of Agriculture, Ottawa, Ontario 


Schinia pallicincta Smith (1906, p. 24) feeds in the larval stage on the 
heads of Desert Marigolds, Baileya pauciradiata Harv. and Gray and B. 
multiradiata Harv. and Gray. All specimens of pallicincta in the Canadian 
National Collection were taken on the California deserts in areas between 
Ocotillo, San Diego Co. and Mono Lake, on dates between the middle of 
March and the first of June. 

The species is obviously closely related to Schinia miniana (Grote, 
1881, p. 175) which was described from New Mexico. The latter species 
is larger and more brightly coloured, however, and the hind wings are 
rosy red rather than smoky brown. Two specimens of miniana in the 
Canadian National Collection from the Big Bend area of western Texas 
were taken in the heads of Baileya multiradiata. It is possible that 
pallicincta and miniana represent only well-defined races of a single spe- 
cies. Life history data on miniana, and a study of series of specimens 
from intermediate localities will undoubtedly elucidate the situation. 


Behaviour 


The full globular eyes of Schinia pallicincta suggest that the species is 
primarily nocturnal, and indeed it is often taken at light; however, it is also 
frequently found during the hours of daylight flying among clumps of its 
food or ovipositing in the heads. The eggs are deposited in both the buds 
and blossoms, but those buds which have opened sufficiently to expose 
the upper ends of the still tightly closed florets are greatly preferred to 
smaller buds or open heads. The eggs are inserted among the florets. 

Five captive females deposited a mean of 40.3 eggs, and the maximum 
deposited by a single individual was 74. The majority of eggs hatched on 
the sixth day after deposition. 

The newly hatched larva bores into an adjacent floret and feeds on the 
contents. In the early stadia, the larva tunnels within the florets. In the 
median stadia the larva feeds on both seeds and florets, and in the ulti- 
mate stadium, the whole contents of the receptacle except for the ray 
florets are consumed. When feeding in the heads of Baileya multiradiata, 
the inner ray petals are drawn together by the late-stadium larva to form 


30 JourNAL OF THE LEPIDOPTERISTS SOCIETY 


VoLuME 26, NuMBER 1] 31 


a nest in which it remains concealed; the habit was never noted among 
larvae feeding in the heads of Baileya pauciradiata. 

On the completion of feeding, the larva makes its way to the ground 
and tunnels into the soil to pupate. When the larva tunnels into the sandy 
soil characteristic of the dunes areas in which Baileya pauciradiata is 
abundant it forms a delicate silken tube leading down to the pupal cell. 
Evidence of this tube could not be found in cases in which larvae pupated 
in heavier soil. 


Descriptions of Stages 


The following descriptions of immature stages were based on the prog- 
eny of five females taken in the sand dunes area west of Indio, California. 
The specimens were taken flying about or resting on the heads of Baileya 
pauciradiata. The larvae were reared individually at room temperature 
on the heads of B. pauciradiata. Rearing techniques employed were those 
described by Hardwick (1958). The estimate of variability following the 
means for various values is the standard deviation. 


Adult (Figs. 1, 3). Head and thorax varying from bright yellow to light fawn. Ab- 
domen usually paler than thorax and with a greyish tone. Forewing light fawn marked 
with white. Transverse anterior line broad, white, straight or weakly excurved. Basal 
space pale fawn, suffused with yellow in specimens with yellow vestiture on thorax. 
Transverse posterior line broad, white, weakly excurved around cell, then essentially 
straight to inner margin.. Median space pale fawn, rarely with a brown median shade. 
Orbicular and reniform spots not defined. Subterminal line usually indicated only 
as an elongate dark mark at costal margin. Fused terminal and subterminal spaces 
concolorous with median space. Fringe fawn, usually checkered with darker scaling. 
Hind wing dark smoky-brown, usually becoming paler toward base, and rarely with 
a pink iridescence. Fringe cream, often with a yellow basal line. Underside of fore- 
wing light fawn, usually heavily overlaid with brown through central area; central 
brown area often tinged with pink marginally; fringe pale fawn. Hind wing pale 
fawn, variably marked with brown, or pink and brown, in basal and median areas; 
fringe fawn. 

Expanse: 19.1 + 0.9 mm (16 specimens ). 

Egg. Very pale yellow when deposited and remaining essentially unchanged until a 
few hours before hatching when head capsule becoming visible through chorion. 

Dimensions of egg: length, 0.784 + 0.040 mm; diameter, 0.361 + 0.010 mm (5 
eggs). 

Incubation period: 6.1 + 0.3 days (121 eggs). 

First-Stadium Larva. Head blackish-brown. Prothoracic and suranal shields dark 
smoky-brown. Trunk pale yellow or cream. Spiracles with light- to medium-brown 
rims. Thoracic legs dark smoky-brown. 


< 


Figs. 1-8. Schinia pallicincta Sm. and its food plant. 1, Adult, La Quinta, River- 
side Co., Calif. 2, pupae; 3, adult resting in sunflower head; 4, food plant, Baileya 
pauciradiata Haw. and Gray; 5, 6, left lateral aspect of ultimate-stadium larvae; 7, 8, 
dorsal aspect of ultimate-stadium larvae. 


32 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 9, 10. Apical abdominal segments of pupae. 9, Ventral; 10, right lateral. 


Head width: 0.259 + 0.004 mm (20 larvae). 

Duration of stadium: 3.4 + 0.8 days (40 larvae). 

Second-Stadium Larva. Head dark chocolate-brown or black. Prothoracic shield 
medium to dark smoky-brown, often with a cream or pale-grey median line. Suranal 
shield medium smoky-brown. Trunk cream or creamy-grey, often with a median 
dorsal, and a pair of subdorsal light yellow lines. Spiracles with medium-brown rims. 
Thoracic legs dark smoky-brown. 

Head width: 0.454 + 0.042 mm (20 larvae). 

Duration of stadium: 2.5 + 0.8 days (40 larvae). 

Third-Stadium Larva. Head dark chocolate-brown or black. Prothoracic shield 
medium to dark chocolate-brown, usually with a median and a pair of submarginal 
greyish-yellow longitudinal lines. Suranal shield medium to dark smoky-brown, often 
with three pale yellowish-grey longitudinal lines. Maculation of trunk usually poorly 
defined. Mid-dorsal band greyish-yellow. Subdorsal area paler than mid-dorsal band, 
often with a darker median shade. Supraspiracular area greyish-yellow, margined 
ventrally by a cream line. Spiracular band light yellow. Ventral region yellowish- 
grey. Spiracles with medium-brown rims. 

Head width: 0.656 + 0.053 mm (25 larvae). 

Duration of stadium: 2.2 + 0.8 days (40 larvae). 

Fourth-Stadium Larva. Head varying from medium chocolate-brown through dark 
brown to black; dark mottling usually evident on lighter heads. Prothoracic shield 
medium to dark smoky-brown, with three longitudinal lines of pale yellow or grey. 
Suranal shield light to medium smoky-brown, commonly with three pale longitudinal 
lines. Mid-dorsal band pale yellow, yellowish-grey, or greenish-grey. Subdorsal 
area with a median band somewhat paler than mid-dorsal band, and marginal lines 
of pallid yellow. Supraspiracular area concolorous with median band of subdorsal 
area, with a median line of pale yellow. Spiracular band pale yellow or cream. Ventral 
region pale yellowish-grey. Spiracles with medium- to dark-brown rims. Thoracic 
legs light to dark smoky-brown. 

Head width: 1.16 + 0.07 mm (20 larvae ). 

Duration of stadium: 2.6 + 0.8 days (40 larvae). 

Fifth-Stadium Larva (igs. 5-8). Head shades of orange-brown, suffused and 
mottled with darker brown. Prothoracic shield varying from pale fawn marked with 
brown to uniform dark blackish-brown; shield with three longitudinal lines of pale 


VOLUME 26, NUMBER 1 33 


yellow. Suranal shield paler than prothoracic shield, with three longitudinal, cream 
or pale-yellow lines. Mid-dorsal band varying from pale yellow-fawn to dark smoky- 
brown. Subdorsal area consisting of a median band concolorous with or somewhat 
paler than mid-dorsal band, and marginal lines of pale yellow. Supraspiracular area 
concolorous with median band of subdorsal area. Spiracular band pale yellow, cream 
or sometimes almost white; often a pale smoky-brown line through middle of spiracular 
band. Suprapodal area pale-yellow or pale greyish-yellow. Mid-ventral area usually 
paler than suprapodal area. Spiracles with dark-brown or black rims. Thoracic legs 
varying from dull yellow to orange-brown, variably suffused with dark smoky-brown. 

Head width: 1.72 + 0.11 mm (23 larvae). 

Duration of stadium: 5.1 +1.2 days (40 larvae). 

Pupa (Figs. 2,9, 10). Spiracles borne in shallow depressions of the cuticle. Spiracu- 
lar sclerites narrow. Anterior marginal areas of abdominal segments 5, 6 and 7 con- 
spicuously pitted. Proboscis terminating at or slightly anterior to apexes of wings. 
Cremaster consisting of two rather short setae borne on a peculiarly shaped prolonga- 
tion of the tenth abdominal segment; prolongation of the tenth segment truncated in 
profile and usually flattened on the ventral surface. 

Length from anterior end to posterior margin of fourth abdominal segment: 6.17 + 
0.39 mm (25 pupae). 


ACKNOWLEDGMENTS 


I am grateful to Mr. John E. H. Martin of this Institute for the photo- 
graphs accompanying this paper, and to my associate Mr. Eric Rockburne 
for measuring larval structures and drawing the cremaster area of the 


pupa. 


LITERATURE CITED 


Grote, A. R. 1881. Moths collected by Prof. Snow in New Mexico, with list of 
Eudriini. Papilio 1: 174-178. 

Harpwick, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed 
species of Schinia (Lepidoptera; Noctuidae), with notes on the Heliothidinae. 
Can. Ent. Suppl. 6. 

SmiruH, J. B. 1906. New Noctuidae for 1906-No. 1. Jour. N.Y. Ent. Soc. 14: 9-30. 


PROTECTIVE FUNCTION OF SOUND PERCEPTION AND GREGARIOUSNESS 
IN HYLESIA LARVAE (SATURNIIDAE: HEMILEUCINAE ) 


While in residence at the Tropical Science Center field station on the Osa Peninsula 
of Costa Rica (1.8 miles west of Rincon), I was able to make some observations on a 
colony of Hylesia larvae which suggested a very probable function for their gregarious 
behavior and ability to perceive sound. 

I first discovered a large aggregation of these larvae (approximately 330 individuals ) 
in an oval mass on the trunk of a medium-sized tree, Trema micrantha (Linnaeus), 
on 3 April 1971. The mass was located on the north side of the tree about 1 m above 
the ground and was about 60 cm in length vertically and 18 cm in width. I acci- 
dentally became aware of the ability of the larvae to perceive sound when I shouted 
in their direction from a distance of about 10 meters. I was amazed to see the entire 


> 


surface of the mass “jump.” Each of the larvae responded to the sound of my voice 


34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


at the same instant and in the same manner, a violent jerking of the anterior third 
of the body, so that the head, thorax and anterior portion of the abdomen were arched 
upward or sideward. 

I tested this reaction numerous times and ways by altering the pitch and loudness 
of my voice and determined to my satisfaction that the action was due to sound and 
not to air movement. The larvae reacted only to very sharp and relatively high pitched 
sounds of high intensity. Normal conversation did not cause any movement. I further 
tested the response by playing music (Strauss waltzes) from a tape recorder in the 
immediate vicinity of the mass (within 1 meter) and noted that the larvae responded 
in the same manner to loud, sharp portions of the music. 

I observed the mass for 5 days. Each day I noted fewer individuals until the 8th 
of April, at which time there were no longer any present. Several larvae at the 
beginning had freshly molted, apparently transforming from the penultimate to the 
final instar. On several occasions during the day, larvae were seen on the ground 
crawling away from the tree, apparently in search of pupation sites or possibly food. 
The mass was present on the tree trunk only during the day, migrating at night to 
another place which was never determined. 

This jerking behavior of gregarious lepidopterous larvae in response to sound has 
been observed and recorded by only a few authors. Minnich (1936, J. Exper. Zool. 
72: 439-453) studied the reaction in the larvae of Nymphalis antiopa and records 
several other species of Lepidoptera which have been noted to behave similarly. 
None of these accounts, however, offers an explanation for the function of the behavior. 
Some additional observations which I made on this occasion suggested to me that 
sound perception coupled with the massing of these larvae served as a protection from 
parasites. 

The larvae were being parasitized by two species, a yellow and black chalcidine 
wasp and a tachinid fly. The former, on approaching the mass and hovering over it 
prior to landing and oviposition, elicited the jerking movement. The anterior portions 
of the caterpillars being thrown dorsally had the effect of warding or fending off the 
approaching wasps. The denseness of the hairs of the richly branched scoli furnished 
a barrier to the penetration of the parasites. The larvae in the center of the mass were 
especially well protected since their flanks were not vulnerable to lateral attacks. The 
tachinid flies did attempt to approach the larvae from the side by crawling along the 
bark but were warded off in a similar fashion by lateral jerking. When approaching 
the mass from above, these flies also elicited the dorsal arching. The high pitched 
whining of the wings of the approaching or hovering parasite seemed to be of the 
correct quality and intensity to elicit the jerking response. The effectiveness of this 
defensive maneuver is enhanced by the fact that the larvae are densely grouped and 
heavily clothed with spines and hairs. This behavior would offer much less protection 
to individual larvae than to larvae en masse. 

This protective behavior was not completely successful, however. I observed ovi- 
position by the chalcidine wasps numerous times. Still, the attacking females experi- 
enced great difficulty in gaining access to the host’s skin through the hair and spine 
network and violent defensive movements of the mass. 

The spines of the scoli are highly urticating to human skin and may also function 
more effectively in inhibiting attacks from vertebrates by being forced into the skin 
through this same jerking motion. 

A cinematographic record of these observations is on file with Alan Landsburg 

Productions, Hollywood, California (Production 1106-02, rolls 157, 158 and 166a). 

My thanks are extended to that organization for the opportunity to be in the study 

area and to Julian P. Donahue for assistance in identifying the caterpillars. Unfortu- 

aoe it was impossible to determine the species of Hylesia; several are common at 
1e site. 


Cuartes L. Hocur, Natural History Museum, Exposition Park Los Angeles 
California 90007. 


VoLUuME 26, NuMBER 1 35 


STUDIES ON THE CATOCALA (NOCTUIDAE) 
OF SOUTHERN NEW ENGLAND 
II. COMPARISON OF COLLECTING PROCEDURES 


CHARLES G, KELLOGG AND THEODORE D. SARGENT 


Department of Zoology, University of Massachusetts, 
Amherst, Massachusetts 01002 


Adult nocturnal Lepidoptera are collected by several very different 
methods. Brower (1947) has described some of the more common 
methods—baits (unless indicated otherwise, “baits” in this paper refer to 
some kind of sugar mixture) and lights during the night, and hunting 
resting moths during the day. Each of these collecting methods has been 
modified so that numerous light traps, bait mixtures, etc. are used. Re- 
cently parabolic moth sheets (McFarland, 1966) and collapsible bait 
traps (Platt, 1969) have been added to the list of collecting methods from 
which lepidopterists can choose. With growing interest and knowledge in 
insect pheromones, many entomologists are now using traps baited with 
virgin females cr synthetic sex pheromones (e.g. Saario, Shorey & Gaston, 
1970). Considered together with differences in time of collecting and 
differences in ecological placement of the trap, collecting procedures are 
widely divergent. 

How do these various collecting procedures compare when sampling 
the same area? Hamilton and Steiner (1939) and Robinson and Robinson 
(1950) have previously compared various trapping methods as to effec- 
tiveness of capture. The former, interested in controlling the Codling 
Moth (Carpocapsa pomonella (Linnaeus) ), a noctuid pest of orchards, 
compared bait and light traps and found that light traps captured more 
moths per trap, but that the percentage of females was much smaller than 
in the bait traps. The Robinsons have compared various light sources 
and suggest that the spectral content of the light is not important within 
limits, although the power and surface brightness of the source does affect 
trap efficiency. Others report that the kind of light (mercury, tungsten, 
etc. ) determines not only the species attracted, but also the sex (Edwards, 
1962). 

The importance of a trap’s location in sampling populations has also 
been noted. Hamilton and Steiner (1939) found that traps located at the 
margin (border rows) of an orchard averaged more than twice the num- 
ber of moths per trap than those located in the interior. Holbrook, Beroza 
and Burgess (1960) reported differences in effectiveness of pheromone- 
baited traps (Gypsy Moth—Porthetria dispar (Linnaeus) ) with terrain 


36 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


and local growth. Williams (1939) reported that an elevated light trap 
(35 feet above the ground) collected not only a greater number of species 
and individuals, but also a larger percentage of females than a trap lo- 
cated at ground level. This effect was stronger in some species than in 
others. Saario et al. (1970) confirmed differences in capture of one 
species (Trichoplusia ni (Hiibner) ) with pheromone-baited traps at dif- 
ferent heights from the ground. 

The time of collecting also appears to be an important factor. Williams 
(1935, 1939), Hamilton and Steiner (1939), Hutchins (1940), and Graham 
et al. (1964) have demonstrated differences in activity peaks related to 
time of night. These studies also suggest that activity patterns may be a 
function of temperature, humidity, wind, cloud cover, etc. However, 
Saario et al. (1970) could not find correlations relating daily capture 
variations (one species at pheromone trap) with nightly variations in tem- 
perature, relative humidity, or full moonlight. But Shorey (1966) has 
noted a greater range in copulation timing under naturally fluctuating 
outdoor conditions, and points out that some of these variables (e.g. 
humidity) are difficult to assess in nature because of great variation 
within microenvironments. Edwards (1962) and Saario et al. (1970) 
have shown that the time of median capture relative to sunset may vary 
with moth age and the season. 

The present study represents an attempt to compare collecting pro- 
cedures used in sampling members of a single genus (Catocala). Direct 
comparisons of collecting procedures at a single location (but not always 
the same season) are made by means of rank correlations of species. 
Rank correlations of species collected at different localities, using both 
similar and different collecting procedures are also listed, although they 
may be less meaningful than those obtained simultaneously at one loca- 
tion. 

Studies on Catocala are generally limited to the turn of the century 
(Bailey, 1877; Bunker, 1874; French, 1880; Johnson, 1880, 1882; Rowley, 
1908, 1909; Rowley and Berry, 1909-1914), but recently some comparisons 
within the genus have been made when different sampling methods were 
involved (Sargent and Hessel, 1970). Hopefully the data in this report 
may aid in interpreting prior comparisons, as well as comparisons which 
might be made in the future. 


METHODS 


A total of approximately 11,750 records of individual adult Catocala 
were obtained from four localities in southern New England: (1) West 
Hatfield, Mass. (1622 records, 1969-70, CGK); (2) Pelham, Mass. (544 


VOLUME 26, NUMBER 1 om 
TaBLE 1. Summary of collecting procedures and data. 

Time Time No. No. 

Area Year Method? Season Night? Indiv. Species 
West Hatfield 1970 RCN) 7/13-8/26 Dusk—Dawn 2, 24 
RUE (1B )© 8/23-9/26 Dusk—Dawn 692 26 
Bait 7/13-9/26 < 2400 hrs. 198 19 
1969 Bait 7/20-9/26 < 2400 hrs. 459 21 
Leverett 1970 RT 7/28-10/16 > 2300 hrs. 1161 Bul 
UV July—Oct. < 2300 hrs. 169 23 
Spots July—Oct. < 2300 hrs. 91 8) 
Bait July—Sept. < 2300 hrs. 85 8 
1969 UV July—Aug. < 2300 hrs. Al 18 
Spots July—Aug. < 2300 hrs. 36 10 
Bait July—Aug. < 2300 hrs. 188 13 
1968 Spots July—Sept. < 2300 hrs. 30 9 
Bait July—Sept. < 2300 hrs. 309 15 
1967 Spots July—Sept. < 2300 hrs. onli om 

& Bait 

Pelham 1966 Bait July—Sept. < 2300 hrs. 294 alk 
1965 Bait July—Sept. < 2300 hrs. 195 20 
1964 Bait July—Sept. < 2300 hrs. 50 16 
Washington 1970 RT Entire® Dusk—Dawn 886 30 
1969 RT Entire Dusk—Dawn 579 28 
1968 RT Entire Dusk—Dawn ADA 29 
Ley ~~ Wee Entire Dusk—Dawn IES 35 
1965 RT Entire Dusk—Dawn 553 32 
1964. RE ‘Entire Dusk—Dawn 530 30 
1963 RT Entire Dusk—Dawn 306 31 
1962 RT Entire Dusk—Dawn 1412 29 
1961 RT Entire Dusk—Dawn TS) 33 


2 RT—Robinson Trap; W—woods; F—field. 

> <—_before; >— after. Times are approximate as given and constitute the majority of the records. 
© Woods and field sites totaled had 965 individuals of 31 different species. 

4 Trap was operated both before and after seasons of Catocala flight. 


records, 1964-66, TDS); (3) Leverett, Mass. (2471 records, 1967-70, 
TDS ); (4) Washington, Conn. (7116 records, 1960-70, Sidney A. Hessel). 
Brief descriptions of these localities, and comments regarding collecting 
procedures in each, follow. A summary of collecting procedures and data 
is in Table 1. 


(1) West HATFIELD, Mass. 


Description: The West Hatfield (Hampshire County ) site lies 2.2 miles west of the 
Connecticut River and 5 miles north of Northampton. Farm, woods, and swampland 
lie between the site and the Connecticut River. To the west are the foothills of the 
Berkshires. Collecting was done in mixed deciduous woodlands in an area locally 
referred to as “The Rocks.” At the collecting site there is a 140 foot rise in elevation 
within 800 feet (U.S. Geological Survey Maps). 

Collecting procedures: Collecting was done at bait (a brown sugar-cooking wine- 


38 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


grape juice mixture painted on 20 trees along a trail at “The Rocks”) in 1969 and 
1970 on a daily basis from approximately 15 July to 20 September. Only nights of 
heavy rain, and about a half dozen single day absences, were missed for the two 
years. The mixture was usually applied fresh every night about a half hour before 
dusk, and trees were checked regularly every half hour until about midnight. On 
nights of much Catocala activity, the trail was checked more frequently and until 
0200 hours, at which time there usually appeared to be a lapse in feeding by the 
moths. Occasional collecting after midnight rarely added any individual records on 
nights of poor Catocala activity. Daily records were kept on the species, sex, and time 
of activity. 

In 1970 collecting was done at one Robinson mercury vapor light trap. The trap 
was turned on at dusk and left running until shortly after dawn at which time the 
contents were examined, and the species and sex of each individual Catocala recorded. 
The trap was operated every night, regardless of weather or absence. Attempts were 
made early in the season to check contents at intervals during the night, but the 
activity of the trapped moths made this unfeasible. 

From 13 July to 22 August, the trap was located in the woods about 30 yards from 
the wood’s edge atop a rock ledge. The trap could be seen for some distance within 
the woods, although view was restricted in some directions due to neighboring rock 
ledges. The trap was in view of nearly every tree on the sugar trail. From 22 August 
to 21 September the trap was located in a new situation about 250 yards to the 
northeast. Here the trap was in an open field about 50 yards from the edge of the 
main woods. A row of pine trees was immediately behind the trap. From 23-26 
August a second Robinson Trap was borrowed and traps were run at both locations 
simultaneously. 


(2) LEvERETT, Mass. 


Description: The Leverett (Franklin County) location is 4 miles east of the Con- 
necticut River and 7.5 miles northeast of the collecting site at West Hatfield (USGS 
Maps). Collecting was done on a level area at the top of a hill. At the fringe of the 
Pelham Hills, this area consists primarily of mixed deciduous woodland similar to 
that at West Hatfield. There is also some vegetation typical of earlier seral stages of 
old field succession within the area. 

Collecting procedures: Catocala were taken from 1967-70 at bait (brown sugar- 
beer mixture), at several 150-watt Westinghouse outdoor spotlights, and at rest. The 
data for sugar and lights were not separated in 1967. One 15 watt black light fluo- 
rescent tube was added in 1969 and 1970, and one Robinson mercury vapor light 
trap was added in 1970 (beginning 28 July). Collecting was done on a daily basis 
from 1 July to 1 September with only occasional 1-2 day absences. All collecting 
procedures (except bait) were continued until mid-October in 1970. 

The Robinson Trap was located in a small open area only 10 yards from the edge 
of the woods. It was shaded through approximately 90 compass degrees by a house 
but was visible from all trees on the sugar trail. Records for the trap were kept Seay 
night except for five days in late August when the trap was being used in West 
Hatfield. The trap was running only from 2300 hours to dawn until 10 September 
when it was left running from dusk to dawn for the remainder of the season (also 
running all night on 28-29 August when CGK kept the records for the Leverett 
location). The bait trail was checked and the other light sources were usually kept 
running until 2300 hours. Sex and time of activity data were kept beginning in 1968. 


(3) PELHAM, Mass, 


Description: The Pelham (Hampshire County) site, which is 2.5 miles east of 
Amherst, is 4.2 miles south-southeast of the Leverett site, and 8.5 miles east of the 
West Hatfield site (USGS Maps). Collecting was done in an acre of woods in a 


VoLuME 26, NumMBER | 39 


residential area. The soil is sandy and vegetation is that found in a pitch pine (Pinus 
rigida Mill.) community. Ornamental trees and shrubs, including various Rosaceae 
and honey locust ( Gleditsia triacanthos L.) are common. 

Collecting procedures: Collecting was done during the summers of 1964-66 and 
was limited to bait. Individuals were not routinely sexed, and times of capture were 
not noted. 


(4) WaAsHINGTON, CONN. 


Description: This collecting area ( Litchfield County ) has been described in Sargent 
and Hessel (1970). The mixed deciduous woodlands here have more Juglandaceae 
representatives, particularly butternut (Juglans cinerea L.), than localities in the 
Amherst area 65 miles to the Northeast. 

Collecting procedures: Records of Catocala here were predominantly taken at one 
Robinson mercury vapor light trap with a few records taken at one 15 watt black 
light fluorescent tube. The lights were in operation all night from mid-March to 
mid-November of each year, with continuous records provided from 1961-70 (except 
1966). Data on sex were not routinely kept. 


The Catocala were identified as keyed and described in Forbes (1954) 
and foodplants were also taken from that source. Certain similar species 
(C. gracilis Edwards, C. sordida Grote; C. crataegi Saunders, C. mira 
Grote, C. blandula Hulst) were not always identified as to species. C. 
gracilis and C. sordida were distinguished by TDS and CGK in 1970 and 
C. crataegi, C. blandula and C. mira by CGK in 1969-70 (collected speci- 
mens only ), TDS in 1970 and S. A. Hessel since 1961. 

The rank correlation of species was found using the Spearman test 
(Siegel, 1956) with the correction factor for tied ranks being used in every 
case. The Spearman coefficient was also used to find the probability 
under a Student's t distribution that correlations between ranks were due 
to chance. In cases where certain species were not distinguished in one of 
the samples being compared, a single rank was given to the total number 
of individuals of these species in both samples. 


RESULTS AND DISCUSSION 
Comparison of Collecting Procedures 


The rank correlations of species collected at a single location by dif- 
ferent procedures, as well as by similar procedures in different years, are 
listed in Table 2. Whenever possible, correlations (of different proce- 
dures) are from comparisons of samples taken during the same season. 
The correlations are grouped according to location—West Hatfield, 
Leverett, and Washington—and are arranged within each group in de- 
creasing order of similarity of species ranks. 

The following observations derived from data in Table 2 seem most 
interesting. At both Leverett and West Hatfield, there were higher 
degrees of similarity between samples taken at bait in two consecutive 


40 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


TaBLE 2. Rank correlations of species across different procedures (or seasons) at a 
single locality. 


Procedures (or Seasons) 


Compared 

hee A B N r, 2, t P, 

W. Hatfield 1969 1970 we} 653 < Ol 3.92 < .0005 
Bait 

W. Hatfield RT Bait Be) A69 < .O1 2.91 < .005 
1970 

Leverett Spots UV Tl 841 < il Gell < .0005 

1969-70 
Leverett UV, Spots RT 29 778 < {Ol 6.46 < .0005 
1968-70 1970 

Leverett 1969 1970 14 OT < .05 2.28 < .025 
Bait 

Leverett UV, Spots Bait 28 oon Ss (U5 1.83 <<—.05 

1968-70 
Leverett RT Bait 29 238 05 1.29 > .10 
1970 1968-70 

Leverett Light Bait 30 193 > (05 1.04 lO 
1970 (total ) 

Washington 1965 1969 33 831 < Ol 8.31 < .0005 
RT 

Washington 1963 1967 36 ipl, < Ol 5.91 < .0005 
RT 


N—Number of different species for combined procedures. 
r.—Spearman Rank Correlation Coefficient. 
P,. —Probability associated with Spearman Coefficient. 


8 
t—Student’s t value. 
P,—Probability associated with Student’s t value. 


years than there were between bait and lights within a single year. The 
correlation was statistically significant for West Hatfield but only margin- 
ally so for Leverett. At Leverett the highest correlation occurred between 
samples taken at spot lights and a UV light summed over two years, with 
samples taken at UV and spots combined also showing a high correlation 
with the Robinson Trap. This is not surprising if the spectral composition 
of a light source is not important (Robinson and Robinson, 1950). 

The higher degree of similarity between spots and UV, if significant, 
may be explained by the simultaneous operation of these light sources, 
while the Robinson Trap was sampling activity during a different time of 
the night. The higher degree of similarity between UV and spots com- 
bined and bait, than between the Robinson Trap and bait may be similarly 
explained. 


VoLUME 26, NUMBER 1 


Al 


TABLE 3. Rank correlations of species across different localities using similar and 
different collecting procedures. 


Procedure Area A Area B N i Po t Py 
Lights WH L 31 843 <0): 8.32 < .0005 
1970 
RT—1970 WH L ol 811 <a Oil 7.46 0005 
Totals WEL L 33 193 < dl 7.25 0005 
1970 
Lights WH L 25 851 <i Hell < .0005 
7/13-8/22 ( woods ) 
1970 
Lights WH L 29 .669 < {Hl 4.68 < .0005 
8/27-9/21 ( field ) 
1970 
Bait WH L 25 A95 <0 2.73 <0 
1968-70 
Bait le L 26 715 <en Oil! 5.01 < .0005 
1964—66 1968-70 
Bait WH Ie 30 .262 > .05 1.44 <0 
1969-70 1964—66 
RT WH W 35 049 <A OII! 3.80 < .0005 
1970 1967 
1970 WH&L W 35 A92 <0) 3.25 < .005 
RT—1970 WH W 33 O01 << Oil 3.23 < .005 
Lights Is W 34 A44 < 2.80 < .005 
1970 
RT—1970 L W 32 435 <A 2.65 << Oil 
Totals WHEL W 36 349 PAT < .025 
to 1969 
1970 WHEéL W 37 231 1.40 <0) 
No RT 


Symbols same as in Table 2. 
Localities are designated as follows: Leverett (L), Pelham (P), West Hatfield (WH) and Wash- 
ington (W). 


It is equally interesting that a very high degree of correlation occurred 
between two separated years of sampling by a Robinson Trap at a single 
location (Washington ), even when there was nearly a fourfold difference 
in total moths taken (e.g. 1967 vs. 1963). 


42, JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Comparison of Localities 


Table 3 lists rank correlations of Catocala species collected at different 
localities, using both similar and different collecting procedures. The 
interpretation of these correlations may be somewhat equivocal. When 
comparing across localities (and seasons ), presence or absence of correla- 
tion might be due to either similarities or differences in 1) collecting pro- 
cedures, or 2) the populations being sampled. 

Samples taken at light sources at West Hatfield and Leverett during 
the same year appear to be statistically identical. This is not surprising 
since the vegetation is quite similar at the two locations. However this 
identity is barely maintained when comparing the populations taken at 
bait from the same two locations. 

The populations sampled by light at Washington and both localities in 
the Amherst area during the same year have a high degree of similarity, 
although less than that between West Hatfield and Leverett. The de- 
crease in similarity is not surprising, considering the greater abundance 
of Juglandaceae species as well as the singular presence of bayberry 
(Myrica pensylvanica Loisel) at Washington. However, the similarity 
existing in populations is not as evident when different collecting proce- 
dures have been used over different years (combined data before 1970 
which is presented by Sargent and Hessel, 1970, as well as 1970 analysis of 
combined Amherst areas without Robinson Trap vs. Washington ). 

Pelham and West Hatfield both seem to be better baiting areas than 
Leverett (number of species as well as number of individuals) although 
the high degree of similarity at bait occurs between Pelham and Leverett, 
with Pelham and West Hatfield having a correlation which is not even 
statistically significant. This can apparently be explained by a disparity 
between moths within the major foodplant groups which are active at 
bait in these localities—Salicaceae and Juglandaceae feeders predomi- 
nating at West Hatfield, and Rosaceae and Ericaceae feeders predomi- 
nating at Pelham, and to a great extent at Leverett as well. Since species 
within these foodplant groups are relatively similar in abundance at light 
in West Hatfield and Leverett, this further suggests that collecting at bait 
and light measures different kinds of activity. 

It is impossible to determine which collecting procedures give a better 
representation of the population present in a locality without actually 
knowing what this population is, independent of the collecting procedures. 
Since all collecting procedures appear to miss species which appear to be 
reasonably common in a given locality (e.g, C. cara Guenée and C. 
amatrix (Hubner) at light and several Juglandaceae feeders at bait), 
a combination of collecting procedures would seem to be the best choice 


VoLuUME 26, NuMBER 1 43 


when sampling Catocala populations. Generally speaking, light trap sam- 
ples seem to be more consistent from year to year and yield larger samples 
(both in terms of species and individuals ) than other procedures. 


Location of Robinson Trap 


The location. of the light trap appears to be an important consideration 
if the researcher is interested in assessing the number of moths in a given 
area. Data to support this contention were obtained when the trap oper- 
ated in West Hatfield was moved from a woods to a field site in the mid- 
dle of the 1970 season. The trap operated at Leverett was kept at the 
same location for the entire season and thus served as the control in this 
experiment. Since the effective range of the mercury vapor lamp trap is 
reputed to be about 100 yards (Robinson and Robinson, 1950), the nor- 
mal movement of adult Catocala to be greater than 200 yards (Brower, 
1930), and the vegetation of the woodlands to be roughly homogeneous 
across both West Hatfield trap sites, one can assume that the populations 
being sampled at the two West Hatfield trap sites were virtually identical. 

From Table 3 it can be seen that samples from both West Hatfield sites 
shared a close correlation with the Leverett trap, a fact which was not 
surprising considering the wooded location of the control area. However, 
more surprising was the difference in numbers captured at the two loca- 
tions. Because Catocala are active in the woods, one might intuitively 
expect that a trap located there would collect more moths than the same 
trap located in a field 50 yards from the wood’s edge. However, Fig. 1, 
which graphically depicts the average number of species and individuals 
captured per day over five-day intervals during the 1970 season, reveals 
that the trap located in the field was much more effective than the same 
trap located in the woods. 

Because the control trap was operating continuously and simultaneously 
with the experimental trap and showed no major seasonal differences in 
moth activity, it dramatically reinforces the conclusion that seasonal dif- 
ferences are not important factors in the increased effectiveness at the 
field site. The four-day interval when traps were operated at both wood 
and field sites lends further support. It is interesting that the field and 
control traps show nearly identical curves for activity of species and indi- 
viduals. The greater effectiveness of the field trap over the control in the 
mid-portion of the season can be explained both by the greater abundance 
of a single species (C. ilia (Cramer) ) at the West Hatfield location, and 
a difference in collecting procedures (field trap was operated all night 
whereas the control trap was operated only after 2300 hours). The near 
identity of activity in the field and control traps during the latter portion 


44 JourNAL OF THE LEPIDOPTERISTS’ SOCIETY 


@—e LEVERETT 

0o---0 W. HATFIELD 
(Woods) 

o---0 W. HATFIELD 
(Field) 


74) 

Lid 
OG 
we 
fe) 
= 74) 
=) 
Zz 
> 
al 
ft 
a 40 B 
i - 
O 
<x 
neo 
Lid 
a 
Tse 

~ 

= 26 

> 

a) 

Zz 

=r £0 

Qa. 
oO” 
7/28- 8/22 - 9/16- 
8/1 8/26 9/20 
FIVE-DAY PERIODS 
Fig. 1. Average daily capture of Catocala species (A) and individuals (B) in a 


Robinson Trap over successive 5-day periods at three locations in 1970. 


VoLUME 26, NUMBER 1 45 


W. HATFIELD 


MB Leverett 


300 A 
200 
(Va) 
= 
<x 
= 
Q 100 
> 
a 
Zz 
a F S J E 
u 300 
Oo 
72) 
ce 
w 200 
o . 
= 
=) 
Zz 


100 


FOODPLANT GROUPS 


Fig. 2. Numbers of individuals of each Catocala foodplant group captured in 
Robinson Traps in West Hatfield (A, woods, 13 July—26 August; B, field, 22 August—21 
September) and Leverett (A, 28 July-26 August; B, 22 August-21 September) in 
1970. Foodplant groups are: Fagaceae (F), Salicaceae (S), Juglandaceae (J), 
Ericaceae (E), Myricaceae (M), and Rosaceae (R). 


of the season also corresponds to the period when both traps were oper- 
ated all night. 

Examination of Fig. 2 points out that the increased effectiveness of the 
field over the woods site relative to the control is true for all foodplant 
groups with the exception of the Myricaceae. Also of the twenty species 
which were active both before and after the experimental trap was moved, 
80% (i.e., all but four) had an increase in numbers relative to the control 
when the trap was relocated in the field. 

One naturally wonders why the trap located in the field was more effec- 
tive than the same one located in the woods. The answer may be found 
with reference to the work of Robinson and Robinson (1950). They con- 
cluded that a light source is not an attractant to moths; rather activity at 
light merely reflects activity of moths within the “inner dazzle sphere” of 


A6 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


FOODPLANT GROUPS 
ies (a ae *# SALICACEAE 
o——e JUGLANDACEAE 
o—-—e FAGACEAE 
o-—---e ROSACEAE 


PERCENT OF TOTAL CAPTURED 


1969 1970 1970 1969 1970 1970 
BAIT BAIT LIGHT BAIT BAIT LIGHT 


Fig. 3. Percentages of total individuals of some “common” Catocala species cap- 
tured at bait and a Robinson Trap in West Hatfield in 1969 and 1970. Species 
fluctuating greatly with collecting procedures (to the left) include: 1) ilia, 2) cara, 
3) amatrix, 4) amica, 5) habilis, and 6) palaeogama. Species relatively constant across 
collecting procedures (to the right) include: 7) retecta, 8) ultronia, 9) concumbens, 
10) epione, and 11) grynea. 


the light source. If this is true, one would suspect that large trees and rock 
ledges around the trap would form shadow cones within the “dazzle 
sphere,” enabling moths to escape. The shadow areas cast in this zone 
from the trap located in the field are much smaller and might very well 
account for the difference in effectiveness. The control trap was located 
near the wood’s edge, and it might be expected that shadows interrupting 
the “dazzle sphere” of this trap were not as great as those cast by the trap 
in the woods, and might have been intermediate between those cast by 
the traps at the field and woods sites. Attractant theories may also pro- 

ide an answer if one views shadow cones as breaking areas of attraction. 


VoLUME 26, NuMBER 1 A7 


Williams (1939) has commented on the surprising success of a trap in an 
open and unsheltered location previously thought not to be a particularly 
good location for insects. 


Collecting Procedure Comparison by Species 


An examination of the samples taken by various collecting procedures 
suggested that species feeding on certain foodplants were much more com- 
monly collected by one procedure than another. Further, relative differ- 
ences in activity at bait and light were not uniform for all species feeding 
on the same foodplant. These differences are graphically presented in 
Fig. 3. Data used in this figure are from the records of CGK from West 
Hatfield. 

In general, it appears that very different collecting procedures, e.g., 
bait and light, are not sampling identically from the same population. 
Further, there appears to be consistency (in terms of species rank correla- 
tions ) across seasons in samples taken by the same collecting procedures. 

Fig. 3 illustrates the percentage of total Catocala collected for several 
common species within the genus. Generally, percentage differences 
between two years at bait are minimal for each species, whereas differ- 
ences in the same year between bait and light are often dramatic. For 
example, C. ilia comprised about 50% of all Catocala collected at bait 
(and greater than 90% of all Fagaceae feeders at bait), but was domi- 
nated by another Fagaceae feeder, C. amica (Hubner), at light. Similarly 
C. cara comprised nearly 50% of all Salicaceae feeders at bait, yet was 
uncommonly collected at light. On the other hand, C. concumbens 
Walker, which was relatively uncommon at bait, was the most common 
Salicaceae feeder in the light trap. Common species within the same 
foodplant group (defined as all Catocala feeding on that foodplant), 
generally appear to parallel each other in activity at bait and lights (e.g., 
C. cara and C. amatrix; C. paleogama Guenée and C. habilis Grote). 
Three Ericaceae feeders, C. gracilis, C. sordida and C. andromedae 
(Guenée ), were commonly taken at lights, but only one individual of the 
three species was taken at sugar for the two years. These results can be 
essentially replicated with data collected by TDS in Leverett. 


SUMMARY 


Catocala populations taken by various collecting procedures at four 
localities in southern New England were compared using species rank 
correlation coefficients. Highest correlations were obtained when the 
same procedure was used and when collecting was done during the same 
time of night. These correlations were greater for light sources than for 


48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


baits. A comparison of samples taken in a light trap located at a field 
and woods site showed a similar degree of correlation to a control, but 
there was much greater effectiveness of the trap at the field site. 


ACKNOWLEDGMENTS 


This study was done in partial fulfillment of the Master's degree require- 
ments by the first author, with funds provided by a National Science 
Foundation Traineeship. 

The many records generously provided by Sidney A. Hessel from 
several years collecting at one site using the same collecting procedure 
are gratefully acknowledged. 

Sincere thanks are also given to Yale University for loan of one of their 
Robinson Traps, and to L. Vollinger and K. Fisher for use of their land 


for collecting. 


LITERATURE CITED 


BamLEy, J. S. 1877. Catocalae taken at sugar at Center, N.Y. Canad. Ent. 9: 215— 
218. 

Brower, A. E. 1930. An experiment in marking moths and finding them again 
(Lepid.: Noctuidae). Entomol. News 41: 10-15. 

1947. Methods for collecting underwing moths (Catocala). The Lepi- 
dopterists’ News 1: 19-20. 

Bunker, R. 1874. Notes on collecting Catocalas. Canad. Entomol. 6: 25-26. 

Epwarps, D. K. 1962. Laboratory determinations of the daily flight times of 
separate sexes of some moths in naturally changing light. Can. J. Zool. 40: 511— 
530. 

Forses, W. T. M. 1954. Lepidoptera of New York and Neighboring States. III. 
Noctuidae. Cornell Univ. Agric. Exp. St., Memoir 329, 433 pp. 

FRENCH, G. H. 1880. Notes on Catocala hunting. Canad. Entomol. 12: 241-242. 

GraHaM, H. M., P. A. Gricx, anp D. F. Marrin. 1964. Nocturnal activity of 
adults of six lepidopterous pests of cotton as indicated by light-trap collections. 
Ann. Entomol. Soc. Amer. 57: 328-332. 

Hamitton, D. W. ann L. F. STErNER. 1939. Light traps and codling moth control. 
J. Econ. Entomol. 32: 867-872. 

Horsrook, R. F., M. Beroza, anp E. D, Burcess. 1960. Gypsy moth (Porthetria 
dispar) detection with the natural female sex lure. J. Econ. Entomol. 53: 751- 
756. 

Hurcnins, R. E. 1940. Insect activity at a light trap during various periods of the 
night. J. Econ. Entomol. 33: 654-657. 

eon, J. S. 1880. Early appearances of Catocalas. Canad. Entomol. 12: 137— 


1882. Catocalae taken in the vicinity of Frankford, Pennsylvania. Canad. 
Entomol. 14: 59-60. 

McI’artanp, N. 1966. A moth sheet. J. Res. Lepid. 5: 29-36. 

we A, tee A lightweight collapsible bait trap for Lepidoptera. J. Lepid. Soc. 
23: 97-101. 

Ropinson, H. S. AND P. J. M. Ropinson. 1950. Some notes on the observed behav- 
ior of Lepidoptera in flight in the vicinity of light sources together with a descrip- 


tion of a light-trap designed to take entomological samples. Entomol. Gaz. 1: 
3-20. 


VOLUME 26, NUMBER 1 A9 


Row ey, R.R. 1908. Notes on Catocala. Entomol. News 19: 115-120. 

1909. Another season with Catocalae. Entomol. News 20: 127-135. 

Row.ey, R. R. anp L. Berry. 1909. Notes on the study of some Iowa Catocalae. 
Entomol. News 20: 12-18. 

1910. Notes on the life stages of Catocalae; a summer’s record and inci- 

dental mention of other Lepidoptera. Entomol. News 21: 441-455. 

1910. Further study of the Catocalae. Entomol. News 21: 104-116. 

1912. A dry year’s yield of Catocalae (Lepid.) 1911. Entomol. News 23: 

207-214. 

1913. Last year’s work with Catocalae and other Lepidoptera. Entomol. 

News 24: 197—205.. 

1914. 1913 as a Catocala year (Lepid.). Entomol. News 25: 157-167. 

Saario, C. A., H. H. SHorEy, AND L. K. Gastron. 1970. Sex pheromones of 
noctuid moths. XIX. Effect of environmental and seasonal factors on captures 
of males of Trichoplusia ni in pheromone-baited traps. Ann. Entomol. Soc. Amer. 
63: 667-672. 

SARGENT, T. D. AND S. A. HEssEL. 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. 

Suorey, H. H. 1966. The biology of Trichoplusia ni (Lepidoptera: Noctuidae). 
IV. Environmental control of mating. Ann. Entomol. Soc. Amer. 59: 502-506. 

SIEGEL, S. 1956. Nonparametric Statistics for the Behavioral Sciences. New York: 
McGraw-Hill Book Company. 312 pp. 

WitiiaMs, C. B. 1935. The times of activity of certain nocturnal insects, chiefly 
Lepidoptera, as indicated by a light trap. Trans. Roy. Entomol. Soc. London 83: 
523-555. 

1939. An analysis of four years captures of insects in a light trap. Part 1. 

General survey; sex proportion phenology; and time of flight. Trans. Roy. 

Entomol. Soc. London 89: 79-131. 


THREE BUTTERFLY SPECIES (LYCAENIDAE, 
NYMPHALIDAE, AND HELICONIIDAE) NEW TO 
TEXAS AND THE UNITED STATES?! 


Roy O. KENDALL? 
Route 4, Box 104-EB, San Antonio, Texas 78228 


The author (1970) gave five species of Rhopalocera new to Texas and 
the United States which were probably introduced through Hurricane 
Beulah of 1967. Three more species are now added. Time and additional 
research would be required to determine the specific ecological factors 
influencing permanent residence should any of these become established 
north of the Rio Grande. 


1 Contribution No. 206. Bureau of Entomology, Division of Plant Industry, Florida Department 
of Agriculture and Consumer Services, Gainesville. 

2 Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, Florida 
Department of Agriculture and Consumer Services. 


50 


JOURNAL OF 


THE LEPIDOPTERISTS SOCIETY 


Figs. 
male, 


Dorsal and ventral surfaces of 


VoLUME 26, NuMBER 1 pill 


Strymon albata sedecia ( Hewitson ) 


Thecla sedecia Hewitson 1874, Ent. Mo. Mag. 11: 105 (TL: “Mexico”); ibid. 
1877, Ill. Diurn. Lep. Lycaenidae: 198, pl. 18 figs. 637, 638; Godman & Salvin 
1887, Biol. C.-Amer., Rhop. 2: 94: Draudt 1920, in Seitz, Gross-schmett. Erde 5: 
808, pl. 159 k; Hoffmann 1941, An. Inst. Biol. Mexico 11: 719 (#744); Comstock 
& Huntington 1963, J. New York Ent. Soc. 71: 116. 

Strymon albata sedecia: Clench 1967, J. Lepid. Soc. 21: 183 (TL: restricted to 
Mazatlan, Sinaloa, Mexico). 

This subspecies, formerly known only from Mexico and Guatemala, 
is now represented by four examples from Texas. Three of these, in fair 
to poor condition, were taken by Michael A. Rickard at the Santa Ana 
National Wildlife Refuge, Hidalgo Co.: 9 November 1968 (1), 24 Novem- 
ber 1968 (1), and 14 December 1968 (1). The latter male is in the 
author's collection. The fourth example, a fresh male (Figs. 1 & 2), was 
taken by John E. Hafernik, two miles east of Brownsville, Cameron Co., 
10 July 1970. 

Based on these data, this species seems to be established in Texas at 
least temporarily. It may have been introduced by Hurricane Beulah of 
1967. Once we have reared it and know more about its life history a 
more precise evaluation can be made of its residence status. 


Chlosyne rosita browni Bauer 
Chlosyne rosita Hall, 1924. Entomologist 57: 241-243, 2 figs. TL: Western Guate- 


mala. 
Chlosyne rosita browni Bauer, 1960. J. Lepid. Soc. 14 (2): 148-154, 2 figs. TL: 
E] Salto, San Luis Potasi, Mexico. 

This subspecies was described from 36 specimens from Mexico (San 
Luis Potosi, Nuevo Leon, and Tamaulipas). It is now represented in the 
United States by a good series (42¢, 9?) of field-caught specimens plus 
nine (56, 42) reared examples, all from Santa Ana National Wildlife 
Refuge, Hidalgo Co., Texas. Bauer (1960), in describing this subspecies, 
suggested that it might be found in the United States. 

On 24 October 1970, the author and Mrs. Kendall joined our very good 
friend Dr. J. W. Tilden at the Santa Ana National Wildlife Refuge for a 
weekend of collecting. On this day while collecting near the Dicliptera 
Trail, the author saw an adult but missed catching it. A few minutes later 
Mrs. Kendall took a fairly good male (possibly the same specimen). On 
the following morning, the insect was common along the Dicliptera Trail. 
Fourteen examples (136, 12) were collected. On 26 October, same 
location, we took seventeen more (13¢ 42). 

Adults collected were mostly worn and were found mainly at two spots 
along the trail where Dicliptera brachiata (Pursh) Spreng var. alternata 
Gray and Dicliptera vahliana Ness (Acanthaceae) grow. At one spot both 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


24 MM 


45 mM 


= 
= 
NY 
NY 


VoLUME 26, NuMBER 1] 53 


plant species were growing together. The first named is a perennial, the 
latter an annual. At the last spot, in particular, males were observed 
flying back and forth low over these plants, apparently searching for 
virgin females. The females taken were sitting on these plants, leisurely 
flexing their wings in a preoviposition manner. 

All four females taken 26 October were kept alive for egg production in 
the laboratory. Female Nos. 1, 2 and 3 each deposited a single multi- 
cluster of eggs beneath a leaf of D. alternata 27 October; females 1 and 
2 were then placed in a killing jar. Female No. 3 deposited another 
cluster on 29 October and two more small clusters 31 October. No. 4 
female deposited clusters 28 and 30 October. Both remaining females 
were placed in the killing jar 31 October. All egg clusters were deposited 
beneath leaves in a typical Chlosyne/Phyciodes manner. 

Eggs started hatching about 0600 on 5 November. Most first instar 
larvae were lost, reason undetermined, but the absence of fresh foliage 
suspected. A few remaining larvae were offered fresh foliage of Siphono- 
glossa pilosella Torr., which they readily ate. Later instars accepted D. 
alternata, the most likely preferred oviposition substrate. Pupation oc- 
eumed-o Wil (3), 7-XIl (1), 8-XII (3), and 10-XII (3). Adults 
emerged: 13-XII (26, 12), 14-XII (1¢), 15-XII (2¢), 17-XII (22), 
and 18-XII (12) for a total of 56, 42. Preserved immatures include 
one small cluster of eggs, all first instar larvae that died, one last instar 
larva, and one pupa. Also, one small cluster of eggs was furnished Dr. 
Alvah Peterson for color photographing. 

Following our lead, Tilden collected the same area during the period 
25 October-17 November and took twenty (16¢, 42) examples. Those 
taken near the end of the period were badly wom, and several additional 
examples were released after capture. At the time we collected these 
specimens, we were under the impression they represented Chlosyne 
janais Drury. After returning to the laboratory and reexamining the 
specimens, the author realized this insect was not C. janais. Tilden kindly 
paid us a departing visit on 18 November at which time we discussed the 
matter. Upon returning to California, Tilden soon discovered by checking 
the literature and specimens in the California Academy of Sciences Mu- 
seum that our insect was C. r. browni. He also discovered among his 
catch three examples (24, 12) of C. janais which had been taken in 


< 


Figs. 3-4. Chlosyne rosita browni Bauer. Dorsal and ventral surfaces of male 
and female. 


Figs. 5-6. Chlosyne janais Drury. Dorsal and ventral surfaces of male and female. 


54 JourNAL oF THE LEPIDOPTERISTS SOCIETY 


Figs. 7-8. Eueides cleobaea zorcaon (Reakirt). Dorsal and ventral surfaces of 
female. 


another part of the Refuge. On the chance this confusion may have hap- 
pened to other collectors, the two species are illustrated (Figs. 3-6). 
The similarity is sufficiently great as to suggest a mimic/model relation- 
ship. Superficially, the larvae are more easily separated than the adults, 
although there should be no confusion when the adults of each are 
compared. 
Based on available collection records, it would appear this subspecies 
perhaps four broods. It probably has an inherent larval diapause, 


VoLUME 26, NuMBER 1 55 


manifested in a few immatures only, which may be triggered by tempera- 
ture. Additional field research is necessary to determine this. Further, it 
is unknown whether the species is a permanent resident in Texas. 


Eueides cleobaea zorcaon ( Reakirt ) 


Eueides Hiibner, 1816. Verz. bekannt. Schmett. (1): II; type species Nereis dianassa 
Hiibner [1806]. Samml. exot. Schmett. 1: pl. [8], selected by Scudder, 1875, 
Proc. Acad. Arts Sci., Boston 10: 169. 

Eueides zorcaon Reakirt, 1866. Proc. Acad. Sci. Phila. V. [18], 243, no. 12. (TL: 
Near Vera Cruz, Mexico); Fassl, A. H., 1909, Jugendzustande Trepischer Tag- 
falter. Soc. Ent. 24: 105-107 (mentions larval foodplant, egg and larva); Seitz, 
A., (1913), Vol. 5, p. 398 [pl. 80g], (ranging through Central America as far 
west as Mexico); Wolcott, G. H., 1923, Insectae Portoricensis, J. Dept. Agr. 
Puerto Rico 7 (1): 1-318 (records Passiflora sp. as larval foodplant); Seitz, A., 
1924, Die Gross-Schmetterlinge der Erde. Vol. V. Die Amerikanischen Tagfalter. 
Stuttgart, 1141 pp. (describes egg, generic larva, and records Passiflora sp. as 
larval foodplant). Wolcott, 1936, Insectae Portoricensis, J. Dept. Agr. Puerto 
Rico 20: 1-627 (again records Passiflora as larval foodplant). 

Eueides cleobaea zorcaon: Neustetter, H., 1929, Lepidopterorum Catalogus (makes 
zorcaon a subspecies of cleobaea Geyer); Hoffmann, C., 1940, Catalogo Siste- 
matico y Zoogeografico de los Lepidopteros Mexicanos. An. Inst. Biol. Mex. 11 
(2): 639-739 (gives range as warm and hot regions of the Gulf Coast and 
Chiapas); Ross, Gary N., 1964, Life history studies on Mexican butterflies, J. 
Res. Lepid. 3 (4): 207-229 (describes the egg, larva [1-5 instars], and pupa; 
illustrates, in black and white, adult [upper side], egg, 5th instar larva, head 
capsules of Ist and 5th instar larvae, and pupa [dorsal and latteral views] ). 


This subspecies, previously known only from Central America and 
Mexico, is now represented in the Nearctic fauna by 13 examples, all 
from Texas. One of these is illustrated (Figs. 7-8). It would appear 
from collection dates that zorcaon has perhaps three broods when, from 
time to time, it becomes temporarily established north of the Rio Grande. 

In 1966 the author examined three examples of this subspecies in the 
Panther Junction Museum, Big Bend National Park, Brewster Co., Texas. 
They were collected in the park by Rollin H. Baker, 15 July 1937 (1) 
and 22 July 1937 (2). 

In 1968, thirty years later, this insect was distributed over a wide area 
in Texas, but it has not been seen since. Its reintroduction and temporary 
residence probably resulted from Hurricane Beulah of 1967. Known ex- 
amples collected in Texas during 1968 are: Near Skidmore, Bee Co., 
20 April (3), leg M. A. Rickard; same location and date, (2), leg Roy 
Jameson; two other specimens were sighted at this location. On the same 
day Rickard saw one example at Lake Corpus Christi State Park, San 
Patricio Co. San Antonio, Bexar Co., 21 April (12 ), leg Glenn Y. Belyea. 
Big Bend National Park, Brewster Co., 11 June (1), leg David A. Easterla; 
this specimen was feeding on blossoms of Acacia greggi Gray growing in 
the Chisos Mountains basin. McAllen, Hidalgo Co., 4 May (22), leg Dr. 


56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


J. Bolling Sullivan III. Santa Ana National Wildlife Refuge, Hidalgo 
Co., 24 June (sight record), 25 June (22 ), leg J. R. Heitzman. 


ACKNOWLEDGMENTS 


I wish to thank William T. Krummes, regional director, Bureau of Sport 
Fisheries and Wildlife, U.S. Department of the Interior, Albuquerque, 
New Mexico, for providing the necessary permits to conduct basic re- 
search on the Lepidoptera in the Santa Ana National Wildlife Refuge. I 
also wish to thank Carrell Ryan and Wayne a Shifflett of the Santa Ana 
Refuge for their excellent cooperation in the conduct of this research. 

I am also indebted to Harry K. Clench, Cyril F. dos Passos, William 
D. Field, Paddy McHenry, Dr. J. W. Tilden, and Dr. Howard V. Weems, 
Jr., for providing certain references and for reviewing the MS in whole 
or in part. Special thanks go to my good friend and fellow lepidopterist 
André Blanchard for taking time from his own research on Texas Lepi- 
doptera (Heterocera) to photograph the species here recorded and illus- 
trated. 


LITERATURE CITED 


KENDALL, Roy O. 1970. Three hairstreaks (Lycaenidae) new to Texas and the 
United States. J. Lepid. Soc. 24 (1): 59-61. 

1970. Lerema ancillaris (Hesperiidae) new to Texas and the United 

States. J. Lepid. Soc. 24 (4): 266. 


MORE NEW MOTHS FROM TEXAS (NOCTUIDAE) 


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


This is a follow-up of the Blanchard 1966, 1968 and 1970 papers. 


Oncocnemis cottami, A. Blanchard, new species. 


Head: Black; scales on upper half of front, vertex, and base of antennae long, raised 


and whitish tipped; antennae simple; palpi rough scaled, whitish basally, blackish 
distally; second segment longest, not quite reaching middle of front; third segment 
very short, porrect. 

Collar: Black basally, dark gray on top; the white band in between is itself divided 
in two by a thin black line. 

Thorax: Disc and patagiae brownish gray; sordid white below; foretibia short, 


armed with 1 strong claw; each segment of foretarsus contrastingly dark basally, 
vhitish distally: m ddle and hind tibiae loosely clothed with brown and whitish 


s( ale S al d long 


VoLuME 26, NuMBER 1 57 


Abdomen: Brownish gray on top, lighter beneath. The male has a pair of hair pen- 
cils in two grooves, one on each side at base of abdomen. 

Pattern of maculation: Background of forewing a mixture of white and brown scales 
appearing as a powdery, light brownish gray; lighter basally near costa; darker basally 
between Cu and inner margin; transverse lines obsolete; a fusiform, slightly diffuse, 
black fascia runs along Cu from base to middle of wing; a thicker, fusiform, black 
fascia runs in the cell and beyond, almost from base to outer margin between M2 and 
M3; the space between these two fasciae is so dark as to give the appearance of only 
one big fascia, very thin at its ends, but covering about a fourth of the width of the 
wing in its middle; vein ends slightly darkened; darker, longer, intervenular dashes; 
a thin, black subcontinuous terminal line; fringe yellowish at extreme base, checkered 
distally, dark brown between vein ends; some specimens show the black outline of the 
claviform, immediately under the black fascia; orbicular and reniform obsolete. Hind- 
wing of male iridescent white, blackish at vein ends and along a thin terminal line; 
fringe basally yellowish, white distally, with a thin brownish dividing line. Hindwing 
of female similar except for soiled veins and presence of a diffuse soiled band along 
outer margin. Undersurface of forewing white, brownish along costa, near apex, and 
along Cu; fringe as above. Undersurface of hindwing white, slightly soiled along 
costa; fringe white. 

Length of forewing: 13 to 14 millimeters. 

Male genitalia: As in Fig. 1 and la. 

Female genitalia: As in Fig. 2. 

Holotype: Male, Texas, Big Bend National Park, Basin, altitude 5500 feet, 10 May 
1966, genitalia on slide A.B. 1050, deposited in the National Museum of Natural 
History, (No. 68147). 

Paratypes: All taken in or around the Chisos Mountains of Big Bend National Park: 
Basin, 7 July 1964, 14; 10 July 1964, 14; 4 Aug. 1964, 14; 29 Aug. 1965, 14; 29 
June 1965, 1¢, 19. Oak Spring, 9 July 1964, 36 6, 299; 5 Aug. 1964, 19; 31 
Aug. 1964, 1¢. Green Gulch, 27 June 1965, 1¢; 25 Mar. 1971, 1¢; 28 Mar. 1971, 
2292. Government Spring, 27 Mar. 1971, 1¢. Paratypes will be deposited in the 
National Museum of Natural History, in the American Museum of Natural History, 
in the British Museum (Natural History) and in Dr. J. G. Frandemont’s collection. 


Oncocnemis cottami is very close to O. atricollaris Harvey. The trans- 
verse lines of atricollaris are generally well marked, sometime only partly 
traceable, rarely completely obsolete; the part of the reniform which falls 
below the upper black fascia generally shows as a pure white small pupil: 
the white median band of the collar appears narrower because its top 
half is much soiled with brown scales. If all three of these characters 
appear doubtful, one has to rely on the genitalia to remove all doubt. 
The valves of the male genitalia of atricollaris are broader, rounder, and 
present a well developed corona (Fig. 3); those of cottami have no trace 
of a corona. The female genitalia are also different, the ductus bursae of 
atricollaris goes directly into the bursa copulatrix, whereas it is much 
contorted in cottami. Distribution: O. cottami is known only from the 
Chisos Mountains of Texas. O. atricollaris, which is known from a large 
territory including parts of Colarado and Arizona and most of Texas, is 
either absent or quite rare in the Chisos Mts. O. nigrocaput Smith, another 
species of the same group, described from Colorado, also extends to Texas 


58 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


|| 
| 


tf | 
/)) 
Y/ 
LY 


bf a 


MA | 
HL 
h | Hi My 
lH 


i Hy) i } 


Wed 
SWB, 


Ab 
Hi 


VoLUME 26, NuMBER 1 59 


it must also be quite rare as we have only two males, both taken 27 March 
1971 at Government Spring in Big Bend National Park. 

I take great pleasure in dedicating this new species to Dr. Clarence 
Cottam, Director of the Rob and Bessie Welder Wildlife Foundation. This 
is a privately endowed, non-profit organization, supporting educational 
and research activities, situated near Sinton in the San Patricio County of 
Texas. Dr. Cottam always made us feel welcome there. 


Oncocnemis heterogena, A. Blanchard, new species. 


Head: Front covered with short, raised, black, white tipped scales; vertex with 
rough squamation of longer, black, ochreous tipped scales; tongue strong; palpi 
ascending, exceeding front by half eye diameter, first and second segments clad with 
rough scales, mixed on underside with much longer hairlike scales; first segment 
ochreous below, blackish laterally; second segment with more black than ochreous 
scales; third segment, the shortest, blackish. Antennae simple, dark reddish brown. 

Collar: Wide, smooth, pinkish ochreous, more or less darkened by blackish scales. 

Thorax: Disc and patagiae smooth, brownish black; whitish below. Legs blackish; 
segments of metatarsi distally ringed with pinkish ochreous; foretibia armed with claw. 

Abdomen: Ochreous above, whitish below. The male has a pair of hair pencils in 
two grooves, one on each side at base of abdomen. The female presents a tuft of 
very dark reddish brown scales, below, at the tip of the abdomen. 

Pattern of maculation: Maculation of forewing on upperside produced by mixture 
in variable proportions of scales of essentially two colors only: black and same pinkish 
ochreous as collar; these two colors about evenly mixed in background; three deep- 
black spots on costa: one adnate to and distad of t.a. line, one adnate to and basad 
of t.p. line, and one marking anterior origin of rather indistinct median shade; basal 
line pinkish ochreous, internally bordered by fine black line, extending from costa to 
cubital vein; t.a. line pinkish ochreous, outcurved between veins, generally slanted a 
little outwardly, with faint, discontinous indication of black borders more definite 
near inner margin; t.p. line pinkish ochreous, narrow, outwardly convex and well 
beyond reniform between costa and M3, broader, lighter in color and outwardly 
concave between M3 and inner margin; s.t. line pinkish ochreous, very irregular and 
discontinuous; terminal line thin, black, varying from almost continuous to nearly 
obsolete; fringe grayish ochreous, checkered with darker gray between vein ends on 
better marked specimens; orbicular elongated, with dark center, outlined in pinkish 
ochreous; reniform with dark center, inwardly bordered with pinkish ochreous; these 
discal spots quite variable in intensity and sometimes obsolete; on some specimens the 
light color of the broader posterior part of the t.p. line extends into the subterminal 
space, contrasting with the always darker posterior part of the wing between transverse 


< 


Figs. 1-7. Male and female genitalia. Scales shown equal 1 mm. Figs 1-2. 
Oncocnemis cottami: 1 and la, ¢ type, Big Bend National Park, Basin, slide A.B. 
1050; 1, aedeagus omitted, la, aedeagus. 2, 2, Big Bend National Park, Oak Spring, 
slide A.B. 2384. Fig. 3. Oncocnemis atricollaris Harvey, Lake Brownwood State 
Park, part of right valve, slide A.B. 1055. Figs. 4-5. Oncocnemis heterogena: 4 and 
4a, 6 type, Big Bend National Park, Green Gulch, slide A.B. 1059; 4, aedeagus 
omitted, 4a, aedeagus. 5, 9, Big Bend National Park, Basin, slide A.B. 1354. Figs. 
6-7. Paramiana callaisata: 6 and 6a, ¢ type, Guadalupe Mountains National Park, 
Pine Spring Canyon, slide A.B. 1138; 6, aedeagus omitted, 6a, aedeagus. Fig. 7 
2, Guadalupe Mountains, Pine Spring Canyon, slide 1377. 


? 


JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


60 


VOLUME 26, NUMBER 1 61 


lines. Hindwing white, darkened on veins and along diffuse narrow marginal band, 
somewhat wider and darker in the female than in the male; fringe yellowish basally, 
white distally, with narrow brownish median line. Undersurface of forewing smoky, 
darker along costa, varying to sordid white along inner margin; fringe varying from 
smoky to whitish and checkered. Undersurface of hindwing white, darker on veins, 
near apex and along narrow terminal line; fringe white. 

Length of forewing: Male 15 to 16 millimeters; female 15 to 17 millimeters. 

Male genitalia: As in Fig. 4 and 4a. 

Female genitalia: As in Fig. 5. 

Holotype: Male, Big Bend National Park, Green Gulch, 27 August 1965; genitalia 
on slide A.B. 1059, deposited in the National Museum of Natural History (No. 68148). 

Paratypes: Big Bend National Park, Basin, 4 August 1964, 19; 4 Sept. 1964, 19; 
29 Mar. 1965, 19; 25 Aug. 1965,5¢ ¢, 22 9; 28 Aug. 1965, 1¢; 14 May 1966, 1¢; 
7 April 1967, 16, 12. Big Bend National Park, Pine Canyon, 1 April 1965, 1¢; 8 
April 1967, 1¢. Big Bend National Park, Green Gulch, 3 April 1965, 14, 39 9; 27 
Aug. 1965, 1 ¢; 5 April 1967, 19; 25 Mar. 1971, 19. Sierra Diablo Wildlife Manage- 
ment Area, 22 & 23 June 1965, 36 6, 19; 20 May 1968, 11¢ ¢, 82 9; 5 to 7 June 
1969, 63 g, 222; 14 & 15 July 1969, 76 4, 529; 1 Sept. 1969, 22 2; 31 Mar. 
1970, 26 6,19; 31 Aug. and 1 Sept. 1970, 44 6,19. 

It is remarkable, but probably not meaningful, that although the Davis Mountains lie 
between the Chisos Mountains of the Big Bend and the Sierra Diablo, this list does not 
include a single specimen from the Davis Mountains. It may more likely mean that 
our collecting spots, limited as they are to the few locations accessible by car, where 
our traps appear reasonably safe during the night, do not satisfactorily represent the 
Davis Mountains. 

Paratypes will be deposited in the National Museum of Natural History, in the 
American Museum of Natural History, in the British Museum (Natural History) and 
in Dr. J. G. Franclemont’s collection. 


This species is quite variable, the transverse lines vary from well defined 
and contrasty, as they are in the holotype, to fuzzy and nearly obsolete; 
Oncocnemis heterogena appears to resemble O. homogena Grote most 
closely in the pattern of maculation and size, but it may be easily sepa- 
rated because the basal part of the forewing is not noticeably paler than 
the median area, and the t.a. line is differently colored than in homogena. 
In the latter species the t.a. line is uniformly dark brown and black, 
strongly contrasting with the remainder of the wing. In heterogena the 
t.a. line is composed of a thinner, less intensely colored line of brown 
scales bordered basally by a line as wide or wider of ochreous scales. 


Paramiana callaisata, A. Blanchard, new species. 


Head: Retracted; front bulging, dusky, smoothly scaled; vertex roughly clad with 
white tipped, black scales; eyes large, slightly elliptical; palpi short, concolorous with 
front, ascending, barely exceeding front; first and second segments rough scaled, 
third segment smooth, shorter than second; antennae finely, shortly pubescent in male, 
simple in female. 


< 


Figs. 8-10. Male holotypes: 8, Oncocnemis cottami; 9, Oncocnemis heterogena; 
10, Paramiana callaisata. 


62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Collar: Disc of thorax and patagiae covered with raised, white tipped, black scales, 
forming a high tuft on metathorax. Legs dusky, with loose, long hairs. 

Abdomen: Fuscous above and below, with weak tuft on first segment. 

Pattern of maculation: Maculation of forewing consisting mostly of hard-to-trace, 
transverse bands and fuzzy patches of black and green; basal band, when not com- 
pletely obsolete, reduced to its inner black line, reaching from costa to anal vein; t.a. 
band substantially reduced to its fine, black, discontinuous outer border, starting on 
costa about one fourth distance from base to apex, slightly bulging between veins 
from costa to anal vein, squarely outcurved on anal vein and reaching inner margin 
almost at its middle; t.p. band poorly defined, median line paler than background, 
inner black line consisting of intervenular, finely confluent, black spots, outer line 
almost obsolete, in course arising on costa above reniform, outwardly bent just below 
costa to well beyond reniform, then parallel to outer margin, except for broad, shallow 
concavity in fold; s.t. line consisting of a succession of elongated, intervenular, black 
spots sprinkled in their center with green scales; terminal line reduced to intervenular, 
small, triangular, black spots; a white spot at vein ends; fringe concolorous with back- 
ground, checkered, with a darker median line; orbicular small, green, outlined in 
black; reniform large, squarish, mottled yellowish green and black, broadly changing 
to bluish white at outer corners and along lower side; a black spot along inner margin 
midway between base and a.m. band; rest of area between basal and a.m. band well 
sprinkled with green scales; a.m. outer and p.m. inner lines enlarged in fold, forming 
there two black patches more or less connected by darker background; a large black 
patch basad of reniform; a green patch distad of p.m. band in fold; four to five white 
dots on costa, about equally spaced in its outer half. Hindwing of male light brassy 
ochreous, heavily sprinkled with brown scales forming two extramedial bands; terminal 
line brown, accented between veins; fringe concolorous with background, with medial 
brown dividing line. Hindwing of female more heavily sprinkled with brown scales. 
Undersurface of forewing dusky; with darker p.m. line; a terminal brown line; fringe 
concolorous, checkered. Undersurface of hindwing dusky, darker along costa, an 
extramedial line in continuation of that of the forewing. 

Length of forewing: 18 to 19 millimeters. 

Male genitalia: As in Figs. 6 and 6a. 

Female genitalia: As in Fig. 7. 

Holotype: Male, Guadalupe Mountains, Pine Spring Canyon, altitude 5700 feet, 28 
August 1967; genitalia on slide A.B. 1138, deposited in the National Museum of 
Natural History (No. 68149). 

Paratypes: Guadalupe Mts., Pine Spring Canyon, 28 Aug. 1967, 146, 692 2. Davis 
Mountains, Mt. Locke (McDonald Observatory), altitude 6000 feet, 25 Aug. 1967, 
364; 30 Aug. 1969, 18; 6 Sept. 1969, 1034, 1299; 27 Aug, 1970. 
263, 329; 29 Aug. 1970, 36 3, 299. Paratypes will be deposited in the 
National Museum of Natural History, in the American Museum of Natural History, in 
the British Museum (Natural History) and in Dr. J. G. Franclemont’s collection. 


The forty four specimens before me show very little variation. Sepa- 
rating Paramiana callaisata from other species in the same genus is a very 
simple matter since its maculation resembles that of no other previously 
described species that I have been able to compare it with in the National 
Museum. The distinctive whitish contrasting reniform spot separates it 
from all the other species of the genus. Paramiana laetabilis Smith also 
has a contrasting, large reniform spot, but it is blue in fresh unfaded 


specimens. The broad blue subterminal band of Iaetabilis is absent in 
the new species. 


VoLUME 26, NuMBER 1 63 


ACKNOWLEDGMENTS 


I wish to thank Dr. E. L. Todd of the Agricultural Research Service of 
the U.S. Department of Agriculture for revising the manuscript and for 
many useful suggestions. I also wish to extend my appreciation to the 
personnel of the Big Bend National Park, the Guadalupe Mountains 
National Park, the State-owned Wildlife Management Areas of Texas and 
the McDonald Observatory for authorizing us to collect at these places. 


LITERATURE CITED 


BLANCHARD, A. 1966. A new species of Glaucina (Geometridae) from Texas. J. 
Lepid. Soc. 20(4): 247-250. 

1968. New moths from Texas (Noctuidae, Tortricidae). J. Lepid. Soc. 

92.(3): 133-145. 

1970. Observations on some Phycitinae (Pyralidae) of Texas with descrip- 

tion of two new species. J. Lepid. Soc. 24(4): 249-255. 


A POSSIBLE CASE OF MIMICRY BETWEEN LYCAENID 
BUTTERFLIES (LYCAENIDAE) 


Mimicry is a relatively common phenomenon among lepidopteran insects. Familiar 
North American examples have been studied extensively (e. g. Brower 1958, Evolution 
12: 32-47, 123-136, 273-285). Recently, Downey (1965, J. Lepid. Soc. 19: 165- 
170) suggested a mimicry complex with several blues (Plebejinae, especially Plebejus 
icarioides Boisduval) serving as models for the noctuid moth Caenurgina caerulea 
Grote. Another possible case of mimicry with blues as models involves the male Blue 
Copper (Lycaena heteronea Boisduval). The striking superficial resemblance of the 
dorsal wing surface of the latter and blues has been noted often (Ehrlich and Ehrlich 
1961, How to Know the Butterflies; Comstock 1927, Butterflies of California; Brown 
et al. 1957, Colorado Butterflies). The difference in coloration of the male L. 
heteronea from that of other coppers immediately suggests mimicry. 

I suggest that the male of L. heteronea is a mimic of Glaucopsyche lygdamus 
Doubleday and possibly other sympatric blues. The dorsal coloration of L. heteronea 
is nearly identical to that of G. lygdamus. The geographical range and flight period 
of the copper appear to be entirely within those of the blue. 

Furthermore, Batesian mimicry is indicated by at least three sets of data. The 
dorsal resemblance is nearly perfect. Batesian mimics tend to be more perfect than 
Miillerian (Ford 1964, Ecological Genetics). The food plant of the copper is 
Eriogonum; those of G. lygdamus include Lupinus and Astragalus among other 
legumes (Ehrlich and Ehrlich, op. cit.). Lupinus contains alkaloids (Robinson 1968, 
The Biochemistry of Alkaloids) and many Astragalus contain selenium; both are 
known to be poisonous to vertebrates (Kingsbury 1964, Poisonous Plants of the 
United States and Canada). If these compounds are incorporated into insect tissues, 
such insects may be poisonous or unpalatable to their predators. The mimic appears 
to be less common than its model (Comstock, op. cit.; Brown et al., op. cit.). In 
June and July, 1963, in Garfield County, Colorado (Coffee Pot Springs, White River 
Plateau, 10,000 feet), I found G. lygdamus to be about ten times as common as L. 
heteronea; the latter was ecologically sympatric with the former and could not be 
distinguished in the field. 


64 JouRNAL OF THE LEPIDOPTERISTS’ SOCIETY 


If this is found to be a true case of mimicry, it is apparently the only one among 
butterflies involving only the male (the female L. heteronea is a typical copper). 
Other cases of Batesian mimicry in butterflies are restricted to the female, or both 
sexes are involved (Ford, op. cit.). Also of interest is that L. heteronea is nearly 
allopatric with another blue mimic (C. caerulea), overlapping only in eastern Cali- 
fornia (Downey, op. cit.). Additionally, the several species of lupine feeding blues 
may form a large Miillerian complex. 

I hope that these observations will stimulate further research on mimicry in these 
butterflies, both in the field and the laboratory. For example, I have seen remains 
(wings) of blues at mud puddles. It would be of interest to have counts of each 
species represented and to compare these with their abundance. 

I thank W. L. Nutting for comments on the manuscript. Collecting in Colorado 
was supported in part by an N. S. F. undergraduate research assistantship through 
the Institute of Arctic and Alpine Research of the University of Colorado. 

GreorcE T. Austin, Department of Biological Sciences, University of Arizona, 
Tucson (present address: Box 1102, Las Vegas, Nevada 89101). 


THE USE OF CHLOROETHANE FOR IMMOBILIZATION 
OF FIELD SPECIMENS 


Chloroethane (ethyl chloride) is a colorless, volatile liquid used in the medical 
profession as a topical anesthetic. Its low vapor pressure causes quick evaporation at 
normal atmospheric temperatures and cooling of any object in contact with it. 

In 1969 I began using chloroethane to “freeze” butterflies caught in the field. 
The spray is easily directed at the specimen in the net and almost immediately 
immobilizes it. Although many species do not regain activity after adequate immobili- 
zation, a killing jar is still used because some species do become active several minutes 
post-exposure. 

The chloroethane immobilization allows for better specimens to be brought back from 
field collecting. It lessens the battering of specimens in net and killing jar and prevents 
their escape in the transfer from the net. The substance has not caused any residue 
on wings or any pigmentation changes in any species to date. 

Chloroethane is supplied in four-ounce glass bottles with fine, medium or course 
nozzles. I have found the fine nozzle to be adequate for most Lepidoptera and the 
medium nozzle to be adequate for even the largest. The four-ounce fine nozzle will 
do 250-300 specimens and the medium nozzle 150-200 specimens. If the bottles are 
stored in a refrigerator when not in use the chloroethane evaporates very slowly and 
may be used over many weeks with almost no loss between usages. I have found 
that wrapping the bottle in crumpled tinfoil is also useful. This acts as an insulator, 
keeping the fluid cooler longer, lessening evaporation and so allowing the chloroethane 
to be used for more specimens. The tinfoil also protects the glass bottle from breakage 
should it be dropped. 

The main disadvantage to chloroethane is that it is flammable. It burns with a light 
green flame and liberates hydrogen chloride as a by-product. Except for the flamma- 
bility, chloroethane is a relatively safe chemical when compared to other poisons used 
in entomology. 

Chloroethane is a prescription drug and some difficulty may be encountered in 
obtaining it for non-physicians. Presently it may be obtained only from hospital 
pharmacies with a prescription from a physician. If sufficient interest is expressed 
from collectors, manufacturers may make it available to biological supply houses for 
use in entomology. 

Raymon CastitoniA, Loma Linda University, Loma Linda, California. 


VoLuME 26, NuMBER 1 65 


MIDGES SUCKING BLOOD OF CATERPILLARS 
(DIPTERA: CERATOPOGONIDAE ) 


Lepidopterous larvae have been known to be attacked by bloodsucking midges since 
De Geer (1752, Mem. Hist. Insectes 1: 327) wrote, “Nos grandes Chenilles sont 
aussi incommodées par ces petits Moucherons; sur le corps dune de ces Chenilles 
jen vis cing ou six, qui avoient enfoncé leur petite trompe dans la peau, & oui étoient 
occupés a sucer la liqueur verdatre, qui est le sang de la Chenille.” In all the 
lepidopterous literature appearing in the two hundred years since that time, however, 
there are so few accounts of these tiny parasites that it would appear that they are 
extremely rare. I believe that such is not the case, and that these midges have been 
overlooked because of their small size (less than 2 mm long), their rapid darting 
flight, and probably their nocturnal habits. 

In 1956 (Ann. Ent. Soc. Amer. 49: 356-364) I reviewed all the published records 
and available museum specimens of these caterpillar parasites and concluded that all 
pertained to Forcipomyia fuliginosa (Meigen), a common species with nearly world- 
wide distribution. I have just finished going over the museum material and literature 
again, and although I can now recognize several distinct local species of caterpillar 
parasites in tropical America, F. fuliginosa still remains as the main species attacking 
lepidopterous larvae around the world. It also occasionally attacks other soft-bodied 
insects such as sawfly larvae, dragonflies, and meloid beetles. 

Several observers have noted outbreaks of these bloodsuckers in sufficient numbers 
to indicate that on occasion they may have a role in reducing caterpillar populations. 
Hagen (1883, Ent. Mo. Mag. 19: 254) called attention to their attacks on Pieris 
menapia Felder in Washington Territory, and Baker (1907, U.S. Dept. Agr. Bur. 
Ent. Bull. 67: 117) reported their depredations on an outbreak of Melanchroia 
geometroides Walk. in Cuba. Mayer (1955, Ztschr. fiir Ang. Zool. 1: 95-107) 
suggested that the death of the caterpillars was probably not caused directly by the 
feeding of such small midges, but that their habits fitted them well to act as vectors 
of polyhedral viruses or other diseases. 

The purpose of this note is a plea to lepidopterists to make a special effort to look 
out for these minute caterpillar pests during their field work. I suspect that night-time 
observations might give us more data on the abundance of these midges and their 
habits. In some situations it is very possible that these tiny flies may have some role 
in biological control, particularly in bringing a virus infection from “wild” hosts in 
a field or orchard margin into a caterpillar population on a crop plant. 

Wits W. WirtH, Systematic Entomology Laboratory, ARS U. S. Department of 
Agriculture c/o U. S. National Museum, Washington, D. C. 20560. 


NOTES AND NEWS 


It is a pleasure to announce that Dr. Alexander B. Klots and Dr. Eugene Monroe 
have been overwhelmingly approved by the membership as honorary life members of 
the Lepidopterists’ Society. 

The new editor of the Journal asks that prospective authors carefully read the 
Notice To ContrisuTors (inside back cover), as several changes in editorial policy 
have been instituted. 


66 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


OBITUARY 


PERCY HEATH HOBART GRAY 
1891-1971 


It is with deep regret that we record the death on October 11th, 1971, 
of Dr. P. H. H. Gray, a Charter Member of the Lepidopterists’ Society, 
and for some years a member of the Entomological Society of Canada 
and the Entomological Society of Quebec. 

He was born in Woodland Heights (Richmond), Virginia, on November 
21st, 1891, and studied in England at Oxford, (M.A.) and London Uni- 
versity (D.Sc.). In 1920 he joined the staff of the Bacteriology Depart- 
ment, Rothamsted Experimental Station, England, and in 1929 became 
Professor and Chairman of the Department of Agricultural Bacteriology, 
Macdonald College of McGill University, Ste. Anne de Bellevue, P.Q., 
Canada. During the course of almost 30 years prior to his retirement in 
1957, his research output was considerable, concerning mainly the ac- 
tivities of soil microorganisms towards cellulose, urea and pesticides and 
a great many scientific papers were published in this connection. For 
many years Dr. Gray guided numerous students who trained in micro- 
biology and who now hold important positions across Canada. 

During this period, Dr. Gray found time for and took a great interest 
in entomology, developing a good local collection of Lepidoptera, col- 
lected for the most part in his garden at Baie dUrfe, P.Q., and in the 
surrounding woods and fields. His careful collecting produced several 
species not previously recorded for the Province of Quebec. He published 
some 25 papers on Lepidoptera. He was also a proficient classical scholar, 
and his quiet wit was at its best in many verses which he composed and in 
his letters to friends. 

Shortly after retiring he moved to Nova Scotia, purchasing a home 
outside Digby, where he resided and continued with his interest in ento- 
mology until the time of his death. Our deepest sympathy is extended 
to his wife and son. 


BIBLIOGRAPHY (ENTOMOLOGICAL) OF P. H. H. GRAY 


1949. Alfred Russell Wallace on butterfly hunting. Lep. News 3: 80. 

1949. An economical storage box. Lep. News 3: 104. 

1951. An apparatus for incubating lepidopterous larvae or pupae in nutrition and 
environment tests. Lep. News 5: 35. 

1951. Results of humidity tests with Papilio pupae. Lep. News 5: 67. 

1951. A multiple reflector light trap. Lep. News 5: 72. 

1952. The weights of fresh and dried butterflies. Lep. News 6: 73. 


1953. A note on the colors of pupae of Pieris rapae developed under artificial con- 
ditions. Lep. News 7: 5. 


VOLUME 26, NUMBER 1 67 


1953. 
1953. 
1954. 
1954. 
1954. 
1954. 


1955. 
1955. 


1955. 
1957. 
1959. 
1960. 
1960. 
1961. 
1962. 
1965. 
1967. 


1968. 


Correlations between “pupal volume” and wing radius and weight in butter- 
flies. Lep. News 7: 48. 

Aids to distinguish between females of the “Winter-Moths”’, Alsophila pometaria 
and Operophtera bruceata (Geometridae). Lep. News 7: 127. 

Effects of humidity during growth of Pieris rapae larvae. Lep. News 8: 88. 
1952 captures in Quebec. Lep. News 8: 102. 

Notes on the occurrence of Alsophila pometaria (Geometridae) at Baie d’Urfe 
and Macdonald College, Quebec, in 1953. Lep. News 8: 140. 

Aristotle’s description of the life history of a butterfly. (Psyche) Lep. News 
8: 145. 

An apparatus for weighing small insects. Lep. News 9: 58. 

Aphomia gularis (Zeller) (Lepidoptera: Pyralidae) at Baie d’Urfe, Quebec. 
GansEnt.o7: 239: 

An apparatus for the rapid sorting of small insects. The Entomologist 88: 
OS: 

Mold-free relaxing bottles for Lepidoptera. Lep. News 11: 54. 

Dwarf Pierine butterflies. The Entomologist 92: 1154. 

A method for estimating the wing radius in Lepidoptera. Lep. News 14: 63. 
Dwarf Pierine butterflies: a supplementary note. The Entomologist 93: 180. 
A comparison of scales on wings of Colias species: (Lepidoptera: Pieridae). 
The Entomologist 94: 125. 

‘Giant’ scales on the wings of Thecla species (Lepidoptera: Lycaenidae). 
The Entomologist 95: 76. 

A method of estimating the area of a butterfly’s wing. The Entomologist 98: 
64, 

Correlations between characters in species of Colias (Lepidoptera: Pieridae). 
The Entomologist 100: 62. 

Noviculoid scales in the fore wing terminal borders of Colias species: (Lepi- 
doptera: Pieridae). The Entomologist 101: 175. 


A. C. SHEPPARD, Lyman Entomological Museum, Macdonald Campus of McGill 
University, Ste. Anne de Bellevue 800, P.Q., Canada. 


EDITORIAL COMMITTEE OF THE JOURNAL 


Editor: THEopore D. SARGENT, Department of Zoology, 
University of Massachusetts, Amherst, Massachusetts 01002 


K. S. Brown, J. M. Burns, R. H. Carcasson, J. P. DoNAHUvE, J. F. 
Gates CLARKE, R. O. KENDALL, J. H. Masters, L. D. MILLER, 
A. P. Piatt, J. R. G. TURNER 


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, 844 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, 1961la, 1961b) and all must be listed alphabetically 
under the heading LirERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural Selection and Heredity. 2nd. ed. Hutchinson, 
London. 209 p. 
1961a. 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: 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 814 
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 article’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 FiIcuREs, 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 approx- 
imate 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. 

Page Charges: Authors with grant or institutional funds are requested to pay a 
charge of $24.00 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 
case of authors who have no grant or institutional funding, as it is not intended that 
any author should pay this charge from personal funds. However, all authors will 
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: Dr. C. V. Covell, Department of Biology, University of 
Louisville, Louisville, Kentucky 40208. 


ALLEN PRESS, INC. eueo LAWRENCE, KANSAS 
usr 


CONTENTS 


NOTES ON THE BALANOTES (MeEyrick) Group OF OIDAEMATOPHORUS 
WALLENGREN WITH DESCRIPTION OF A NEW. SPECIES ( PTER- 
OPHORIDAE). Everett D. Cashatt 


ANNOTATED LIST OF THE BUTTERFLIES OF INDIANA, 1971. Ernest M. 
Shull and F. Sidney Badger 0. 


Tue Lire History oF SCHINIA INTRABILIS (NoctuwaE). D. F. Hard- 


Wich Lv eA ES OS a rr 


THE Lire History or SCHINIA PALLICINCTA (NoctuiAE). D. F. 
Hardwick) 020 SOS Se eee a Ls ae 


STUDIES ON THE CATOCALA (NocrumAE) OF SOUTHERN NEW ENGLAND 
II. CompPaARISON OF CoLLECTING Procepures. Charles G. Kellogg 
and Theodore D. Sargent 


THREE BUTTERFLY SPECIES (LYCAENIDAE, NYMPHALIDAE, AND HELICONT- 
IDAE ) NEw To TEXAS AND THE UNrTeED States. Roy O. Kendall — 


More New Motus rrom Texas (Noctuwar). André Blanchard 


GENERAL NOTES 
Natural inter-breeding of close nymphalid groups. Raymond J. Jae —.- 


Protective function of sound perception and gregariousness in Hylesia larvae 
(Saturniidae: Hemileucinae), Charles L. Hogue ae 


A possible case of mimicry between lycaenid butterflies ( Lycaenidae ). 
George T. Austin 


The use of chloroethane for immobilization of field specimens. ei Hingis 
Castilonia 


Midges sucking blood of caterpillars (Diptera: Ceratopogonidae). Willis 
W. Wirth 


OBITUARY 3 ee i ae 


13 
24 


29 


ae ee ee 


Volume 26 1972 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 


17 May 1972 


THE LEPIDOPTERISTS’ SOCIETY 
EXECUTIVE COUNCIL 


Lioyp M. Martin (Prescott, Ariz.) President 

J. F. Gates CLarxeE (Washington, D.C.) President-elect 
S. A. Az (Nagoya, Japan) Ist Vice President 

Kerry S. Brown (Rio de Janeiro, Brasil) Vice President 
H. A. FREEMAN (Garland, Texas) Vice President 

S. S. Nicouay (Virginia Beach, Va.) Treasurer 

LEE D. MiiueEr (Sarasota, Fla.) Secretary 


Members at large (three year term): R. B. Dominick (McClellanville, S.C.) 

B. MATHER (Clinton, Miss.) 1972 1973 

M. Ocara (Osaka, Japan) 1972 J. P. DonanueE (Los Angeles, Calif.) 1973 

E. C. Wexuinc (Merida, Mexico) 1972 J. M. Burns (Cambridge, Mass.) 1974 

ANDRE BLANCHARD (Houston, Texas ) R. H. Carcasson ( Vancouver, B.C.) 1974 
1973 M. C. Nietson (Lansing, Mich.) 1974 


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 eight numbers of the News each year. 


Active members—annual dues $10.00 
Student members—annual dues $5.00 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $15.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia 23455. 


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


by Cyr. F. pos Passos 


Price, postpaid: Society members—$5.00, others—$7.50; uncut, unbound signatures 
available for interleaving and private binding, same prices; hard cover bound, mem- 
bers—$8.00, others—$10.00. Revised lists of the Melitaeinae and Lycaenidae will be 
distributed to purchasers free (separately with paper covered copies and unbound 
signatures, bound in with hard covered copies ). 


The Lepidopterists’ Society is a non-profit, scientific organization. The office of 
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520. 
Second class postage paid at Lawrence, Kansas, U.S.A. 66044. 


ey! 


yee 


= 


Color Plate. 


Examples of freeze-dried larvae (natural size). 1. Eacles imperialis 
imperialis (Drury), on Liquidambar stryaciflua, L. 2. Callosamia securifera (Maas- 
sen). 3. Darapsa myron (Cramer), on Vitis sp. 4. Hyalophora cecropia (L.), on 


Liquidambar stryaciflua, L. 5. Ewmorpha fasciata (Sulzer ) 


, on Ludwigia leptocarpa 
(Nuttall). 


JOURNAL OF 


Tue Leprpoprerists’ SOCIETY 


Volume 26 3 1972 Number 2 


PRACTICAL FREEZE-DRYING AND VACUUM 
DEHYDRATION OF CATERPILLARS 


RIcHARD B. DOMINICK 
The Charleston Museum, Charleston, South Carolina 29401 


The purpose of this paper is to describe a method of preserving larvae 
of Lepidoptera and other soft-bodied insects whereby a lifelike appear- 
ance can be maintained, the process completed in a short time, the inter- 
nal anatomy preserved, and the required equipment purchased within 
the budget of a small laboratory, even of the make-shift variety. 

Papers on the subject have been published by Meryman (1960, 1961), 
Blum & Woodring (1963), Woodring & Blum (1963), and Harris (1964, 
1968 ). But the equipment used by these workers is rather involved and 
expensive. Flaschka & Floyd (1969) developed a method that requires 
only a tightly stoppered jar with desiccant, and a home freezer. But the 
simplicity and low cost are countered by extremely long drying times. 
The method described here combines a minimum of drying time with 
an intermediate cost of a few hundred dollars or less. The description 
will be given in such a way that alternatives and modifications may be 
applied that allow further reduction of the expenses. For example, the 
two-stage vacuum pump rated at a vacuum of 0.1 micron is ideal, but a 
cheaper pump will be found satisfactory as long as it will pull down to 
about 100-200 microns. The degree of vacuum is one of the factors that 
has a direct bearing on drying time. The gas ballast recommended for 
museum systems is unnecessary here because of the relatively small 
amount of moisture encountered. In any case one must try to be sure 
that water vapor is not permitted to reach the pump oil, and the purpose 
of a gas ballast is to remove large quantities of water vapor without dam- 
age to the pump. By the same token, dirty oil should be changed. 

Provided the specimens are in close enough proximity to the desiccant 
(or vapor trap) to lie within reasonable reach of the mean free path of 
the water vapor molecules, several factors influence the time required 


70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


for, and the efficiency of, the drying. The two most important are tem- 
perature and degree of vacuum. Their appropriate choice allows manip- 
ulation of the process within wide ranges. The method of Flaschka & 
Floyd (1969), while simple, effective and inexpensive, cannot provide 
much leeway. The temperature of a home freezer is fixed and usually 
slightly lower than ideal, and no vacuum is applied. Consequently a dry- 
ing time of about 100 days for a medium sized larva (e.g. Malacosoma 
americanus) results. Appropriate selection of temperature and vacuum 
can improve the situation to a remarkable degree. With a vacuum of 0.1 
micron at 25° F., such a caterpillar will be adequately dry in 48 hours, 
finishing off for a few hours at room temperature. In general the speci- 
men should be frozen before applying vacuum to prevent distortion in 
the freeze-dry process, and I have found the ideal temperature to be 
about 20-25° F. rather than the lower temperatures recommended in the 
literature. Such temperature guarantees that the frozen state is main- 
tained while leaving leeway for opening the freezer door, but is not so 
low as to seriously curtail the rapidity of molecular motion. In principle 
there is a slight cooling effect due to the sublimation process itself (simi- 
lar to the cooling experienced when water evaporates from one’s skin), 
but the heat exchange involved is so small that the phenomenon can be 
disregarded. It becomes a factor of importance only when working with 
large museum specimens (mammals, reptiles, etc.) where it is one of the 
conditions that lead to prolonged drying time. 


Some Theoretical Considerations 


The following discussion is presented in the hope that it may give the reader 
enough understanding of the underlying processes to help him perform the job more 
efficiently and enable him to use his ingenuity when adapting the method to his own 
needs and budget. 

Water in both liquid and solid form (ice) exerts a vapor pressure that is a func- 
tion of the temperature. This means that if brought into a confined chamber, water 
or ice will evaporate and increase the total gas pressure by an amount equal to the 
vapor pressure at the prevailing temperature. In relation to the total gas pressure 
present, one refers to the partial pressure of water vapor. It is important to realize 
that this concept holds only as long as there is some water or ice still left. Then, 
when the required partial pressure is reached, a state of dynamic equilibrium is estab- 
lished; in other words, the number of water vapor molecules per unit time evaporating 
from the surface of the water or ice is exactly equal to the number of molecules con- 
densing on that surface from the gaseous phase. If water vapor is removed from the 


chamber (e.g. by pumping out, freezing out, or binding on a drying agent) more 
water vapor will evaporate in order to restore the partial pressure to the level dictated 
by the temperature. The more molecules of other gasses 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. If, however, a vacuum is applied and the air molecules re- 
noved, the evaporation of water and dissipation of its vapor can proceed with less 
obstruction, and thus more efficiently. It may be noted in passing that no gasses other 


VoLUME 26, NUMBER 2 a1 


than water vapor are produced from the specimen by the drying process in sufficient 
quantity to merit serious consideration. A further process is involved, namely that of 
bringing the water from within the specimen to the surface where it can evaporate. 
This process is called diffusion, and its rate is a function of molecular motion and 
thus also of temperature. 

So one tries in freeze-drying to operate at as high a temperature as practical while 
still keeping at all times below freezing. In vacuum dehydration, one depends on 
the internal pressure of the specimen, the comparative toughness of the integument, 
and the rapidity of drying for the maintenance of the shape of the specimen, though 
in this latter process there is the inherent danger of overstretching. 

For a closer look at this situation it is helpful to introduce the concept of the mean 
jree path length, which is defined as the average distance a gas molecule can travel 
before colliding with another gas molecule (bumping into the wall does not count 
as a collision). The mean free path length depends, therefore, on the extent of 
evacuation (of air). For water vapor at a pressure of 10 mm Hg the mean free path 
is 0.0034 mm; at 10 microns, it is 3.4 mm; at 1 micron it is 34 mm (Meryman, 1961). 
It can thus be seen that in either system the advantage of operating in a vacuum is 
enormous. If the vacuum is adequate, water vapor molecules leaving the drying speci- 
men. can travel almost without obstruction to the desiccant where they are held. 

It should be kept in mind that the sole function of the pump is to create a vacuum 
rapidly and efficiently. Once the vacuum is attained the stop-cock (G) is closed to 
keep the leak-proof desiccator evacuated, and the pump is shut off. It does not run 
continuously in order to pump out the water vapor as it forms, nor does it circulate 
an air stream which carries away the humidity. The drying of the specimen is 
achieved exclusively by the molecular motion of the water vapor passing from the 
specimen to the desiccant. 


The Desiccant 


While in theory the cold trap removes water vapor from the specimen 
more efficiently, the equipment and operation are considerably more 
complex. A desiccant is perfectly adequate for the purpose at hand. Silica 
gel (SiOz) or calcium sulfate (CaSO,) impregnated with an indicator 
such as cobalt chloride do very well. Both chemicals with the indicator 
are blue when able to adsorb water and pink when exhausted. Both can 
be regenerated by spreading in a pan and drying in an oven at about 
3090-400° F. for about 2 hours. Calcium sulfate is the slightly more effec- 
tive drying agent but silica gel will be found equally satisfactory. It is 
advisable not to mix the two agents because much of the efficiency of 
the more effective of the two will be lost, since it will first dry out the 
less effective, until finally the higher vapor pressure dictated by the latter 
will prevail. It is preferable to use the desiccant in granules of about 
#8 mesh (roughly 3-5 mm in diameter). Smaller granules are difficult 
to manage cleanly, and a certain amount of fine dust is produced in any 
case which must be kept off the greased portion of the desiccator and its 
lid (a ring of paper cut to size and laid on the greased surface helps), 
otherwise the vacuum will not hold. The dust should also be wiped from 
the inside of the vessel to prevent the formation of a deposit on the speci- 
men that may be difficult to remove. 


72, JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


FREEZE-DRYING VACUUM DEHYDRATION 


Refrigerator 


Vacuum pump 


\ 


Manometer 


115 volt outlet 


= 


“4 
\ 
SS 
4 
: 
= 


Shown with door removed Rubber pad 


Fig. 1. Apparatus for freeze-drying and vacuum dehydration. Explanation in text. 


Replenishment of the desiccant may be necessary before the caterpil- 
lars are dry if they are large or numerous. A freshly charged desiccator 
should be on hand in the freezer and the transfer of specimens accom- 
plished with reasonable rapidity so that no thawing occurs. It is gener- 
ally best not to put too much material in the works at one time. Meryman 
forsees in an unpublished paper the difficulties likely to beset the over- 
enthusiastic student who might be tempted to freeze-dry specimens the 
size of a physics professor, leading to excessive drying time and inferior 
results. 


Basic Equipment 


My equipment (Fig. 1) consists of a small (20 x 20 x 20 inch outside 
measurement) standard refrigerator with its normal ice-making compart- 
ment. By dint of wrapping a bit of tape around the capillary tube (CT) 
of the thermostat in the ice-cube compartment, the temperature of the 
whole refrigerator can be lowered to about 20° F. A hole (H) is drilled 
through the wall of the refrigerator (but not leading into the ice-cube 
compartment nor any other place where damage to the refrigeration coils 
might result). Through this hole is passed a length of copper tubing (T) 
of 4 or % inch outside diameter, which is then firmly fixed (e.g. epoxy 
glue or bracket) to the refrigerator wall. The outside diameter of the 
tubing must closely approximate that of the glass tubing of the stop-cocks 
(G) in order to achieve air-tight connections. This length of tubing (T) 
is extended about an inch inside the refrigerator, sufficient to attach a 
short sleeve (S) about 2 inches long of tightly fitting heavy flexible plas- 
tic tubing. A piece of plywood is placed alongside the refrigerator, and 


VoLUME 26, NuMBER 2 We 


securely and permanently fastened thereto are the vacuum pump and 
manometer, the vacuum-tight brass line valves, and the permanent con- 
necting copper tubing and fittings; these include such additional hook- 
ups as may be wanted to perform the vacuum dehydration (B). A rub- 
ber mat under the plywood will absorb any vibration from the pump and 
cushion any screws or bolts appearing on the under surface of the ply- 
wood. | 

It is worth emphasizing that all joints and valves, stop-cocks and desic- 
cator lids—in other words the entire system—must be as absolutely leak- 
proof as possible at high vacuum. Silicone grease, or even vaseline, on 
meticulously cleaned stop-cocks, desiccator lids, and around the joints 
connected by the plastic sleeves (S), will help ensure tight fits. The 
permanent copper joints depend on proper flaring for sealing; but Duco 
cement or similar sealer will help after the joints are made. Teflon sealer 
on any screw-in joints (as with the manometer ) likewise is of advantage. 
Not shown in the figure, but easily incorporated if desired, are filters 
(glass wool is a good material or vacuum filters may be purchased) to 
prevent foreign material from entering valves or pump. 

Various vacuum gauges are available which give accurate absolute 
readings, such as the McLeod or Pirani gauges. A simple double column 
U-shaped mercury manometer is cheaper and fully adequate since an 
absolute reading is unnecessary. When the two columns of mercury be- 
come stabilized in relation to one another, the system is evacuated as far 
as it will go. One can judge the degree of vacuum achieved by assuming 
an approximation of the manufacturer’s rating of the pump employed. 
Valve (C) leading directly to the pump is then closed, and any leak will 
be registered in degree according to the rapidity with which the mercury 
column moves. It is a good practice in any case to check the vacuum in 
the desiccators periodically to be on the safe side. A well-sealed desic- 
cator, of course, can be evacuated and after closing stop-cock (G) be 
removed from the system to be replaced by another desiccator contain- 
ing another lot of specimens. 

I have used four types of desiccators, all of which hold both specimens 
and desiccant. The first (Fig. 1) is a sleeve-top desiccator made of heavy 
“Pyrex” glass (DA) whose sleeve-top incorporates a stop-cock (G) with 
hose connection. They come in different sizes and the small refrigerator 
shown will easily accommodate two of the 8-inch diameter vessels. The 
bottom is filled with desiccant, and a disc (or preferably a shallow bas- 
ket) of fine screen wire cut to size is placed over the chemical. The lid 
and stop-cock of this desiccator require greasing. A second type of con- 
tainer is made of polycarbonate plastic with a rubber gasket beneath its 


74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


lid. The lid is made of opaque plastic and contains the outlet for the 
stop-cock. It is slightly taller than the first, and the gasket, being rubber, 
must not be greased. It is available from the Nagle Sybron Corporation 
in Rochester, New York. Both these containers are very satisfactory, be- 
ing large enough to contain foodplant as well as larvae, and holding the 
vacuum well. A third type of desiccator is also commercially available 
and is the drying unit for “Drierite,” a trade name for calcium sulfate. It 
is sold containing a charge of the desiccant labelled as approximately 1% 
lbs. of #8 mesh calcium sulfate with a capacity for 50 grams of water. It 
is made of “Plexiglas” and contains its own filter. After slightly widening 
the spring coil attached to the metal screw-on top, one can insert one of 
the plastic pill vials readily available at any pharmacy. The vial contains 
the larva, and its open end is covered with a bit of cheesecloth fastened 
on by a rubber band. The vial is inserted into the desiccator so that the 
open end is in almost direct contact with the desiccant. The metal top 
with its rubber gasket (ungreased) is tightly replaced, and a stop-cock is 
added to the lower tubing connection while the upper tubing connection 
remains sealed, and the unit attached to the vacuum system as before. 
The purpose of the plastic vial, as well as of the screen wire in the previ- 
ous examples, is to prevent the specimens from becoming indented by 
loose granules of desiccant or other matter, a phenomenon which can 
occur even when the larva is thoroughly frozen. Still another desiccator 
(DC) can be home-made from any thick-walled wide mouth jar, say 
about 8-10 inches high. It is a good idea to wrap it carefully with trans- 
parent tape to prevent fragments of glass from flying around in case of an 
implosion. The mouth of the jar is fitted with a heavy bevelled rubber 
stopper into which a hole has been bored for the snug fit of one arm of a 
glass stop-cock (G). Again, the jar is partly filled with desiccant and the 
specimen(s) within the vial inserted as before and the vacuum applied. 
Desiccators must, of course, be transparent so that it is possible to watch 
both the specimens and the color of the desiccant. 

Whichever desiccator is used, it is a good idea to keep two in the 
freezer ready for immediate use. In operation, the outlet from the stop- 
cock (G) receives a short tight sleeve (S) of the flexible plastic tubing, 
in turn connected by a suitable length (L.) of copper tubing to the fixed 
end of the tube (T) that runs into the freezer. The other end of tube 
(T’), of course, ends up at the pump. Attaching any desiccator to the 
vacuum system is reduced to simplicity itself by the use of an intermedi- 
ate length of copper tubing (L) of any length and curvature desired, 
‘onnecting it by the sleeves (S). Any desiccator must be able to with- 

outside pressure of up to one atmosphere, and some may be 


VOLUME 26, NUMBER 2 75 


ordered that can be fitted with a steel wire screen that acts as a guard 
against flying fragments in the event of an implosion. 

Outside the freezer, a small number of line valves are added. One (A) 
is located near the end of the copper tubing leading to the freezer. An- 
other (B) is located at such point as a vacuum dehydration line is desired. 
It may be found advantageous to have two or more of these. Valve (C) 
shuts off the pump alone, for the pump, when not in operation, will grad- 
ually bleed air back into the system and this valve allows the pump to be 
segregated from the entire system while leaving the manometer con- 
nected. The last valve (D) shuts off the manometer alone. Its purpose 
is to protect the manometer. This valve should be left either just cracked 
open, or closed and used only when actually measuring the vacuum. 
Otherwise a sudden loss of vacuum in the system (as may easily happen 
if one forgets to shut off the appropriate valve when removing a desic- 
cator) may cause the mercury to blow through its glass top. 


Procedure 


Let us now proceed through a freeze-dry operation step by step. First 
one must kill the larva. While this can be done in boiling water, a cyanide 
jar or by other methods, I prefer to place it in some suitable container 
and simply let it freeze for two hours or so. One can let the caterpillar 
crawl along its foodplant, and with luck and if the animal is not too ac- 
tive, it may remain grasping the leaf or stem in a natural posture. Should 
it freeze in an undesirable position, let it thaw sufficiently (including its 
interior) before attempting to manipulate it, or the specimen will crack. 
Then refreeze it in such position as you choose. The desiccator is assumed 
to have been prepared and below freezing. The weight of the frozen 
larva is quickly recorded on a balance that will weigh to about “%o of a 
gm. One can thus later determine the point at which a weight loss of 75% 
is reached. This represents approximately 90% of the total water content, 
and at this point the process is sufficiently advanced to allow removal and 
permanent storage in the collection. Since the ambient air always con- 
tains some moisture, the desiccator should not be opened until it has 
warmed to room temperature. Otherwise contact of the frozen larval sur- 
face with the warm moist air will be apt to ruin the appearance of the 
specimen. I make it a practice to leave it thus at room temperature, i.e. 
in a state of vacuum dehydration, for 24 hrs. One can hasten this process 
by use of a heat lamp provided the heat is administered with restraint. 
Of passing interest is the fact that about 5% of the water content of the 
larva is chemically bound, and therefore unavailable for evaporation. 
After a bit of practice it will be possible to judge with reasonable accu- 


76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


racy when the specimen is ready without having to bother with weighing 
every individual. 

The frozen specimen is placed in the desiccator after weighing, the lid 
tightly sealed, and the container returned to the freezer, connected to the 
vacuum line, and the vacuum applied. The weighing and transfer of the 
larva should of course be accomplished as rapidly as possible so as not to 
allow thawing. When the vacuum pull-down is complete, close Valve 
(A) and the stop-cock (G), disconnect the desiccator from the system 
and move it to another part of the freezer in order to be ready for the 
next desiccator. During the drying process keep a check on the vacuum, 
and on the color of the desiccant, replacing the latter if necessary. If the 
foodplant is to be included as well, it too is preserved by the process. 
Contrary to what one might expect, dried specimens will not resorb wa- 
ter even in a humid climate, but it is important to remember that they 
are attractive to pests such as dermestids—and even to squirrels, as one 
collector discovered to his distress. 

Another possibility is illustrated by the case of a caterpillar that has 
been attacked by a stink bug or other sucking creature. One can hasten 
to place a flaccid carcass of this sort, if fresh, in a desiccator without prior 
freezing and apply the vacuum. The empty skin will expand just like a 
toy balloon. A larva that has sickened and become flabby can sometimes 
be preserved in a similar manner. If there is danger of overstretching, 
the vacuum may be held at any desired level by closing the stop-cock and 
letting the specimen freeze at that level of vacuum. Then, when thor- 
oughly frozen, full vacuum may be applied. 

If a spreading board is cut into sections short enough to fit into a 
desiccator, the occasional mounted butterfly or moth can be rapidly dried. 
Often their abdomens may be soft and tender, so to avoid unsightly ex- 
pansion freeze-drying is the method of choice. I have successfully ap- 
plied this method on occasion to fresh specimens during the course of 
photographing plates for Moths of America North of Mexico, when a 
freshly caught specimen happened along that could be used to better 
advantage than the museum specimen originally chosen. There is scope 
for numerous experiments of this sort. 

Meryman (1960) recommends that freezing be done at modest tem- 


peratures as in a home freezer, rather than at very low temperatures. 
Such relatively high temperatures cause the water content of the larva 
to freeze in large extracellular crystals that easily sublimate. As they 


grow, water is removed from both intracellular and extracellular locations 
without damage to the tissues or cells. Sudden very deep freezing will 
se the formation of small intracellular crystals as well, causing possible 


VoLUME 26, NUMBER 2 ih 


histological damage and increasing the drying time due to a slower rate 
of sublimation. However, the quick-freeze method provides the outstand- 
ing advantage of ensuring a lifelike position on the foodplant as shown 
on the color plate. Liquid nitrogen (used by cattle breeders among 
others) may be tried (temperature —320° F.) provided one is familiar 
with the proper procedures and precautions. Or one can use dry ice 
(temperature -110° F.) in alcohol or other compatible liquid, or a low 
temperature freezing unit. Another good agent is a quick-freeze aerosol 
such as ethyl chloride, or “Cryokwik” (a trade name for an aerosol mix- 
ture of fluorinated hydrocarbons ). 

Some of the larger larvae have comparatively thick integuments, and 
as the integument dries it presents an ever-increasing barrier to the pas- 
sage of water vapor molecules from within the specimen to without. 
However, with a fresh larva the increase in drying time is negligible, and 
such large caterpillars as Hyalophora cecropia or Citheronia regalis will 
be weil dried in 72 hours. It has been reported by Blum & Woodring 
(1963) that larvae frozen for weeks or months without attention to dry- 
ing tend to distort and toughen, thus making the process less satisfactory. 
In such cases, as with the tough covering of pupae, holes punched through 
the integument will shorten the drying time. The same authors state that 
some, but not all, greens eventually tend to fade or change color due to 
chemical differences in the chromatophores. So far in the year and a half 
I have used this process, no noticeable change has occurred, but only 
time will tell. 

Vacuum dehydration, which is carried out at room temperature, will 
be found satisfactory at times, and here the selection of appropriate speci- 
mens is of importance, as thin-skinned insects will stretch or even burst. 
The two advantages of this method are the greater speed of drying and 
the fact that no freezer is required. As a practical matter one can dis- 
count the latter since the majority of specimens will turn out more satis- 
factorily if freeze-dried at least in the initial stages of the procedure. 
With vacuum dehydration, of course, the vacuum is essential, since the 
drying process must proceed more rapidly than the enzymatic activity 
which at room temperature promotes rapid putrefaction. 

As for pinning the specimens, an anal wire wrapped around a pin may 
be used as is usually done with inflated larvae; or a pin may be inserted 
directly through the dorsum after drying. A drop of cement is generally 
required for stability. As a rule, specimens should not be pinned before 
drying, as chemical action in some cases will tend to cause a dark spot 
around the pin. On the other hand, some very small larvae will have to 
be pinned while only semi-frozen since they may break when dry. 


78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Larvae prepared by either process can be reconstituted for dissection 
or histological study (Van Cleave & Ross, 1947; Harris, 1964). To quote 
from a personal communication to the author from Mr. Harris: “We used 
from 0.5% to around 2% tribasic sodium phosphate and usually incubate 
at about 37° C. Be careful not to overdo things. I would point out the 
following: As far as histology is concerned it should not be necessary to 
use the reconstituting agent as no shrinkage should have taken place, and 
all that is necessary is to place the specimen in a warm preservative for 
the usual process of dehydration and embedding, etc. However, there 
are times when one is not quite sure and it may be safer to use the agent 
before further treatment. If you are able to look at a copy of Man, an 
article by A. Sanderson on The Study of Mummified and Dried Human 
Tissues published about 1959, you will find many interesting points that 
could be applied to entomology, etc.” I myself have not yet attempted 
reconstitution. 

I shall close by wishing the reader many happy hours with flaring tools 
and wrenches, and success in his endeavours. 


SUMMARY 


The principles involved in both freeze-drying and vacuum dehydration 
lie in the permanent removal of water vapor that evaporates from the 
tissues of the specimen. A method is presented of setting up a reasonably 
flexible apparatus on the basis of equipment at a cost that should be 
within the reach of most small laboratories and many private collectors. 
I have confined myself to the more practical matters wth enough theory 
to enable the reader to gain a reasonable understanding of what he is 
doing in order to prepare natural and lifelike specimens of larvae and 
similar insects, and to enable him to use his own ingenuity in improving 
or varying the apparatus according to his inclination, needs and budget. 
One or two theoretical matters are mentioned that are not applicable to 
the system described, because they will be found in the literature and 
might cause confusion when so encountered. 


ACKNOWLEDGMENTS 


[ should like to extend my warm thanks to Dr. Hermann A. Flaschka, 


who has given much enthusiastic help and has steered me safely through 
some of the more dreadful pitfalls of physical chemistry. He has also 
taken considerable time, effort and care in reading the manuscript and 
has offered much by way of constructive criticism. The helpful coopera- 
tion of my colleague, Charles R. Edwards, is also appreciated. He shares 


in the daily work in the laboratory and grins over my shoulder and comes 
0 the rescue when my own machinations with valves, wrenches and 


VoLUME 26, NUMBER 2 72 


plumbing end up in disaster. I should also like to extend grateful thanks 
to Mr. R. H. Harris, of the Experimental Laboratory of The British Mu- 
seum (Natural History) in London, for the cheerful, stimulating and 
willing help and advice he has shared from his store of expert knowledge 
concerning the processes described. 


LITERATURE CITED 


Bium, M. S. ann J. P. Wooprinc. 1963. Preservation of larvae by vacuum dehy- 
dration. J. Kansas Entomol. Soc. 36: 96-101. 

FiascukaA, H. A. AND J. FLoyp. 1969. A simplified method of freeze-drying cater- 
pillars. J. Lepid. Soc. 23: 43-48. 

Harris, R. H. 1964. Vacuum dehydration and freeze drying of entire biological 
specimens. Experimental Laboratory, British Museum (Natural History), London. 
Ann. & Mag. Natur. Hist. Ser. 13, vii: 65-74. 

. 1968. A new apparatus for freeze-drying whole biological specimens. Med. 
Biol. Ill. 18: 180-182. 

MeryYMAN, H. T. 1960. The preparation of biological museum specimens by freeze- 
drying. Curator 3: 5-19. 

1961. The preparation of biological museum specimens by freeze-drying: 
II Instrumentation. Curator 4: 153-174. 

VAN CLEAveE, H. J. AND J. A. Ross. 1947. A method for reclaiming dried zoological 
specimens. Science 105: 318-321. 

Wooprinec, J. P. anp M. S. BLum. 1963. Freeze-drying of spiders and immature 
insects. Ann. Entomol. Soc. Amer. 56: 138-141. 


A NEW SPECIES OF THE GENUS PYROMORPHA 
HERRICH-SCHAEFFER (PYROMORPHIDAE ) 


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


Some male specimens of this species have been in my collection for 
over five years. I have delayed describing it, hoping that I would take at 
least one female. In this I have been disappointed. As I have a sizeable 
series of males I am offering this description of it. 


Pyromorpha caelebs A. Blanchard, new species 


Male (Fig. 1): Head black, closely scaled, except on vertex where some long scales 
project forward between antennae or lean against their scapes. Tongue strong. Labial 
palpus short, filiform. Maxillary palpus vestigial. Antenna bipectinate, of about 35 
segments, black, closely scaled above; each pectination slightly swollen near apex, 
tapering to base, bearing two rows of cilia. Collar, thorax, patagiae and abdomen 
black. Legs slender, closely scaled, black except yellow inner side of foreleg, some 
yellow scales distally on midfemur; one pair of terminal, rudimentary spurs on mid 


80 JouURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Pyromorpha caelebs, Holotype. 


and hindtibiae. Forewing above thinly scaled, semitranslucent, black except for 
ochreous-yellow fascia anteriorly limited on costa, posterior limit of which extends 
along basal two-thirds of cubital vein, slides in cell to radial vein, and follows Ry, to 
costa; distal half of fascia well defined, basal half less definite because of sprinkling 
of yellow scales between cubital and first anal veins; fringe consisting of somewhat 
irregularly planted, bluish-black, narrow scales. Hindwing above black; very light 
and narrow sprinkling of yellow scales along costa; fringe similar to that of forewing. 
Forewing beneath as above, posterior margin of yellow fascia more diffuse. Hindwing 
beneath as above except for well-marked, yellow fascia between costa and discal cell. 
Venation of wings as shown in fig. 2; Rg and R4 of forewing either separate or connate. 

Length of forewing: 10 to 12 millimeters. 

Genitalia: As represented in Figs. 3, 4 and 5. Uncus absent or possibly represented 
by mucrones on each half of tegumen; these separate except where they meet strongly 
sclerotized gnathos. Juxta oval with small indentation under aedeagus. Dorsal part 
of anellus a complicated assemblage of two spiny pads, one on each side of aedeagus, 
tied together back of it by narrow transtilla, supported on each side from ninth ab- 
dominal segment by two sclerotized, contorted arms. 

Female: Unknown. 

Holotype: Male, Fort Davis, Texas, Hospital Canyon back of Historical Fort, 18 
May 1971, deposited in the National Museum of Natural History. Type n°. 71981. 

Paratypes: (All males). Fort Davis, Texas, Hospital Canyon, 17 May 1966, six; 
ll June 1969, ten; 18 May 1971, thirteen. Davis Mts., Mt. Locke, McDonald Ob- 

ervatory (6700’), 25 May 1968, four; 10 June 1969, five. Fort Davis, one mile north 
of city near Limpia Creek, 21 May 1971, four. All these paratypes collected in 15 
t fluorescent traps by A. and M. E. Blanchard. Paratypes will be deposited in the 

| Museum of Natural History, in the American Museum of Natural History 

the British Museum (Natural History). Some will remain in my own col- 

he mn the National Museum a specimen labeled only “Kerrville, Texas” 
to be conspecific; but it was never spread, its wings are wrinkled and, 

ould be an interesting location, I prefer not to make it a paratype. 


VoLUME 26, NUMBER 2 81 


Fig. 2. Venation of wings; Figs. 3, 4 & 5: male genitalia: 3, posteroventral view, 
valves widely open, aedeagus omitted; 4, aedeagus, lateroventral view; 5, left side 
view, valves and aedeagus removed. The linear segments represent one millimeter. 


The male of Pyromorpha caelebs is close to that of P. dimidiata Herrich- 
Schaeffer (1855). They are about the same size and the colors of the 
background and fascia are close. They can be separated by the following 
combination of characters: The costa of the forewing of caelebs is nearly 
straight in its basal two-thirds, that of dimidiata is much more arched. 
The forewing of caelebs is narrower; the ratio of its length to its midwidth 
being approximately 2.5 instead of about 2.25 for dimidiata. The radial 
veins R3 and Ry, of the forewing of caelebs are either connate or separate 
instead of shortly stalked. The ratio of the length of the longest pectina- 
tions of the antenna to the width of the flagellum is about 2.5 instead of 


82 JouRNAL OF THE LEPIDOPTERISTS’ SOCIETY 


about 4; the flagellum is also somewhat stouter. The yellow fascia of 
caelebs is narrower because the costa is less arched, and in its basal half 
it barely exceeds the cubital vein instead of following the first anal all 
along its basal half. It is difficult to define exactly how the genitalia dif- 
fer; the juxta of caelebs is oval or very slightly cordate, that of dimidiata 
definitely cordate (concave lateral margins); the gnathos is longer and 
its base narrower, but the great thickness of the genitalia relative to their 
width makes it difficult to present on a single conventional preparation a 
satisfactory view of all the organs and particularly the gnathos. 


ACKNOWLEDGMENTS 
I wish to thank the administration of Fort Davis N.H.S. for the authori- 
zation to collect in Hospital Canyon and Mr. Curtis D. Laughlin for let- 
ting us set our traps at Mt. Locke on the McDonald Observatory grounds. 
I am grateful to Dr. R. W. Hodges for his interest and help and for revis- 
ing the manuscript. 


LITERATURE CITED 


HERRICH-SCHAEFFER. 1855. Samml. aussereur. Schmett., Pl. 43, fig. 222. 


THE LIFE HISTORY OF SCHINIA CITRINELLUS 
(NOCTUIDAE) 


D. F. Harpwick 
Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario 


Schinia citrinellus (Grote & Robinson, 1870, p. 180) feeds in the larval 
stage on Croton californicus Muell-Arg. C. californicus is distributed 
from southern California to Arizona (Munz, 1963), but citrinellus occurs 
from the southern California deserts eastward at least to central Texas 
(Brazos Co.), so presumably other species of Croton serve as its food 
plant in the more eastern areas of its range. Schinia citrinellus has two 
annual flight periods, one in the spring and one in late summer and early 
‘all. These probably correspond closely with the blossoming periods of its 
host plants. The spring flight period on the deserts of southern California 

xtends between the end of March and the middle of June. 


Behaviour 


vnid citrinellus is evidently an exclusively nocturnal species, and no 
vas noted during daylight hours on the deserts of southern 


VoLUME 26, NUMBER 2 83 


California at a time when moths were being taken in moderate abundance 
in a light trap. The eggs are deposited among the small buds or within 
the partially open buds at the apex of the plant. The rather unusual 
opaque white eggs of citrinellus are difficult to detect in the hairy, pale- 
grey vestiture of the Croton buds. 

A single wild-caught female deposited a total of 176 eggs in captivity. 
The majority of these hatched on the fourth day after deposition, a few 
on the fifth day. 

The fully formed larva has the habit of resting for a considerable period 
after it has chewed an exit hole in the chorion but before it has made its 
escape from the shell. After freeing itself from the egg, the larva makes 
its way down through the blossom or bores through the petals of the bud 
to reach the orange glands located on the surface of the receptacle; the 
first-stadium larva feeds almost exclusively on these glands. In later 
stadia, other floral parts are consumed. The small seed capsules are fed 
on to a limited extent but the more mature stone-like fruits are never 
attacked. The last-stadium larva feeds preponderantly on the buds, but 
unlike most heliothidine larvae, will also feed on the leaves of its host 
plant. At the cessation of feeding, the larva makes its way to the ground 
and tunnels into the soil to pupate. 


Descriptions of Stages 


The ultimate-stadium larva and the pupa of Schinia citrinellus have 
previously been described by Comstock (1931). The immature stages 
on which the following descriptions are based, constituted the progeny 
of a single female taken at Thousand Palms, near Indio, California. The 
durations of stadia listed are those obtained from rearings maintained at 
room temperature. Rearing techniques employed were those outlined by 
Hardwick (1958). The estimate of variation following the means for 
various values is the standard deviation. 


Adult (Fig. 1). Head, thorax and forewing dull creamy-yellow; museum specimens 
often rather intensely yellow as a result of the action of the killing agent. Abdomen 
cream or creamy-grey. Forewing almost immaculate, usually with a vague dark 
reniform and subterminal band; these evidently only the transmission of dark mark- 
ings from the underside of the wing. Forewing usually with a few scattered dark 
points, the most consistently present being at the apico-costal margin of the reniform 
spot. Fringe concolorous with remainder of wing. Hind wing white with a brown 
outer-marginal band containing a white median shade; outer band often evanescent 
near anal angle. A brown discal spot usually present. Fringe white. Underside of 
forewing white with brown orbicular and reniform spots, and subterminal band. 
Usually a series of brown sagittate marks at outer margin of wing, these occasionally 
fusing to form a brown marginal band. Costal and outer areas of wing often suffused 
with yellow. Underside of hind wing white or pale cream, often with a brown spot 
on disc, and another near anal angle of wing. 


84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-5. Schinia citrinellus (Grt.) and its food plant. 1, Adult, Palm Springs, 
Calif.; 2, food plant, Croton californicus Muell.-Arg.; 3, 5, dorsal aspect of ultimate- 
stadium larvae; 4, ventral aspect of pupae. 


Expanse: 26.1 + 14 mm (34 specimens). 
Egg. Of a peculiar white opacity unusual in heliothidine eggs; showing little change 


until a few hours before hatching when mouth parts and head capsule become visible 
through chorion. 


Dimensions of egg: length, 0.593 + 0.030 mm; diameter, 0.451 + 0.033 mm (20 
eggs ). 

Incubation period: 4.1 + 0.3 days (157 eggs). 

First-Stadium Larva. Head black. Prothoracic and suranal shields dark smoky- 
brown. Trunk creamy-grey. Spiracles with light- to medium-brown rims. Thoracic 
legs medium smoky-brown. 

Head width: 0.300 + 0.013 mm (25 larvae ). 

Duration of stadium: 4.1 + 0.3 days (46 larvae ). 

Second-Stadium Larva. Head pale pinkish-cream, mottled with pale fawn dorsally. 
Prothoracic and suranal shields white or pale cream, mottled with light green. Macula- 


VoLUME 26, NUMBER 2 85 


Figs. 6, 7. Apical abdominal segments of pupa. 6, Ventral; 7, right lateral. 


tion of trunk well defined. Mid-dorsal band greyish-green. Subdorsal area with white 
marginal lines and a greyish-green median band. Supraspiracular area greyish-green, 
usually with a white or pale cream, irregular and broken, median line. Spiracular 
band narrow, white or pale cream, often discontinuous. Suprapodal area light greyish- 
green. Mid-ventral area pale grey. Spiracles with light-brown rims. Setae of trunk 
light brown with white or cream bases; setal bases interrupting maculation of trunk. 
Thoracic legs pale pinkish-cream. 

Head width: 0.558 + 0.026 mm (25 larvae). 

Duration of stadium: 3.1 + 0.5 days (46 larvae). 

Third-Stadium Larva. Head cream, mottled with fawn. Prothoracic shield light 
green, mottled with cream, and usually with three cream longitudinal lines. Suranal 
shield light green, mottled with cream or white. Maculation of trunk complex. Mid- 
dorsal band greyish-green or light olive-green, with a white or cream, often discon- 
tinuous, median line. Subdorsal area with marginal white lines and a median band 
concolorous with mid-dorsal band; median band usually with a discontinuous white 
median line. Supraspiracular area green, with a discontinuous white median line. 
Spiracular band narrow, white or cream, occasionally discontinuous. Suprapodal area 
greyish-green, mottled with white or cream. Mid-ventral area pale grey, suffused with 
green. Setae of trunk with enlarged cream or white bases that interrupt the general 
pattern. Spiracles with light- to medium-brown rims. Thoracic legs cream suffused 
with pale green. 

Head width: 0.92 + 0.04 mm (25 larvae). 

Duration of stadium: 3.1 + 0.5 days (46 larvae). 

Fourth-Stadium Larva. Head pale cream, variably mottled dorsally with grey or 
fawn. Prothoracic shield white or pale cream, marked with dark green; usually a 
median and a pair of submarginal lines free of mottling. Suranal shield white, mottled 
with light green. Trunk greyish-green with numerous, frequently irregular and broken, 
longitudinal white lines; setal bases large and white. Mid-dorsal band medium to 
dark greyish-green with a discontinuous median white line. Subdorsal area white with 
a pair of discontinuous, greyish-green median lines. Supraspiracular area green, heav- 
ily and irregularly mottled with white. Spiracular band irregular, white or pale cream. 
Suprapodal area green, of a more greyish appearance than dorsum, mottled with white 
or pale cream. Mid-ventral area pale grey suffused with pale green. Spiracles with 


86 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


light-brown rims. Thoracic legs pale cream, suffused with brown distally and with 
green proximally. 

Head width: 1.64 + 0.09 mm (25 larvae). 

Duration of stadium: 3.5 + 0.7 days (46 larvae). 

Fifth-Stadium Larva (Figs. 3, 4). Head flesh-coloured, inconspicuously mottled 
with pale fawn. Arcs free of mottling diverging upward and outward from centre of 
face. Prothoracic shield pale cream or white, mottled with pale greenish-grey; a 
median and a pair of submarginal lines free of mottling. Suranal shield pale greyish- 
green, mottled with white or pale cream. Maculation of trunk highly unusual, and 
with normal zonation of dorsum difficult or impossible to distinguish. Dorsum creamy- 
white with a confusing array of circles, bars, and undulating lines of greyish-green. 
Spiracular band creamy-white. Suprapodal area greyish-green mottled with cream. 
Mid-ventral area paler green than suprapodal area. Spiracles with light-brown rims. 
Thoracic legs cream, weakly suffused with green. 

Head width: 2.52 + 0.08 mm (11 larvae). 

Duration of stadium: 4.9 + 0.8 days (46 larvae). 

Pupa (Figs. 4, 6, 7). Uniform mahogany brown. Spiracles on abdominal segments 
5, 6 and 7 borne in shallow oval pits; spiracular sclerites moderately projecting. 
Anterior marginal areas of abdominal segments 5, 6 and 7, with a narrow band of 
fine but rather conspicuous pitting. Proboscis terminating well anterior to apexes of 
wings. Cremaster consisting of an elongate conical prolongation of the tenth abdom- 
inal segment bearing spically four elongate slender setae. 

Length from anterior end to posterior margin of fourth abdominal segment: 9.2 + 
0.3 mm (14 pupae). 


ACKNOWLEDGMENTS 


I am grateful to Mr. John E. H. Martin of this Institute for assistance 
in the field and for the photographs accompanying this paper. I appreci- 
ate the help of my associate, Mr. Eric Rockburne, who measured the 
immature stages and drew the cremaster area of the pupa. 


LITERATURE CITED 


Comstock, J. A. 1931. Studies in Pacific Coast Lepidoptera. Bull. So. Calif. Acad. 
Sci. 30; 15-20. 

Grote, A. R. ann C. T. Rospinson, 1870. Description of American Lepidoptera, 
No. 5. Trans. Amer. Entomol. Soc. 3: 176-189. 

Harpwick, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed 
species of Schinia (Lepidoptera:Noctuidae), with notes on the Heliothidinae. 
Can. Entomol. Suppl. 6. 116 p. 


whee? - A. 1963. A California Flora. University of California Press, Berkeley. 
68] p. 


LIFE HISTORY NOTES ON CALLOSAMIA SECURIFERA (SATURNIIDAE) 

On 29 August 1970, three 2nd instar larvae 
era (Massen) were found on a low shrubb 
giniana L.. (Sweet Bay). The larvae were found approximately eight miles north of 
McClellanville, Charleston County, South Carolina. The area is typical of coastal South 
Carolina, with many of the swampy pine forests where M. virginiana is common. The 


and nine eggshells of Callosamia securif- 
y example of its foodplant, Magnolia vir- 


VoLuME 26, NUMBER 2 87 


a 


Fig. 1. Callosamia securifera (Massen): A, ova shells; B, 5th instar larva; C, 
cocoon; D, adult male. 


88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


McClellanville area is of special interest, as it is one of the few areas in the US. 
where all three species of Callosamia fly sympatrically: C. angulifera Walker in as- 
sociation with Liriodendron tulipifera L. (Tulip Tree), C. promethea Drury in associa- 
tion with Symplocos tinctoria L’Hérit (Horse Sugar), and C. securifera in association 
with M. virginiana. 

The three larvae were found singly on the undersides of the leaves, resting on the 
midrib. The eggshells were all on the underside of one leaf, located near the tip in 
a small cluster. Perhaps the ova are usually deposited in this manner, but this is only 
conjecture, as no other eggs have been found. Even after stripping the shrub to the 
ground, there was no trace of the other six larvae. Evidently the attrition rate to 
parasites and predators is quite high. Interestingly, these three larvae seemed to have 
lost the usual gregarious tendencies so prevalent with early instar Callosamia larvae, 
as they were all found some distance apart on the plant. The larvae from previous 
laboratory-reared broods of the three Callosamia species invariably remained gregari- 
ous until third instar. 

According to all known records, this is the first instance of C. securifera larvae being 
found in the natural state, although the cocoons are often found in the McClellanville 
area during the winter months. Since F. M. Jones’s original description of the adults 
and larvae (1909, Entomol. News 20: 49-52), little has been published on C. securif- 
era. It is now known to be a distinct species and not a subspecies of C. angulifera as 
believed previously. The totally different foodplant, sympatric flight with C. angulif- 
era, distinctive reproductive habits, and unique cocoon leave no doubt that C. securif- 
era is indeed a valid species. 

The larvae were reared outdoors on caged examples of M. virginiana, but unfortu- 
nately only one of the three was reared through to an adult. One larva escaped, while 
a second fell prey to a stinkbug in late 5th instar. (However it was vacuum-freeze 
dried and preserved for the accompanying figure.) Due to normal limits of variation, 
the larvae are impossible to distinguish from those of C. angulifera. Hopefully more 
careful observation of these two species in the larval stages will yield some distinguish- 
ing characters. The single remaining larva spun a cocoon in early October of 1970. 

The cocoon is perhaps the most distinctive aspect of the life history of C. securifera. 
The always large, baggy, pendant cocoon is wrapped in several leaves and attached 
securely to a twig of the foodplant. When freshly spun the cocoon is of a beautiful 
silvery tan but fades somewhat with weathering. The cocoon from the last remaining 
larva, an excellent example, is shown on the accompanying figures. While the length 
of the cocoon’s peduncle varies, as does the irregular shape, it never resembles the 
conical, compact cocoon of C. promethea or the loosely woven dark brown cocoon of 
C. angulifera. 

A male emerged from the cocoon on 22 March 1971, shortly after sunrise. The 
premature emergence (4—6 weeks early) of the adult was puzzling, since it was reared 
under natural conditions and left outdoors throughout the winter until it emerged. 
The normal flight period for the first brood of C. securifera is from late April into 
early May, depending on weather conditions. There is a second brood in early to mid- 
August. These late summer adults of C. securifera, like those of summer brood C. 
angulifera, are much darker than the spring adults. 

More careful notes were not taken on the life history of these first wild larvae, in 
anticipation of another, more detailed paper on this species by Dr. Richard B. Domi- 
nick and Mr. Charles R. Edwards of the Charleston Museum, Charleston, South Caro- 
lina. These men have spent the better part of two years thoroughly studying the 
habits and life history of C. securifera, Their paper, now in preparation, will have a 
wealth of previously unknown facts concerning the life history and ecology of this 
rare ind he autiful silk moth. I extend my sincere thanks to these close friends who 

00k time out from an unbelievably busy schedule to photograph the figures 


irticle. 


mL D. VAn Buskirk, 4512 47th S.W., Seattle, Washington 98116. 


VOLUME 26, NUMBER 2 89 


THE LIFE HISTORY OF SCHINIA JAEGERI (NOCTUIDAE) 


D. F. Harpwick 
Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario 


Schinia jaegeri (Sperry, 1940) feeds in the larval stage in the heads of 
Orcutt’s Aster, Machaeranthera orcuttii (Vasey & Rose) and of the Mecca 
Aster, M. cognata (Hall). Both asters are endemic to the Colorado Desert 
of southern California and to areas of northern Baja California (Munz, 
1963). The distribution of jaegeri is probably coincident with the dis- 
tribution of its two food plants. 

Northward of the Colorado Desert jaegeri is replaced by the closely 
related Schinia ligeae (Smith) which feeds in the larval stage on the 
Mojave Aster, Machaeranthera tortifolia (Gray) (see Hardwick, 1971). 
The two species of moths are obviously closely related: their patterns of 
development are much the same, the adults are structurally very similar, 
and the pupae are essentially indistinguishable. 

In areas of the Colorado Desert where its food plants are abundant and 
in early blossom, adults of jaegeri may usually be found without great 
difficulty. The species is in flight from the last week of March to the end 
of April. 


Behaviour 


Schinia jaegeri is active only at night. During the daylight hours, the 
adults may be found resting either on the buds and blossoms or among 
the foliage of the two species of Machaeranthera on which the larva feeds. 
The eggs are laid either on the exterior of the unopened bracts of these, 
or between the florets of the open blossom. Occasionally eggs are de- 
posited within the throats of individual florets. In Split Mountain Can- 
yon, San Diego Co., a number of eggs deposited in exposed positions on 
the outside of the buds of Orcutt’s Aster were found to be parasitized by 
a species of Trichogramma. 

Of the few females confined in captivity, the maximum number of eggs 
deposited by any individual was 19. 

The majority of eggs hatch on the seventh day after deposition. Larvae 
hatching from eggs deposited within the head immediately attack the 
florets. Larvae hatching from eggs deposited on the unopened bud gen- 
erally enter the bud at its extreme apex. Occasionally a larvae works its 
way downward under the bracts and attacks the bud at its very base by 
boring into the fleshy tissues of the receptacle. The survival rate among 
such receptacle—boring larvae is probably not very high. Among a large 


90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


number of aster buds that were dissected, very few penetrations through 
the receptacle to the seed layer were noted. 

Larvae generally remain within the initial head until they have reached 
one of the median stadia. Usually during the fourth stadium, the larva 
quits the first head and enters a second which it bores into from the top. 
Feeding is usually completed within the second head. 

Forty-six of 52 individually reared larvae matured in five larvae stadia, 
the remainder in four. 

The ultimate-stadium larva assumes a decided reddish tone on the dor- 
sum a day or so before it enters the earth to form its pupal cell. The ma- 
jority of the year is spent as a pupa a few inches below the surface of 
the soil. 


Description of Stages 


The following descriptions of immature stages are based on the progeny 
of three females taken at Biskra Palms, near Indio, California, and on 
eggs dissected from the Mecca Aster in Painted Canyon, Mecca, Calif., 
and from Orcutt’s Aster in Split Mountain Canyon, San Diego Co., Cali- 
fornia. Rearing techniques employed were those outlined by Hardwick 
(1958). The estimate of variation following the mean for various values 
is the standard deviation. 


Adult (Figs. 1, 2). Head, thorax and abdomen clothed with cream vestiture. 
Forewing cream, vaguely marked with white and occasionally shaded with pallid 
fawn. Transverse anterior line rarely defined, white when present, strongly angular 
outwardly, the apex of the angle resting on the cubitus stem. Basal space uniform 
cream. Transverse posterior line rarely defined, white, excurved around cell, then 
angling inward to trailing margin. Median space narrow, occasionally suffused with 
pallid fawn; orbicular spot not defined, reniform spot occasionally evident as a narrow, 
whitish, arc. Subterminal space occasionally suffused with pallid fawn. Fringe and 
terminal space concolorous with basal space. Hind wing usually cream, occasionally 
pallid fawn; fringe cream. Underside of both wings shining cream. 

Expanse: 29.7 + 2.4 mm (58 specimens). 

Egg (Fig. 3). Pale cream when deposited and remaining unchanged during the 
next day. Egg assuming a slight pinkish tone on second day; then darkening to a 
light orange during third and fourth days. Anterior half of egg gradually becoming 
darker in colour than posterior half; during the fifth and sixth days, anterior half 
reddish-brown and posterior half light greyish-orange. Egg turning grey on day of 
hatching, and head capsule becoming visible through chorion a few hours before 
hatching. 


Dimensions of egg: length, 1.66 + 0.07 mm; diameter, 0.84 + 0.04 mm (26 eggs). 

Incubation period: 7.2 +0.6 days (27 eggs). 

First-Stadium Larva. Head medium orange-brown to blackish-brown. Prothoracic 
and suranal shields dark smoky-brown. Trunk usually a translucent purplish-brown 
when larva hatches, becoming light yellow or cream after feeding. Spiracles with 
medium- to light-brown rims. Thoracic legs smoky-brown. 

Head width: 0.A7T7 + 0.016 mm (23 larvae). 

Duration of stadium: larvae maturing in 5 stadia, 4.7 + 1.4 days (46 larvae); 


~ 


larvae maturing in 4 stadia, 5.3 + 0.6 days (6 larvae). 


VoLUME 26, N 91 


Figs. 1-5. Schinia jaegeri (Sperry) and its food plants. 1, Adult, Split Mountain 
Canyon, San Diego Co., Calif.; 2, a pair of adults, presumably freshly broken from 


copula, on head of Mecca Aster; 3, egg on bud of Mecca Aster; 4, Orcutt’s Aster; 5, 
ventral aspect of pupae. 


Second-Stadium Larva. Head orange-brown, variably mottled and reticulated with 
slightly darker brown, but usually inconspicuously so. Prothoracic shield medium 
chocolate-brown to dark smoky-brown, usually with a light median line. Suranal 
shield smoky-fawn to dark smoky-brown. Trunk cream or greyish-cream. Spiracles 
with light- to medium-brown rims. Thoracic legs smoky-brown. 

Head width: 0.76 + 0.04 mm (19 larvae). 

Duration of stadium: larvae maturing in 5 stadia, 2.5 + 0.7 days (46 larvae); 
larvae maturing in 4 stadia, 3.3 + 0.9 days (6 larvae). 


92 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


dd 
UG 


» lee HE: oe 


Yy 


Y> 
MYM YW YP 


YY : y ; ¥ y | hh 
MA 


Figs. 6-9. Schinia jaegeri (Sperry) and its habitat. 6, Painted Canyon, Mecca, 
Calif., with clumps of the Mecca Aster; 7-9, right lateral and dorsal aspects of ulti- 
mate-stadium larvae. 


Antepenultimate-Third-Stadium Larva. Head orange-brown, with darker-brown 
reticulation and shading dorsally. Prothoracic shield smoky-fawn, marked with black- 
ish-brown and with a median and a pair of submarginal, cream lines. Suranal shield 
smoky-brown, often with three, cream or fawn, longitudinal lines. Trunk cream or 
greyish-cream, usually with 2 pairs of pale lines on dorsum. Spiracles with dark- 
brown or black rims. Thoracic legs smoky-fawn to medium smoky-brown. 

Head width: 1.10 + 0.05 mm (33 larvae). 

Duration of stadium: 2.4 + 0.6 days (46 larvae). 


schinia jaegeri (Sperry), apical abdominal segments of pupa. 10, 
lateral. 


VoLUME 26, NUMBER 2 93 


Penultimate-Stadium Larva. Head orange, shaded and reticulated with medium 
brown. Prothoracic shield very dark brown or black, with a white or cream median 
line and usually with a similar submarginal line on either side. Suranal shield dark 
smoky-brown with three longitudinal lines of white or cream, the median line often 
evanescing. Mid-dorsal band light grey or creamy-grey. Subdorsal area paler than 
mid-dorsal band, margined on either side by a pale-yellow line. Supraspiracular area 
and spiracular band concolorous with subdorsal area, separated from each other by 
a pale-yellow line. Ventral region pallid grey. Spiracles with black rims. Thoracic 
legs smoky-fawn. 

Head width: fourth-stadium larvae maturing in five stadia, 1.60 + 0.06 mm (37 
larvae); third-stadium larvae maturing in four stadia, 1.32 + 0.06 mm (6 larvae). 

Duration of stadium: fourth stadium of larvae maturing in five stadia, 2.6 + 0.3 
days (46 larvae); third stadium of larvae maturing in four stadia, 3.7 + 0.9 days (6 
larvae). 

Ultimate-Stadium Larva (Fig. 7-9). Head orange, often with a pair of dark-brown 
ares on vertex. Prothoracic shield light brown, marked with black, and with three 
longitudinal bands of white or cream. Suranal shield medium to dark smoky-brown, 
with a cream submarginal line on either side and with at least a partial cream median 
line. Mid-dorsal band medium slate-grey, margined by pale-yellow bands. Subdorsal 
area paler grey than mid-dorsal band. Supraspiracular area somewhat paler than sub- 
dorsal area, margined on either side by a pale-yellow line; ventral marginal line usu- 
ally broken. Spiracular band concolorous with subdorsal area, shaded in middle of 
each segment with pale creamy-grey, and margined ventrally by an irregular pale 
band. Suprapodal and mid-ventral areas light grey. Spiracles with black rims. 
Thoracic legs cream, tinged with orange. 

Head width: 2.23 + 0.13 mm (20 larvae). 

Duration of stadium: fifth stadium of larvae maturing in five stadia, 9.6 + 1.8 
days (46 larvae); fourth stadium of larvae maturing in four stadia, 9.0 + 1.4 days 
(6 larvae). 

Pupa (Figs. 5, 10, 11). Essentially indistinguishable from that of Schinia ligeae 
(Smith) (see Hardwick, 1971). Anterior marginal areas of abdominal segments 
often forming a more prominent ridge than in ligeae. Lateral cremaster setae present 
in some pupae of ligeae, absent from all pupae of jaegeri examined. 

Length from anterior end to posterior margin of fourth abdominal segment: 9.1 + 
0.5 mm (25 pupae). 


ACKNOWLEDGMENTS 


I am grateful to Mr. John E. H. Martin of the Entomology Research 
Institute for the fine photographs accompanying this paper, and to my 
associate, Mr. Eric Rockburne, for measuring larval structures and draw- 
ing the cremaster area of the pupa. 


LITERATURE CITED 


Harpwick, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed 
species of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae. 
Can. Entomol. Suppl. 6. 116 p. 

1971. The life history of Schinia ligeae (Noctuidae). J. Lepid. Soc. 25: 
274-277. 

Munz, P. A. 1963. A California Flora. University of California Press, Berkeley. 
1681 p. 

SPERRY, GRACE H. 1940. A new noctuid from the desert region of southern Cali- 
fornia. Can. Entomol. 40: 147-148. 


94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


STUDIES ON THE CATOCALA (NOCTUIDAE) OF SOUTHERN 
NEW ENGLAND. III. MATING RESULTS WITH C. RELICTA 
WALKER 


THEODORE D. SARGENT 


Department of Zoology, University of Massachusetts 
Amherst, Massachusetts 01002 


Successful matings of Catocala moths have apparently been rarely 
achieved in captivity. I am aware of no published accounts of such mat- 
ings in the North American literature. Considering, however, the many 
genetic, ecological, and evolutionary questions posed by this vast assem- 
blage of moths, successful matings should provide results of wide interest. 
Consequently, I have begun attempts to mate Catocala, and this paper 
reports initial successes with C. relicta Walker. 

C. relicta was selected for initial study, as its close European ally, C. 
fraxini Linnaeus, has been mated in captivity (e.g. Cockayne et al., 1937- 
38), and further, C. fraxini and C. relicta have been successfully hybrid- 
ized (Meyer, 1952). 

The present report is based on studies conducted from 1969-1971 in 
Leverett, Massachusetts. During that period a total of 541 adult C. relicta 
were reared from eggs, and 25 matings were observed. 


METHODS AND MATERIALS 


C. relicta occurs in three forms in the northeast (Beutenmiiller, 1903; 
Forbes, 1954): typical, with clouding of black scales between am. line 
and base of forewing, and between pm. and st. lines (Fig. la, e, f); 
clara, with these areas largely white (Fig. lc, d); and phrynia, with a 
nearly even dusting of black scales over the entire forewing (Fig. Ib). 
The present mating studies involved only typical and clara moths, al- 
though wild-caught specimens from Leverett have been approximately 
30% typical, 60% clara, and 10% phrynia (1968-71, 43 records). 


Rearing Procedures 


The larval stages and pupa of C. relicta have been previously described 
in detail by Clark (1888) and Rowley & Berry (1910). 
in the present study, eggs were obtained by placing wild-caught or 
nated adult females into paper bags which were hung outdoors for sev- 
| days. The eggs were then transferred on small pieces of paper into 
tood jars which were left outdoors to overwinter. Larvae hatched 
'-May (initial hatch from 8-18 May over the three years ) and were 


VoLUME 26, NUMBER 2 95 


amen as (eras Vs 
cms. 


Fig. 1. Specimens of C. relicta: a. P: wild-caught typical female; b. wild-caught 
phrynia male; c, d. lightest and darkest clara obtained in F: crosses; e, f. lightest and 
darkest typical obtained in F, crosses. 


immediately supplied with fresh poplar leaves (Populus tremuloides ). 
At 1-2 weeks of age the larvae were transferred individually into pint, 
plastic ice-cream containers. Herein each larva was provided with a 
poplar twig for resting and a dry, curled beech leaf (Fagus grandifolia) 


96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Mating success with C. relicta, 1969-70. 


No. No. Percent 
Situation Paired Mated Mated 
LOCATION 
Indoors BA 20 95% 
Outdoors 5 4 80% 
CAGE 
Small 6 5 83% 
Large 10 10 100% 
Cylinder 10 9 90% 


for pupating. Fresh poplar leaves were added, and frass removed, each 
day until pupation. Pupae were usually formed in the beech leaves and 
occasionally in the fresh poplar foliage (initial pupation from 15-22 June). 
Adults generally eclosed about one month later (initial eclosion from 15- 
18 July). 

C. relicta did particularly well under these rearing conditions, with 80- 
90% survival from egg to adult being commonly obtained. 


Mating Procedures 


The plastic containers housing the individual pupae were checked each 
day for adults. When a male and female were available for pairing, they 
were transferred from these containers into one of three different types 
of cages: small, homemade of aluminum window screening on a wooden 
frame (approx. 20 X 23 x 23 cm.); large, obtained from Ward’s Natural 
Science Est. (14W 7500), made of nylon mesh on a plywood frame (ap- 
prox. 28 x 25 x 41 cm.); cylinder, homemade from aluminum window 
screening, rolled into a cylinder (18 cm. diam. xX 38 cm. high) and cov- 
ered at both ends with cardboard pie plates. These cages were then 
either placed outdoors (on stumps, and hung from limbs) or indoors (base- 
ment, approx. 15°C., with small windows allowing some natural light). 
All paired moths were provided with an opportunity to feed from small 
pieces of sponge which were soaked daily with a honey and water 
solution. 

The cages were checked by flashlight (covered with a sheet of red 
cellophane ) at intervals during the night. At these times the behaviors 
of all moths were carefully noted, but prolonged observations of courtship 
were not attempted, as the flashlight seemed to distract and disturb the 
moths to some extent. 


Mating was considered achieved only when a pair was observed in 


VoLUME 26, NUMBER 2 97 


Fig. 2. C. relicta pair in copulation on side of large cage—female above, male 
below (approximately natural size). 


actual copulation. On the day following this observation the female was 
bagged for eggs and the male killed and retained as a specimen. When 
the female died she was also retained—male, female, and eggs being 
labeled with a common symbol. 


RESULTS AND DISCUSSION 


Behavior 


Of 26 pairs of moths which were carefully observed during the sum- 
mers of 1969 and 1970, 24 (92%) were observed in copulation, and 20 
of these matings resulted in fertile eggs (i.e. produced larvae). Mating 
success was high in all three types of cages, and both outdoors and in- 
doors (Table 1). 

Courtship behavior, though not observed in detail, seemed generally 
similar to that described for other noctuids (e.g. Shorey, Andres & Hale, 
1962; Birch, 1970). The females almost invariably adopted a “calling” 
posture shortly after dusk, and maintained this posture (unless mating 
occurred ) for most of the night. In this calling posture a female elevated 
the abdomen (from either a horizontal or vertical surface) above the 
plane of her partially spread wings (the wings being vibrated rapidly 


98 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


20 


10 


OF PAIRS 


NO. 


2100-  2200- 2300- 
2200 2300 2400 


HOURS OF THE NIGHT 


Fig. 3. Time of mating (Eastern Daylight Savings Time) for 25 pairs of C. 
relicta observed during the summers of 1969-71. 


during high intensity calling), and protruded the pheromone-producing 
gland beyond the tip of her abdomen. Just prior to mating, considerable 
male activity (walking and flying about the cage) was noted, presum- 
ably involving behaviors similar to those described by Birch (1970). 
Copulation then quickly ensued, usually on the side of a cage (on three 
occasions under the roof), with the female the uppermost moth. After 
initial contact the hindwings of both moths were visible beneath their 
partially spread forewings, but shortly thereafter the wings were closed 
as in the resting posture, with the female’s forewings overlapping those 
of the male (Fig. 2). Pairs remained in copulation for from 2-13 hrs., a 
longer period than that reported in other noctuids (Shorey et al., 1962; 
Birch, 1970). 

Visual stimuli were apparently not required for courtship and mating, 
as successful copulations were achieved in essentially total darkness 
(basement). This agrees with prior observations on other noctuids 
(Shorey, 1964; Shorey & Gaston, 1970). 

Although females were observed in the calling posture during all hours 
of the night, matings were initiated during a relatively short period (Fig. 
3). These observations suggest that males were chiefly responsible for 
the timing of mating; and although daily cycles in male responsiveness 


VOLUME 26, NUMBER 2 99 


PAIRS 
oO 


NUMBER OF 


I VAVAVAVA 


NIGHT OF MATING 


Fig. 4. Night following pairing during which mating occurred in 25 pairs of C. 
relicta. Actual ages of the moths from night of eclosion are given within each box 
(male/female ). 


to female pheromone are known in noctuids (Shorey & Gaston, 1964, 
1965), the timing of mating is chiefly controlled in these species by a 
circadian rhythm of female pheromone release (Sower, Shorey & Gaston, 
1970, 1971). The possibility then of chiefly male timing in C. relicta 
would seem to warrant further investigation. 

The night during which mating occurred following pairing of the moths 
varied widely, and fertile eggs were obtained from matings involving 
males and females ranging from 1-16 days of age (Fig. 4). Some vari- 
ability in the occurrence of mating was expected in view of prior studies 
on reproductive age in noctuids (Shorey, McFarland & Gaston, 1968; 
Shorey, Morin & Gaston, 1968), but I am not aware of prior reports of 
delays in mating extending to two weeks or more in caged moths which 
are apparently capable of mating much earlier. 

A totally unexpected result obtained in this study was the failure of 
calling females to attract males to their cages. This occured even though 
C. relicta were being captured in the study area at bait and in light traps, 
and despite the release of scores of unmated (marked) males into the 
study area. While many factors are known to reduce the effectiveness 
of pheromone attraction (Saario, Shorey & Gaston, 1970; Sharma, Shorey 
& Gaston, 1971), none of these would seem to account for the present 
total failure. Variable numbers of caged females were observed calling 


100 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


every night over one-month periods during the summers of 1969 and 
1970, and these periods included considerable variation in environmental 
factors such as humidity, temperature, wind, and moonlight. In addition, 
the height of the cages was varied between 0 and 6 ft. above the ground, 
and on occasion fresh poplar foliage was added to the cages (in the event 
that the female pheromone plus a chemical from the foodplant was neces- 
sary to attract males—as reported for the saturniid, Antheraea polyphemus 
(Riddiford & Williams, 1967) ). 

One possible explanation for this situation would involve postulating a 
non-chemical communicating stimulus which attracts males from a dis- 
tance and which caged females are incapable of producing. Such a 
stimulus could conceivably be auditory (perhaps produced in free flight), 
and be analagous to the visual stimuli known to initiate courtship be- 
haviors in butterflies (e.g. Magnus, 1963; Brower, Brower & Cranston, 
1964). A significant role of auditory stimuli in directing the approach 
of courting moths has recently been reported in Achroia grisella (Fabr. ) 
(Pyralidae) (Dahm et al., 1971). Perhaps the males caged with the 
females in the present study were in close enough proximity to bypass 
any long-distance communication and to proceed immediately with 
chemically mediated courtship stages. Whatever the explanation, this 
matter warrants close study in the future. 


Genetics 


All C. relicta discussed in this report are descendents of a wild-caught 
female taken in Leverett in 1968 (Fig. la). This typical female produced 
90 eggs, from which 85 progeny (F,) were reared: 42 typical (20 ¢ 4, 
22 22) and 43 clara (27 64,16 22). This 1:1 phenotypic ratio sug- 
gested that a single Mendelian factor was responsible for the difference 
between the typical and clara forms, and that the cross had involved a 
homozygote x heterozygote for that factor. This suggestion was con- 
tirmed through subsequent crosses of the F,, the typical allele proving 
to be dominant (Table 2). 

The alleles were given the symbols C (dominant) and c (recessive), 
with CC and Cc representing respectively homozygous and heterozygous 
typical individuals, and cc representing homozygous clara individuals. 
Homozygous and heterozygous typicals were not phenotypically distin- 
guishable with certainty, there being considerable and rather continuous 
variation among the typical progeny of all Cc x Cc crosses. However, 
the darkest typicals were obtained in these crosses (e.g. Fig. If), aad 
these may have been homozygous individuals. The extent of variation 

mong typical and clara moths in the progeny of all F, crosses is illus- 


VoLUME 26, NUMBER 2 101 


TABLE 2. Summary of C. relicta crosses, 1969-71. 


Matings Progeny 
clara typical Totals 
Brood La slat e abalal eae TE oh ees SES 
Number Male Female Male Female clara typical 


clara x clara (cc X cc) 


08-70 25 19 = - 42, B 
09-70 8 5 = = 13 = 
11-70 5 a) = a 7 we 
02-71 10 12 = = 22. = 
Totals 84 0 
clara < typical (cc xX Cc) 
01-69 7 16 20 22, 43 42, 
01-70 8 ii 4 ih 15 et 
03-70 8 6 6 8 14 14 
05-70 9 11 10 q 20 y/ 
06-70 5 8 4. 9 1s 13 
07-70 9 1 6 6 16 1 
Ole 5 5 8 6 10 14 
04-71 6 10 9 6 16 15 
Totals : 147 138 
clara < typical (cc x CC) 
03-71 _ — 39 51 0 90 
typical x typical (Cc x Cc) 
02-70 4 2, 1) 2 6 2.4 
04-70 6 2 6 12 8 18 
10-70 5 6 13 2 11 15 
Totals 25 57 


trated in Fig. 1 (c, d and e, f)—no intermediates between the forms 
were obtained. 

All F,; crosses produced phenotypic ratios which did not differ signiti- 
cantly from those expected in one-factor crosses involving autosomal 
alleles and complete dominance (chi-square goodness-of-fit tests ). How- 
ever, the totals in Cc X Ce crosses actually fitted a 2:1 (typical : clara) 
ratio better than a 3:1 (chi-squares of 0.29 and 1.33 respectively), sug- 
gesting that the homozygous dominant genotype may have been lethal 
in some cases (perhaps in the presence of certain modifiers). Close 
examination of the data reveals a marked deficit of typical females in 
brood 10-70 (Table 2). The two typical females obtained in this brood 
were rather light, whereas five of the 13 typical males were extremely 
dark. It may be that lethality here is similar to that described in situ- 


102 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


N 58) 129" 158 Pails) = 59 18 4 


PERCENT MALES 


1 2 3 4 5 6 7 
THREE- DAY INTERVALS 


Fig. 5. Percentage of males eclosing over successive three-day intervals of the 
eclosion period for each brood (summed over 16 broods, 1969-71). The number of 
individuals eclosing is given above each point on the graph. 


ations involving the sex-limited “alba” gene in Colias butterflies (Reming- 
ton, 1954), but the present numbers are rather small, and further study 
is needed. 


Sex Ratios 


The over-all sex ratio in moths from 16 broods (Table 2) was 275 ¢ ¢ 
and 266 2 2, with only one brood (the previously noted 10-70) departing 
significantly from a 1:1 ratio (chi-square 3.84, P = 0.05 in that case). 

There was a consistent tendency for females to eclose before males 
(Fig. 5), with the sex ratio for eclosion days 1-10 summed for all broods 
being 188 6 ¢ and 211 2 2 (47% male), and for eclosion days 11-20 being 
57 64 and 55 22 (61% male). These ratios are significantly different 
(chi-square, 2 X 2 contingency test, 8.03; P < 0.01). 

In most Lepidoptera, males tend to eclose before females, and this 
certainly is true in C. fraxini (Cockayne et al., 1937-38). The significance 
of the present reversal of the usual trend is not yet apparent. 


SUMMARY 


Twenty-five matings were observed in caged pairs of C. relicta during 
‘the summers of 1969-71 in Leverett, Massachusetts. All of these matings 


VOLUME 26, NUMBER 2 103 


were initiated between 2100 and 2400 EDST, and it appeared that males 
were chiefly responsible for this timing. Successful matings were ob- 
tained with males and females ranging from 1-16 days of age. Progeny 
reared from these matings revealed that the expression of the typical and 
clara forms in the adults is largely controlled by a single gene, with the 
allele for the typical pattern being dominant. The possibility of some 
lethality among homozygous typical individuals was suggested. Over-all 
sex ratios were close to 1:1, but females tended to eclose before males. 


LITERATURE CITED 


BEUTENMULLER, W. 1903. Notes on some species of Catocala. Bull. Amer. Mus. 
Nat. Hist. 19: 505-510. 

BircH, M. 1970. Pre-courtship use of abdominal brushes by the nocturnal moth, 
Phlogophora meticulosa (1...) (Lepidoptera: Noctuidae). Anim. Behav. 18: 
310-316. 

Brower, L. P., JANE V. Z. BROWER AND FLORENCE P, CRANSTON. 1965. Courtship 
behavior of the queen butterfly, Danaus gilippus berenice (Cramer). Zoologica 
50: 1-39, 7 plates. 

Ciark, H. L. 1888. Preparatory stages of Catocala relicta Walk. Canad. Entomol. 
20: 17-20. 

CockaynE, E. A., C. N. Hawkins, F. H. LEEs, B. WHITEHOUSE AND H. B. WILLIAMS. 
1937-38. Catocala fraxini, L.: A new British record of capture and breeding. 
Entomologist 70: 241-246, 265-272; 71: 13-17, 35-38, 54-59. 

Daum, K. H., D. Mryrer, W. E. FINN, V. REINHOLD AND H. ROLLER. 1971. The 
olfactory and auditory mediated sex attraction in Achroia grisella (Fabr.). Natur- 
wissenschaften 58: 265-266. 

Forses, W. T. M. 1954. Lepidoptera of New York and Neighboring States. III. 
Noctuidae. Cornell Univ. Agri. Exp. St. Memoir 329. 

Macnus, D. B. E. 1963. Sex limited mimicry. II. Visual selection in the mate 
choice of butterflies. Proc. XVI Intern. Congr. Zool. 4: 179-183. 

Meyer, J. H. 1952. Ein neuer Catocala-hybrid. Zeit. Wiener Entomol. Gesell. 37: 
65-71, taf. 9. 

RemincTon, C. L. 1954. The genetics of Colias (Lepidoptera). Adv. Genet. 6: 
403-450. 

RippiFrorpD, L. M. anp C. M. WitiiAMs. 1967. Volatile principle from oak leaves: 
role in sex life of the polyphemus moth. Science 155: 589-590. 

Rowtey, R. R. anp L. Berry. 1910. Further study of the Catocalae. Entomol. 
News 21: 104-116. 

SaArio, C. A., H. H. SHorEy anp L. K. Gaston. 1970. Sex pheromones of noctuid 
moths. XIX. Effect of environmental and seasonal factors on captures of males 
of Trichoplusia ni in pheromone-baited traps. Ann. Entomol. Soc. Amer. 63: 
667-672. 

SHARMA, R. K., H. H. SHorry AND L. K. Gaston. 1971. Sex pheromones of noctuid 
moths. XXIV. Evaluation of pheromone traps for males of Trichoplusia ni. J. 
Econ. Entomol. 64: 361-364. 

SuHorEy, H. H. 1964. Sex pheromones of noctuid moths. II. Mating behavior of 
Trichoplusia ni (Lepidoptera: Noctuidae) with special references to the role of 
the sex pheromone. Ann. Entomol. Soc. Amer. 57: 371-377. 

, L. A. ANDRES AND R. L. HALE, JR. 1962. The biology of Trichoplusia ni 

(Lepidoptera: Noctuidae). I. Life history and behavior. Ann. Entomol. Soc. 

Amer. 55: 591-597. 


104 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


. AND L. K. Gaston. 1964. Sex pheromones of noctuid moths. III. Inhibi- 
tion of male responses to the sex pheromone in Trichoplusia ni (Lepidoptera: 
Noctuidae). Ann. Entomol. Soc. Amer. 57: 775-779. 

. AND L. K. Gaston. 1965. Sex pheromones of noctuid moths. V. Circadian 

rhythm of pheromone-responsiveness in males of Autographa californica, Heliothis 

virescens, Spodoptera exigua, and Trichoplusia ni (Lepidoptera: Noctuidae). 

Ann. Entomol. Soc. Amer. 58: 597-600. 

. AND L. K. Gastron. 1970. Sex pheromones of noctuid moths. XX. Short- 

range visual orientation by pheromone-stimulated males of Trichoplusia ni. Ann. 

Entomol. Soc. Amer. 63: 829-832. 

, S. U. McFARLAND AND L. K. Gastron. 1968. Sex pheromones of noctuid 

moths. XIII. Changes in pheromone quantity, as related to reproductive age 

and mating history, in females of seven species of Noctuidae (Lepidoptera). 

Ann. Entomol. Soc. Amer. 61: 372-376. 

, K. L. Morin ann L. K. Gastron. 1968. Sex pheromones of noctuid moths. 
XV. Timing of development of pheromone-responsiveness and other indicators 
of reproductive age in males of eight species. Ann. Entomol. Soc. Amer. 61: 
857-861. 

Sower, L. L., H. H. Shorey anp L. K. Gaston. 1970. Sex pheromones of noctuid 
moths. XXI. Light-dark cycle regulation and light inhibition of sex pheromone 
release by females of Trichoplusia ni. Ann. Entomol. Soc. Amer. 63: 1090-1092. 

1971. Sex pheromones of noctuid moths. XXV. Effects of temperature 

and photoperiod on circadian rhythms of sex pheromone release by females of 

Trichoplusia ni. Ann. Entomol. Soc. Amer. 64: 488-492. 


OBSERVED MATING BETWEEN PIERIS RAPAE AND PIERIS PROTODICE 
(PIERIDAE) 


Late in the afternoon of 5 July 1971, I explored a weed-infested lemon and avocado 
orchard in Goleta Valley (Santa Barbara County, California), looking for Nathalis iole 
(Boisduval). N. iole had once been locally common in lemon groves in rural areas 
around Santa Barbara, but apparently had not colonized this orchard. The only 
butterflies sighted that afternoon were Danaus plexippus (Linnaeus), Vanessa carye 
(Hubner), Pieris rapae (Linnaeus) and Pieris protodice (Boisduval & LeConte). The 
latter two species were flying around the wild mustard in great abundance, fluttering 
close to the ground, and often landing—obviously getting ready to settle down for the 
night. 

At approximately 1700 I noticed a copulating pair of Pieris land a few feet in front 
of me. Upon closer observation, I noticed that they were two different species, a 
male P. rapae and a female P. protodice. The female protodice was the flying partner. 
They were netted, pinched and carefully placed in an envelope. Still in copulation 
I later mounted them on a piece of cardboard and placed them in my collection. 

When two close species are found together in abundance, such interspecific matings 
are possibly not as rare as one might suspect. I have previously observed mating be- 
tween these two species in the Santa Barbara area. About a decade ago, in a field 
across from Arroyo Burro Beach State Park, I noted a pair in copulation. Considering 
that these species are very common throughout the United States, and are usually 
ignored by local lepidopterists, many such matings could go unnoticed. 

RicnArp C. Prresrar, 5631 Cielo Avenue, Goleta, California 93017. 


VoLUME 26, NUMBER 2 105 


SPECIES DIVERSITY IN CATOCALA (NOCTUIDAE) IN 
THREE LOCALITIES IN NORTH AMERICA 


D. F. OWEN 
Department of Animal Ecology, University of Lund, Sweden 


Moths of the genus Catocala (Noctuidae) are abundant in deciduous 
woodland in many parts of North America, and in some localities between 
30 and 40 species may occur together. The larvae of almost all the spe- 
cies of Catocala feed on the leaves of trees, especially trees belonging to 
the Salicaceae, Juglandaceae, Fagaceae, and Rosaceae. Each species of 
moth tends to be confined to a rather small range of tree species. Many 
species of Catocala are extremely similar, but there appears to be no evi- 
dence of natural hybrids (Sargent & Hessel, 1970). The most obvious 
difference between species is the color and pattern of the hindwing 
which, at least during daylight, is only apparent when a moth is startled 
from its resting place on a tree trunk. It would seem likely that the diver- 
sity in the pattern of the hindwing is in some way related to the predatory 
behaviour of birds, although exactly how this is achieved is not known 
(Sargent, 1969). 

Sargent & Hessel (1970) have published information showing the rela- 
tive abundance of species of Catocala at two localities in the United 
States. In 1961 I obtained a rather similar sample from a single locality 
in Michigan, and my aim in this paper is to compare the diversity of 
species in the three samples. But before discussing the estimates of diver- 
sity it is necessary to outline briefly the different methods used to obtain 
the three samples: 

Sample 1 consists of 2009 moths obtained at several localities in Frank- 
lin and Hampshire Counties in north-central Massachusetts. Specimens 
taken at bait (a mixture of brown sugar and beer painted on tree trunks ) 
comprise 84% of the sample, while 12% were taken at light, and the re- 
maining 4% were found resting on tree trunks. The specimens were ob- 
tained in the summers of 1964-69, baited and light-trapped specimens 
being obtained mostly before midnight. Very few specimens were col- 
lected after early September, a month in which many species of Catocala 
are common. Possibly some individuals were recorded more than once as 
most of the moths were released after being examined, although Sargent 
& Hessel (1970) are of the opinion, on the basis of recaptures of color- 
marked individuals, that recaptures were few. 

Sample 2 consists of 5806 specimens from a single site in Litchfield 
County, west central Connecticut, about 65 miles south-west of Amherst, 


106 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Estimates of species diversity in three samples of Catocala. 


Sample N S a and Standard Error 


1. Franklin and 
Hampshire Counties, 


Mass. 2009 oo 5.61 + 0.40 
2. Litchfield 

County, Conn. 5806 39 5.62 + 0.33 
3. Livingston 

County, Mich. 1331 30 5.45 + 0.43 


Massachusetts. All specimens were taken at lights, mostly from a single 
mercury vapor trap operated from dusk to dawn in the summers of 1961- 
65, 1967, and 1969. Trapping continued through September, and there is 
again the possibility that a few individuals may have been recorded 
more than once. 

Sample 3 consists of 1331 specimens taken at a mercury vapor light 
operated on the Edwin S. George Reserve, Livingston County, Michigan, 
during the summer of 1961. The light was operated on almost every 
suitable night throughout the season and all specimens captured were 
killed. The light was not normally operated after about 1 a.m. 

All three samples are from areas of deciduous woodland mixed with 
farmland and abandoned farmland. The essential point about the three 
samples is that number 1 differs markedly from numbers 2 and 3 by the 
method used to obtain the moths. The three localities are geographically 
separate from one another, but in view of the distribution and abundance 
of Catocala in North America it is likely that in at least some of the spe- 
cies there is gene flow between the areas. 

Numerical estimates of species diversity are possible when both the 
number of species and the number of individuals per species are known. 
In estimating species diversity in the three samples I have used the 
method proposed by Fisher, Corbet & Williams (1943), amplified in 
Williams (1964). The method involves the assumption, which can be 
tested, that the distribution underlying the number of species represented 
by different numbers of individuals is approximately a logarithmic series. 
The distribution is defined by two parameters: x, a property of sample 
size only, and a, a property of the populations sampled, which can be 
regarded as a measure of diversity. The method of estimating a and its 
standard error is given in Fisher, Corbet & Williams (1943), and rough 
estimates can be read from the nomogram given in Williams (1964). 
Observed values of N, the number of individuals, and S, the number of 
species, are required. 


VOLUME 26, NUMBER 2 107 


TABLE 2. The three commonest species of Catocala in three different localities. 


Locality Species % of Total Sample 
il ilia 34 
ultronia 16 
crataegi* 5 
2) . palaeogama 155 
residua 15 
habilis 11 
3 amica 31 
epione 12 
concumbens 6 


* This includes records of Catocala blandula and Catocala mira. 


The results for each of the three samples are given in Table 1. The 
three computed values of a are similar and the standard errors are small. 
There is no significant difference between them, and the result is thus in 
accordance with the view that species diversity within a group of orga- 
nisms is an intrinsic property of the environment. The three samples 
differ in the way in which they were obtained (the difference being 
greatest between sample 1 and samples 2 and 3) but this does not appear 
to have affected the value of a. Many of the species taken are common 
to all three samples, but the relative frequency of most of the species in 
each locality is quite different. All three samples are characterised by a 
small number of abundant species and many relatively rare species, but 
a rare species in one sample is sometimes among the commonest in an- 
other. Table 2 shows the three most abundant species in each of the three 
samples. With the possible exception of Catocala crataegi,! which was 
not positively identified in sample 3, each of the species shown in Table 
2 occurred at all three localities, but in each case the three most abun- 
dant species are different in each of the three samples. These differences 
are presumably associated with local ecological conditions, possibly with 
the relative availability of larval foodplants in the three localities, but the 
important point is that in each locality only three species comprise 55%, 
41%, and 49%, respectively, of the total samples. This is the familiar 
result obtained whenever a group of species is sampled. But despite the 
variations in abundance of the species in the three localities, almost iden- 
tical values of species diversity are obtained. This suggests that species 
of Catocala are partitioned out by the local environment in such a way 
that a constant diversity is maintained. It would be of considerable 
interest to know exactly how this is achieved. 


1 Catocala mira, which was tentatively identified in sample 3, is easily confused with Catocala 
crataegi, and it is possible that what I have called mira is in reality crataegi. 


108 JouRNAL oF THE LEPIDOPTERISTS SOCIETY 


SUMMARY 


Samples of moths of the genus Catocala from three localities in the 
United States have values of species diversity that are not significantly 
different. It is suggested that this result supports the view that diversity 
is an intrinsic property of the environment and that although the indi- 
vidual species differ in their relative abundance in different places, an 
essentially identical pattern of diversity occurs in three widely separated 
yet ecologically similar localities. 


LITERATURE CITED 


Fisuer, R. A., A. S. Corser anp C. B. Wittiams. 1943. The relation between the 
number of species and the number of individuals in a random sample of an 
animal population. J. Anim. Ecol. 12: 42-58. 

SARGENT, T. D. 1969. A suggestion regarding hindwing diversity among moths of 
the genus Catocala (Noctuidae). J. Lepid. Soc. 23: 261-264. 

SARGENT, T. D. anv S. A. Hesset. 1970. Studies of the Catocala (Noctuidae) of 
southern New England. 1. Abundance and seasonal occurrence of the species, 
1961-1969. J. Lepid. Soc. 24: 105-117. 

WituraMs, C. B. 1964. Patterns in the Balance of Nature. Academic Press, London 
& New York. 324 p. 


McFARLAND MOTH COLLECTION DONATED TO THE NATURAL 
HISTORY MUSEUM OF LOS ANGELES COUNTY 


Noel McFarland has been one of the more prominent students of North American 
moths for many years. When he moved to Australia he donated his splendid collec- 
tion of 8,420 specimens to the Natural History Museum of Los Angeles County. 

By far the most significant portion of the collection is 3,315 beautifully preserved 
larvae, in alcohol, and the accompanying copious notes on foodplants, behavior, and 
larval descriptions. The larvae of many species, such as Saturnia albofasciata, are 
represented in few, if any, other collections. 

In addition to the larvae, there are 5,105 adult moths, many of which bear distinc- 
tive labels to associate them with rearing notes and preserved larvae. Included are 
most of the 278 species of moths taken during extensive collecting in Los Angeles 
County that resulted in “The moths (Macroheterocera) of a chaparral plant associa- 
tion in the Santa Monica Mountains of Southern California,’ (1965, J. Res. Lepid. 
4: 43-74). In addition, there are a number of specimens from Kansas, and numerous 
moths from the edge of the Mojave Desert adjacent to the San Gabriel Mountains. 

Earlier, in 1963, McFarland donated to the Museum 860 specimens of moths from 
Benton County, Oregon. These represent a large portion of the 360 species that he 
collected during a 20-month study that was the basis of his unpublished 1963 Master 
of Science thesis at Oregon State University, “The Macroheterocera (Lepidoptera) of 
a Mixed Forest in Western Oregon.” 

These generous donations are a valuable addition to the Museum’s extensive hold- 
ings of western Lepidoptera, while the larvae will form a strong nucleus for the 
collection of immatures. 


JULIAN P, DONAHUE, Assistant Curator of Entomology, Natural History Museum of 


} 


ingeles County, 900 Exposition Boulevard, Los Angeles, California 90007. 


VoLUME 26, NUMBER 2 109 


POLYMORPHISM IN PAPILIO GLAUCUS L. (PAPILIONIDAE): 
MAINTENANCE OF THE FEMALE ANCESTRAL FORM 


SALLY K. MAKIELSKI 


Department of Biological Sciences, Loyola University, 
New Orleans, Louisiana 70118 


The Papilio glaucus L. female is dimorphic. One form is dark and 
resembles P. (Battus) philenor L.; the other resembles the yellow male. 
The inheritance of female color is controlled by a Y-linked gene (Clarke 
& Sheppard, 1962). Yellow mothers produce yellow daughters and dark 
mothers produce dark daughters. Consequently the two morphs are in 
direct competition with each other; and, heterozygote advantage cannot 
be the mechanism which maintains the dimorphism. 

The dark form of P. glaucus is thought to belong to the mimetic assem- 
blage which surrounds P. philenor. Other members are P. troilus L., P. 
polyxenes (Fabricius), Limenitis arthemis astyanax (Fabricius), and the 
females of Speyeria diana (Cramer). The main evidence that the dark 
form of glaucus mimics P. philenor is the correspondence between the 
occurrence and abundance of the two. In general (Brower & Brower, 
1962) the proportion of mimics is highest in the middle Atlantic States 
where philenor is common. This proportion decreases in a southerly and 
northerly direction as philenor decreases. 

One hypothesis for explaining the maintenance of the dimorphic fe- 
male population in P. glaucus is that the males show a mating preference 
for the yellow females. Burns (1966) has tested this hypothesis by count- 
ing the number of spermatophores present in the bursa copulatrix of the 
two morphs. He obtained data from two samples of females collected 
during the summer of 1965; one in the vicinity of Mountain Lake Biologi- 
cal Station, Giles County, Virginia, and the other in Baltimore County, 
Maryland. In each sample the mean number of spermatophores per fe- 
male was higher in the yellow morph than in the dark morph (Mt. Lake, 
yellow morph-2.08 spermatophores, dark morph-1.69 spermatophores; 
Baltimore County, yellow morph-1.88 spermatophores, dark morphs-1.54 
spermatophores ). 

The present study was suggested by the relatively small difference be- 
tween the mean number of spermatophores per female (Mt. Lake sam- 
ple, 0.39; Baltimore sample, 0.34) found by Burns. It seemed that further 
information about the insemination frequency in P. glaucus would be of 
interest. 

Two hundred P. glaucus females were collected during August and 


110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Observed frequency distribution, expected frequency distribution (num- 
bers in parentheses), and mean number of spermatophores in wild-caught females of 
Papilio glaucus. 


Number of Spermatophores Mean Number of 


Specimens.» 2 a ee Spermatophores/ 
Morph (No. ) 0 1 2) 3 Female 
Mimic 128 2 65 51 10 1.54 
(2756) (65292) an (aS 4) (ne OS) 
Yellow 2, Dh 38 30 DY 1.44 
(1.44) (37.08) (29.16) (4.32) 
Total 200. 4 103 81 2, 1.51 


September of 1966, 1967 and 1968 from two valleys in Albemarle County, 
Virginia; one in the Blue Ridge Mountains (elevation 800’), the other 
approximately 24 miles east of the Blue Ridge (elevation 590’). The 
butterflies were found in abandoned fields in which thistle was growing. 
Streams and larval foodplant (Liriodendron and Prunus) were located 
nearby. 

The females were either dissected immediately after they were caught 
or were frozen alive in a moist chamber and dissected later. 

A comparison of the mean number of spermatophores per female (Ta- 
ble 1) shows that the yellow females (x = 1.44) were inseminated less 
frequently than the mimic females (x = 1.54). However, this difference 
is not significant (x(2)? = 2.07; .50 > P > .30; categories 0 and 1 sper- 
matophores combined). Statistical analysis also shows that there was no 
heterogeneity with respect to locality (x(1)2 = .34; .80 > P > .70). 

The results, then, show that there is no difference in the frequency of 
insemination between the mimetic and yellow females in the population 
studied. These results conflict with those of Burns (1966) and with some 
but not all of the data collected by Levine (1970) from females found in 
the vicinity of Mountain Lake, Virginia. 

The reason for the discrepancy between the two sets of data is not 
clear. It may be related to the altitude at which the butterflies are found, 
the relative frequency of the model in relation to the frequency of P. 
glaucus, or the relative frequencies of the two female morphs. The ratio 
of dark to yellow morphs is 6:1 in the Mountain Lake area and 2:1 in 
Albemarle County, while the yellow butterflies are preferentially insemi- 
nated at Mountain Lake and are inseminated equally as frequently as the 
mimics in Albemarle County. Thus, it is also possible that, as in the case 
of Drosophila pseudoobscura (Ehrman, 1967; Spiess, 1968), female mat- 

ng advantage occurs in P. glaucus. That is, that where the morph is rare 


VoLUME 26, NUMBER 2 JG 


it is inseminated more frequently. This hypothesis can be tested by ob- 
taining data on the relative frequency of insemination of mimic females 
in populations where the proportion of the mimics is small. 

While the mechanism which maintains the dimorphism in P. glaucus 
is not yet clearly understood, the evidence collected to date shows that 
regional differences exist in the frequency with which the two female 
morphs are inseminated. 


ACKNOWLEDGMENT 


This research was done while working in the laboratory of Dr. J. J. 
Murray, Jr., in the Department of Biology, University of Virginia, Char- 
lottesville, Virginia. My thanks to Dr. Murray for his encouragement and 
for the use of his laboratory facilities. 


LITERATURE CITED 


Brower, L. P. Anp J. VZ. Brower. 1962. The relative abundance of model and 
mimic butterflies in natural populations of the Battus philenor mimicry complex. 
Ecology 43: 154-158. 

Burns, J. M. 1966. Preferential mating versus mimicry: disruptive selection and 
sex-limited dimorphism in Papilio glaucus. Science 153: 551-553. 

CLARKE, C. A. AND P. M. SHEeppArD. 1962. The genetics of the mimetic butterfly 
Papilio glaucus. Ecology 43: 159-161. 

EuHRMAN, L. 1967. Further studies on genotype frequency and mating success in 
Drosophila. Amer. Natur. 101: 415-424. 

Levin, M. P. 1970. The biology of Papilio glaucus L. as it relates to polymorphism 
and mimicry. Doctoral dissertation, Virginia Polytechnic Institute, Blacksburg, 
Virginia. 

Spiess, E. B. 1968. Low frequency advantage in mating of Drosophila pseudo- 
obscura karyotypes. Amer. Natur. 102: 363-379. 


PSEUDOPHILOTES BEURET, 1958 


In a recent article of mine about Glaucopsychie piasus I called attention to the 
fact that Philotes sonorensis, the type species of Scudder’s genus, is generically dif- 
ferent from the rest of the genus found in North America. This requires a different 
name for enoptes and its allies. Such a name has been proposed. It is Pseudophilotes 
Beuret, 1958, with European baton as its type species. Baton is cogeneric with 
enoptes, et al. 

The probable reasons for this generic name being overlooked by American taxono- 
mists are two: the Zoological Record citation made no mention of nearctic members 
in Pseudophilotes; the paper in which the name was proposed is in a journal rarely 
seen in North America. The full citation for the description of Pseudophilotes is: 
Beuret, H., 1958, “Zur systematischen Stellung einiger wenig bekannter Glaucopsychidi 
(Lep., Lycaenidae)” Mitt. ent. Ges. Basel (N.F.) 8: 61-79, 1 pl., 12 figs.; 8: 81-100, 
13 pls. The original description begins on page 100. 

I wish to thank Dr. Lionel G. Higgins for calling this publication to my attention. 

F. MARTIN Brown, Fountain Valley Rural Sta., Colorado Springs, Co., 80911. 


112 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


JUNIPERUS (CUPRESSACEAE) SPECIATION AND THE 
RANGES AND EVOLUTION OF TWO CALLOPHRYS 
(LYCAENIDAE) 


KuRT JOHNSON 
Museum of Natural History, University of Wisconsin, Stevens Point 54481 


It seems popularly assumed that the allopatric distributions of Callo- 
phrys (Mitoura) siva Edwards and C. (M.) gryneus Huebner are deter- 
mined by the ranges of the several “species” of Juniperus which are the 
larval food-plants. Klots (1951) reports C. (M.) gryneus as feeding on 
Juniperus virginiana L. while Brown (1957, and in correspondence) re- 
ports C. (M.) siva utilizing several western junipers like J. scopulorum 
Sarg., J. utahensis Engelm., and J. occidentalis Hock. There is no other 
discussion in the literature known to this author which indicates opposing 
evidence to this popular assumption. Remington & Pease (1955) seem to 
sense the confusion within Juniperus taxonomy and report C. (M.) gryneus 
as feeding simply on “Juniperus sp.” They also establish the usability of 
Swamp White Cedar [Chamaecyparis thyoides (L.)] for C. (M.) gryneus. 

Until recently there has been no definitive work on the speciation of 
Juniperus in the areas of the United States including the distributions of 
C. (M.) siva and C. (M.) gryneus. Van Haverbeke (1968) has produced 
a detailed computer analysis of morphological and lipid characters (the 
latter analyzed by infrared spectroscopy ) of Juniperus spp. in which he 
develops indices of hybridization between J. scopulorum and J. virginiana 
in the Missouri River Basin. The important problem is whether the bi- 
hybrid swarm of Juniperus he describes is best characterized as two spe- 
cies or simply variation within one clinal situation obscured through years 
of evolution. In his conclusions, through the analysis of statistical dis- 
tributions, he presents several evaluations which strongly support the 
presence of two species (J. scopulorum and J. virginiana), Although no 
extreme parental types were discovered east or west in his study areas, 
he favors the conclusion that J. virginiana and J. scopulorum possess in- 
tegrity enough to warrant his interpretation of data as percentage of each 
of these species’ germ plasm in each study area. The data and conclusions 
within his study are important in making comments on factors determin- 
ing the distribution of the two species of Callophrys considered in this 
paper. 

in recent years C, (M.) siva has been reported on the Great Plains in a 
number of populations. Johnson (1972) reports C. (M.) siva in eight 
Nebraska counties, a few east of the 100th meridian. John S. Nordin (in 


VoLUME 26, NUMBER 2 LES 


-.....____ MISSOURI RIVER BASIN 
é SSS ee 
SR \---- ">>, z-BASIN BOUNDARY 
” ! \ 
as leg 67+6 esa ae 
pee Os G94 4 pe OSE ees, 
pe act alg “7043 *; 
67+6 + ‘ 
i] : ‘) 
cae. 59+5 opel i 
wy 61+5 = 68+5 65+ 6 4 . = 
GSS Rong el a 
‘ 1+10 
‘ z pales 20 Oe 5944 
lys svn ee 67+4) 78 +8 SO66 
. BaMO SOaer 52+6 > 
| ‘. 
ete 7Ot4-@@ (7844 3740 
Mo Be e 38+4 wee 
% 6643 e 52+4 0+4 + pS 
| ‘eaika | gato “get 0 
ita ce. Oe LS ! 
! 63+4 | @ 4349 ABs 
‘ etek, 63+8 = 36+4 38+7 : 
N oS 5515 e a 
e ‘ 
N 55+6 42+7 38+2 he 
fs \  61+6 oo S 
iimiig ig ee ! 48+6 us See aan 
F ae Y 4ata ae) iat Re Sit 
| San 2 4444 34+4° ¢ 
I y ! pe \ 
60+8 se pe a SS I ; 
ina 7) OO 38+0 t 
BC 57+ ce 
! : Pau aes ! 38+5 27+4 3444 
Soe aS aee -~ 38+4 
! N ‘ N aL 
ppc ad oe 2 eee 1 294+4 
I ee a i 
Sai 
SCALE SLA : 
560 0 50 100 150 i SQL een 
Cee eee MILES 


30+4 where: 30 = Percent J.scopulorum germ plasm 
+4 = Range in percent 
(x J.scopulorum x virginiana 30 %) 


Fig. 1. Locations of known Callophrys (Mitoura) siva populations on the Great 
Plains superimposed above the plus-or-niinus sign of a Van Haverbeke computation. 
Note that one population (70 + 4%) is located by an arrow and that the single 
Wyoming population corresponds to no Van Haverbeke computation. In the figure a 
computation such as 30 + 4 means 30% plus or minus 4% J. scopulorum “germ 


plasm.” 


Counties in which C. (M.) siva populations occur, with the general Van Haverbeke percentage 
given to facilitate more rapid association, are listed below. Two counties named together indicate 
juniper areas running across county boundaries; county locations of particular areas mentioned in 


the text are noted. 
North Dakota: Billings County (63%); South Dakota: Pennington County (Black Hills area) 


(70%), Meade County (52%), Custer County (63%); Wyoming: Weston County (none); Ne- 
braska: Dawes County (63%), Cherry County (43%), Brown and Rock Counties (Long Pine 
Recreation Area) (36%), Banner and Scotts Bluff Counties (55%), Thomas County (Nebraska 


National Forest at Halsey) (42%), Keith County (48%). 

correspondence) reports the species at several locations in western and 
central South Dakota, and he and F. Martin Brown (in correspondence ) 
report specimens from North Dakota. All of these populations occur ( ex- 
cept for the Black Hills of South Dakota) in scattered escarpments or in 
areas of Juniperus on bluffs and buttes on the plains. The geographical 
locations of these populations can be seen in Fig. 1. 


114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1 superimposes the location of known populations of C. (M.) siva 
upon Van Haverbeke’s figure of computer percentages of hybridization 
based on the sum of all his analyses of characters. The dots representing 
populations of C. (M.) siva are located directly above the plus-or-minus 
sign of the corresponding Van Haverbeke computation except where 
located by an arrow (see explanation of figure). 

A great amount of variation in the type of juniper being utilized by C. 
(M.) siva is apparent, varying from 70 + 4% J. scopulorum in the Black 
Hills of South Dakota to 36 + 4% J. scopulorum in the Long Pine area of 
north-central Nebraska. Though this information is interesting, it gains 
its greatest import when compared to hybrid indices for Juniperus popu- 
lations occurring into the range of C. (M.) gryneus. These indices are 
given elsewhere in Van Haverbeke’s paper. Indices for Juniperus spp. in 
northeast and central Missouri, within the range of C. (M.) gryneus (the 
species is also reported in Nebraska by Klots, 1951, but was not validated 
by Johnson, 1972) range up to 38 + 4% J. scopulorum, and in areas of the 
eastern United States upwards to 39% J. scopulorum. The latter, how- 
ever, requires caution due to the small amount of sampling in extreme 
eastern areas. 

In the intervening area between the allopatric ranges of C. (M.) siva 
and C. (M.) gryneus there is considerable overlapping in the identity of 
the type of Juniperus available for food-plant use. The question emerges 
whether there is a threshold at which C. (M.) siva and C. (M.) gryneus 
segregate in their use of food-plants. Such segregation could be in reac- 
tion to morphological characters or chemistry. If segregation exists there 
must be some type of selection of juniper in the areas where the integri- 
ties of juniper used by the two species overlap. Such a selection might 
not be made by the female imago but might simply be due to larval mor- 
tality, or sterility of resulting adults, when ova are laid on unacceptable 
plants. However, it is equally possible that there may be no segregation 
due to food-plant type between these two butterfly species and that some 
other factor is keeping them allopatric in distribution. This indicates the 
possibility that the two species may eventually become sympatric. 

The populations of C. (M.) siva in north-central Nebraska are newly 
discovered, and therefore their length of establishment is not known. 
‘lowever, the species was never reported in these areas in the early litera- 
ture on Nebraska Rhopalocera. C. (M.) siva could have reached eastern 
outposts like the Nebraska National Forest at Halsey and the Long Pine 
“ecreation Area in Nebraska by import of Juniperus seedlings planted in 
those areas by the Forest Service in fairly recent years. This opens the 


VoLUME 26, NUMBER 2 115) 


possibility of germ plasm “pollution” of the Juniperus populations by the 
introduction of western imports. 

There are two possibilities emerging from these data: the sympatry of 
C. (M.) siva and C. (M.) gryneus and its ramifications, or the segrega- 
tion of the two (though the types of juniper available for utilization, 
according to Van Haverbeke’s evaluation, seem to overlap). It would be 
desirable to experiment with the food-plant preferences of C. (M.) siva 
and C. (M.) gryneus in the laboratory. It is, however, the purpose of this 
paper to point out that the new evidence of Juniperus hybridization pre- 
sented by Van Haverbeke has important bearing on lepidopterists’ evalua- 
tion of the distribution of species of butterflies feeding on that taxon of 
plants. If C. (M.) siva and C. (M.) gryneus have evolved into two dis- 
tinct species through isolation of their gene pools, it seems that some bar- 
rier to their sympatry must have been imposed at some time. If J. 
scopulorum and J. virginiana once had extreme parental type distribution 
areas, which are now introgressing spatially east and west, there might 
be some evidence of a former barrier. Within these two former distribu- 
tion areas of Juniperus parental types might have been the harbors of the 
populations which evolved what we call C. (M.) siva and C. (M.) gryneus. 
Perhaps fluctuations in distributions and barriers in the history of Junip- 
erus left the Callophrys ranges as “relicts’ of former ranges of more 
typical juniper parental types. If, as Van Haverbeke suggests, J. vir- 
giniana might be an eastward evolutionary manifestation of J. scopulorum, 
there is a vague possibility that the original gene pool of C. (M.) gryneus 
utilized Swamp White Cedar and later adapted to Juniperus spp. as a 
food-plant. 

Certainly the current taxonomic status of Juniperus species in the cen- 
tral United States has great import on the considerations of reasons for 
the respective distributions of C. (M.) siva and C. (M.) gryneus, and 
reminds lepidopterists to be more careful in forming generalizations 
about insect distributions because of their food-plants until more fully 
aware of the complexities in the plant groups themselves. 


ACKNOWLEDGMENTS 


Dr. John S. Nordin (Webster, South Dakota), Dr. F. Martin Brown 
(Colorado Springs, Colorado), and Mr. Glen Viehmeyer (North Platte 
Station, University of Nebraska) provided additional data for the study. 
Dr. John C. Downey (University of Northern Iowa, Cedar Falls) and 
Miss Doris Gates (Chadron State College, Chadron, Nebraska) made 
helpful suggestions concerning the project, and Mr. Leonard Running 
(Custer, South Dakota) accompanied the author on collecting trips. Dr. 


116 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


David F. Van Haverbeke (University of Nebraska, Lincoln) generously 
allowed complete access to his Juniperus study including the reproduc- 
tion of one figure. 


LITERATURE CITED 


Brown, F. M., D. Err anp B. Rorcer. 1957. Colorado Butterflies. Denver Mus. 
Natur. Hist., Denver. 368 p. 

Jounson, K. 1972. The butterflies of Nebraska. Lepid. Res. Found. (in press). 

Kiors, A. B. 1951. <A Field Guide to the Butterflies of North America, east of the 
Great Plains. Houghton Mifflin Co., Boston, xvi + 349 p. 

REMINGTON, C. L. AND R. W. Pease, JR. 1955. Studies in food-plant specificity, 1. 
The suitability of Swamp White Cedar for Mitoura gryneus (Lycaenidae). 
Lepid. News 9: 4-6. 

Van HAVERBEKE, D. F. 1968. A population analysis of Juniperus in the Missouri 
River Basin. Univ. Nebr. Studies, No. 38. 82 p. 


ACCIDENTAL OCCURRENCE OF AGLAIS URTICAE 
(NYMPHALIDAE) IN NOVA SCOTIA 


On November 7 1970, John Godfrey of the faculty of Dalhousie University in 
Halifax, unpacked a crate of books shipped by sea from England. A butterfly flew 
out of the crate and was captured and brought alive in a plastic box to FWS by 
George Halverson, a neighbor of Mr. Godfrey’s. The specimen, a fresh male of Aglais 
urticae (Linnaeus), could not be killed until the following day, by which time the 
wings were worn and chipped. 

According to Mr. Godfrey the crate was packed and closed in Oxford, England on 
August 23 1970, and left England by sea in mid-October. The time elapsed between 
Bee and unpacking was 76 days. Mr. Godfrey did not notice a pupal skin in 
the crate. 

The interesting aspect of this occurrence is our knowledge of how the butterfly 
arrived in Nova Scotia. Had it escaped and then been collected outdoors in Halifax, 
its presence would have been quite baffling. 

We know of no other North American records of this species. The specimen has 
been deposited in the Lepidoptera collection at the Nova Scotia Museum. 

FREDERICK W. Scortr AND BArry Wricut, Nova Scotia Museum, 1747 Summer 
Street, Halifax, Nova Scotia. 


BUTTERFLIES FEEDING ON A DEAD BOBCAT 


On 6 July 1970, Mr. S. K. Dvorak and I captured the following butterflies, all 
males, imbibing the juices of a dead, decaying, young bobcat along a roadside: 
Cercyonis oetus (Boisduval) (1), Speyeria zerene conchyliatus (J. A. Comstock) (3), 
Speyeria callippe near nevadensis (Edwards) (1), and Euphydryas chalcedona (Dou- 
bleday), ssp. (1). The location was ca. 2 road mi. NE of SW entrance to Lava Beds 
National Monument (road from McCloud), Modoc National Forest, Siskiyou Co., 
Calif. Instances of butterflies at carrion are given by Payne & King (1969, J. Lepid. 
Soc. 23: 191-195). 

OAKLEY SuHiELDs, Department of Entomology, University of California, Davis, 
California 95616. 


VoLUME 26, NUMBER 2 1a 


MALE GENITALIA OF LEPIDOPTERA: MORPHOLOGY AND 
NOMENCLATURE IV. NOTES ON TUXEN’S “TAXONOMIST’S 
GLOSSARY OF GENITALIA IN INSECTS”: SECOND 
ENLARGED EDITION 
A. SIBATANI 
30 Owen St., Lindfield, N.S.W., 2070, Australia 


I have found that Parts II and III of this series (Okagaki et al., 1955; 
Ogata et al., 1957) are not referred to in the recently published Second 
Enlarged Edition of “Taxonomist’s Glossary of Genitalia in Insects” edited 
by S. L. Tuxen (1970). I have also found that the nomenclature which 
we proposed in one of those papers was not included in the very extensive 
and useful Glossary but was nevertheless employed to define another 
term in the same Glossary. The situation is thus a source of further con- 
fusion. I therefore wish to rectify this point in the present paper. Also 
Alexander B. Klots (1970) in his text of Lepidoptera (p. 124) of this new 
edition commented on the nomenclature for the subdivision of the valva, 
and referred to Part I of this series (Sibatani et al., 1954). I have a dif- 
ferent opinion on his points, and judge that a note on his text may be 
useful. 


The Scaphium and “Sociuncus” 


Ogata et al. (1957) proposed that the dorsal appendages belonging to 
the 10th abdominal segment of male Lepidoptera, which may be sub- 
divided into the median uncus (sometimes divided medially) and lateral 
socii should be called collectively the scaphium according to the original 
terminology of Gosse (1882) in the absence of any other suitable name. 
We made this proposal because we considered the socii and uncus as two 
similarly weighted and not mutually exclusive derivatives of a morpho- 
logical unit structure, and also because sometimes it is difficult to assign 
portions of this unit to either uncus or socii. The term scaphium sensu 
Ogata et al. has since been extensively used by Japanese authors (Shirdzu 
and Yamamoto, 1959; Shirézu, 1960; Inoué and Kawazoé, 1964, 1965; 
Kawazoe and Wakabayashi, 1969), but it is not referred to in this sense 
in Tuxen’s Glossary, let alone mentioned in the text. However, the Glos- 
sary lists on p. 295: 

“Okonze (russian) ¢ Lep. Kusnezov 1924 (!) teste Shirdzu and Yamamoto 1956 
in Lycaenidae. Lateral membranous area between tegumen and scaphium” (Italic and 
iby AS. ). 

Shirézu and Yamamoto (1956) were using the nomenclature outlined 
by Ogata (1950) in a paper which briefly summarized the conclusion as 


118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


published later in this series, and in which the scaphium was defined 
in the sense of Ogata et al. (1957), and not of Pierce (1909) or of most 
other authors. The term okonze (to be spelled “okontse” according to 
the currently employed transliteration of Cyrillic) was also mentioned 
in that paper by Ogata (1950). 

Since, however, Tuxen’s book has fixed the meaning of scaphium in 
Pierce’s sense (p. 326), the way to minimise the confusion is to drop the 
use of scaphium as proposed by Ogata et al. (1957), and I so advise 
those who have so far followed the latter. This does not, however, alevi- 
ate the need of a collective term for the morphological unit covering the 
uncus and socii, as the recent extensive use by Japanese authors of the 
term scaphium in this sense testifies. I therefore suggest a new term 
sociuncus to replace the scaphium in the sense of Ogata et al. (1957). 
This is a synthesis of the words socii and uncus, and I hope that it is self- 
explanatory. It may be argued that the need for such a term would be 
readily dissolved by an expression like “uncus-socii complex.” However, 
contrary to what such an expression would seem to imply, the sociuncus 
is a structure of primary morphological significance, and the uncus and 
socii are its secondary derivatives and not vice versa. In this sense, the 
situation is not comparable with that of frequently used expressions like 
“harpe + ampulla region” (Shir6zu and Yamamoto, 1956) or “harpe- 
ampulla complex” (Inoué and Kawazoé, 1965) of the valva, because in 
the latter case, the harpe and ampulla are of primary significance, which 
may secondarily become united. 

There are several other names proposed in Part III of this series which 
[ think Tuxen’s Glossary should have included, and therefore I attach a 
“Proposed Addenda” to it at the end of this paper. They include two 
unlisted names (cochlear and fenestrula) which have been extensively 
used by Japanese authors, as well as certain other terms which are now 
involved in confusion for which I am mainly responsible. 


Subdivision of the Valva 


The second point I would like to discuss briefly concerns Klots’s state- 
ment (p. 124) in Tuxen’s book: 


“... these findings [recognition of “six” (actually seven) fundamental regions by 


Sibatani et al. (1954) in the valva—A.S.] do not agree in part with the very im- 
portant conclusions of Forbes (1939) who studied the valval musculature, especially 
arding the so-called “clasper,’ which in some groups has a separate musculature 
which would seem to infer a separate origin. Still more disparate are the conclusions 
of Birket-Smith (1965) about many structures, particularly of the valvae, based on 
\dies of the structure and their musculature in Lithosiinae. This author introduces 


yely new nomenclature, which deserves very careful consideration.” 


VoLUME 26, NUMBER 2 119 


It is hard to understand the first half of this statement. The subdivision 
of the valva as proposed by Sibatani et al. (1954) was based on an exten- 
sive study of the musculature, and as far as the more specialised families 
of Lepidoptera are concerned, it supported the findings, if not the inter- 
pretation, of Forbes (1939). Especially difficult to follow is Klots’s com- 
ment on the musculature of “clasper” (= harpe), because both in Forbes 
(1939) and Sibatani et al. (1954) the “clasper” or harpe was virtually 
defined by its insertion at the base of Muscle 5 (Flexor of harpe), which 
arises from the base of the valva (usually sacculus). 

It is true that Forbes mentioned some anomaly of Muscle 5, inserting 
mostly into juxta in place of the sacculus in an unidentified and unillus- 
trated species of Geometridae, noting that the “clasper” is missing in this 
instance. I suspect that here he was dealing with a form of Boarmiinae 
having the furca (Okagaki et al., 1955), in which the sacculus is modi- 
fied into a juxta-like structure (= furca) and Muscle 5 inserted in the 
harpe has a reversed orientation of the motion transfer as Forbes himself 
noted. Therefore, I do not see any discrepancy between Forbes (1939) 
and Sibatani et al. (1954). 

In more primitive groups of Lepidoptera (Hepialidae, Incurvariidae, 
Tortricidae) Muscle 5 is missing. Instead, Muscle 6 (Protractor of 
aedoeagus ) may insert in the base (Tortricinae) or the tip (Incurvariidae 
and Olethreutinae ) of the valva. The origin (or terminus) of this muscle 
attached to the cephal end of the aedoeagus is very variable even among 
less primitive families of Lepidoptera, ranging from tegumen or vinculum 
(the typical case) to various parts of the valva and/or juxta. It is there- 
fore conceivable that Muscle 6 may occasionally be mistaken for Muscle 
5 or its derivative. 

It is also important to notice that the harpe (and its Muscle 5) appears 
predominantly in more specialised groups and not in primitive forms, so 
that its derivation must be only secondary. This concept would be useful 
for determining the homology of the valva with more general structures. 
I shall discuss this subject more fully in a separate paper of this series. 

Birket-Smith (1965) divided the valva of Lithosiinae into the following 
portions: basis valvae (including processus momenti and supravalva), 
ala valvae, plica centripetalis and valvella. A careful examination of his 
text and figures clearly indicates that the musculature in this group is 
essentially the same as observed by Forbes (1939) and myself (Sibatani, 
in prep. ). Only in minor special groups unusual sclerites like those called 
the valvella with its unusual muscles seem to appear. However, such 
cases can be recurrently observed in widely scattered groups of 
Lepidoptera. 


120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The division of the main body of the valva into basis valvae and ala 
valvae seems to have been worked out from the particular pattern of 
sclerotisation of the valva in Lithosiinae, but this should have a more 
functional rather than fundamentally morphological significance. Thus 
the following assignment of synonymy is easily made. Assignment similar 
to that used in the Glossary of Tuxen (1970) is marked with an asterisk. 

Ala valvae = Sacculus + harpe + their corresponding wall on the outer 
surface of valval sclerite. 

Basis valvae (s.str.) = Basal portion of the outer wall of the valva. 

Plica centripetalis = Ampulla. 

Processus momenti = Unconnected transtilla, acting as apodeme for 
muscular attachment.* 

Supravalva = Cucullus. 

Valvella = A structure peculiar to the group studied. 

Therefore, Birket-Smith’s new system of nomenclature does not enforce 
a drastic revision of the nomenclature for the division of the valva ap- 
plicable to more specialised groups. Since the subdivision proposed by 
Sibatani et al. (1954) was concerned solely with the structures appearing 
on the mesal surface of the valva, and if the subdivision of the outer 
surface is called for, Birket-Smith’s terms of basis valvae and ala valvae 
might prove useful. However, the variation among different families 
is so enormous that I doubt that a generally workable system can be 
derived from his terminology. 


Proposed Addenda to Glossary 


Only those terms which the author and his former collaborators are 
responsible for naming and interpreting are listed. 

Cochlear (-is, ia) é Lep. Ogata 1950, Ogata et al. 1957. Median process 
of gnathos. > Brachia. 

a (-ae,-ae) 3 Lep. Ogata et al. 1957. Dorsal median hyaline 
part of conjunctival membrane between sociuncus and tegumen. > 
Lateral fenestrula. 

Syn. Okontse (= Okonze) Kuznetsov (= Kusnezov ) 1916. 
— 6 Lep. Inoué and Kawazoé 1964 in Hesperiidae. Entire sclerotised or 
unsclerotised conjunctival membrane between tegumen and sociuncus. 
> Okonze Ogata 1950 nec Kuznetsov (= Kusnezov ) 1916. 

Furc/a (-ae,-ae) & Lep. Okagaki et al. 1955, Pierce 1914 in Geometridae 
(Ourapteryx, Plagodis, Cepphis, Epione etc. nec Ennomos ). Sacculus 
becoming an independently movable process between juxta and harpe- 
valvula area of valva; sometimes asymmetric and even unilateral. 


Fenestrul 


' 


VoLUME 26, NuMBER 2 PALI 


Okontse (russian) ¢ Lep. Kuznetsov (=Kusnezov) 1916. = Fenestrula 
Ogata et al., Okonze Ogata 1950 (part.). 

Okonze (russian) é Lep. Kusnezov 1916 = Okontse Kuznetsov 1916. — 
6 Lep. Ogata 1950. Entire conjunctival membrane between sociuncus 
and tegumen. — Fenestrula, Lateral fenestrula. 

Lateral fenestrula ¢ Lep. Inoué and Kawazoé 1965 in Riodinidae and 
Lycaenidae. Lateral hyaline parts of the conjunctival membrane be- 
tween sociuncus and tegumen at the base of brachia. > Fenestrula. 
Syn. Lateral window Shirdzu and Yamamoto 1956. 

Lateral window ¢ Lep. Shirézu and Yamamoto 1956 = Lateral fenestrula. 

Scaphium ¢ Lep. Ogata et al. 1957, Gosse 1882 = Sociuncus. 

Sociunc|us (-i, -i) ¢ Lep. Sibatani 1972 (this paper). Dorsal appendages 
of tenth somite as a morphological unit; probably homologous with 
pygopods; may be further divided into uncus and socii. 

Syn. Scaphium Ogata et al. 1957 nec Pierce 1909. 


SUMMARY 


A new term “sociuncus” was introduced for the entire dorsal appendage 
of the 10th somite of male Lepidoptera, of which the uncus and socii are 
parts. The subdivision of the valva was reexamined in the context of 
Klots’s view in Tuxen’s “Taxonomist’s Glossary of Genitalia in Insects,” 
Second Enlarged Edition (1970), with the conclusion that no alteration 
is necessary for the system proposed in a previous paper of this series. 
The “Glossary” was supplemented by a “Proposed Addenda” to it, which 
lists some names treated inadequately or missing therein. 


ACKNOWLEDGMENT 


My hearty thanks are due to Mr. D. P. Sands, Newport, N.S.W., for his 
critical reading of the manuscript. 


LITERATURE CITED 


BirkET-SMiTH, J. 1965. A revision of the West African eilemic moths, based on 
the male genitalia. Papers Fac. Sci., Ser. C, Zool. No. 1, Haile Selassie I Univ., 
Addis Ababa. 161 p. 

Forses, W. T. M. 1939. The muscles of the lepidopterous male genitalia. Ann. 
Entomol. Soc. Amer. 32: 1-10. 

Gossr, P. H. 1882. On the clasping organs ancillary to generation in certain groups 
of the Lepidoptera. Trans. Linn. Soc. London (2) 2: 265-345, pl. 26-33. 
INovE, S. AnD A. Kawazort. 1964. Hesperiid butterflies from South Vietnam (1). 

Tyo to Ga (Trans. Lepid. Soc. Jap.) 15: 34-50. 

1965. Riodinidae, Curetidae and Lycaenidae (Lepidoptera: Rhopalocera ) 
from South Viet-Nam. Nature and Life in Southeast Asia 4: 317-394, pl. 1-20. 

KawazoE, A. aNnD M. WaxkaBayaAsHI. 1969. Two new species of the Chrysozephyrus 


122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Shirézu and Yamamoto (Lycaenidae) from Formosa. Ty6 to Ga (Trans. Lepid. 
Soc. Jap.) 20: 1-10; pl. 1. 

Kuots, A. B. 1970. Lepidoptera, in “Taxonomist’s Glossary of Genitalia in Insects” 
(ed. S. L. Tuxen), 2nd ed. Munksgaard, Copenhagen. p. 115-130. 

Kuznetsoy (= Kusnezov), N. J. 1916. Contributions to the morphology of the 
genital apparatus in Lepidoptera. Some cases of gynandromorphism. Rey. russe 
dentomol. 16: 151-191. 

Ocata, M. 1950. Studies on the male genitalia of Japanese Hesperiidae (Lepidop- 
tera) II. (in Japanese) Trans. Kansai Entomol. Soc. 15 (2): 33-47. 

Ocata, M., Y. Oxapa, H. OxAcaAxr AnD A. SrBATANI. 1957. Male genitalia of 
Lepidoptera: morphology and nomenclature. III. Appendages pertaining to the 
tenth somite. Ann. Entomol. Soc. Amer. 50: 237-244. 

OxacAkI, H., A. SrpaATANI, M. Ocata AND Y. OxApA. 1955. Male genitalia of 
Lepidoptera: morphology and nomenclature. II. Morphological significance of 
sacculus and furca. Ann. Entomol. Soc. Amer. 48: 438-442. 

Prerce, F. N. 1909. The Genitalia of the Groups Noctuidae of the Lepidoptera of 
the British Islands. Liverpool. 88 p., 32 pls. 

SHtmrROzu, T. 1960. Butterflies of Formosa in Colour (in Japanese). Hoikusha, 
Osaka: ASIN pe 16) pls: 

SHimROzU, T. AND H. YAMAMOTO. 1956. A generic revision and the phylogeny of 
the tribe Theclini (Lepidoptera: Lycaenidae). Sieboldia 1: 329-421, pl. 35-85. 

SIBATANT, A., M. OcaTta, Y. OKADA AND H. Oxacaxr. 1954. Male genitalia of Lepi- 
doptera: morphology and nomenclature. I. Divisions of the valvae in Rhopaloc- 
era, Phalaenidae (= Noctuidae) and Geometridae. Ann. Entomol. Soc. Amer. 
47: 93-106. 

Tuxen, S. L. (Ep.). 1970. Taxonomist’s Glossary of Genitalia in Insects. Munks- 
gaard, Copenhagen. 359 p. 


ANOTHER LARVAL FOODPLANT FOR EUPHYDRYAS PHAETON 
(DRURY) (NYMPHALIDAE) 


During the first week of June, 1971, I was collecting in a meadow in lower Orange 
County, New York State, where I had collected Euphydryas phaeton (Drury) in other 
years. This year the season was about one week late and very few of the checkerspots 
were flying. Numerous mature larvae were identified, however, (about two dozen ) 
feeding on scattered bushes of the arrow-wood (Virburnum recognitum L.). This 
shrub was fairly plentiful in this field. Usually there were two to three larvae on a 
bush, feeding on the uppermost leaves. Though turtlehead (Chelone glabra L.) was 
also present in this field, no caterpillars were seen feeding on it. 

It is to be noted that a few days earlier in Andover, Sussex County, New Jersey, a 
at larva of this same butterfly was collected on White Ash (Fraxinus americana 

In 1969, Joseph Muller reported a new larval foodplant for Euphydryas phaeton, 
namely Pentstemon hirsutis (L.) (J. Lepid. Soc. 23: 48). Apparently there are several 
plants which are accepted by mature phaeton larvae. As phaeton is known to form 


colonies on turtlehead in its earlier instars, perhaps its diet becomes more catholic 
with maturity. Perhaps also the requirements of the larger caterpillars may outstrip 
| . . . . . . 

the availability of the original foodplant, forcing a change. 


jouHN J. Bown, M.D., 249 Mountain Avenue, Ridgewood, New Jersey 07450. 


VoLUME 26, NUMBER 2 123 


BOOK REVIEWS 


CENTURIE DE LEPIDOPTERES DE L'ILE DE CuBA, by Ph. Poey. 1832. I-XII + (1) — (4) 
+ [1-50], 20 coloured plates. Reprinted “1970” [1971] by E. W. Classey Ltd., 
Hampton, Middlesex. Distributed exclusively in North America by Entomological 
Reprint Specialists, P.O. Box 77971, Dockweiler Station, Los Angeles, California 
90007. Price $30.00 U.S. 


This is another in the series of excellent reproductions of significant entomological 
books being produced by E. W. Classey Limited. The original has long been exces- 
sively rare. Though it contains descriptions and figures of only 20 species, the work 
having lapsed after the publication of the first two “decades” of the “Centurie,” the 
selection is of such common, interesting or striking forms that the work has com- 
manded widespread attention. Such well-known species as Eumaeus atala, Phoebis 
orbis, Cocytius duponchel and Eurema dina were described here for the first time 
and five genera were described as new, including the pyraustine genus Syllepis, of 
which I have had the pleasure of describing several new species in recent years, and 
Acrolophus, now recognized as the type-genus of the family Acrolophidae. 

The reproduction appears excellent, though I do not have a copy of the original 
available for comparison. Care has been taken to work from an example with a good 
set of plates; one or two inaccuracies have been introduced in the course of partly 
correcting deterioration in the original pigments, but these are carefully noted in the 
introduction. The letterpress is clear and clean, though the texture of the paper is 
somewhat unpleasantly smooth and plastic-like. The introduction by C. F. Cowan 
is concise but scholarly and informative and has been combined harmoniously by the 
typesetters with the main text. The cloth binding is neat and attractive, but some 
will find the chrome yellow dust jacket, with white bands and red lettering and 
ornament, somewhat overpowering. 

The price may seem rather high for a volume dealing with so few species, but 
considering the historical importance and previous scarcity of the work and the good 
quality of the reproduction, the market will undoubtedly justify the publisher's 
judgment. 

EuGENE Munroe, Entomology Research Institute, Canada Department of Agricul- 
ture, Ottawa. 


A MONOGRAPH OF THE ITHOMIIDAE (LEPIDOPTERA). PArT IV. THE TrIsE NAPEO- 
GENINI Fox, by Richard M. Fox and Herman G. Real. 1971. 368 p., 352 fig. Memoirs 
of the American Entomological Institute, Number 15. 


The Napeogenini exhibit Mullerian mimicry and parallel evolution. The relation- 
ships of individual species are so complicated that identification by non-specialists 
has been almost impossible in the past. This reference book provides keys to all of 
the species and subspecies and should open up this tribe of butterflies to the non- 
specialist. To somebody who has not previously worked with the Ithomiidae, the 
keys may appear complicated at first, but considering the insects that they deal with, 
I think they should prove more than satisfactory to most users. 

The monograph treats 7 genera, 104 species and over 300 subspecies. One genus, 
Aremfoxia, is described as new, along with 12 new species and 30 new subspecies. 
Each species and subspecies is described and keyed, and most are depicted in black 
and white plates. Annotation includes citations of original descriptions, synonymies 
and distributional data. Biological data (life histories, behavior, habitats, etc.) are at 
a minimum. 

To fully appreciate this monograph, some knowledge of the extenuating circum- 
stances surrounding its publication is required. With the manuscript about 75% com- 


124 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


pleted, Richard Fox died suddenly on 25 April 1968. The task of completing the 
monograph and preparing it for publication was left to his widow, Jean W. Fox, and 
Herman G. Real, a graduate student who had just arrived in Pittsburgh to study 
under Fox. Further complications resulted when Mrs. Fox died on 10 March 1970 
with the final manuscript still being drafted. About this same time, Real returned 
to California and the task of final proof-reading fell largely upon George E. Wallace. 
In view of these circumstances, the overall quality of the treatment is remarkable. 
The problems presented by a combination of posthumous and joint authorship are 
fairly well overcome, by having those parts of the text that were not written by Fox 
clearly indicated as such. New taxa described are variously credited to Fox, to Real, 
or to Fox and Real. 

The four parts in this series of monographs on the Ithomiidae have been published 
in four different publications. A very high level of excellence was obtained in part 
three (the Mechanitini) and by comparison this publication does not measure up too 
well. It is published by offset printing from typewritten plates and does not have 
the slick appearance of letter press printing. Range maps and other illustrative ma- 
terials were eliminated by economic pressures and the annotation is less extensive. 
These shortcomings do not, however, detract from the scientific usefulness of the 
publication. 

Under the circumstances, this is a remarkable book and it will be a valuable refer- 
ence to those dealing with Ithomiidae for many years. With four more tribes of 
Ithomiidae remaining to be revised, it is hoped that Mr. Real will take up the chal- 
lenge, add his life’s work to Dr. Fox’s and complete this series. 

Joun H. Masters, Lemon Street North, North Hudson, Wisconsin. 


LEePIpopTERA GENETICS, by Roy Robinson. 1971. Int. Ser. Monogr. in Pure and Appl. 
Biol., Zool. Div., Vol. 46: 687 p., 63 tables and 18 figs.; hardbound. Pergamon Press 
Ltd., Headington Hill Hall, Oxford, Eng. $26.50. 


This work encompasses an exhaustive review of the literature dealing with the 
genetics of Lepidoptera published prior to 1966. The author admirably fulfills his 
purpose “to provide a systematic account on a worldwide basis of genetic and karyo- 
logical studies with Lepidoptera species.” The book is intended as a reference for 
“any person who is interested in the variation or breeding of butterflies and moths” 
(i.e.: the amateur, as well as the professional and the specialist). 

The book includes a rather rambling, wide-ranging (but informative) Introduction, 
in which such diverse subjects as 1) color and pigmentation, 2) seasonal and environ- 
mental influences, 3) breeding techniques, 4) genetic and sexual abberations, 5) sex 
determination mechanisms, 6) hybridization, and 7) procedures of taxonomic nomen- 
clature are briefly discussed. Then follow review chapters on the basics of Lepidoptera 
Genetics, Elementary Biometry, Population Genetics and Polymorphism, Industrial 
Melanism, and Mimicry. Each chapter is well-written and lucid, and is more or less 
self-contained. All are thoroughly referenced. 

The next two lengthy chapters present an encyclopedic listing of all species of 
Rhopalocera and Heterocera about which any genetic information has been published. 
Many species are superficially treated, merely having been included in the book to 
indicate the completeness of the literature survey. So little is actually known about 
the genetics of some of them that they could have been omitted without detracting 
1 the book at all. The arrangement of species is alphabetical, and the genetics of 
| developmental stage are discussed in turn, whenever information is available. 
the author points out, many of the postulated genetic mechanisms are specula- 
tive, because of the sparseness of data upon which they are based. The most thor- 

1 ated genera include Colias, Erebia, Heliconius, Maniola, Papilio, and Pieris 
imong the butterflies, and Abraxas, Anagasta, Arctia, Biston, Bombyx, Celerio, 


VOLUME 26, NUMBER 2 125 


Choristoneura, Deilephila, Ectropsis, Galleria, Luffia, Lymantria, Panaxia, Philosamia, 
Solenobia, Sterrha, and Zygaena among the moths. (The above list is representative, 
and not all-inclusive. ) 

The final chapter on Karyology of Lepidoptera consists mainly of a 27-page table, 
listing alphabetically by genera, all species for which the haploid chromosome num- 
ber has been determined. This table includes the references for each observation. 
The chapter concludes with a number of comments and observations dealing with 
meiotic theory under topics such as “polyploidy and the fusion/fragmentation con- 
cept, chiasmata frequency, supernumerary chromosomes, and sex chromatin.” 

One of the most important parts of the book is its comprehensive 60-page Bibliog- 
raphy. All references I examined are accurately cited. Author, subject, and species 
indexes also are included, and represent a most useful feature of the book. The tables 
and figures are presented clearly, and usually can be interpreted without reference to 
the text. Very often they summarize information published in greater detail elsewhere, 
in order to indicate the trends shown by the data. However, the book does lack a 
list of tables and figures. 

The author's style, although somewhat verbose, provides interesting reading. The 
book is clearly worded and his statements are for the most part accurate. The author 
has gone to considerable length to point out to the reader both flaws in the existent 
data and important problems in need of further investigation. For a work of this size 
the book contains remarkably few typographical errors. 

The organization of the book is perhaps its greatest drawback. In his discussions of 
topics such as Polymorphism, Industrial Melanism, and Mimicry, the author has 
tended to follow an historical approach in reviewing the literature. The result of this 
is a somewhat lengthy “hodge-podge” presentation. I should think that the author’s 
rather dreary alphabetical listings of genetic information on the Rhopalocera and 
Heterocera could have been better supplanted by family groupings or some other 
more scientific format. 

Unfortunately, certain portions of the text already seem to have been somewhat 
outdated by research done since 1966. Nevertheless, there are several references cited 
bearing 1967 and 1968 dates. 

Concerning the author’s purpose, as stated in the Preface to the book, I believe this 
work will provide a very handy reference for the specialist and the professional, but 
I think that the biometric, statistical, and genetic theories and methods contained in 
the book are so complex that they will be difficult for one not previously trained in 
these areas to master. Nevertheless, the book does “fill an important gap in the ento- 
mological literature,” by bringing together and summarizing in readily comprehensible 
form a considerable amount of information on the genetics of Lepidoptera. 

Austin P. Puatr, Department of Biological Sciences, University of Maryland Balti- 
more County, 5401 Wilkens Avenue, Catonsville, Maryland 21228. 


THE BUTTERFLIES OF WISCONSIN, by James A. Ebner. Milwaukee Public Museum 
Popular Science Handbook No. 12. 205 p. Available from the Milwaukee Public 
Museum for $5.00. 


This book has an impressive appearance, is well printed and has an excellent for- 
mat, reminiscent of F. M. Brown’s Butterflies of Colorado. A total of 139 species is 
treated, including ten hesperids considered questionable for Wisconsin, and Limenitis 
arthemis being regarded as a distinct species from Limenitis astyanax. Each of these 
is illustrated with black and white photographs and discussed in about a page of 
text. The book is well indexed and includes the usual introductory chapters on butter- 
fly morphology, taxonomy and collecting techniques. 

On reading the book, it becomes apparent that the author has not collected exten- 
sively statewide in Wisconsin and as a result information regarding the northern and 


126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


western counties is quite sparse. Many species, especially bog inhabiting ones, that 
are locally abundant in northwest Wisconsin are given very ineffectual treatment. 
Polygonia satyrus is omitted entirely, although being fairly common from Sawyer 
county northward. Satyrium caryaevorus is not mentioned, although the photographs 
of Satyrium calanus (page 64) appear to be erroneously identified as caryaevorus. 
Colias interior vividior Berger, the only butterfly with a Wisconsin type locality, re- 
ceives no mention at all. 

The author has not attempted to delineate Wisconsin subspecies, stating that, “Sub- 
specific designations for some Wisconsin butterflies have been omitted awaiting study 
and clarification.” However, with only a few species given subspecific designations, 
five are incorrectly cited. Coenonympha tullia inornata is treated as C. inornata ben- 
jamini; Oeneis jutta ascerta is treated as O. jutta ridingiana; Agraulis vanillae incarnata 
is treated as A. vanillae nigrior; the unnamed Wisconsin population of Lycaeides 
argyrognomon is treated as_ L. argyrognomon scudderii; and Boloria selene is treated 
under atrocostalis, which does occur in northern Wisconsin, however southern Wis- 
consin populations, and very obviously the specimen figured as atrocostalis, are sub- 
species myrina. 

This is the first publication to treat all of the butterflies of Wisconsin on a state- 
wide basis. It is a book which has been needed for a long time and which will be in 
use for many years. It is unfortunate that more extensive and detailed “statewide” 
collecting records could not have been. included. 

Joun H. Masters, Lemon Street North, North Hudson, Wisconsin. 


NOTES AND NEWS 
Errata 


In the paper by E. M. Shull and F. Sidney Badger, “Annotated List of the Butter- 
flies of Indiana, 1971” (Vol. 26 (1): 13-24), the family name LyCAENIDAE was inad- 
vertently omitted on p. 18 (between Swamp Metalmark and Coral Hairstreak). Also, 
on p. 17, the subspecific name of Pieris napi should be oleracea (not oleracoa). 


i re ge et eee —S 


i 2 


thee 


ee aS > 
CAPO 


EDITORIAL COMMITTEE OF THE JOURNAL 


Editor: THEODORE D. SarcENT, Department of Zoology, 
University of Massachusetts, Amherst, Massachusetts 01002 


K. S. Brown, J. M. Burns, R. H. Carcasson, J. P. DoNAHUE, J. F. 
Gates CLARKE, R. O. KENDALL, J. H. Masters, L. D. MILLER, © 
AP. Peatr,; J: KR. CG. TorRNeR 


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, 844 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, 1961a, 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 814 
< 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 article’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 approx- 
imate 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. 

Page Charges: Authors with grant or institutional funds are requested to pay a 
charge of $24.00 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 
case of authors who have no grant or institutional funding, as it is not intended that 
any author should pay this charge from personal funds. However, all authors will 
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: Dr. C. V. Covell, Department of Biology, University of 
Louisville, Louisville, Kentucky 40208. 


ALLEN PRESS, INC. eee LAWRENCE, KANSAS 
USA 


CONTENTS 


PRACTICAL FREEZE-DRYING AND VACUUM DEHYDRATION OF CaTeRpr.- 


LARS. Richard. B. Dominick ee 


A NEw SPECIES OF THE GENUS PYROMORPHA HERRICH-SCHAEFFER 
(PyromMorPHwae). André Blanchard | 


Tue Lire History or SCHINIA CITRINELLUS (NocrumaE). D. F. 


Hardwick 2 oe | 


Tue Lire History oF SCHINIA JAEGERI (Nocrumae). D. F. Hardwick 


STUDIES ON THE CATOCALA (NOCTUDAE) OF SOUTHERN NEW ENGLAND. 
III. Matinc Resutts witu C. RELICTA WALKER. Theodore 
D. Sargent 


Species Diversity IN CATOCALA (NOCTUIDAE) IN THREE LOCALITIES IN 
NortH America. D. F. Owen 


POLYMORPHISM IN PAPILIO GLAUCUS L. (PAPILIONIDAE): MAINTE- 
NANCE OF THE FEMALE ANCESTRAL Form. Sally K. Makielski 


JUNIPERUS (CUPRESSACEAE) SPECIATION AND THE RANGES AND EVOLU- 
TION OF Two CALLopuHRys (LYCAENDAE). Kurt Johnson 


MALE GENITALIA OF LEPMOPTERA: MORPHOLOGY AND NOMENCLA- — 


TURE IV. Notes oN TUXEN’s “TAXONOMIST’S GLOSSARY OF 
GENITALIA IN INSECTS’: SECOND ENLARGED EDITION. A. 
Sibatani 


GENERAL NOTES 


Life history notes on Callosamia securifera (Saturniidae). Michael D. 
Van Buskirk oo a 


Observed mating between Pieris rapae and Pieris protodice (Pieridae). 
Richard C.. Priestaf 00 


McFarland moth collection donated to the Natural History Museum of Los 

Angeles county. Julian P. Donahue 2...) 
Pseudophilotes Beuret, 1958. F. Martin Brown 
Butterflies feeding on a dead bobcat. Oakley Shields _...- 


Accidental occurrence of Aglais urticae (Nymphalidae) in Nova Scotia. 
Frederick W. Scott and Barry Wright bs 


Another larval foodplant for Euphydryas phaeton (Drury) (Nymphalidae). 
John J. Bowe 


Book REvIEws 


116 


j 
A 


Volume 26 1972 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 


29 September 1972 


THE LEPIDOPTERISTS’ SOCIETY 
EXECUTIVE COUNCIL 


Lioyp M. Martin (Prescott, Ariz.) President 

J. F. Gates CLarxe (Washington, D.C.) President-elect 
S. A. Az (Nagoya, Japan) Ist Vice President 

Kerru S. Brown (Rio de Janeiro, Brasil) Vice President 
H. A. FREEMAN (Garland, Texas) Vice President 

S. S. Nico.ay (Virginia Beach, Va.) Treasurer 

LEE D. Mixer (Sarasota, Fla.) Secretary 


Members at large (three year term): R. B. Dominick (McClellanville, S.C.) 

B. MATHER (Clinton, Miss.) 1972 1973 

M. Ocata (Osaka, Japan) 1972 J. P. DonanveE (Los Angeles, Calif.) 1973 

E. C. Wetuinc ( Merida, Mexico ) 1972 J. M. Burns (Cambridge, Mass.) 1974 

ANDRE BLANCHARD ( Houston, Texas ) R. H. Carcasson (Vancouver, B.C.) 1974 
1973 M. C. Nretson (Lansing, Mich. )1974 


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 $10.00 
Student members—annual dues $5.00 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $15.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia 23455. 


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


by Cyru F. pos Passos 


Price, postpaid: Society members—$5.00, others—$7.50; uncut, unbound signatures 
available for interleaving and private binding, same prices; hard cover bound, mem- 
bers—$8.00, others—$10.00. Revised lists of the Melitaeinae and Lycaenidae will be 
distributed to purchasers free (separately with paper covered copies and unbound 
signatures, bound in with hard covered copies ). 


The Lepidopterists’ Society is a non-profit, scientific organization. The office of 
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520. 
Second class postage paid at Lawrence, Kansas, U.S.A. 66044. 


JOURNAL OF 


Tue LepIpopreERISTS’ SOCIETY 


Volume 26 1972 Number 3 


THE GENUS ZESTUSA (HESPERIDAE) IN EL SALVADOR 
WITH DESCRIPTION OF A NEW SPECIES 


STEPHEN R. STEINHAUSER 
Apartado 109, San Salvador, El Salvador 


The genus Zestusa Lindsey, 1925, a replacement name for Plestia 
Mabille, 1888, which is a junior homonym of Plestia Stal, 1871, includes 
two species: Z. dorus (Edwards), 1882 and Z. staudingeri (Mabille), 
1888, the type. Z. staudingeri is further subdivided by Evans (1952) into 
two subspecies: the nominate and elwesi (Godman & Salvin), 1893. Ap- 
proximate geographic ranges of the three forms have been reported as 
follows: 


Z. dorus—Southwestern U. S. to northern Mexico (Sonora) 
Z. s. elwesi—Mexico (Vera Cruz) 
Z. s. staudingeri—Mexico (Chiapas) to Guatemala 


Very little has been published relating either directly or indirectly to 
the butterflies of El] Salvador. Distribution data for Zestusa (= Plestia) 
published by Godman & Salvin (1893), Draudt (1921) and Evans (1952) 
do not mention El Salvador, nor were any Zestusa species listed by Franz 
and Schroder (1954). The recent discovery in E] Salvador of two Zestusa 
species, s. staudingeri and a new species described below, therefore con- 
stitutes a new distribution record for the genus. 

Both of these insects were found in the cloud forest of the Hacienda 
Montecristo near the summit of Cerro Miramundo which marks the com- 
mon corner of El Salvador, Guatemala and Honduras. Fifteen males and 
one female of the new species and 53 males of Z. s. staudingeri were 
taken during late February and March, 1970 through 1972 at 2300 m. 

The examples of Z. s. staudingeri from Miramundo match the figure in 
Godman & Salvin (1893) very closely and fit quite nicely in the key of 
Evans (1952) although some specimens show traces of the hyaline spots 
in spaces 4 and 5 of the forewing that are typical of Z. s. elwesi, perhaps 


128 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1 & 2. Upper and under side of Zestusa levona Steinhauser, holotype male, 
Hacienda Montecristo, Cerro Miramundo, Cloud Forest, El Salvador, elev. 2300 m. 27 
February 1971 (S. R. & L. M. Steinhauser). Natural size. 


indicating a clinal variation between the two forms. The figure of elwesi 
in Seitz (Draudt, 1921) is somewhat exaggerated; the yellow discal and 
submarginal markings of the upper side of the hindwing as illustrated 
are entirely too regular and sharply defined. Z. s. staudingeri is not 
figured. The male genitalia of the Miramundo specimens differ some- 
what from the sketch in Evans (1952) which is undoubtedly of elwesi as 
there were no examples of staudingeri in the British Museum (Natural 
History ) at the time he prepared his catalogue. Unfortunately he did not 
illustrate the uncus, thus implying that it is nearly identical to that of 
Z. dorus which is illustrated; Godman & Salvin’s figure shows only the 
lateral view. Because of this I have included a sketch of the genitalia of 
staudingeri from Miramundo (Fig. 4). The principal difference from 
elwesi lies in the greater caudal production of the cuiller and generally 
narrower clasp of staudingeri. The genitalia of 31 males examined showed 
very little individual variation. 

The new species, which I am pleased to name levona in honor of my 
wife, a very able and most enthusiastic collector, is described below. 


Apart from substituting penis for aedeagus, the morphological and struc- 
tural terms of Evans (1952) have been adopted. 


Zestusa leyona Steinhauser, new species. 
MALE 


Wing Measurement: Forewing, base to apex, 20 mm (range of 19-21 mm in 15 
type specimens ). 


Upperside (Fig. 1): Forewing with pronounced costal fold; color dark brown with 
red-bronze reflection, slightly darker toward termen with bronze hairs on basal and 
discal areas; fringe dark grey-brown. Small white hyaline spots in discal area: two 
(upper and lower) usually separate but occasionally narrowly conjoined in space 2 
just distad origin vein 3; sometimes minute dot space 1b immediately behind outer 


VOLUME 26, NUMBER 3 129 


Fig. 3. Male genitalia of Zestusa levona, paratype; specimen not dissected. Ven- 
tral view (left) shows relative position of opened clasps to uncus, gnathos and penis. 


edge of lower spot space 2; narrow straight dash space 3 normal to vein 3 well sepa- 
rated distad from spots space 2; generally three pre-apical spots in straight line nearly 
normal to costa spaces 7 (may be absent) 8, 9 and fourth (may be minute or absent) 
space 6, displaced distad; may be minute upper spot space 4 behind spot in space 6; 
upper cell spot basad of spots in space 2, usually extending more than halfway across 
cell but may be minute; small spot near base space 11 behind costal fold. 
Hindwing with short (+ 5 mm) tail along prolongation vein 1b; cell weakly closed, 
vein 5 present but vestigial, marked by well defined crease. Color same dark brown 
as forewing but with diffused yellowish-ochreous discal stripe, more sharply defined 
basad, crossed by dark veins and zig-zag dark line distad of origins veins 4 and 6. 
This pale discal area tapers from vein 6 or 7 where very diffuse to vein 1b where 
terminates in more sharply defined point just forward of base of tail. Fringe at inner 
anal angle dark brown, sharply contrasting with pale yellow to white fringe of tail 
and rear half termen; on forward half of termen checkered with dark brown at vein 
ends and grading to completely brown at apex. Fringe on tails long (2 mm). Costal 
area thinly covered dark brown hairs; dorsum densely covered pale tawny hairs. 


Underside (Fig. 2): Forewing dark brown; same hyaline spots as upper side sur- 
rounded by diffuse haloes of slightly darker brown; scattered ochreous-yellow super- 
scaling more concentrated along costa above vein 12, beyond end of cell in spaces 4 
and 5 forming vague paler spot, and in space 1b forming indistinctly divided spot 
behind and basad of hyaline spot in 1b. May be additional small white spot on costa 
in space 12 adjacent to spot in 11. 

Hindwing as upper side but discal pale stripe more sharply defined, pale yellow, 
extending from vein 8 to vein 1b, widest (4 mm) in 5 and 6, containing four small 
dark brown spots in spaces 3—6, those in 5 and 6 offset distad. Detached diffuse 
pale spot space 7 basad of discal stripe and yellow super-scaling in basal wing area. 


Body: Head, thorax and abdomen dark brown; thorax and abdomen densely clothed 
tawny hairs above, yellow-tawny hairs beneath; abdomen beneath ringed pale yellow 
at each segment. Head above with mixed pale yellow and dark brown hairs. Palpi 
porrect, hairy, dark brown above with some pale yellow hairs and scales, white be- 
neath with some black hairs; cheeks white. 

Legs brown with pale scaling, densely hairy; mid and hind tibiae smooth; mid 
tibiae single pair spurs; hind tibiae two pairs. Antennae longer than half costa, 
reaching beyond origin vein 11; shaft black, plain below, some yellow checkering 
near base above; club broadly arcuate, pale yellow beneath; nudum 19-22 brown. 


130 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


M5 \ | | 
| \ | } \ 
| \ \ \ i | : 
} / if 
/ ) , ! 
) (ih ara hed | 
\ / Nas aN \ | | 
\ fi Y / \, \ | I 
\ / 
; Ve. (4 


/ 


\ { 
] \ | WY wa 
Wey i Ze 
=) | 


Fig. 4. Zestusa staudingeri staudingeri (Mabille), male genitalia of specimen from 
Cerro Miramundo, El Salvador, 18 March 1971 (S. R. & L. M. Steinhauser ). 


Genitalia (Fig. 3): Uncus weakly bi-lobed, narrow; gnathos laterally sclerotized, 
non bi-furcate; penis relatively broad and blunt. Clasps symmetrical, cuiller project- 
ing to sharp serrated dorsal tooth, its ventro-caudal apex right angled and sharp; valva 
with short inward-pointing dorsal projection. 


FEMALE (Fig. 5) 


Wing Measurement: Forewing, base to apex, 22 mm. Both wings slightly broader 
than male. 


Upperside: As male but forewing spots all present and larger. No fold. 


Underside: As male but forewing spots as above with additional faint sub-apical spot 
in 5; hindwing as male but pale discal stripe wider (5 mm). 


Body: Head, thorax, abdomen, legs as male; antennae as male, nudum 22. 


Type Material: Holotype, male, Hacienda Montecristo, Cerro Miramundo, Cloud 
Forest, El Salvador, elev. 2300 m, 27 February 1971 (S. R. & L. M. Steinhauser), No. 
H-1124, in the U.S. National Museum, Washington, D.C. Fourteen male paratypes, 
same location (type locality) and collector, 7 March 1970 (Nos. H-223, H-224); 27 
February 1971 (No. H-1125); 13 March 1971 (No. H-1168); 18 March 1971 (No. 
H-1234); 4 March 1972 (No. H-4185); 5 March 1972 (No. H-4186); 25 March 1972 
(Nos. H-4288, H-4289) and 26 March 1972 (Nos. H-4291 through H-4295) are 
deposited as follows: one in the American Museum of Natural History, New York; 
one in the Carnegie Museum, Pittsburgh; one in the British Museum (Natural His- 
tory), London and 11 in my own collection. The allotype female, same location and 
collector, 26 March 1972 (No. H-4290) will remain in my own collection. 


Z. levona is strikingly different in habitus (Figs. 1, 2,5) from the three 
previously known forms of Zestusa and can be recognized immediately 
by the pale yellow stripe and prominent, white-fringed tail of the hind- 
wing. The male genitalia (Fig. 3) have a slightly broader and more pro- 
nouncedly lobed uncus than staudingeri (Fig. 4); the cuiller is ventrally 

ight angled rather than smoothly rounded and the valva is longer and 
is an inwardly curved dorsal projection. 


VOLUME 26, NUMBER 3 131 


Fig. 5. Zestusa levona Steinhauser, allotype female, upper and under side; Hacienda 
Montecristo, Cerro Miramundo, Cloud Forest, El Salvador, elev. 2300 m. 26 March 
1972 (S. R. & L. M. Steinhauser). Natural size. 


Biological Notes 


The isolated cloud forest of Cerro Miramundo covers an estimated 20 
sq. kms. of the frontier zone between E] Salvador, Guatemala and Hon- 
duras, extending down from the peak (2418 m) to about 2100 m. Being 
the sole remaining patch of virgin cloud forest in El Salvador, it affords 
a unique site to explore the distinctive and restricted insect fauna associ- 
ated with this environment. Of the 114 butterfly species so far known 
from there, 35 have not yet been found elsewhere in E] Salvador. Be- 
cause of the extremely dense forest growth, collecting is virtually impos- 
sible except in clearings or at the edge of the forest. 

The observed flight season of Zestusa at Miramundo does not extend 
beyond February, March and April; despite good collecting conditions 
and careful search, none were found on 23, 24 January or 8, 9 May 1971. 
Unfortunately it was not possible to visit the area between 18 March and 
8 May 1971, and the earliest good weather conditions encountered after 
January 1971 were on 27 February. Only one collecting trip (7 March) 
was made between January and November 1970. No Zestusa were found 
12-15 February 1972 but collecting was good for both species during 2-5 
and 25-26 March 1972. 

Although both species were found in the same limited area at the same 
time, their habits are quite different. Z. levona was observed only either 
in flight (approx. 7 m) or visiting flowers. One female and 14 males 
were taken on the pale lavender blossoms of a tree of the family Aster- 
aceae identified by T. F. Hall as Ageratum sp. close to rugosum Coulter. 
Standley and Calderon (1925) describe rugosum as generally less than 
one meter high, whereas this tree grows up to five meters. One male was 
taken from the purple flowers of another tree not yet identified. Z. s. 
staudingeri on the other hand, was never found visiting flowers. One 


132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


specimen was taken while drinking from a damp spot on a vertical rock 
face (tuff) along with a male hesperiid, Doberes anticus sobrinus (God- 
man & Salvin), 1895. Aside from this, it was observed only in a “territorial 
defense” position on leaves or bare twigs of various trees along the road, 
whence it frequently sallied forth in a rather leisurely “inspection” flight, 
covering up to 20 meters in each direction before returning to its observa- 
tion post, or in rapid attack flight against other passing skippers. It never 
was seen to attack other insects or even butterflies of other families 
though many flew past. It frequently could be induced to investigate a 
net waved to and fro resulting in its capture, although it often found my 
head a more interesting subject, circling so close as to make capture 
impossible. 

Despite these differences in habits and the much less extensive nudum 
of the antennal club (21 segments instead of the 34 mentioned by Evans 
(1952) as a character of the genus), the porrect palpi, very hairy legs, 
broadly curved antennal club and general aspect of the genitalia place 
levona quite clearly in the genus Zestusa. 


ACKNOWLEDGMENTS 


I am very grateful to Col. S. S. Nicolay for his helpful advice and sug- 
gestions and to Mr. H. A. Freeman for confirming Col. Nicolay’s opinion 
that Z. levona had not been described. I am also indebted to Dr. Thomas 
F. Hall for identifying the tree whose flowers are so attractive to levona, 
to Dr. Sam Breeland for his critical review of this manuscript and to Don 
Eimesto Freund, owner of the Hacienda Montecristo, for allowing us 
access to his property to collect insects. 


LITERATURE CITED 


Draupt, M. 1921. MHesperidae (sic). In Macrolepidoptera of the World, A. Seitz, 
ed. Vol. 5. Stuttgart. 

Evans, W. H. 1952. A Catalogue of the American Hesperiidae in the British Mu- 

_ seum (Natural History). Part 2. Pyrginae. Section 1. London. 

FRANZ, E, AnD H. ScurOpER. 1954. Tagfalter (Lep. Rhopalocera) aus El Salvador. 
Senck. biol. Band 35, Nummer 1/2, Seite 75-87. Frankfurt am Main. 

GopMaN, F’. D. anp O. Satvin. 1887-1901. Biologia Centrali-Americana. Insecta. 
Lepidoptera — Rhopalocera 2: 244-637. 

STANDLEY, P. C. ann S. CALDERON. 1925. Lista Preliminar de Plantas de El Salva- 
dor. Publ. del Ministerio de Instruccion Publica de la Republica de El Salvador, 
C, A. 


VOLUME 26, NUMBER 3 133 


SOME OBSERVATIONS ON THE LEPIDOPTERA OF 
BROMELIADS 


Carios R. BEUTELSPACHER 
Instituto de Biologia, Apdo. Postal 70-233, México 20, D.F. Mexico 


The bromeliads are mostly tropical plants, with leaves generally ar- 
ranged in rosettes which permit the accumulation of water and detritus; 
these form a microhabitat suitable for many organisms, principally insects, 
whose different interrelations constitute special ecosystems of great bio- 
logical interest (Picado, 1913; Smith, 1938). 

Compared to other groups of insects inhabiting these plants, Lepidop- 
tera are scarce. Thus, in his masterly work “Les Bromeliacees epiphytes 
considerees comme milieu biologique,’ Picado (1913) reports only two 
species: Valentinia bromelia Walsing. (Blastobasidae) from Cordoba, 
Veracruz, México, and Acrolophus pallidus Moschler (Acrolophidae ) 
from Costa Rica. He reports the larvae as living among the leaves of 
Aechmea and other large bromeliads. Biezanko (1961) reports four moths 
from bromeliads in Rio Grande do Sul, Brazil: Castnia acraeoides Gray, 
C. boisduvali Walker, C. garbei Fortterle and C. satrapes catharina Preiss, 
collected in Tillandsia aéranthos (Loisel) L. B. Smith, Bromelia anti- 
acantha Bertol and Ananas comosus (L.) Merr. 

In Mexico I have collected the following species in bromeliads—ar- 
rangement according to Hoffmann’s Catalogue (1940): 

Melinaea imitata Bates (Danaidae). A chrysalis was found in a leaf of 
Aechmea bracteata (Swartz) Griseb (Fig. 1.), at the Tropical Biological 
Station “Los Tuxtlas,’ near Sontecomapan, Veracruz. This species we 
consider only occasional, since larvae have not been found feeding on 
this plant, although all known Ithomiidae feed on Solanaceae as larvae. 

Napaea eucharilla picina Stichel (Riodinidae) (Figs. 7-8). Caterpil- 
lars were found eating the leaves of Aechmea bracteata (Swartz) Griseb, 
and Aechmea nudicaulis var. nudicaulis L. B. Smith, collected at the “Los 
Tuxtlas” Station. 

Caria domitianus ino Godm. & Salv. (Riodinidae). Larvae eating the 
leaves of Tillandsia caput-medusae E. Morren, collected at San Francisco 
Acuitlapan, Guerrero; they hatched on 14 June 1971. 

Thecla basalides Geyer (Lycaenidae) (Figs. 2-4). The caterpillars are 
a serious pest of pineapple (Ananas comosus (L.) Merr.), causing great 
losses of this crop; they enter the inflorescence and cause an abnormal 
development, besides facilitating the entry of bacteria and fungi which 
cause rotting. I have also found these larvae eating the fruits of another 


134 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Aechmea bracteata (Swartz) Griseb, a giant bromeliad from Veracruz. 


bromeliad, Aechmea bracteata (Swartz) Griseb (Fig. 1), in southern 
Veracruz. 

Thecla hesperitis Btlr. & Dre. (Lycaenidae). I found larvae eating the 
leaves of Tillandsia caput-medusae E. Morren, which I collected at 
Acahuizotla, Guerrero; they hatched on 20 June 1970. 

Automeris janus metzli (Sallé) (Saturnidae). A cocoon was found in 
an Aechmea bracteata var. pacifica Beutelsp. at E] Rincon, Guerrero. It 
hatched on 18 August 1971; but this, as well as the next species, is con- 
sidered a casual occurrence. 

Urania fulgens Walk. (Uranidae). A cocoon was found among the 
leaves of Aechmea bracteata (Swartz) Griseb, at the “Los Tuxtlas” Bio- 
logical Station. 

Ammalo megapyrrha Walk. and Ecpantheria sp. (Arctiidae) found 
among leaves of the same Aechmea bracteata in San Luis Potosi and in 
southern Veracruz. 

Acrolophus vigia Beutelsp. (Acrolophidae) (Figs. 5-6). This species 
was described from larvae cultivated in the laboratory and reared to ob- 
tain adults (Beutelspacher, 1969). The larvae were found in Aechmea 


VoLUME 26, NUMBER 3 sp 


Figs. 2-4. Thecla basalides Geyer (Lycaenidae). 2, egg on fruits of Aechmea 
bracteata (Swartz) Griseb; 3, pupa, lateral; 4, female. 


136 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 5-6. Acrolophus vigia Beutelsp. (Acrolophidae). 5, male; 6, female. 
Figs. 7-8. Napaea eucharilla picina Stikel (Riodinidae). 7, pupa; 8, male. 


VoLUME 26, NUMBER 3 ISI 


bracteata (Swartz) Griseb, in Veracruz and Yucatan, as well as in Aechmea 
mexicana Baker and Vriesia gladioliflora (Weindl.) Ant. in the “Los 
Tuxtlas” region, Veracruz, and in Vriesia chiapensis Matuda, from Bochil, 
Chiapas. The caterpillars feed on bromeliad leaves, and are semi- 
aquatic in their habits. The adults emerged in February. 

I would like to express my thanks to Dr. Allan R. Phillips for English 
translation of this work. 


LITERATURE CITED 


BEUTELSPACHER, B. C. 1969. Una especie nueva de Acrolophus Poey, 1832, de 
Bromeliaceas. (Lepidoptera, Acrolophidae). An. Inst. Biol. Univ. Nal. Autén. 
México, Ser. Zool. 40 (1): 43-48. 

BiEZANKO, C. M. 1961. Contribucao ao conhecimento da fisiografia do Rio Grande 
do Sul. XIV. Castniidae, Zygaenidae, Dalceridae, Eucleidae, Megalophygidae, 
Cossidae et Hepialidae da Zona Missioneira do Rio Grande do Sul. Arq. Entomol. 
Ser. B. Escola de Agronomia “Eliseu Maciel.” 

HOFFMANN, C. C. 1940. Catalogo Sistematico y Zoogeografico de los Lepiddpte 
ros Mexicanos. 1* parte, Papilionoidea. An. Inst. Biol. Univ. Nal. Auton. México 
11 (2): 639-739. 

Picapo, C. 1913. Les Bromeliacées epiphytes, considerées comme milieu biologique. 
Bull. Sci. Fr. Belg. 47: 215-360, Pl. 6-34. 

SmirH, L. B. 1938. North American Flora (Xyridales) Bromeliaceae. N. Y. Bot. 
Garden 19 (2): 61-228. 


THE EFFECT OF CAUTERIZING 
THE MNPPM OF THE PUPA OF THE 
MONARCH BUTTERFLY (DANAUS P. PLEXIPPUS) (DANAIDAE) 


F. A. URQUHART 


Scarborough College and the Department of Zoology, University of Toronto, 
Toronto, Canada 


The pupa of the monarch butterfly possesses well-defined surface pig- 
mented areas (color plate, e) which, owing to the configuration of the 
lamellae of the cuticle plus the presence of a yellow epidermal pigment, 
imparts to them the appearance of golden spots and hence they have 
been referred to as “gold spots” in the literature. Since “gold spots” is 
not descriptive of these structures the terminology prismatic pigmented 
maculae (PPM) has been suggested (Urquhart & Tang, 1970). 

Since the PPMs are constant in number and position and further since 
it was suspected that they may perform specific functions, terms have 
been applied to each pair, the designation of such terms referring to 
morphological structures of the developing imago (Urquhart, 1960). 


138 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


As to the possible function of the PPMs, it was suggested that perhaps 
they acted as “light receptors” controlling in some manner the develop- 
ment of the imago (Urquhart, 1960). However, it was later indicated that 
light did not appear to have any effect on either the emergence time or 
the morphological structures of the adult butterfly (Petersen, 1964; 
Taylor, 1964). 

By utilizing a micro-cauterizing technique (Urquhart & Dampney, 
1969) the tissues of the lateral ulnar (LU), lateral notal (LN) and 
median ocular (MO) PPMs were destroyed and it was found (Urquhart 
and Tang, 1970) that certain areas of the wing became faded (with 
respect to the treatment of the LUPPM and LNPPM) or that the head 
lacked scales (with respect to the treatment of the MOPPM). 

In the present report the tissues of the median notal MNPPM (Fig. 1) 
were destroyed. Since the MNPPMs possess a much thinner layer of 
cuticle, compared to the other maculae which have been previously 
studied, one series of 25 pupae were treated for 3 secs. fulgurating time 
and a second series for 5 secs. Both maculae of a pair were treated in 
some pupae and only one of the pair in others. This was done to compare 
results of the effect of cauterization on one specimen thus eliminating 
the possibility of individual variation. Twenty-five pupae were used as 
controls in which an area remote from the MNPPM, but still within the 
notal region, was cauterized. 

Of the 25 specimens cauterized for 3 secs., none exhibited wing fading, 
thus indicating that the time period was not sufficient to destroy the 
tissue of the PPM. 

Of the 25 pupae cauterized for 5 secs. fulgurating time, 23 adults 
emerged and all but two of them indicated varying degrees of wing 
fading. Fourteen specimens indicated fading in the mesothoracic wings, 
mostly in the cubital area (Color plate, a- in this particular example only 
the left MN was treated ); one specimen (Color plate, b) indicated fading 
in both meso- and metathoracic wings; six specimens (Color plate, c) 
indicated marked fading of the metathoracic and slight fading in the 
cubital area of the mesothoracic wings; two specimens indicated no 
discernible fading (Color plate, d). No fading was indicated in the 
controls. 

It has been previously suggested that the PPM might govern wing 
pigmentation for fairly well-defined areas of the body ( Urquhart & Tang, 


=> 


Color Plate: a, b, c, d—effect on wing pigmentation of cauterization of the 
ViNPPM of the pupa; e—pupa of the monarch butterfly showing the PPMs (“gold 


ots’) on one side. 


—> 


VOLUME 26, NUMBER 3 139 


a 
YaN 
SA ae NS Ea LU 
‘ S652 
( + 
tee Baste LING 
; oO Ge 
Naf bn 
| jr \ 
/ — 
Sy: \ 
| = ~ aN \ | 
\e =. ey 
y Wl at Abd 
—— 


Fig. 1. Abbreviated designations of the PPM: LU, lateral ulnar; LN, lateral 
notal: MN, median notal; Abd, abdominal; Df, dorsal frontal; MO, median ocular; 
VF, ventral frontal; LO, lateral ocular; MU, median ulnar; Al, alar. 


1970). However, the present experiment would seem to indicate that, 
although the hind wings became faded as a result of cauterizing the 
MNPPM, which did not occur with cauterizing the LUPPM and LNPPM, 
the degree of variation is such as to presuppose an interrelationship be- 
tween the various PPMs. Thus, one pair of PPMs may govern wing pig- 
mentation in a certain area of the wing, as in the case of the LU and 
LNPPM, while others, although primarily governing pigmentation in 
another area, such as the MNPPM and the metathoracic wings, may also 
affect other areas, such as the mesothoracic wings. 

It is not known in what manner the PPMs influence the pigmentation 
of the scales of the wings nor is it known for what period of time the 
PPMs remain active in controlling scale pigmentation. In our experiments 
we have timed the period of cauterization within 24 hours after the ap- 
pearance of the pupa from the larval skin because our histological studies 
indicated that the pigment disappeared from the cell cytoplasm after 24 
hours and also the morphology of the tissue cells changed so as to be 
indistinguishable from the surrounding epidermal tissue. 


140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


It may be conjectured that PPMs found in the pupae of other families 
of Lepidoptera will exhibit results similar to those indicated in the present 
and previous papers. 


ACKNOWLEDGMENTS 


The present investigation, which is part of a large project dealing with 
the ecology of the monarch butterfly, was supported by grants from the 
National Geographic Society of the United States and the National Re- 
search Council of Canada. 

The colour photographs appearing with this paper were taken by 
David Harford, photographic technician of Scarborough College. 


LITERATURE CITED 


PerersEeN, B. 1964. Humidity, darkness and gold spots as possible factors in pupal 
duration of monarch butterflies. J. Lepid. Soc. 18(4): 230-232. 

Taytor, R. L. 1964. The metallic gold spots on the pupa of the monarch butterfly. 
Entomol. News. 72(10): 253-256. 

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

& P. Dampnery. 1969. Microcauterization to maxilectomize lepidopterous 

larvae by fulguration. Can. J. Zool. 47(6): 1416-1417. 

& A. P. S. Tanc. 1970. The effect of cauterizing the PPM (“gold spots” 

of authors) of the pupa of the monarch butterfly (D. plexippus). J. Res. Lepid. 

9(3): 157-167. 


CERCYONIS PEGALA BLANCA, A “MISSING TYPE” IN THE 
EVOLUTION OF THE GENUS CERCYONIS (SATYRIDAE) 


THomas C, EMMEL 
Department of Zoology, University of Florida, Gainesville 32601 


AND STERLING O. MATTOON 
2109 Holly Avenue, Chico, California 95926 


ach of the smaller three species of the nearctic Cercyonis has a very 
distinctive whitish race which is adapted to the arid alkaline flats of the 
western deserts of the United States. Cercyonis meadi alamosa Emmel & 
mmel occurs in salt flats of the isolated San Luis Valley of south-central 
Colorado at 8,500 feet elevation (Emmel & Emmel, 1969). Cercyonis 
sthenele paulus Edwards is a white-marked form occurring in extensive 
populations throughout the western portions of the Great Basin between 
ockies and the Sierra Nevada. Cercyonis oetus pallescens Emmel & 


VOLUME 26, NUMBER 3 141 


Emmel, an extraordinary form with whitish ventral surface, inhabits 
the dry alkaline meadows of the Reese Valley in Nevada (Emmel & 
Emmel, 1971). At the time of this last-mentioned publication, it was 
thought that the closest analogous phenotype in the large Cercyonis 
pegala complex was C. pegala gabbi Edwards from Utah, which has light 
undersurfaces. : 

However, in a remote part of Nevada in late summer 1970, one of us 
(SOM) discovered a heretofore-unsuspected form of Cercyonis pegala 
Fabricius, which matches for dramatic adaptation of ventral white colora- 
tion the previously mentioned smaller Cercyonis. The purpose of this 
paper is to formally describe the new set of populations and to report the 
unique life history of this subspecies. 


Cercyonis pegala blanca Emmel & Mattoon, new subspecies 


Holotype, male: Expanse, 47.8 mm. Forewing length, 16.6 mm. Forewings, 
superior surface: Identical to pattern and coloration of C. pegala gabbi, being dark 
brown with a faint yellow ring around each of two major ocelli. Both major ocelli 
pupilled with white scales. Occasionally one or two additional minor ocelli, lacking 
pupils. Hindwings, superior surface: Dark brown, with two or more (as many as 
five) well-marked submarginal ocelli. Large ocellus in cell Cu: is almost always 
pupilled with white scales. Forewings, inferior surface: Brown areas (except dark 
brown striations) present in other pegala subspecies are here covered with whitish 
or light tan scaling. Yellowish rings around major ocelli are broad but not joined 
as in C. pegala ariane f. stephensi [2] Wright (see Emmel, 1969). Hindwings, in- 
ferior surface: Entire wing except for dark brown striations is heavily suffused with 
silvery white scaling. Six marginal ocelli present in all males collected. Head, thorax, 
and abdomen: Dark brown on dorsal surfaces, whitish on ventral surfaces. Genitalia: 
As in other C. yegala (Emmel, in prep.). 

Allotype, female: Expanse, 55.5 mm. Forewing length, 31.1 mm. Superior 
surface: Generally similar to the variable dorsal phenotype found in C. pegala 
ariane t. stephensi | 2], with a broad yellow submarginal band always found on the 
forewing and usually present on the secondaries also. The ground color usually 
quite tannish rather than a dark brown as in the male. Forewing ocellation varying 
from two to five ocelli, but usually only two major ones and these are always pupilled. 
Hindwing ocellation very well developed, with as many as six black ocelli (M: and 
Cu: ocelli pupilled). Inferior surface: Both forewings and hindwings heavily suffused 
with whitish or silvery white scaling, obscuring all brown areas except principal 
striations. Forewing-band area is light tan on most specimens. Head, thorax, and 
abdomen: As in male, though lighter on dorsal surface corresponding to lighter color 
of wings. 

Holotype male: Nevada, Hwy. 140 at Dufurrena Ranch—Chas. Sheldon Antelope 
Range, T45N, R26E, Humboldt County, August 21, 1970, S. O. Mattoon, collector. 

Allotype female: Nevada, Hwy. 140 at Dufurrena Ranch—Chas. Sheldom Antelope 
Range, T45N, R26E, Humboldt County, August 11, 1970, S. O. Mattoon, collector. 
Paratypes: 20 ¢¢, 10 2° 9, same locality and date as holotype, S. O. Mattoon, 
collector; 8 ¢ 6, 20 2 2, same locality and date as allotype, S. O. Mattoon, col- 
lector. 

The holotype and allotype will be deposited in the Florida State Collection at the 
University of Florida. Pairs of paratypes will be deposited in the following institu- 
tions: American Museum of Natural History, Allyn Museum of Entomology, Cali- 


142 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-6. (1) Cercyonis pegala gabbii male, dorsal surface, from Salt Lake City, 
Salt Lake Co., Utah, 3 July 1965, Kenneth B. Tidwell, collector; (2) Cercyonis 
pegala blanca Emmel & Mattoon, dorsal surface of holotype male, from the type 
locality, Humbolt Co., Nevada; (3) Cercyonis pegala blanca Emmel & Mattoon, dorsal 
surface of allotype female; (4-6) ventral surfaces of the above specimens, respec- 
tively. 


fornia Academy of Sciences, and the Los Angeles County Museum of Natural His- 
tory. The remainder of the paratypes are being retained by the authors for further 
study. The subspecific name, blanca, is Spanish for “white.” 

This subspecies differs from previously described forms of Cercyonis 
pegala primarily in its extraordinary white scaling on the undersides of 
both wings. It is of special evolutionary interest because its development 
of white scaling matching its unusual white alkaline-flat environment 
parallels the phenetic changes that have occurred in Cercyonis meadi, 
sthenele, and oetus under similar environmental regimes. Each represents 
a terminal point as offshoots of the main stocks of the respective species, 
and while the existence of C. pegala blanca was previously unsuspected 
it is gratifying to find such a development to complete that particular 
picture of adaptive phenetic change in the genus Cercyonis. 


Description of the Habitat 


This butterfly inhabits the swampy alkaline area around the Dufurrena 
Kanch sub-headquarters of the Charles Sheldon Antelope Range in 
tlumboldt County, northern Nevada. Thousand Creek Spring feeds this 

ea in a former lake basin surrounded by lava flows. The wet area 


VOLUME 26, NUMBER 3 143 


Fig. 7. The egg of Cercyonis pegala blanca Emmel & Mattoon: (a) dorsal end, 
showing micropylar region; (b) ventrolateral view; (c) dorsolateral view. 


vegetation is dominated by lush grasses and willows, while wild rye, 
Elymus cinereus, dominates the slightly drier adjacent terrain. The latter 
species of tall bunch grass is used as a roosting site for Cercyonis pegala 
blanca, while one or more of the shorter grasses and sedges in the wet 
areas are apparently used for oviposition. These latter grasses include 
Eleocharis palustris, Juncus balticus, J. lescurii, and Beckmannia syzi- 
gachne, and the sedge Carex nebrascensis. Aside from the springs area 
the vegetation dominating this arid basin is sagebrush (Artemisia con- 
fertifolia and A. truncata). 


Life History 


Living females collected in August 1970 were induced to oviposit in 
the laboratory of S. O. Mattoon at Chico, California, and the eggs were 
maintained and larvae reared on commercial-brand blue grass in an out- 
door greenhouse there. Techniques were as described by Emmel (1969) 
in general. All measurements and descriptions of the stages are by the 
second author. 

It is particularly noteworthy that the life history of this Nevadan 
Ceryonis pegala subspecies has only five larval instars (at least under 
our experimental conditions ), whereas Colorado and coastal California 
Ceryonis pegala boopis exhibited six larval instars under constant- 
temperature conditions (Emmel, 1969). We plan to investigate further 
this variation in instar number. As in all other Cercyonis, this new sub- 
species diapauses and overwinters in the first larval instar; when the 
larvae hatch several weeks after oviposition, they crawl to the base of a 
grass clump and enter diapause without feeding. 


EGG. SHAPE: Egg 1.1 mm in height, 1.0 mm maximum width; somewhat conical, 
broader and flatter at base with top more rounded. Adorned laterally by numerous 
vertical ridges (approximately 19) which intersect and traverse approximately four 
progressively smaller ridges approaching the top of egg, producing a somewhat 
scalloped appearance. The vertical ridges terminate basally just below the widest 


144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


a 


Wi 


YY 


Fig. 8. The five larval instars of Cercyonis pegala blanca: (1) first, (2) second, 
(3) third, (4) fourth, and (5) fifth instars. Head capsule (a), dorsal view of larva 
(b), and lateral view of larva (c). 


portion of the egg and do not traverse another series of similar but more numerous 
basal concentric ridges. COLOR: White at oviposition, turning tan within three 
days, and lightly marked between the ridges by a sparse mottling of irregularly sized 
orange-brown spots and splotches during later development. 

LARVAL STAGES. First Instar. HEAD CAPSULE: Average width, 0.64 mm, 
height, 0.65 mm (20 larvae). Ground color purplish brown (light brown after 
ecdysis). Surface retiform, depressions between ridges more darkly pigmented. 
Antennae (in all instars) light amber, basal area green, distal extremity becoming 
darkened. Labial and maxillary palpi translucent, darkened distally. Mandibles 
translucent, cutting edge with broad margin of black. Labrum translucent, notch 
black. Facial suture margins darkened. Ocelli black. Head capsule setae sparse, 
piniform, all oriented anteriorly then most curving slightly ventrally (in all instars), 


VOLUME 26, NUMBER 3 145 


LONGITUDINAL 9 
STRIPES 


DORSAL 


ADDORSAL 


SUBDORSAL 


SUPRASPIRACULAR 


SUBSPIRACULAR 


SUBVENTRAL 


oe Alls 
Blend [PEK 
Ot, Lat a 
Say Pee: 


, ZA 2-5 # 2 iE ae a ey 
Zak \ ee ee ee : 
* Sf: Djs? Se od J: Fj—=za2 ~~ S 


Fig. 9. Map of setal arrangement and pigmented stripes described in text for 
first-instar larva of Cercyonis pegala blanca. 


setae long (averaging three times the width of large ocelli), translucent, granular 
(in all instars), darkened at juncture with head with minute purplish brown chalazae, 
and in a constant bilaterally paired arrangement. TRUNK: Average length 5.0 mm. 
First abdominal proleg segment width, 0.85 mm (20 larvae). Ground color of 
diapausing larva light tan to light brown, turning grass green within 24 hours after 
feeding. Body segments annulated, five annuli per thoracic and six per abdominal 
segments, marked by dorsal, addorsal, subdorsal, supraspiracular, and subventral 
purplish brown longitudinal stripes, and laterally by a prominent greenish white 
subspiracular stripe. Randomly distributed whitish green pigment spots showing 
through integument are more concentrated bordering stripes. Thoracic legs faint 
purplish brown, darkened distally. Claw darkened at juncture with tarsus. A pair 
of spinose, opposed setae extend beyond claw in all instars. Abdominal prolegs grass 
green; crochets in a uniordinal uniserial latero-series, colored brownish black. The 
anal prolegs with purplish brown pigment patch (sometimes obscure) on lateral 
aspect. Spiracular openings on distal extremity of rusty orange, globe-tipped stalks. 
Globe with minute transverse ridges running from opening to stalk. Body armed 
bilaterally by five longitudinal rows of long spiniform setae, some knob-tipped on 
cephalic margin of first abdominal segment. Setae barely discernable without 
magnification. Setal arrangement (Fig. 9) is as follows. Row I: Located dorsad 
of the addorsal stripe. Comprised of posteriorly-oriented, horizontally-aligned pairs 
on each abdominal segment and one seta anteriorly oriented on each thoracic segment, 
except headed by a group of four (sometimes knob tipped) on cephalic margin of 
first thoracic segment. Row II: Single, anteriorly oriented, spiniform, except paired 
and sometimes knob tipped on first thoracic segment (row restricted to thoracic 
segments only), located on addorsal line. Row III: Ventrad of subdorsal line. All 
spiniform, single, anteriorly oriented except posteriorly oriented on abdominal seg- 
ments nine and ten, and paired on caudal and first thoracic segments. Row IV: In 
subspiracular line, all spiniform (except sometimes knob tipped on first thoracic 
segments), paired, and posteriorly oriented except on thoracic segments two and 
three. Row V: Located subventrally at leg bases. All posteriorly oriented and paired, 
except singular on thoracic segments two and three and abdominal segments seven, 
eight, and nine. Five long, spiniform, ventrally or posteriorly oriented setae arise 
from within the perimeter of anal proleg color patch. Caudal segment blunt. Long, 
spiniform, posteriorly oriented, bilaterally paired setae project from conical purplish 


146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


brown chalazae on the posterior of caudal segment. The longest pair arise dorso- 
laterally and appear to extend caudally the tapering body line. 

Second Instar. HEAD CAPSULE: Average width 1.01 mm, height 0.90 mm 
(20 larvae). Ground color grass green with faint purplish brown cast (light brown 
after ecdysis). Capsule surface features as in first instar, except surface reticulations 
more irregular, the depressions not darkened. Facial sutures only faintly darkened 
with two dark spots on attachment margin of labrum. Head capsule setae more 
numerous, minute, strongly knob tipped in cephalic region, averaging one-half the 
width of large ocellus, becoming more spiniform and longer around frontal portion. 
Setae not darkened at juncture with chalazae as in first instar. Chalazae white 
(slightly larger than in first instar) and more bulbous. TRUNK: Average length, 
7.8 mm; first abdominal proleg segment width, 1.13 mm. Ground color grass green. 
In second through fifth instars, integument surface textured with a multitude of 
minute circular smooth convex areas producing translucent frosted appearance, more 
pronounced early in instar and lessening as skin tightens. The longitudinal dorsal, 
addorsal, subdorsal, and supraspiracular solid purplish-brown stripes more obscure 
than in first instar. Subventral stripe dashed and purplish brown dorsad of leg bases. 
Subspiracular striple along the somewhat protruding extreme lateral aspect of trunk 
well defined and solid white. Body segments more noticeably annulated. Thoractic 
legs brownish green, tarsal claw and distal half of tarsus darkened. Tibia, femur, and 
proximal half of tarsus amber brown, coxae green. Abdominal prolegs green, crochets 
more numerous. Setae on all leg bases strongly knob tipped from minute green 
chalazae, becoming progressively more numerous and spiniform from smaller chalazae 
distally. The more distal setae sometimes darkened at attachment with chalazae. 
Spiracles with stalk much reduced to absent; often, globe incomplete distally, ex- 
posing hollow interior with spiracular opening at base. Body setae now numerous, 
minute (much shorter than first instar, not discernable without magnification in 
second through fourth instar), irregular but in somewhat poorly defined longitudinal 
rows. Most strongly knob tipped and posteriorly arching from minute green chalazae 
except some anteriorly arching; setae longer, spiniform, darkened at juncture with 
chalazae along cephalic margin of first thoracic segment and along lateral aspect 
of all thoracic segments. Caudal segment now divided into two posteriorly-projecting, 
fleshy-rose-colored, short, conical forks, each covered with posteriorly-oriented, short, 
strongly knob-tipped, slightly rose colored setae. 

Third Instar,. HEAD CAPSULE: Average width, 1.47 mm; height, 1.42 mm 
(20 larvae). Ground color grass green with very slight brownish cast (brown after 
ecdysis). Head capsule very similar to second instar, except setae more numerous, 
chalazae more prominent, bulbous, and white. TRUNK: Average length, 13.5 mm; 
width at first abdominal proleg segment, 2.00 mm. Ground color grass green, 
modified especially on dorsal and lateral surfaces between longitudinal stripes by 
whitish pigmented chalazae of next instar showing through integument. Annulae 
more strongly pronounced, giving the appearance of minute whitish transverse rings. 
Longitudinal striping limited to dorsal, subdorsal, and subspiracular lines. Dorsal 
stripe faint purplish green, central portion almost obscured. Subdorsal stripe area 
lacks underlying pigment spots, thus appearing translucent grass green with faint 
yellowish white stripe along dorsal margin. Subspiracular stripe solid white, well 
defined on the less pronounced lateral aspect of body. Thoracic and abdominal legs 
grass green, setae as in second instar, thoracic legs darkening distally with tarsus 
and claw amber brown. Crochets basically biordinal uniserial lateroseries (an ap- 
parent biserial condition sometimes results from approximately three of the shorter 
central crochets being slightly offset from uniserial line, but not enough to be con- 
sidered biserial). Crochets amber brown, anal proleg series in a biordinal uniserial 
mesal semicircle. Spiracles rusty orange and each comprised of a prominent circular 
peritreme raised along central caudal margin into fleshy incurving nipple-like conical 
vrojection. Peritreme larger, transverse, and eliptical on first thoracic and eighth 


VOLUME 26, NUMBER 3 147 


S 


GRASS 


GREEN TRANSLUCENT 


WHITE 


Fig. 10. Details of shape of head capsule chalazae and setae on (1) first instar, 
(2) second instar, (3) third instar, (4) fourth instar, and (5) fifth instar larvae. 
Stippled area in (1) is light purplish brown; white areas are pearly white unless 
otherwise indicated. 


abdominal segments. Structure of first thoracic spiracle reversed with raised pro- 
jection on cephalic margin. Body setae as in second instar except more numerous 
and randomly distributed, darkened setal bases not as extensive except ventrally 
where spiniform setae arise from more bulbous white chalazae along leg bases and in 
groups on corresponding segments lacking legs. Caudal segment as in second instar. 

Fourth Instar. HEAD CAPSULE: Average width, 2.21 mm; height, 2.27 mm 
(20 larvae). Ground color grass green (green after ecdysis). Surface with reticula- 
tion of ridges connecting chalazae. Ridges less pronounced ventrally. Setae more 
numerous than in third instar; blunt and spiniform to knob tipped dorsally, becoming 
predominantly knob tipped ventrally, all arising from slightly more conical, white 
chalazae. Facial sutures more heavily margined brown, otherwise head capsule as in 
third instar. TRUNK: Average length, 21.9 mm; first abdominal proleg segment 
width, 2.94 mm (20 larvae). Ground color grass green. Body setae mostly blunt 
and spiniform dorsally, knob tipped laterally becoming shorter and strongly knobbed 
ventrally, all from minute conical green chalazae. Annulations strongly pronounced, 
integument and setae producing whitish pubescence. Ground color modified late in 
instar dorsally and laterally, appearing yellowish white due to coloration of developing 
chalazae beneath integument. Longitudinal striping as in third instar, except dorsal 
stripe grass green, sometimes faintly bordered yellowish white. Subdorsal stripe 
lightly defined yellow to greenish white, sometimes bordered dorsad with grass 
green. Legs as in third instar, except setae are knob tipped on leg bases, becoming 
progressively more spiniform terminally and in corresponding areas on segments 
without legs. Some chalazae mostly on lateral aspect of thoracic leg bases darkened 


148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 11. Dorsal and ventral views of the three pupal color phases of Cercyonis 
pegala blanca: (a-c) black-and-white form (d-e) green form; (f-g) intermediate 
form. 


terminally. Abdominal proleg crochets a uniordinal biseria] lateroseries; crochets on 
anal proleg a uniordinal biserial penellipse opened caudally. Spiracles as in fourth 
instar except larger with raised projection smaller in proportion to diameter of 
peritreme. Caudal segment as in third instar except extreme posterior extremity of 
segment between forks adorned with small patch of purplish brown microspines. 

Fifth Instar. HEAD CAPSULE: Average width, 3.21 mm; height, 3.35 mm (20 
larvae). Ground color grass green (splitting along frontal suture with cast skin 
remaining attached, brown after ecdysis). Surface and setae as in fourth instar, 
except setae slightly longer, spiniform along dorsal cephalic margin. Chalazae large 
and bulbous, pearly white, sometimes becoming green basally. Most longer setae 
have bases expanded into a collar at attachment with chalazae. Mandibles with 
blackish brown color of cutting edge margin comprising one-third of mandible, central 
one-third cream colored, outer one-third translucent. Labrum translucent to cream 
colored distally, notch usually darkened. Bases of maxillary and labial palpi green, 
otherwise amber brown. Facial sutures heavily margined amber brown with brownish 
urved line connecting bases of in line ocelli. TRUNK: Average length, 33.2 mm 
20 larvae; from 29.5 to 38.0 mm). Average width at first abdominal proleg segment, 
> mm. Ground color grass green modified by strongly defined segment annula- 
id frosted integument surface, and by pubescence of the predominantly spini- 


VOLUME 26, NUMBER 3 149 


form setae (barely discernible without magnification), to give trunk a whitish over- 
cast, transversely ringed appearance. Chalazae greenish white and conical. Dorsal 
and lateral areas of trunk between longitudinal striping sometimes yellowish green, 
resulting from yellowish pigment spots under integument. Longitudinal striping as in 
fourth instar, except white of subdorsal and subspiracular stripes more pronounced. 
Abdominal prolegs as in fourth instar, except setae all long and spiniform arising 
from conical chalazae. Chalazae darkened distally along lateral aspects of legs. 
Crochets uniordinal and ttriserial (third row sometimes very limited). Anal proleg 
crochets form uniordinal triserial penellipse opened caudally. Spiracles all larger, 
transverse, and eliptical, otherwise as in fourth instar. Caudal segment setae on forks 
spiniform and a translucent rose color. Chalazae have distinct rose colored collar at 
juncture with setae. Amber brown microspines cover most of caudal extremity 
between forks. 

PUPA. GREEN FORM: Entirely grass green, unmarked by any trace of black 
and white pattem. BLACK AND WHITE FORM: Pupal case totally marked by 
black and white longitudinal stripes. Around abdomen, striping evenly spaced and 
widest at thorax tapering to cremaster. The stripe located addorsally continued 
anteriorly adjacent to the dorsal meson white stripe terminating at the head. The 
subdorsal black stripe follows the dorsal margin of wing case to head, and the 
ventral meson black stripe divides at juncture of wing cases following the inner 
margin of antennae; they are nearly reconnected by a transverse black mark which 
crosses the head anterior to the crest of the eyes. Wing cases marked with irregular 
black dashes oriented obliquely to, and some appearing as extensions of, the 
longitudinal abdominal striping. INTERMEDIATE FORMS: Ground color of all 
pupae green. The extent to which black and white markings overlay the green 
highly variable, the green often being incompletely masked; thus all gradations of 
black, white, and green occur between the two extreme color forms. Black and 
white, however, modified simultaneously in that both tend to increase and decrease 
together. Occasionally, black pigment totally absent, exposing the green ground 
color which is still silhouetted by a remaining trace of white. 


ACKNOWLEDGMENTS 


This research has been supported in part by NSF Grant GB-8442 and 
the Division of Sponsored Research, University of Florida. We would 
like to thank in particular Research Horticulturist Robert L. Smith, of the 
U. S. Plant Introduction Station, Chico, California, for generously allow- 
ing the use of photographic equipment and darkroom facilities for photog- 
raphy of the life history stages. 


LITERATURE CITED 


EMMEL, T. C. & J. F. Emmen. 1969. A new subspecies in the Cercyonis meadi 
group (Satyridae). J. Lepid. Soc. 23: 161-164. 

1971. An extraordinary new subspecies of Cercyonis oetus from central 
Nevada (Lepidoptera: Satyridae). Pan-Pacific Entomol. 47: 155-157. 
EMMEL, T. C. 1969. Taxonomy, distribution and biology of the genus Cercyonis 
(Satyridae). I. Characteristics of the genus. J. Lepid. Soc. 23: 165-175. 


150 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


A NEW SUBSPECIES OF LYCAEIDES ARGYROGNOMON 
(LYCAENIDAE) FROM THE EASTERN CANADIAN FOREST ZONE 


Joun H. MASsTERs 
P.O. Box 7511, St. Paul, Minnesota 55119 


It has been recognized for some time that the populations of Lycaeides 
argyrognomon from Southeast Manitoba, Ontario, Minnesota, Wisconsin 
and (probably ) the upper peninsula of Michigan represent an undescribed 
subspecies distinct from scudderi (Edwards ) with which they are usually 
allied. Griewisch (1953) collected long series of L. argyrognomon in 
Oconto and Marinette Counties, Wisconsin 1-15 July 1952, sent samples 
to Vladmir Nabokov at Harvard for determination, and quoted Nabokov’s 
comments on them as follows: 

“The Minnesota thing, which I described and figured, but did not 
name, because of scantiness of material (Bull. Mus. Comp. Zool., 1949, 
p. 505, Pl. 5, fig. 54, male, Pequot, Minn.) is a subspecies of argyrognomon 
(Bergstrasser, Tutt), which I now think is sufficiently distinct from the 
Canadian (north of 50°) argyrognomon scudderi (type loc.: The Pas, 
west of Winnipeg L., Manit.) to warrant a new subspecific name for it. 
It is this form that your specimens belong to, and you should be con- 
gratulated on establishing the interesting Wisconsin range of argyro- 
gnomon. It comes very near to a point where it should fly together with 
melissa samuelis Nab.” 

Griewisch later supplied a long Marinette County series of L. argyro- 
gnomon to Frank Chermock of Baltimore, Maryland. Chermock intended 
to describe these as an argyrognomon subspecies and distributed a num- 
ber of them as “paratypes” bearing a manuscript name. Chermock and 
Griewisch are both deceased and for the last 20 years Nabokov has 
vacillated on doing further work with Lepidoptera. With no other heir 
apparent to carry on their work, I have assumed the responsibility, with 
this paper being the result. 

The collections of Lepidoptera at my disposal probably contain more 
ample material in Lycaeides argyrognomon from the key areas of Mani- 
toba, Minnesota and Wisconsin than any others. After an examination 
and study that has lasted three years, I find myself in full agreement with 
the earlier conclusions that the Eastern Canadian Forest Zone population 
of Lycaeides argyrognomon from southeast Manitoba, western Ontario, 
northeast Minnesota and northern Wisconsin is sufficiently distinct from 
L. argyrognomon scudderi to deserve it’s own subspecific name. I take 
pleasure in naming it in honor of Dr. Nabokov, who first recognized its 


VOLUME 26, NUMBER 3 liso 


distinctness and whose papers on Nearctic Lycaeides (1943, 1944, 1949) 
have provided a background to make this description possible. 


Lycaeides argyrognomon nabokovi J. Masters, new subspecies 


Male (Fig. 1): The same general appearance associated with all populations of 
Lycaeides argyrognomon but of a larger size and slightly more purplish color than 
any other. The expanse of one forewing (base to apex) 14 to 17 mm (average 16 mm). 

Upperside (Fig. 1): Ground color of wings a vivid violet blue. Melanic margins 
(0.75 to 1.25 mm wide) not sharply defined on their basal margins. White fringes 
wide and well pronounced on fresh specimens. Darkened melanic spots, centered 
in the hindwing cells just basad of the melanic margin, present in some individuals. 
Scales along forewing veins darkened, especially limbally which gives the impression 
that their outer ends are swollen. 

Underside (not figured): Markings essentially like the female, but with macules 
not as pronounced and with the orange and metallic elements not as vivid. 

Genitalia (not figured): Typical of L. argyrognomon. Length of forearm of 
falx = 0.43 mm. Length of humerulus of falx = 0.32 mm. Length of uncas lobe = 
0.33 mm. (Average values for three male paratypes dissected. ) 

Female (Figs. 2, 3): Differs from male in being considerably more melanic on 
upper surfaces, confining the violet-blue coloration to the basal areas. The expanse 
of one forewing (base to apex) 14.5 to 17 mm (average 16 mm). 

Upperside (Fig. 2): The purplish-blue coloration confined to the basal third 
of the forewing and the basal half of the hindwing. Remainder of the wings clouded 
with melanic scales that have little or no contrast with the borders. White fringes 
present on fresh specimens. On the hindwings, 4 to 7 oval melanic eye-spots present, 
centered in the cells near the outer margins, and these usually accompanied by an 
orange lunule basally. These macules especially prominent in cells CU: and Cup. 

Underside of forewings (Fig. 3): The ground color varies from a smoky grey 
to a bright white—tending to be lighter and whiter than in L. argyrognomon 
scudderi. A narrow melanic margin, thickened at the vein terminals, often extends 
around the apex to the costal margin. Two rows of submarginal spots, elongated or 
slightly crescent shaped, although never as dark or as well defined as the post- 
median spots. Submarginal spots often surrounded by a tint of orange. A row of 
six very well defined post-median spots, which vary greatly in shape but always 
baroque and non-uniform in appearance. Post-median macule in cell Cu: always 
largest and conspicuous by its shape and position, being elongated and displaced 
basally. Post-median spots ringed in white, but contrast with the background color 
slight and effect not nearly so conspicuous as in L. argyrognomon scudderi. One 
additional macule on the forewing, a large oval or slightly crescent shaped spot at 
the end of the discal cell. 

Underside of hindwings (Fig. 3): Ground color as on the forewings, with rela- 
tively little indication of the blue-green overcast that is usually conspicuous in the 
basal area with other subspecies of L. argyrognomon. Margin separated into triangu- 
lar shaped spots at each vein terminus with only a thin line connecting them. Two 
rows of submarginal spots surrounded by an orange umbra which, on some speci- 
mens, coalesce into an orange band. Inner row characterized by a pronounced 
crescent shape while the outer row possesses metallic green centers basally, most 
pronounced in cells 2V, Cus, Cu: and Ms. A widely disjunct row of eight post-median 
macules, more regular in both shape and size than those on the forewing. Macule 
at the end of the discal cell crescent shaped and relatively thin and narrow. Three 
additional macules in the submedian area. All of the median macules ringed with 
white; however, hardly discernable on some specimens. 

Holotype male: Along Forest Road 122 in section 24 of Township 56 N., Range 
11 W., Lake County, Minnesota (16 July 1966), J. H. Masters, collector. Type 


152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-3. Lycaeides argyrognomon nabokovi new subspecies: 1, male paratype, 
dorsal view; 2, female paratype, dorsal view; 3, female paratype, ventral view. All 
three specimens captured at “McNair,” Lake County, Minnesota, 8 August 1967 by 
J. H. Masters. Approximately 11% times natural size; millimeter scale at lower left. 


wise. 


locality 13 miles north of Two Harbors, near the former refueling stop of “McNair 
on the DM & IR railway, now a part of The Superior National Forest. Expanse of 
forewing 16 mm. To be deposited in Carnegie Museum, Pittsburgh. 

Allotype female: Same data as the holotype. Expanse of forewing 16 mm. To 
be deposited in Carnegie Museum, Pittsburgh. 

Paratypes: 58 males, 34 females, same locality as holotype, various dates, July 
and August 1966, 1967, 1968 and 1970, J. H. Masters and William A. Bergman 
collectors. Paratypes will be deposited in the collections of the United States Na- 


tional Museum, American Museum of Natural History, Los Angeles County Museum, 
Museum of Comparative Anatomy at Harvard, Manitoba Museum of Man and 
‘Nature, and University of Minnesota. The remainder will, for the present, remain 


n the author’s and W. A. Bergman’s private collections. 
Other localities: Paratypes were restricted to specimens from the type locality. 


VOLUME 26, NUMBER 3 L538 


Other populations that I consider belonging to this subspecies include specimens 
from the following localities: Minnesota: Echo Trail, near Ely, Saint Louis County; 
Gunflint Trail, T.64 N., R.l1 W., Cook County; near Pengilly, Itasca County; vic. 
Pequot Lakes, Crow Wing County. Wisconsin: near Crivitz, Marinette County; 
Wambee Lake Area, Oconto County. Manitoba: White Shell Provincial Park; 
Wanipigow River, near Bissett. Ontario: The following localities are cited by Riotte 
(1971) for Lycaeides argyrognomon; Armstrong, Favourable Lake, Minaki and 
Nakina; all of which undoubtedly represent this subspecies. 

Lycaeides argyrognomon nabokovi is one of eleven Nearctic subspecies 
which include anna (Edwards ), ricei (Cross), lotis (Lintner ), alaskensis 
(Chermock), scudderi (Edwards), aster (Edwards), ferniensis (Cher- 
mock ), atrapraetextus ( Field ), sublivens Nabokov, and longinus Nabokov. 
Descriptions and distributional data for all of these can be found in 
Nabokov (1949). DosPassos (1964) also includes kodiak (Edwards) and 
empetri (Freeman) as subspecies under argyrognomon. Nabokov con- 
sidered kodiak as a taxa of uncertain status, probably not belonging to 
Lycaeides; Brown (1970) considered it as a subspecies of Plebejus 
saepiolus. Nabokov considered empetri (type locality Cape Breton Is- 
land) to be a synonym of aster. 

In the past, populations of both Lycaeides argyrognomon nabokovi and 
Lycaeides melissa samuelis have been confused with L. scudderi. The 
specimens figured by Holland (1931, plate XXX, figs. 48, 49) as scudderi 
are actually samuelis. Lycaeides argyrognomon scudderi (type locality 
“between Lake Winnipeg and Cumberland House,” Manitoba; Brown, 
1970) is now restricted to regions west and north of Lake Winnipeg— 
it occurs as far south as Riding Mountain. The barrier between scudderi 
and nabokovi is Lake Winnipeg and the Red River Valley (the area 
formerly occupied by glacial Lake Agassiz), which suggests that the two 
subspecies have been isolated since late in the Pleistocene. Since nabokovi 
is more likely to be confused with scudderi than with any other subspecies 
in the future, it seems pertinent to summarize the differences between 
the two: 

Ground color of ventral surfaces tends to be lighter and whiter in 
nabokovi, which renders the white rings, circling the post-median 
macules, less distinct. Basad blue-green dusting on ventral hindwing 
is much less evident on nabokovi. The post-median macules on the ventral 
forewing of nabokovi are larger and much more irregular in shape and 
position than they are in the case of scudderi. In the case of scudderi, 
they form an almost uniform row of almost equal-sized spots. The 
elongated and distorted macule in cell Cu, is usually diagnostic of 
nabokovi; in the case of scudderi, this macule is much more rounded 
and regular in appearance. This specimen selected by Brown (1970) 
as the neotype for Lycaena scudderi is an exception to this, having the 


154 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


macule in Cu; elongated very much like nabokovi. I have not noted this 
condition, however, in 30 specimens of scudderi that I have examined from 
Riding Mountain, Duck Mountain and The Pas, Manitoba. 

The Lycaeides argyrognomon aster/empetri populations, with which 
nabokovi more than likely intergrades at some point, occur in Newfound- 
land, Nova Scotia, Labrador, Cape Breton Island and northeastern 
Quebec. Brown (1970) confines aster to the Avalon Peninsula of Quebec 
and relegates populations from other parts of this region to empetri. The 
combined aster/empetri populations are small and pale with small, round 
and well separated macules in the post-median series. The post-median 
macules are the same size or smaller than the marginal macules, whereas 
in the case of nabokovi they are significantly larger. The aster/empetri 
populations are characterized by considerable blue-green dusting basally 
on the ventral hindwings, and by very little blue coloration dorsally in the 
female. There is a probable zone of intergradation between these two 
forms in eastern Ontario or Quebec. 

The only species occurring in the same regions with nabokovi, with 
which it might be confused, is Lycaeides melissa samuelis. These two 
cannot be considered as sympatric, however, as nabokovi is confined 
to Canadian Zone forests while samuelis is confined to open prairie 
regions. The ventral wing margin in Lycaeides melissa has a narrow but 
solid and straight terminal line; while in Lycaeides argyrognomon, this 
line is usually broken into triangular shaped spots at the vein terminals. 
If this fails to provide a definitive separation of the two species, dissection 
of the male genitalia will. In L. melissa the length of the forearm of the 
falx is twice as long as the length of the humerulus of the falx while in 
L. argyrognomon it is but one and a half times as long. 


LITERATURE CITED 


Brown, F. M. 1970. The types of Lycaenid butterflies named by William Henry 
Edwards. Part III. Plebejinae. Trans. Amer. Entomol. Soc. 96: 353-433. 

bosPassos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. Soc. 
Mem. 1. 145 p. 

Griewiscu, L. 1953. Lycaeides argyrognomon in Wisconsin. Lepid. News 7: 54. 

Hoittanp, W. J. 1931. The Butterfly Book. Revised edition. Doubleday, Garden 
City, New York. 424 p. 

Nasoxov, V. 1943. The nearctic forms of Lycaeides Hub. (Lycaenidae, Lepidop- 
tera). Psyche 50: 87-99. 

———. 1944. Notes on the morphology of the genus Lycaeides (Lycaenidae, 
Lepidoptera). Psyche 51: 104-138. 

—_—_—_ . 1949. The nearctic members of the genus Lycaeides Hubner (Lycaenidae, 
Lepidoptera). Bull. Mus. Compar. Zool., Harvard 101: 479-541. 

uuorre, J.C. E. 1971. Butterflies and skippers of northern Ontario. Mid-Continent 
Lepid. Ser., Saint Paul 21: 1—20. 


VOLUME 26, NUMBER 3 55 


THE ECOLOGY AND ETHOLOGY OF THE TROPICAL 
NYMPHALINE BUTTERFLY, VICTORINA EPAPHUS. 
I. LIFE CYCLE AND NATURAL. HISTORY 


ALLEN M. YOUNG 
Department of Biology, Lawrence University, Appleton, Wisconsin 54911 


This paper is the first in a series on the neotropical butterfly, Victorina 
(Amphirene) epaphus Latreille in central Costa Rica. Despite its wide- 
spread occurrence throughout the montane and upper premontane tropi- 
cal wet forests of Costa Rica, little is known about its biology. According 
to Seitz (1924) the life cycle of this Central American species is unde- 
scribed. This paper describes the life cycle of this species, and brings 
together various observations on its ecology and ethology, to be explored 
in depth in later papers. Data on larval host plant specificity, oviposition, 
and survival of immatures in natural populations also are given. 


METHOpS 


V. epaphus was studied at Cuesta Angel de Sarapiqui (Heredia Province, 
1000 m elev.) at various times from February 1969 through July 1970. 
Observations were made at the bottom of a steep ravine, in relatively 
undisturbed, primary-growth forest. The study of V. epaphus consisted 
of three approaches: larval host plant studies, field studies, and labora- 
tory studies. 


Larval Host Plant Studies 


The study site is near an extensive strip of the larval host plant, Ruella 
tubiflora H.B.K. var. tetrastichantha (Lindau) Leonard (Acanthaceae ) 
which borders the Rio Sarapiqui and a foot trail at the bottom of the 
ravine. This plant, (a first record for V. epaphus) is an understory form, 
usually between 15-40 inches in height, and confined to very wet soil. 
At Cuesta Angel, it grows in continuous patches along with Gynandropsis 
pulcherrinia Standl. (Capparidaceae), another understory plant of simi- 
lar height and growth form, which is a major larval host plant of Itaballia 
caesia (Pierinae), whose life cycle will be summarized later. Unlike the 
spatial distribution of larval host plants of tropical butterflies belonging 
to the genera Morpho (Young, in prep.), Parides and Battus (Young, 
1972a, b, c), the larval host plant of V. epaphus is characterized by rather 
extensive homogeneous patches, permitting a concentration of reproduc- 
tive effort (Labine, 1968) over small portions of the habitat. 


156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The larval host plant of V. epaphus was in active vegetative growth 
during the study period and floral structures were absent despite the low 
degree of seasonality known for montane tropical wet forests (Richards, 
1952; Roth & Bifano, 1971). Extensive searching revealed that it is the 
only host plant of this butterfly at Cuesta Angel. 


Field Studies 


Field studies consisted of (1) mark-recapture studies on adults, (2) 
analysis of diurnal peak activity periods of adults, (3) observations on 
oviposition behavior, (4) records on the distribution of eggs and larvae 
on individuals of the host plant, including observations on larval ac- 
tivities, and (5) occasional records of predation and parasitism on larvae. 

Mark-recapture study employed a standard sampling procedure in 
which adults were netted on several traversions through the host plant 
patch, marked, and immediately released. This was done at irregular 
intervals throughout the study period, using a marking system modified 
from Ehrlich & Davidson (1960) and in which sexes were differentiated. 
Estimations of adult populations then were made for each day of ob- 
servation, using the Lincoln Index. Eggs, larvae, and pupae also were 
sampled in a standardized manner by walking transects through the 
patch once daily. Individual host plants along the transect were examined 
frequently for eggs and larvae. This sampling technique was very suitable 
since the host plant grew in large homogeneous patches, allowing many 
individuals to be sampled quickly. 


Laboratory Studies 


Broods of eggs were reared, both in the laboratory and out of doors. 
Life stages and developmental time were studied in this manner. Lab- 
oratory cultures were established in tightly closed 8 in by 12 in clear 
plastic bags, prior to each of these receiving several eggs on cuttings of 
Ruella, This method has proven successful with several species of tropical 
butterflies (Young, 1972a, b, c, d, e). Various aspects of larval behavior 
were also noted, such as diurnal pattern of peak feeding, gregariousness, 
and aggressiveness. Laboratory rearing was undertaken to explore also 
the feasibility of using this widespread species for estimation of physiologi- 
cal life tables (Istock, 1970; Young, 1972e) to be eventually compared 
with realized life tables of natural populations. Other interests focused 
upon the suitability of such a widespread tropical species for electro- 
phoretic studies of enzyme variability as a function of geographic and 
opographic distributions of populations. 


VOLUME 26, NUMBER 3 Sy 


RESULTS 
Description of Life Cycle 


The eggs are laid singly in small, loose clusters on apical, unfolding 
leafbuds of Ruella (Fig. 1,A), and occasionally on older leaves. Anywhere 
from 1-6 of the dorso-ventrally flattened (diameter of 0.9 mm) ribbed 
eggs are laid by an individual female during one visit to a plant. When 
first deposited, each egg is pale green but rapidly (10-15 min.) becomes 
dark green between the yellowish ribs (Fig. 1A). Eggs are placed care- 
fully on leaves of Ruella and “errors” in oviposition (Dethier, 1959a, b) 
have not been seen. 

The first-instar larva devours its empty egg shell. The young larva 
then moves to the ventral side of an apical, unfolding leaf to begin feed- 
ing. The first-instar larva is jet black except for the first segment behind 
the head, and the two most posterior segments, which are a dull-orange 
color. The head is shiny jet black, bearing two long slightly barbed 
spines; shorter branched spines are present on all other body segments. 
With the exception of the single pair of head spines, all remaining 
spines are ringed with white at their bases. Each body segment, excluding 
the first behind the head, bears three pairs of these spines. The second- 
instar larva is about 7 mm long. The third instar is similar (Fig. 1,C) 
and about 16 mm long when molting to the fourth instar. 

A drastic change in appearance occurs in the fourth instar. The larva 
is now between 29-35 mm long and the head is still shiny jet black, but 
the body is velvety maroon. The head spines are still long. All other 
spines are now bright yellow and more highly-branched (Fig. 1,D). 
The bases of all body spines are bright yellow. The final, fifth-instar 
larva (Fig. 1,E) is almost identical to the fourth instar except that the 
head spines and the most lateral pair of spines along the body axis are 
red. The expanded basal spots no longer are associated with each spine. 
Of the three pairs of spines on each body segment present in all five 
instars, only the most dorsal pair remains bright yellow; the next pair has 
the lower half a dull yellow and the upper half light orange; the third, 
most lateral pair of spines are entirely red. All spines remain very highly 
branched. The mature larva is between 45-55 mm in length. 

Pupation typically occurs on the undersides of older leaves near the 
ground. The pupa (Fig. 1,F) is about 30 mm long, and pale, translucent 
green, although flecked with tiny black spots arranged in longitudinal 
rows on the thorax and abdomen. There are two prominent orange-based 
black spines originating dorsally from the first and second abdominal 
segments. The head region is slightly forked anteriorly, with projections 
being green below and black above. 


158 


Lip, 
Vii 

Z Vii, 
this 


Hag, 


instar, (D) fourth instar, (E) fifth instar, 


Life cycle of Victorina epaphus: 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


UM iiiig 


(A) eggs, (B) second instar, (C) third 
and (F) pupa. 


VOLUME 26, NUMBER 3 159 


<a 
4 


: ae 


Fig. 2. Adult Victorina epaphus. Dorsal views of female (above) and male 
(below). Mean wingspan length is 7.0+ 3.8 cm (N = 34) with females slightly 
larger than males. 


A lack of sexual dimorphism characterizes the adults of V. epaphus 
(Fig. 2). However, the female generally is larger than the male (by 3-6 
mm) and the cream-colored band separating distal orange areas and 
proximal brown areas of both wings is broader. It is very difficult to 
distinguish sexes in the field. 

The developmental time for V. epaphus (summarized in Table 1), is 
slightly more than one month. Data on survivorship of larvae and pupae 
in the laboratory indicate that this insect can be reared successfully under 
artificial conditions (Table 1). 

Development within egg clusters is highly synchronous, in the field and 
laboratory, with virtually no individuals completing postembryonic de- 
velopment markedly later than other individuals. Breeding is continuous 
throughout the year. 


160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. The developmental time (days) for Victorina epaphus under laboratory 
conditions.* 


Instar Instar Instar Instar Instar 

Egg 1 2; 3 4 5 Pupa Total 
Mean 6.85 4.97 5.05 4.14 2.20 Dimlesy 10.35 Onl 
Sree ae 1k) ae JL ae (Us) +0.9 a 10) ae (4 as)5) ae 4 
Range 6-9 46 4-7 3-6 D5) 2—4 9-11 32-38 
No. of 
Individuals 57 54 54 54 51 48 48 48 
No. which 
Died 0 3 0) 0 3) 3 0 9 


* Times estimated from a total of 57 eggs, representing 10 clusters deposited on 10 individual 
foodplants by a single female within an 18 minute period (1440-1458, CST) on 29 Jan. 
1970 near Cuesta Angel, Heredia Province. All cultures were kept in large (8 X 12 in.) clear 
plastic bags in one large room in San Jose, Costa Rica. Eggs were collected at Cuesta Angel 
immediately following oviposition and brought directly to San Jose for culture. 


Population Size and Mortality 


Estimations of adult population sizes are summarized in Table 2 along 
with frequencies of immature stages. A striking result of this survey is 
that both egg and adult numbers remain constant throughout the year 
despite fluctuations in numbers of larvae and pupae. 

Survivorship from egg through pupa in the field high (Table 3). Pre- 
dation and parasitism on eggs and larvae are probably low. However, the 
large fluctuations in larval numbers and the diminished numbers of pupae 
(Table 2) suggest operation of undetected mortality factors. I have 
consistently failed to rear tachinid or braconid parasites from larvae col- 
lected in the field. 


Taste 2. Weekly frequencies of adults and immature stages of Victorina 
epaphus at Cuesta Angel, Heredia Province, Costa Rica from early February through 
early April, 1969. 


Number of Individuals Seen Per Week 


_Stages_ a Feb. 3 Feb.10 Feb. 17 Feb. 24 Mar.10 Mar.17 Mar.24 Mar. 31 Apr. 7 
Adults 44. 40 43 26 38 59 63 44 49 
Kegs 134 168 159 103 97 107 115 78 iss 
Larvae 109 Ihaliss 1eA6) He 68 89 100 52 102 


Pupae oe, 18 20 a 20 31 18 9 ONS) 
Total 309 341 345 209 25 286 296 NGS 307 


Adult dae a Beare on cCapture-mark-recapture estimates. No observations were made 
the week of March 8. 


VOLUME 26, NUMBER 3 161 


TABLE 3. A summary of some measurements of population parameters of the 
tropical nymphalid, Victorina epaphus.* 


Egg No. of No. eggs No. eggs Larval Pupal Overall % 
Cluster eggs per killed by failing to mor- mor- survivor-= 
No. cluster predation hatch tality tality ship 
1 i 9) 0 1 0 13.8% 
2 5 3 0 0 0 40.0% 
3 8 4 0 1 1 25.0% 
4 6 0) 0 3 0 30.0% 
5 6 0 0 2 0 20.0% 
6 3) 2 0 2 0 20.0% 
fh 0 2 0 3 0 29.0% 
8 8 0 Ih 5 I 12.5% 
9 10 I 0 6 0 30.0% 
10 4 4. - _ 0 0.0% 
1g) 6 0 II 3 0 33.4% 
12 5 0 0 p) 0 60.0% 
1s) ue 3 0 3 it 0.0% 
14 6 Yh 0 2 0 33.4% 
15 6 2 0 1 Ih 33.4% 
16 5 2 0 1 0 40.0% 
1g, 5 2 0 2, 0 20.0% 
18 8 1 0 3 ] 37.5% 
19 6 (OF 1 Z 0 50.0% 
20 5 0 0 2 il 40.0% 
i 7 3 0 3 IL 0.0% 
22 a 0 0 2, 0 71.4% 
3) 5 2, 0 1 0 40.0% 
24 8 a 0 Dy 0 50.0% 
25 6 1 0 p) 0 50.0% 
26 6 0 0 2 0 66.7% 


* These measurements were made in the field on immatures staked out on various individual 
foodplants within a large rectangular area of forest understory. 


Larval Behavior 


Larvae, although diurnal feeders, are difficult to detect in heavily 
shaded forest understory. These observations suggest that V. epaphus 
is a palatable species which employs a protective strategy of crypsis while 
going through ontogeny. Some genera of Acanthaceae contain alkaloids 
and may be toxic (Irvine, 1961), but it remains unknown whether species 
of the genus Ruella are toxic. While predation on adults by birds and 
lizards in the field has not been observed, laboratory feeding studies of a 
related species, V. steneles, indicate non-toxicity (Brower & Brower, 1964). 
But in Costa Rica, larvae of this latter butterfly feed on Justicia in the 
Acanthaceae (Young, 1972e). They invariably feed from the ventral leat 
surface and are therefore usually concealed from potential predators 
foraging from above. Presumably in the dark forest understory, the larvae 
would be difficult to detect by predators on the ground. Such larval 


162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


behavior may also lower parasitic attack by hymenopterans and tachinids 
which lay their eggs directly upon the host. 

Larvae are capable of violent jerking movements of the anterior region 
of the body, presumably as a means of defense against predators. How- 
ever, they do not exhibit these movements collectively, as noted for larval 
sawflies (Prop, 1959). The pupae of V. epaphus are also capable of simi- 
lar movements, possibly a means of deterring parasitic wasps and flies 
(Cole, 1959). The numerous spines on all instars and to a lesser extent 
on pupae, coupled with violent jerking movements, may provide an effec- 
tive predator-parasite deterring mechanism. 

Larvae are never gregarious, despite eggs being laid in loose clusters. 
Immediately following devouring of empty egg shells, larvae disperse 
and never reassociate during ontogeny. Crowded larvae in the laboratory 
are aggressive towards one another. Such behavior may permit effective 
dispersion on host plants. However, larvae never leave an individual 
plant to feed on another one nearby. This is true despite the fact that 
Ruella grows in large patches of high density on the forest floor. Ap- 
parently larval densities per individual host plant never reach levels at 
which overcrowding and aggressiveness result in dispersion from the plant. 


Adult Behavior 


Adults of V. epaphus restrict their flying activities to Ruella patches. 
Here the sex ratio remains close to unity, although adult numbers are low 
(Table 2). 

Presumably courtship occurs near host plant patches and females are 
mated within 5 days after eclosion. Of 38 young females examined, only 3 
contained more than one spermatophore. 

Oviposition generally takes place throughout the day and each female 
performs several oviposition sequences on any given day. Observations 
on young marked females indicate that each individual conducts up to 11 
oviposition sequences within a single Ruella patch before resting. Rest- 
ing periods between groups of oviposition sequences are brief. Each 
female deposits a mean number of 5 + 1.4 eggs per plant. 

Adults sometimes feed collectively in small numbers at moldy leaf 
litter on the forest floor, although they probably also forage at flowers. 
At Bajo la Hondura, this species has been observed to feed at odorous 
small growths of fungi on tree trunks. Collective feeding in this species 
is not structured and individual feeding also prevails. Feeding appears 
to be most frequent during the morming hours. 

Adults are not commonly encountered in second-growth plant com- 
inunities, but appear restricted to primary-growth montane forests. Adults 


VOLUME 26, NUMBER 3 163 


fly low, seldom higher than 10 feet off the ground. It therefore is a 
montane forest species concentrating reproductive and feeding activities, 
as well as general flight activity, to understory habitats. 


DISCUSSION 


The above notes on the biology of V. epaphus may best be integrated 
through consideration of an adaptive strategy for inhabiting understory 
regions of primary-growth forests. Several discussions (Pires, Dobzhan- 
sky, & Black, 1953; Slobodkin, 1970) have emphasized the structural com- 
plexity and stability of primary-growth tropical rain forests. Component 
species of such habitats are predicted to have a long evolutionary history 
integrating them into communities (e.g. MacArthur, 1969). The adapta- 
tions of V. epaphus to the forest understory environment will therefore be 
emphasized in this discussion. 

The strategy of ecological adaptation in a butterfly is molded by the 
spatial distribution and numerical abundance of the host plant (Singer, 
1971). Given the patchy distribution of Ruella in montane wet forest 
understory and the relatively large size (high and numerical abundance ) 
of individual host plant patches, some predictions about the ecological 
traits of V. epaphus (and other herbivores of Ruella) can be formulated. 

V. epaphus can be described as a non-seasonal breeder with reproduc- 
tion occurring throughout the year. Ecological studies of tropical butter- 
flies should focus upon the major adaptations of life cycle to 
seasonal contractions or expansions in the amount of larval host plant 
(i.e., by seasonal differences in vegetative growth rates) in a given type 
of plant community, and at different altitudes. For example, V. epaphus 
is predicted to be a non-seasonal breeder since it is an understory species. 
It has been stressed by botanists (e.g. Webb et al., 1967) that the under- 
story component(s) of primary-growth tropical forests provides a buffered 
environment for many organisms, protecting them from the major local 
vagaries of climate experienced by the canopy. 

The understory represents a non-seasonal environment, and species of 
butterflies which breed there may possess non-seasonal patterns of re- 
production. The strategy is in part a function of host plant selection in 
the understory community, and discrimination of young vegetative growth 
zones (unfolding leaf buds or apical meristems) for oviposition. Also, 
first-instar larvae may be physiologically and morphologically adjusted 
for feeding only on the youngest leaf tissues, in response to both physical 
(leaf shape, thickness, toughness) and chemical (quantitative and/or 
qualitative differences in secondary compounds involved in gustatory 
cues) properties of the leaves. Tropical butterflies may therefore fall 


164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


into three major classes of breeders: wet season breeders, dry season 
breeders, or non-seasonal breeders. The more seasonal breeders are pre- 
dicted for typically second-growth species, although understory habitats 
cannot be excluded entirely, especially for tropical dry forests. 

Oviposition should be very precise in V. epaphus, if selection pressures 
are favoring the evolution of very specialized behavior patterns ( Margalef, 
1968). Errors in oviposition are predicted in species having high fecun- 
dities and colonizing episodes, a good example being the second-growth 
butterfly, Anartia fatima (Nymphalinae) (Young, 1972e). Understory 
species such as V. epaphus concentrate the bulk of their reproductive 
effort (Labine, 1968) over a relatively small portion of the available 
habitat containing substantial numbers of the larval host plant. Adult 
vagility, under such conditions, would be low. At least one temperate 
zone species of pierid associated with second-growth plant communities 
experiences density-related dispersal of gravid females, presumably in 
response to increase in local adult population density (Shapiro, 1970). 
High adult vagility has also been measured in a second-growth satyrid 
(Brussard & Ehrlich, 1970). 

On the other hand, studies of Euphydryas editha (Ehrlich, 1961) 
indicate low vagility. In tropical forest understory, butterflies may be 
less prone to dispersal due to reduced visibility resulting from shading 
and dense foliage. But comparisons of adult numbers with numbers of 
pupae (Table 2) suggest some adult dispersal, assuming negligible mor- 
tality. Nevertheless, a high proportion of individuals remain residential 
at the host plant. Such behavior gives breeding a patchy distribution, 
corresponding to the distribution of host plant patches (although not all 
host plant patches need be colonized by V. epaphus). Both factors, in 
conjunction with either low fecundity or high mortality of immatures, 
prevents over-exploitation and local extinctions of populations of V. 
epaphus. 

Oviposition on tiny unfolding leaf buds may lower rates of predation 
or parasitism upon eggs. Eggs are partially concealed from wandering 
predators such as ants and aerial forms such as predatory Hemiptera. The 
unfolding leaves of Ruella appear much lighter in color than older leaves 
to the human observer. It has been demonstrated under laboratory condi- 
tions for various butterflies, that ovipositing females are capable of 
discriminating shades of green, corresponding roughly to age differences 
in coloration of leaves seen in host plants (Ilse, 1941; Vaidya, 1969). A 
similar discriminatory mechanism presumably occurs in V. epaphus, since 
the majority of eggs are deposited on the very young leaves of Ruella. 
\. major selective force favoring such discrimination may be site selec- 


VOLUME 26, NUMBER 3 165 


tion for oviposition where eggs are least likely to be detected. Certainly 
increased ease of feeding for first-instar larvae on tender leaf tissue may 
also be an important factor (Thorsteinson, 1960). 

The marked variability in time of oviposition is attributed to persistent 
cloud cover over these regions for a substantial portion of each day. 
Persistent cloud cover is a major feature of montane tropical forests 
(Myers, 1969; Roth & Bifano, 1971) and plays an important role in the 
adaptation of various organisms to these habitats. Oviposition in under- 
story butterflies occurs over a wide range of hours since light remains at 
low intensity throughout the day. Thus, potential predators of ovipositing 
females such as understory birds would experience equal difficulty all day 
long in detecting their prey. There would be no hindrance to the 
ovipositing butterflies since they rely mainly upon olfactory and not visual 
cues in locating host plants. 

In lowland tropical forests, oviposition is predicted to occur during 
moments of prolonged cloud cover (which usually takes place during 
the afternoon ) since potential predators would be less of a threat during 
periods of reduced and diffused light. Oviposition in such species that 
have experienced a long evolutionary history involving adaptation to 
living in understory, should be lowest during the morning hours, when 
bright sunlight can filter down through the canopy. Such an oviposition 
pattern has been observed in Victorina steneles, a butterfly which is very 
common in tropical wet lowlands (Young, 1972e). 

The forest understory also provides a suitable habitat for the exploita- 
tion of crypsis as an adaptive strategy for larvae. Not only does shade re- 
duce the hunting success of visual predators, but the feeding positions of 
larvae on the host plant also contribute to protection. Older larvae gen- 
erally feed ventrally along the edges of leaves; they adopt feeding positions 
that apparently optimize food intake per unit time as their positions fit 
very well recent predictions concerning such behavior (Heinrich, 1971). 
This results from progressive movement of larvae down the plant as they 
grow larger; older larvae invariably feed on the lowest (generally oldest) 
leaves. The adaptive significance of this behavior is clear: larvae, as they 
get bigger, are able to take in more plant tissue per unit time and are 
less exposed to potential predators while feeding. Selection may favor the 
evolution of such behavior in a palatable species since it helps to maximize 
escape from predators which forage from above. This hypothesis, how- 
ever, remains to be confirmed. 

If further study reveals that the realized life table (“realized fitness’ — 
Istock, 1970) approaches the physiological life table (“physiological fit- 
ness’) for V. epaphus (Young, 1972e), then the idea that species well- 


166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


integrated into stable tropical communities possess very specialized and 
efficient defense mechanisms against predators, pathogens, and parasites 
would be confirmed (Margalef, 1968; Pianka, 1966). Adult survivorship 
in natural populations of V. epaphus is generally high (Young, 1972e), 
although the reasons for this are not known. A related species, V. 
steneles, enjoys high adult survivorship both in Batesian mimetic associa- 
tion with the heliconiine, Philaethria dido (Brower & Brower, 1964) and 
even when not in mimetic association (Young, 1972d). Recent studies 
of second-growth species of Costa Rican butterflies (Young, 1972a; Young 
& Muyshondt, in prep.) reveal high rates of parasitism on eggs and larvae 
during selected times of the year. High levels of biotic mortality, if 
persistent, suggest lack of integration of herbivorous species into second- 
growth communities. 

The depression in adult numbers during February—April (Table 2) 
may indicate decreased availability of adult food sources associated with 
diminished rainfall during this period. Cuesta Angel experiences a short 
and erratic dry season (“veranillo”). Reduction in monthly precipitation 
may destroy some food sources, such as fermenting patches of fungi on 
trees, or even flowers. During the wetter months, some adults may be 
killed off by heavy rains, although this is apparently constant (Table 2), 
and not highest at the beginning of the wet season, as observed for 
Parides (Papilionidae) on Trinidad (Cook, Frank & Brower, 1971). 
Such an adult mortality factor would be very sensitive to local topographi- 
cal and climatological features, varying greatly at different localities, 
especially along altitudinal gradients (Holdridge, 1967). 

The narrow local host plant specificity of larval V. epaphus is interest- 
ing. Although there have been discussions of mutualistic effects among 
plants and their lepidopterous herbivores (Ehrlich & Raven, 1965), rela- 
tively few studies have dealt with local host plant specificities at the 
species and generic levels, with the notable exception of some temperate 
zone lycaenids (e.g., Downey & Fuller, 1961; Downey & Dunn, 1964), 
Papilio glaucus (Brower, 1958), and Euphydryas editha (Singer, 1971). The 
single host plant of V. epaphus at Cuesta Angel is probably the only one 
that this butterfly exploits in this region of Costa Rica. At Bajo la Hondura, 
a montane region located on the Pacific slopes of the Cordillera, but of 
the same forest type and elevation as Cuesta Angel on the Atlantic 
slopes, V. epaphus exploits a different host plant, Hygrophila guiansis 
Nees (Acanthaceae). Like Ruella tubiflora at Cuesta Angel, this plant 
is characteristic of river edges at a wide range of elevations throughout 
Central America (D, C. Wasshausen, pers. comms.). But Ruella is not 
found at Bajo la Hondura nor is Hygrophila found at Cuesta Angel. It 


VOLUME 26, NUMBER 3 167 


would be interesting to determine whether two distinct strains (or sibling 
species ) of V. epaphus have evolved in Costa Rica, in response to dif- 
ferences in levels of host plant toxicity between the two localities (Cuesta 
Angel and Bajo la Hondura). Clearly experimental feeding studies are 
needed to demonstrate differences in palatability between the two pop- 
ulations. Brower & Brower (1964) found that adults of V. steneles are 
palatable although various members of the Acanthaceae are known to 
contain alkaloids (Arthur, 1954; Irvine, 1961). Levels of toxicity for 
either Ruella or Hygrophila are not known. 

A case of allopatry between V. epaphus and V. steneles may exist in 
Costa Rica. V. steneles is common throughout the Caribbean and Pacific 
lowlands (10-100 m elev.) of Costa Rica, where its major larval host 
plants, various species of Justicia in the Acanthaceae (Young, 1972e), are 
located. Here V. epaphus is generally absent, with the lowest elevations 
of its distribution being the Meseta Central (San Jose Province ) between 
200-500 m. However both species successfully complete development on 
the other's host plants in the laboratory. Therefore, other environmental 
factors, such as physiological adjustments to temperature and ecological 
barriers associated with topography may be important in determining 
altitudinal ranges of both species (Janzen, 1967). In the lowlands, V. 
steneles enjoys high adult survival as an understory species of primary- 
growth forest (Young, 1972d, e), where it represents a major zoogeographi- 
cal expansion of the genus Victorina at low elevations. At higher eleva- 
tions, a similar major adaptive radiation is portrayed by V. epaphus. The 
distribution of these species along altitudinal gradients varies greatly in 
different regions of Central America. In E] Salvador, V. steneles occurs 
at elevations up to about 600 m, and V. epaphus is found at higher eleva- 
tions (Young & Muyshondt, in prep.). 


SUMMARY AND CONCLUSIONS 


(1) The life cycle of V. epaphus is described for the first time, and 
data on developmental time are given. 

(2) V. epaphus is an understory species of montane tropical wet forests, 
where it exploits localized patches of its larval host plant. 

(3) Larval host plant specificity in V. epaphus is very high on a local 
basis, although different host plants are exploited in different parts of its 
extensive geographical range along the Cordillera in Costa Rica. At 
Cuesta Angel, a locality along the Atlantic (Caribbean) slopes, the host 
plant in Ruella tubiflora, while at Bajo la Hondura, a locality of similar 
elevation and forest-type on the Pacific slopes, the host plant is Hygrophila 
guiansis. Both plants are in the Acanthaceae. 


168 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(4) Immatures and adults of V. epaphus have high survival rates. Preda- 
tion and parasitism are low. Such life table characteristics are correlated 
with low fecundity, low adult vagility, and cryptic behavior patterns of 
larvae. 

(5) The genus Victorina in Central America seems to have undergone 
allopatric speciation, splitting into two species, V. epaphus and V. steneles, 
along altitudinal gradients. The former species is characteristic of 
montane forests and plateaus, while the latter species is more character- 
istic of the lowlands. However, larvae of both species can complete 
development successfully on the other’s host plants. 


ACKNOWLEDGMENTS 


This research was funded by N. S. F. Grant GB-7805 ( Daniel H. Janzen, 
principal investigator) and was carried out through the Organization for 
Tropical Studies, Inc., in Costa Rica. Drs. Woodruff Benson, Thomas C. 
Emmel, and Lee D. Miller assisted in identification of the butterfly 
studied, and Dr. D. C. Wasshausen identified larval host plants. Mr. 
Patrick Eagan assisted in location of host plants at Bajo la Hondura during 
1971; this aspect of the research was funded through support from a 
College Science Improvement (COSIP) Grant, GY-4711, awarded to 


Lawrence University. 


LITERATURE CITED 


ArtHur, H. R. 1954. A phytochemical survey of some plants of North Borneo. 
J. Pharmacy & Pharmacol. 6: 66-72. 

Brower, L. P. 1958. Larval foodplant specificity in butterflies of the Papilio 
glaucus group. Lepid. News 12: 103-114. 

& J. V. Z. Brower. 1964. Birds, butterflies, and plant poisons: A study in 
ecological chemistry. Zoologica 49: 137-159. 

Brussarp, P. F. & P. R. Exruicu. 1970. The population structure of Erebia 
epipsodea (Lepidoptera: Satyrinae). Ecology 51: 119-129. 
Core, L. R. 1959. On the defenses of lepidopterous pupae in relation to the 
oviposition behavior of certain Ichneumonidae. J. Lepid. Soc. 13: 1-10. 
Cook, L., M. K. Frank & L. P. Brower. 1971. Experiments on the demography of 
tropical butterflies. I. Survival rate and density in two species of Parides. 
Biotropica 3: 17-20. 

Deruer, V. G. 1959a. Egg-laying habits of Lepidoptera in relation to available 
food. Can. Entomol. 91: 554-561. 

. 1959b. Food-plant distribution and density and larval dispersal as factors 
affecting insect populations. Can. Entomol. 91: 581-596. 

Downey, J. C.& W.C, Futter. 1961. Variation in Plebejus icaroides ( Lycaenidae ) 
I. Foodplant specificity. J. Lepid. Soc. 15: 34-42. 

——— & D. B. Dunn. 1964, Variation in the lycaenid butterfly Plebejus icarioides 

_ If. Additional data on food-plant specificity. Ecology 45: 172-178. 

iumiicn, P. R. 1961. Intrinsic barriers to dispersal in Checkerspot Butterfly. 
Science 134: 1-2. 
— & 5S. HE, Davinson. 1960. Techniques for capture-recapture studies of 
Lepidoptera populations. J. Lepid. Soc. 14; 227-229. 


VOLUME 26, NUMBER 3 169 


& P. H. Raven. 1965. Butterflies and plants: A study in co-evolution. 
Evolution 18: 586-608. 

Hernricu, B. 1971. The effect of leaf geometry on the feeding behaviour of the 
caterpillar of Manduca sexta (Sphingidae). Anim. Behav. 19: 119-124. 
Howiprince, L. R. 1967. Life zone ecology. San Jose, Costa Rica: Tropical Sci. 

Center. 

Insz, D. 1941. The color vision in insects. Proc. Royal Phil. Soc. Glasgow 65: 1-11. 

IrvINE, F.R. 1961. Woody Plants of Ghana. Oxford Univ. Press London. 868 p. 

Istocx, C. 1970. Natural selection in ecologically and genitically defined popula- 
tions. Behav. Sci. 15: 101-115. 

JANZEN, D. H. 1967. Why mountain passes are higher in the tropics. Amer. Natur. 
101: 233-249. 

LaBIngE, P. A. 1968. The population biology of the butterfly, Euphydras editha 
VIII. Oviposition and its relation to patterns of oviposition in other butterflies. 
Evolution 22: 799-805. 

MacArruur, R. H. 1969. Patterns of communities in the tropics. Biol. J. Linn. 
Soc. 1: 19-80. 

Marcater, R. 1968. Perspectives in Ecological Theory. Univ. Chicago Press, 98 p. 

Myers, C. W. 1969. The ecological geography of cloud forest in Panama. Amer. 
Mus. Novitates, No. 2396, 52 p. 

Pranks, E. 1966. Latitudinal gradients in species diversity: a critical review. 
Amer. Natur. 100: 33-46. 

Pires, J. M., TH. DospzHansky, & G. A. BLrack. 1953. An estimate of the number 
of species of trees in an Amazonian forest community. Bot. Gaz. 114: 467-477. 

Prop, N. 1959. Protection against birds and parasites in some _ species of 
Tenthredinid larvae. Arch. Neerl. Zool. 13: 57-66. 

Ricuarps, P. W. 1952. The Tropical Rain Forest. Univ. Press, Cambridge. 217 p. 

Rot, I. & T. M. Brrano. 1971. Morphological and anatomical studies of leaves 
of the plants of a Venezuelan cloud forest. I. Shape and size of the leaves. Act. 
Bol. Venez. 7: 127-156. 

Serrz, A. (ED) 1924. Macrolepidopterra of the world. Vol. 5, American Rhopalo- 
cera. Alfred Kernen Verlag, Stuttgart. 1139 p. 

SHapmo, A. M. 1970. The role of sexual behavior in density-related dispersal 
of pierid butterflies. Amer. Natur. 104: 367-372. 

StnceR, M. C. 1971. Evolution of food-plant preference in the butterfly Euphy- 
dryas editha. Evolution 25: 383-389. 

SLoBopkKIN, L. B. 1970. Tropical Biology in London (book review). Quart. Rev. 
Biol. 45: 373-376. 

TuorsTEInson, A. J. 1960. Host selection in phytophagous insects. Ann. Rev. 
Entomol. 5: 193-218. 

Vaipya, V. G. 1969. Investigations on the role of visual stimuli in the egg-laying 
and resting behavior of Papilio demoleus L. (Papilionidae, Lepidoptera). Anim. 
Behav. 17: 350-355. 

Wess, L. J., J. G. Tracey, W. T. Wint1amMs & G. N. Lance. 1967. Studies in the 
numerical analysis of complex rain-forest communities. I. A comparison of 
methods applicable to site/species data. J. Ecol. 55: 171-191. 

Younc, A. M. 1972a. Mimetic associations in natural populations of tropical 
butterflies. I. Life history and population structure of Battus polydamus in 
tropical dry forest. Rev. Biol. Trop. 19 (in press). 

1972b. Mimetic associations in natural populations of tropical butterflies. 

II. Mimetic interaction of Battus polydamus and Battus bellus. Biotropica 4 

(in press). 

1972c. Mimetic associations in natural populations of tropical butterflies. 

III. Adult population structures of Parides arcas mylotes, P. arcas mycale, and 

P. childrenae childrenae. J. N. Y. Entomol. Soc. 80 (in press). 


170 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


1972d. Interaction of Philiathria dido (Heliconiinae) and Véictorina 
steneles (Nymphalinae) at Stachytarpheta flowers: evidence against mimetic 
association. Act. Biol. Venez. 8 (in press). 

1972e. Strategies of reproduction in some tropical butterflies. Oikos 23 
(in press). 


PIERIS NAPI L. (PIERIDAE) AND THE SUPERSPECIES CONCEPT 


S. R. BOWDEN 
53 Crouch Hall Lane, Redbourn, Herts., England 


This is a subject to be treated at greater length on some future occasion: 
the present note is intended to draw attention to unresolved taxonomic 
questions, which particularly affect the North American populations. 

When the European collector enquires, “Is bryoniae Ochsenheimer a 
separate species?” he may mean either of two different things. He may 
be asking whether ssp. bryoniae is reproductively separate from the other 
Pieris napi L. flying in the same country, or he may want to know whether 
bryoniae + flavescens Wagner + neobryoniae Sheljuzhko and perhaps 
adalwinda Fruhstorfer + hulda Edwards + ...... are reproductively iso- 
lated, severally, from napi + britannica Verity + meridionalis Heyne + 
oleracea Harris + venosa Scudder + .... The answers are likely to be 
different, and only that to the simpler question can be unequivocal. 

Dobzhansky (1970), quoting Mayr and also Amadon, defines a super- 
species as a monophyletic group of closely related and largely or entirely 
allopatric species, or as a group of ..... essentially allopatric taxa that 
were once races of a single species but which have now achieved specific 
status. The components of the superspecies are semispecies or allospecies; 
gene-exchange is still possible among semispecies but not as freely as 
among conspecific populations. Thus the Holarctic Pieris napi-bryoniae 
complex is a perfect example of a superspecies. 

The definition involves “species,” which can itself be defined in many 
different ways. For butterflies, a formally new, though quite theoretical, 
criterion of species status has been given (Bowden, 1972): 


Where the taxa have come into contact, if sympatry is leading to 
increased genetic and sexual barriers between them, the populations 
are to be taken as already belonging to distinct species; if not, not. 


This fixes speciation at a rather earlier stage than the “full speciation” of 
many systematists, and transfers a number of supposed semispecies to the 


VOLUME 26, NUMBER 3 Leffel! 


species category. But of course it still gives no assistance with populations 
which have remained apart geographically, except in so far as captive 
pairings provide evidence of the probable results of cross-breeding in the 
wild. 

The usual tests decide readily enough that Pieris virginiensis Edwards 
is specifically separated from P. napi oleracea, and experiments (Bowden, 
1966, 1972) leave no doubt that it is also distinct from P. n. napi. 

We can be nearly as certain that, in Switzerland, wild bryoniae is re- 
productively almost completely isolated from napi, though fertile hybrids 
are easily obtained in captivity. Experimentally, fertile oleracea xX 
bryoniae F, pairings are obtained as easily as napi X bryoniae, but there- 
after fertility crashes (Bowden, 1972). On the other hand oleracea X napi 
can be carried to the direct F3 at least. These results permit the con- 
clusion that P. n. napi and P. n. oleracea are still conspecific, and will 
interbreed if the Atlantic Ocean is abolished next year. But it is not 
necessary to wait so long to conclude that reproductive barriers would 
in fact go up rapidly—oleracea and European napi are very different 
insects. 

The writer has appreciable breeding experience of only three Nearctic 
subspecies of napi, including virginiensis, the third being marginalis 
Scudder. None of these can be raised on Alliaria, a plant which the 
European subspecies eat readily (Bowden, 1971la, 1971c). The larvae all 
differ from European napi and bryoniae, most conspicuously by the 
absence of bright yellow rings round the spiracles. The pupae vary in 
shape: P. n. marginalis is fairly close to P. n. napi, but virginiensis is 
radically different; P. n. oleracea is intermediate, but tending towards 
virginiensis (Bowden, unpublished). Pupae of the various European 
napi and bryoniae subspecies are, on the other hand, practically indis- 
tinguishable from one another. The relative length of the antennae also 
can be used to differentiate subspecies (Bowden, 1971b). The antennae 
of P. n. oleracea are conspicuously shorter than those of four European 
taxa measured, between which no statistically significant difference is 
found. P. virginiensis is perhaps slightly closer to the European propor- 
tion, but the difference from oleracea was not significant as measured. 
P. n. venosa shows a “European” antenna/wing ratio, as does P. n. hulda. 
P.n. marginalis (Oregon ) is intermediate between oleracea and European, 
and differs significantly from both. 

It is clear that phenotypic differences at three stages combine to 
separate oleracea from napi, even if one disregards the genetically deter- 
mined melanic patterns above and below the wings. The name Pieris 


172 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


oleracea could almost certainly be used in that form without error. But 
the status of other taxa is not so readily determinable. 

Lorkovié (1970) describes a confusing case in south-eastern Europe. 
In parts of Jugoslavia resides a population balcana Lork., which is 
phenotypically like P. napi meridionalis with some characters of ssp. 
pseudorapae Verity, but which karyologically resembles bryoniae and 
also pairs much more willingly with bryoniae than with P. n. napi. In 
the northern Caucasus is another population, balcarica Wojtusiak & 
Niesiolowski, which karyotypically rather approaches balcana but shows 
no reproductive isolation from P. n. napi; morphologically it resembles 
P. n. pseudorapae (Lebanon). Now there is as yet no sufficient evidence 
on which to determine whether Lebanon pseudorapae is specifically or 
only subspecifically distinct from P. napi napi, and its karyotype is un- 
known; hence one cannot decide whether ssp. balcarica is a subspecies of 
napi s.s. or of pseudorapae. 

Where the limits of true species should fall among the groups of sub- 
species will become better known as the relations between subspecies are 
explored in search of the evolutionary paths. But at present only scraps 
of the necessary knowledge are available, and we deceive ourselves if we 
pretend to have more. 

One concludes that for most of the taxa which have ever been included 
in Pieris napi we should continue to use a binomial or a trinomial nomen- 
clature, Genus species subspecies, but that when there is still insufficient 
evidence to decide to what species a particular subspecies belongs, we 
should instead use the superspecies name in parentheses: Genus (super- 
species) subspecies. 

If the subspecies in question constitutes what Kiriakoff (1948) and 
Lorkovié (1953) call a semispecies, this style is identical with theirs. Its 
adoption should imply that a semispecies (while it is considered such) 
cannot be allowed to have its own subspecies, even under the inappro- 
priate guise of “forms.” Indeed, while we are unable to allot a given 
semispecies to a species, we are unlikely to be able to allot further sub- 
species to that semispecies with any degree of certainty. 

If, however, the subspecies is not considered to be a semispecies, but 
is merely a taxon of still uncertain affinities, the same style may be used. 
The resulting slight ambiguity is justified by the rather temporary taxo- 
nomic status of the semispecies: in either case the aim must be final 
reversion to the conventional trinomial. 

Nevertheless, in the Pieris napi group we find a continuous range of 
differentiation, from local populations through subspecies to species, 
which nomenclature cannot fully reflect. The adoption of an arbitrary 


VOLUME 26, NUMBER 3 173 


criterion for specific status will not change this. Names are labels, and 
the amount of biological information that they can be expected to hold is 
limited. 


LITERATURE CITED 


BowpEN, S. R. 1966. Polymorphism in Pieris: “subtalba” in P. virginiensis. 
Entomologist 99: 174-182. 

. 197la. American white butterflies and English food-plants. J. Lepid. Soc. 

25: 6-12. 

1971b. Metrical discrimination of variable butterflies: antenna-length in 

Pieridae. Entomologist 104: 236-239. 

. 1971c. “Pieris napi’ in America: reconnaissance. Proc. Brit. Entomol. 

Nat. Hist. Soc. 4: 71-77. 

1972. “Pieris napi’ in America: genetic imbalance in hybrids. Proc. 
Brit. Entomol. Nat. Hist. Soc. 4: 103-117. 

DoszHANsky, T. 1970. Genetics of the Evolutionary Process. Columbia Univ. 
Press, New York. 505 p. 

Krrtakorr, S. G. 1948. On the so-called “lower” taxonomic categories. Lepid. 
News 2: 3-4. 

Lorxovic, Z. 1953. Semispecies a necessary new taxonomic category. Biol. Glasnik 
7: 236-237. 

1970. Karyologischer Beitrag zur Frage der Fortpflanzungsverhaltnisse 

sudeuropaischer Taxone von Pieris napi. Biol. Glasnik 21(“1968”): 95-136. 


NOTES ON URODUS PARVULA (HENRY EDWARDS) 
(YPONOMEUTIDAE) 


S. W. FRostT 


Frost Entomological Museum, The Pennsylvania State University, 
Department of Entomology, University Park, Penn. 16802 


Urodus parvula (Henry Edwards) is a common species in Florida and 
has been recorded from many localities from Miami north to Jacksonville. 
R. B. Dominick states that the species is also common from March to 
November at McClellanville, South Carolina. A single specimen in the 
Cornell University collection from Okefinokee Swamp, Georgia, is ap- 
parently the only record from that state. This species may occur along the 
Gulf coast towards Texas and northward along the Atlantic coast. Forbes 
(1923) stated, “The northerm record (District of Columbia) is based on 
a single specimen which may have been a stray.” 

Edwards (1881) referred this species to the genus Penthetria, Dyar 
(1898 ) placed it in Trichostibas, and Forbes (1923) assigned it to Urodus. 
Although only one species of Urodus is known from North America, this 


174 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Taste 1. Yearly and monthly occurrence of Urodus parvula in light traps at the 
Archbold Biological Station. 


Number of 
Year greet : Dec. Jan. Feb. Mar. Apr. May Totals 
i 
1958-59 194 I 140 ‘BD 83 — — 1100 
1959-60 WS) 36 153 120 16 _— — 404 
1961 CW) a 13 16 4 — — 33 
1962 53 23 4 — — — Pa, 
1963 68 — — 3 5 —_ — 8 
1964 88 — D 15 23 64 — 107 
1965 We a 14 44 By 56 — AL 
1966 TAS as oi 36 A2 186 — 295 
1967 127 — 128 103 269 649 114 1263 
1968 136 —- 201 44 Din 146 270 688 
1969 97 13 I 2 8 40 1 65 
1970 99 0 1 1 7 63 LIE 83 
Totals 1160 160 710 960 541 1204 396 4244 


genus, according to the U.S. National Museum records, is extensive in 
Central and South America. 

The present studies are based on more than 4,000 specimens captured 
in light traps over a period of twelve years, and some rearings. 

Eggs were not found on the hosts but were obtained in rearing cham- 
bers. They were laid singly and somewhat scattered on the insides on 
the rearing chambers and upon filter papers used to control moisture. 
They are pale yellow, smooth, without sculpturing, 0.74 mm long and 0.37 
mm wide. 

The larva feeds upon red bay leaves, Persea borbonia (L.) Raf., cutting 
narrow irregular areas. This type of injury was often seen but never 
appeared abundant enough to cause serious injury to the trees. The 
larva has also been recorded feeding on southern buckthorn, Bumelia 
reclinata Vent., orange, oak, and hibiscus. The mature larva is 12 mm 
long, noticeably spiny and somewhat colorful. The head is yellow, the 
thorax and abdomen have a yellow line on each side formed by a series 
of irregular yellow spots on the anterior and posterior margins of each 
segment. The strong dorsal setae arise from conspicuous black pinacula. 
The prolegs are brown, each with a lateral yellow spot. The setae of the 
thoracic and prolegs are weak and pale in color. 

The pupa is formed in a lacy cocoon, oval in shape and somewhat 


> 


Migs. 1-3. Urodus parvula (Henry Edwards). 1, adult x 4; 2, cocoon with 
pupa ¥ 3; 3, cocoon showing attachment pedicel and cast pupal skin within x 5. 


VOLUME 26, NUMBER 3 175 


176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Sex ratios of specimens taken in light traps from 1967 to 1970. 


N aber of 

Month See Males Females Totals 
December 10 9 4 13 
January 95 229 102 331 
February Hil 99 ol 150 
March 101 206 95 301 
April es 602 296 898 
May 34 188 198 386 
Totals 430 1333 746 2079 


tapered at the lower end. It is 13 xX 6 mm and formed of regular trap- 
ezoidal meshes supported by a thick thread that extends along the side of 
the cocoon and is attached to the supporting object. 

The adult male and female are similar in color, opaque black, 15 mm 
long with a wing expanse of 22 to 25 mm. The males are easy to recognize 
because of their unusually large genitalia and the possession of an indis- 
tinct short reddish area at the base of each wing near the costal margin. 

Adults were freely attracted to lights. Considerable variation in the 
abundance of the moths was noted from year to year. These variations 
could not be correlated with temperature or precipitation but apparently 
were due to developmental cycles which reached a maximum in 1959 
and 1967. Although light-trap catches extended only from December to 
May, Kimball (1965) has records for June and July and apparently this 
species has a long period of activity. In general the moths seemed to be 
especially abundant throughout the winter and spring months. Males 
were notably more abundant than females. Of 2079 specimens taken in 
light traps from December 1 to April 30, involving 430 nights over several 
years, 1333 were males and 746 were females. 


REFERENCES 


BONNIWELL, J. G. 1918. Notes on collecting in Florida. Lepidopterist 2: 57-60. 

Dyar, H.G, 1898. New American moths and synonymical notes. J. N. Y. Entomol. 
Soc.’ 6: 41. 

1900. Notes on some North American Yponomeutidae. Can. Entomol. 32: 

37-41. 

1913. The larva of Trichostibus parvula. Ins. Insc. Mens. 1: 49-50. 

Epwarps, H. 1881. A new genus and some new forms of North American Zygaeni- 
dae. Papilio 1: 80. 

Forses, W. M. T. 1923. The Lepidoptera of New York and neighboring states. 
N. Y. Agr. Exp. Sta. Mem. 68: 340. 


Frost, S. W. 1963. Winter insect light trapping at the Archbold Biological 
Station, Florida. Fla. Entomol. 46(1): 38-39. 


VOLUME 26, NUMBER 3 Wig 


. 1964. Insects taken in light traps at the Archbold Biological Station, 
Highlands County, Florida. Fla. Entomol. 47(2): 153. 

Ho.tianp, W. J. 1920. The Moth Book. Doubleday, Page and Co. 253 p. 

Kimpatz, C. P. 1965. The Lepidoptera of Florida. V. 1 in Arthropods of F lorida 
and Neighboring Land Areas. Fla. Dept. Agr. p. 113, 292. 


OBSERVATIONS ON FOODPLANT RECORDS FOR 
PAPILIO GLAUCUS (PAPILIONIDAE ) 


MatcoLm P. LEvIN 
Department of Biological Sciences, Stanford University, Stanford, California 94305 


Mary ANN ANGLEBERGER 
Department of Biology, University of Virginia, Charlottesville, Virginia 22900 


Numerous investigators have contributed to a large body of diverse 
data regarding the choice of ovipositional sites and larval foodplant 
records for the Papilio glaucus L. group of swallowtails. These data have 
been compiled and summarized by Brower (1958). The field observa- 
tions described herein were made during the summers of 1968-1971. 
These observations add still another family of plants to the list of known 
ovipositional choices for Papilio glaucus, the Eastern Tiger Swallowtail, 
and clarify the records for two genera of plants found in communities 
associated with populations of P. glaucus in southwestern Virginia. 

During the summers of 1968-1970 more than 12 observations of oviposi- 
tion were noted on Prunus serotina Ehrhart (Wild Cherry) in Giles and 
Montgomery Counties, Virginia. Other observations during this time 
include 6 larvae collected from P. serotina. Although Prunus virginiana 
L. (Choke Cherry) is found in the same locality, larvae reared in the 
laboratory, when given a choice, rejected P. virginiana in favor of P. 
serotina. In this test two groups of second instar larvae (10 progeny from 
each of two females) were placed in rearing dishes containing both 
species of Prunus. During the subsequent 48-hour period no feeding 
damage was observed on P. virginiana. However, the larvae moved about 
freely and were observed to feed on P. serotina. Assuming that these 
plants are equally acceptable one would expect feeding damage to have 
occurred on P. virginiana and that the larvae would be distributed equally 
among the two plant species. These observations (20 on P. serotina: 0 on 
P. virginiana) differ significantly from the expected (chi-square test). 

In contrast, larvae of P. glaucus readily fed on both Magnolia acuminata 


178 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


L. (Cucumber-Tree) and P. serotina when both plants were placed in 
the same dish. Similarly, observations regarding the ovipositional be- 
havior of P. glaucus have also been made on M. acuminata. On two 
occasions in the field, females displayed apparent oviposition on this 
species; moreover, one of the butterflies appeared to have laid at least 
10 eggs in various parts of the tree. Because these butterflies chose the 
uppermost branches, it was impossible to determine that eggs had 
actually been deposited. However, during these observations, the entire 
sequence of ovipositional behavior was normal. Additional evidence 
that M. acuminata is an acceptable foodplant comes from one of the 
author's (MPL) laboratory studies. In the course of rearing P. glaucus, 
M. acuminata was frequently used to elicit oviposition and larvae were 
reared successfully on this deciduous tree. 

Clark & Clark (1951) indicate that Magnolia is a favorite food-plant 
for P. glaucus in the southeast; however, neither they nor Brower (1958 ) 
identifies the species of Magnolia. This seems to be a fairly important 
gap in the data for many magnolias are evergreens, while P. glaucus 
feeds on deciduous trees. The Cucumber-Tree, M. acuminata, is a 
deciduous tree and seems to fall into the general feeding pattern of P. 
glaucus. 

During the summer of 1971 one of the authors (MAA) brought a larva 
into the laboratory from the wild. It was assumed that the larva was P. 
cresphontes Cramer because it had been located on the hop tree, Ptelea 
trifoliata L.. (Rutaceae). Subsequently the authors verified that this was 
a fourth instar larva of P. glaucus. A systematic search of Ptelea at the 
same site, the Sinking Creek area in the vicinity of State Route 700 (Giles 
Co., Virginia), failed to locate other P. glaucus larvae. Further, species 
of plants which are known to be acceptable ovipositional sites for P. 
glaucus larvae were absent from this locality. Orange dog larvae of P. 
cresphontes, however, were found repeatedly. 

A second locality, Spruce Run Valley (Giles Co., Virginia), was also 
examined to determine the relative frequency of Ptelea-feeding. Ptelea 
is abundant along the mouth of this creek; most of the plants are saplings 
and can be examined rather carefully. Again one P. glaucus larva (fourth 
instar ) was collected from Ptelea. This larva was also taken to the labora- 
tory and reared on Ptelea. One additional case of a P. glaucus larva on 
Ptelea has been observed (D.A. West, pers. comm.). It seems unlikely 
that the occurrence of either of the larvae discovered by the authors could 
be due to larval migration. The second larva, like the first, was in an area 
lacking known and suitable ovipositional sites. Finally, in order to obtain 
ome additional information, two P. glaucus larvae (both in the third 


VOLUME 26, NUMBER 3 179 


instar) collected from P. serotina were transferred to Ptelea in the lab- 
oratory. All four P. glaucus larvae appeared to develop normally and 
typical prepupal behavior was observed. The four larvae pupated; of 
these, one died and three are presently in diapause. 

As reported in the literature (Dethier, 1937, 1941, 1953; Hamamura, 
1959; Thorsteinson, 1958, 1960), lepidopterous larvae often require specific 
stimuli to elicit feeding. Moreover, unsatisfactory food plants, those 
lacking appropriate chemotactic or gustatory stimuli, or possessing repel- 
lants, are frequently rejected; thus P. trifoliata seems to provide the 
necessary “token stimulus” and nutritional value for the development of 
P. glaucus larvae. Yet, it seems that the apparent use of P. trifoliata 
as an Ovipositional site is not particularly frequent for our extensive search 
for larvae produced only two. It is probable that the apparent use of 
these plants as an ovipositional choice is of relatively recent origin since 
the observations of P. glaucus on numerous other deciduous trees has not 
escaped observation (Brower, 1958; Clark & Clark, 1951). Nevertheless, 
in this part of Virginia some females may use Ptelea on a regular basis, 
for the trees are found scattered throughout the valleys of Giles and 
Montgomery Counties, Virginia. 

At this time no conclusions can be reached as to the significance of 
the Ptelea-feeding observations. The nature of the foodplant preferences, 
i.e. whether they are induced or hereditary (Jermy et al., 1968), cannot 
be determined without observations from carefully controlled experi- 
ments. As Jermy et al. (1968) point out, the use of field-collected larval 
populations for analyses of feeding preferences may result in faulty con- 
clusions. Thus it would seem that further field observations should be 
undertaken and that a clarification of the alternative hypotheses for food- 
plant preferences is warranted. 


ACKNOWLEDGMENTS 


Support for this work for Levin was in part from a Ford Foundation 
Postdoctoral Fellowship at Stanford and in part from an Ivy Lewis Fellow- 
ship awarded through the Mountain Lake Biological Station. The authors 
also gratefully acknowledge the use of the facilities of the University of 
Virginia's Mountain Lake Biological Station. 


LITERATURE CITED 


Brower, L. P. 1958. Larval foodplant specificity in butterflies of the Papilio 
glaucus group. Lepid. News 12: 103-114. 

Crark, A. H. & L. F. Crarx. 1951. The butterflies of Virginia. Smithsonian Misc. 
CollShIG. No: 7. 

DeTuierR, V. G. 1937. Gustation and olfaction in lepidopterous larvae. Biol. Bull. 
Wan 123. 


180 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


1941. Chemical factors determining the choice of foodplants by Papilio 

larvae. Amer. Nat. 75: 61-73. 

1953. Host plant perception in phytophagous insects. Trans. IX Int. 
Congr. Entomol. 2: 81-88. 

Hamamura, A. 1959. Food selection by silkworm larvae. Nature 183: 1746-1747. 

Jermy, T., F. E. Hanson & V. G. Dernier. 1968. Induction of specific food 
preference in lepidopterous larvae. Entomol. Exp. Appl. 11: 211-230. 

TuorstTEINsON, A. J. 1958. The chemotactic influence of plant constituents on 
feeding by phytophagous insects. Entomol. Exp. Appl. 1: 23-27. 

1960. Host plant selection in phytophagous insects. Ann. Rev. Entomol. 

5: 193-218. 


THE LARVA OF CHAMYRIS CERINTHA (TREITSCHKE) 
(NOCTUIDAE) 


GrorceE L. GODFREY 


Illinois Natural History Survey and Illinois Agricultural Experiment Station 
Urbana, Illinois 61801 


The larva of Chamyris cerintha (Treitschke ) previously was described 
by Coquillett (1881), Forbes (1954), and Crumb (1956). In all instances, 
the descriptions primarily dealt with general body structure and color. 
The notes and illustrations in this paper are designed to describe the 
caterpillar of C. cerintha more thoroughly, especially in respect to the 
mouthparts and chaetotaxy. This is done to further document morphologi- 
cal structures of the larvae of the Noctuidae that I think are of key 
taxonomic importance, as briefly explained earlier (Godfrey, 1971). In 
addition, a habitus drawing of the caterpillar (Fig. 1) is provided to 
facilitate identification of the species. 

The illustrations were drawn to scale by a grid system. The scale lines 
represent 0.5 mm for all figures unless designated differently. The 


terminology and abbreviations are consistent with those used earlier 
(Godfrey, 1970). 


General. Head about 2.5 mm wide. Total length about 32 mm. Abdominal 
prolegs present on third through sixth segments. Head smooth. Body extensively 
covered with minute granules. Dorsal abdominal setae simple, very long. Dorsal 
setae on seventh abdominal segment 6-10 times height of seventh abdominal spiracle; 
setae on eighth segment 19 times height of spiracle on seventh segment. Dorsal 
setae on abdominal segments eight and nine borne on distinct tubercles. 

Head (Fig. 2). Epicranial suture 1.6 times longer than height of frons. Distance 
from frontal seta (F-1) to frontoclypeal suture 0.5 times distance between F-1’s. 
\dfrontal puncture (AFa) and second adfrontal seta (AF-2) posterior to apex of 
| \nterior setae (A 1-3) forming obtuse angle. Lateral seta (L) slightly caudal 


VOLUME 26, NUMBER 3 181 


4 


Figs. 1-5. Chamyris cerintha, Arlington, Va. 1, left lateral aspect of last instar; 
2, frontal aspect of head capsule; 3, left dorsolateral seta arrangement of prothorax; 
4, left aspect of hypopharyngeal complex; 5, oral aspect of left mandible. 


182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of transverse line formed by AF-2’s. First posterior seta (P-1) definitely caudal of 
juncture of adfrontal sutures. Interspaces between ocelli (Oc) 1-2 and 3-4 subequal, 
less than interspace between Oc 2-3. 

Mouthparts. Oral surface of labrum unspined. Hypopharyngeal complex (Fig. 4): 
spinneret tapering, not exceeding tip of Lp-2; stipular seta about 1% length of Lps-1, 
equal to Lp-1, longer than Lps-2 and slightly shorter than Lp-2; distal region covered 
with short, fine spines becoming stouter proximad; proximolateral region with distinct 
row of about 18 large spines. Mandible (Fig. 5) with two closely spaced, large, 
flat, inner teeth; inner ridges indistinct; six outer teeth present. 

Thoracic segments. Segment T-1: seta D-2 caudal of line formed by D-1 and 
XD-2 (Fig. 3); major axis of prothoracic spiracle passing slightly behind seta SD-2 
and both subventral setae (SV 1-2); SD-1 in line vertically with setae D 1-2. Seg- 
ments T 2-3: seta L-1 located above and slightly posterior of L-2. Tarsal claw 
with distinct basal angle. Tarsal setae with parallel sides and rounded tips. 

Abdominal segments. Ab-1: only two subventral setae (SV 1, 3) present; SV-1 
located posterolaterad of line formed by seta V and SV-3. Ab 2-6: three subventral 
setae present. Ab-8: only one seta in each subventral group. Ab-9: seta SD-1 as 
strong as setae D 1-2. Anal and subanal setae no larger than lateral setae on anal 
proleg. Crochets uniordinal. 

Coloration. See Forbes (1954) and Crumb (1956) for the color description. 

Hosts. According to existing records, the caterpillar of cerintha feeds only on 
plants of the family Rosaceae. The recorded hosts are Crataegus sp., Malus sp. 
[apple], Prunus persica (L.) Batsch [peach], Prunus serotina Ehrh. [wild cherry], 
Prunus sp. [wild cherry], Prunus sp. [plum], and Rosa sp. [rose] (Coquillett, 1881; 
Lugger, 1899; Forbes, 1954; Crumb, 1956). 

Material examined: 1 specimen, Arlington, Virginia, July 1949, reared on Prunus 
serotina from oyum from female collected by J. G. Franclemont. Hypopharyngeal 
complex on slide G-0189. 


ACKNOWLEDGMENTS 


I thank Dr. John G. Franclemont, Cornell University, for loaning the 
material used in this study and Dr. W. E. LaBerge, Illinois State Natural 
History Survey, for reviewing the manuscript. 

This research was supported by USDA AGR RMA Grant No. 12-14- 
100-8031(33) awarded to Dr. John G. Franclemont, Cornell University, 
Ithaca, New York, and by funds and facilities supplied by the Illinois 
Natural History Survey, the Illinois Agricultural Experiment Station, and 


the Office of International Agriculture, College of Agriculture, Urbana, 
[linois. 


LITERATURE CITED 


Coguittetr, D. W. 1881. Notes and descriptions of a few lepidopterous larvae. 
Papilio 1: 56-57. 

Crump, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agr. Tech. Bull. 
1135, 356 p. 


forbes, W. T. M. 1954. Lepidoptera of New York and neighboring states. Pt. 3. 


Cornell Univ. Agr. Exp. Sta. Mem. 329. 433 p. 
Goprrey, G. L. 1970. <A review and reclassification of the larvae of the subfamily 
Hadeninae (Lepidoptera, Noctuidae). Ph.D. Thesis, Cornell Univ. 478 p. 
1971. The larvae of Lithacodia muscosula, L. carneola, and Neoerastria 


apicosa. J. Kansas Entomol. Soc. 44: 390-397. 


VOLUME 26, NUMBER 3 183 


Luccer, O. 1899. Fourth annual report of the entomologist to the state experiment 
station of the University of Minnesota, to the Governor, for the year 1898. St. 
Paul, Minn. 280 p. 


MAXIMIZING DAILY BUTTERFLY COUNTS 


KeitH S. Brown, Jr. 
Centro de Pesquisas de Produtos Naturais, U.F.R.J., Rio de Janeiro ZC-82, Brazil 


Early in 1967, a preliminary draft of a most interesting paper by Heinz 
Ebert came into my hands for initial comment. This paper (Ebert, 1970) 
provided me a great impetus toward the systematization of field collecting 
techniques, and the proper keeping of records. It was especially valuable 
in pointing out the great rarity of most species in southern Brazil (and 
probably in most of tropical America), the preponderance of small, 
inconspicuous, and/or highly localized butterfly species in the Neotropical 
fauna, and the advantages of having several collectors in an area at the 
same time to obtain more complete and representative recording of the 
species present. 

I resolved to try to apply the conclusions of Ebert’s paper towards a 
continuing study of the Lepidoptera of the central Brazilian plateau (see 
Brown & Mielke, 1967, 1968; Mielke, 1967, 1968a, 1968b). Initially, a 
complete written list was made of the species (including numbers of 
each sex) that were captured and positively observed in each collecting 
day. The following observation of Ebert (p. 6) provided an initial basis 
for the effort then made to maximize these daily lists: 

“In eastern Brazil the individual frequency of butterflies is generally 
very low. The success of an excursion is exclusively determined by the 
number of species found. The higher the number of species found during 
a trip, the higher the chance to find some regionally (and/or individually ) 
rare species... .” 

This suggested that the maximization of daily species lists of butter- 
flies, a seemingly unscientific goal (though much employed in a sister area, 
ornithology ), could give a large scientific fallout; and this has indeed 
proved to be the case. For the benefit of the butterfly-interested public, 
both amateur and professional, this paper presents a discussion of the 
methods used for maximization and the results obtained, including a 
comparison of various Neotropical collecting areas, both in overall 
Rhopalocera and in individual family or tribal groups. 


184 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Although the collecting methods here described for obtaining maximum 
daily lists differ in many fundamental aspects from those used by Ebert 
in his careful long-term studies of the butterfly frequencies in Brazil 
(which place an emphasis on averages rather than on maximum daily 
lists), the overall results are very similar and further confirm Ebert’s 
conclusions on intensive, extensive, and seasonal abundance of the various 
groups of Brazilian Rhopalocera. This paper thus represents a supple- 
ment to Ebert’s full discussion, written from a different point of view, 
and incorporating data derived principally from about 150 collecting 
days in 1967-1972 (see Table 1 for a selection of these), with maximum 
expenditure of energy, using methods of intensive or large-scale area 
coverage, and efficiency and accuracy of recording, developed and 
sharpened during ten years of taking ornithological censuses. 


METHODS 
Equipment 


It was ascertained early that the routine use of binoculars during the 
day’s collecting (as in ornithology) not only greatly increased the num- 
ber of butterfly species that could be positively recorded in a day, but 
also permitted a beneficial redistribution of energy: most larger common 
species, which could be unequivocally identified even to the sex at long 
distance with binoculars, did not need to be captured, and physical effort 
could be concentrated on collecting the smaller and more unusual species 
present. In general biological terms, the use of binoculars also allowed 
careful field studies of high-flying species and unusually wary forms or 
individuals, rarely mentioned in the lepidopterological literature. In a 
noteworthy case, the female of a new species of Ithomiinae was observed 
while laying eggs high in a tree; this observation led to publication of the 
complete biology of the species together with its original description 
(Brown & d’Almeida, 1970). Many other immature stages have been 
discovered in this same manner with binoculars. Also, the nearly extinct 
Heliconius nattereri was finally relocated, studied, and bred by the 
author following a chance long-distance sighting with binoculars of a male 
in Santa Teresa, Espirito Santo (see Brown, 1970, 1972). 

An eight-foot pole length with a four-foot extension (for more open 
areas), easily manageable in all but the densest woods, is necessary to 
capture the many high-flying species (which generally belong to groups 
of localized and little-known forms, especially Lycaenidae! and Hesperi- 

We use the family Lycaenidae to cover the group often discussed as a superfamily, Lycaenoidea 


in the Neotropics, the Plebejinae, Theclinae, and Riodininae). Likewise, Nymphalidae = Nym- 
ialoidea of many authors. 


VOLUME 26, NUMBER 3 185 


idae ), which can be discovered with binoculars but not always positively 
identified. 

In addition to the binoculars and lightweight net with aluminum pole, 
minimal collecting equipment is kept (forceps, a belt box with envelopes, 
a small poison jar in the pocket ) to permit rapid movements and efficient 
handling of captured specimens. Photographic and breeding equipment 
are left in a central location; early stages discovered are brought back 
there during “slack” collecting hours in the late afternoon. 


Planning 


In attaining a maximum list in a given region, or seeking to use a 
limited number of collecting days most effectively, the lepidopterist 
should work principally in relatively restricted and disturbed wooded 
areas (see below), and move around a great deal in the initial explora- 
tions. The area chosen must be well reconnoitered for paths and roads, 
streams and sandy banks, flowers inside and outside the forest, clearings 
within the woods which catch the sun at various times of the day, and 
accessible forest edges lit by the first and last sun in the day. The latter 
areas will provide the majority of the day’s lycaenids, and the clearings 
and flowers the majority of the Hesperiidae. A partly cleared hilltop will 
provide dozens of additional species throughout the day in both these 
groups, plus many additional and unusual Papilios and Nymphalidae. 
Baited traps (as many as 30) should be placed along trails and on woods- 
edges on the evening or night previous to a collecting day. By using 
traps to capture the many bait-attracted nymphalines and satyrines (a 
number of which enter during the twilight and dawn hours ), the collector 
can focus his attention on the smaller species which do not catch them- 
selves so easily. 


Collecting Period and Distribution of Energy 


In the ideal case, weather and his strength permitting, the collector 
should work from sunup to dark, leaving the woods edge at sundown 
(when the Lycaenidae finally sit down for the night) to capture Bras- 
solini at their suspected or prelocated flyways. Thus the condition of 
maximum effort may be fulfilled, giving the over-all results a measure of 
consistency. “Slack periods” in the early morning and late afternoon, or 
during cloudy spells, may be used for resting, moving between collecting 
areas, and harvesting the traps. In very hot lowland tropical areas (the 
coastal plain in southeast Brazil in summer, much of the Amazon Basin 
near the rivers, or lowland Panama) both butterflies and collectors are 
often driven into inactivity by the heat between 1100 and 1430; but in 


186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


hilly or mountainous regions, both can remain active during most of the 
daylight hours, with peaks in late morning and mid-afternoon. 

A single collector, for best maximization, should move rapidly from 
habitat to habitat (using a vehicle such as a Jeep to cover greater dis- 
tances ), collecting a number of times during the day in each subarea of 
the locality chosen. On the other hand, a party of collectors will get more 
results and tire less during the day if they divide up the locality, each 
covering one biotope or contiguous area very thoroughly during the entire 
day. 

In view of the great preponderance of small-sized Hesperiidae and 
Lycaenidae in the Brazilian butterfly fauna (see Table 1, end), the effort 
to maximize lists requires a heavy concentration on these often ignored 
groups, using binoculars to locate high-perching individuals and spending 
much time collecting the many species that visit flowers. As Ebert (1970) 
has noted, the vast majority of the species in these two families have a 
very low intensive frequency; even in areas where over 300 species in 
each family have been recorded over long periods, the maximum day’s 
list under optimal conditions rarely passes 100 species in either family. 
Casual (non-maximized) collecting usually yields but 10-40 species of 
Lycaenidae or Hesperiidae in a day; concentration on these groups can 
raise the levels to 30-80 or more (see Table 1). 


RESULTS 


The comparison of results obtained in completing the list for the central 
plateau before and after the adoption of these methods is instructive. 
In six weeks’ collecting in the planalto, before the use of the maximization 
method, the author and Nirton Tangerini added only 25 species to the 
total published list (Brown & Mielke, 1967). Using the methods to 
maximize daily lists, and thereby collecting preferentially the less notice- 
able species, the author with Stan Nicolay or Karl Ebert added nearly 
300 species to the list in another six weeks’ collecting in 1967-1970. The 
total list for the planalto is now over 950 species (see Table 1); most of 
the added species are small or inconspicuous forms (Satyrinae, Lycaenidae, 
and Hesperiidae) which probably would have been passed over in casual 
collecting. 

In two weeks’ collecting in Belo Horizonte using these methods in May 
1967 (the first real test of maximization), the author added 145 species 
to the published list for the area (Brown & Mielke, 1967). On one sunny 
day, 260 species of Rhopalocera were positively identified, over 90% of 
these being caught. 


On the local scale, the maximization method could be used to confirm 


VOLUME 26, NUMBER 3 187 


a fact well-known to Neotropical lepidopterists but often misunderstood 
in other areas: the most rich and, surprisingly, often most representative 
collections are to be made in highly restricted areas of partly cut forest, 
surrounded by fields and swamps, preferably with many low flowers, and 
liberally served by paths, roads, and clearings. While the legendary (and 
often imaginary!) areas of vast virgin tropical forest invariably harbor 
many unusual and characteristic species, these tend to occur in very low 
density throughout the forest, and must seek sun at levels unattainable 
by the collector's net. The best overall collecting is always in small varied 
woodlots where most species thrive and remain accessible. 

On the regional scale (see Table 1), the maximization method revealed 
that the largest intensive species frequencies for daily lists (though not 
necessarily for long-term censuses), could be found in the blend zones 
between the cerrado area of the Brazilian planalto and the adjacent 
tropical forests of the southeast coastal mountains (as in Belo Horizonte 
or Pocos de Caldas) or the Amazon Basin to the north. The rarer species 
seemed to be more common in these areas, probably because of the highly 
varied microclimates and floral niches present in the hilly transition areas 
between different faunal regions. 

In the northwestern blend area between the cerrado and the upper 
middle Amazon region in central Mato Grosso, preliminary application 
of these methods in the summer of 1969 suggested that intensive winter 
collecting might give lists approaching the limits of daily butterfly num- 
bers for the Neotropics. Therefore, when in May 1969 an excursion to 
the Chapada de Guimaraes was made in company with Stan Nicolay, full 
plans were developed for intensive collecting with maximization methods, 
both along the way and in the Mato Grosso blend area. The results of 
this trip (see Table 1) amply confirmed the exceptional position of the 
Chapada de Guimaraes in central Mato Grosso with respect to relative 
butterfly abundance. This area produced in one day a list of 361 species 
(307 recorded by the author alone), which, for the time being, stands 
far above the daily lists obtained by us or others in any other area of the 
Neotropics or, indeed, in the rest of the World.” 


2 The late Dr. Richard M. Fox claimed to have collected “about 350” species in a single day’s 
collecting near his field station in Liberia (personal communication). As he reports only 450 
species for the whole country (Fox, Lindsay, Clench & Miller, 1965), we must conclude that, 
unless the percentage of rare and localized species in the Liberian fauna is vastly lower than 
that in the Brazilian fauna, this estimate may be somewhat exaggerated. The author also has 
a letter from John H. Masters indicating that a list of 350 would be attainable in a single day’s 
collecting in the Amazon Basin. We presume that this would be in the blend areas on its western 
edges, where the highland and basin faunas meet and mix. The author has visited a number of 
such areas in Colombia, Peru, Ecuador, and Bolivia; several of them offered the same subjective 
impression of extreme butterfly abundance that is evident in central Mato Grosso. 

A day’s total of 350 should be near the limit for a single collector even in such exceptionally 
rich areas, not due so much to butterfly abundance as to limitations on the speed of moving 
between ecological niches, and observing, collecting, and enveloping specimens. In some excep- 
tionally varied blend areas on the borders of the Amazon Basin, a party of three or four collectors 
might be able to pass the limit of 500 species in a single day. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


188 


toi Oh 6 SIS ORY = ie IE ORY IL = tts SS AL 8) skep GT “Oly wou sysorof [[TYIOOF IOF¥ STV.LOL 


‘ovpriiiodsayy/eepruevoAT FO Ayoreos ‘ovprjeydutAyY JO soURpUNqy ‘uOsves IaMOTT 
[dy ur yved Arepuooas ‘19q0}Q FO sulex AAvay auofaq ‘Sutids AjIVa pue TOJUIA 9}eT UL YeOg ALON 


33S Omr ol Cm ClO) SOG OF ik GORS Co LikS “l= “Te Sr Arp “Joy “ree LO-X-€ 
LOO) Cle ie alte See 40) Ope Ol 6 “6h Go OL "2 = T_T yoy <r89I) L9-X-T 

Ce ear cme laos Ge 40° (0. SOS OL 0 Ol or <Si 8. 1 T- Sar yoy “1vIID L9-XI-0€ “SUIe} 
OSGi ia lt BLCeOe oe S0VS Se Tice I. vi S l= F Fr UBM “Teva 89-XI-SZG -UNOUL FO JOOF 
Sooo Ome nOC iCeOns Oeil =e 6G Tr SCG Te 9S "GS bo 1 =F yoy “1v2aTD 89-XI-61 ye SUIBII}S 2 
OA i eee a Semce An Ol aekce OR I “Sl ko Clay 1 ie “air joy “1vea[D L9-XI-81 smoAUeo ‘UTBITA 
AOC uence ly een en! Ye, Se Gometi= To Of <0 SE cr yoy “Iva[D, 89-XI-91 Apsour “jsar10F 
fee (OS BIL SS hi? O. = I Ws Os 5 GIL he OL OES Arp ‘Joy “IVaI_—s SO-ITA-O€ prumy AAvoy 
On Ole s aeeliee 2 O) eecGew owe GC. SiGe, (6 80 6 Ie aietar yoy “1va[D LO-AI-S1 :o1reuef op 

S/O maa Oat tae 2. 208 Dever het “O° -G.. 6G “E86 = 0 stor yoy “13D LO-AI-TT ONY JO 97¥}S “WI9IOX 

“IOJUIM UL S}SO1F 

eousliedxa YOIyM ‘s}so10f oye1odui9} 9say} JO eUNeF paoNpert YonU AT[eIoUeD ‘IOUTUIMSprUt UL Yyeeg :ALON “dures 
“\so1oy [eordo13qns 

Ciao a Cee Ce Vas (On CG. ew (Se Oo 8° §S)  £  G “ctalsr WABM “TBD 89-II-Z ‘eueieg “equuny 
*soysieul 
4gngie Teale 
‘\ser0F [vordonqns 
:BuRIed “essOIs 

LOCC Cem elie wGe O. Li § 0 SO; 0: 6h Wr 6 G sb ULIBA “19D 89-II-9 eyuod “eypeA BTA 
320 ee O06 O88 O8 & 2001 T Sh Ol O We © ie Gil © USA DAREN) (0i7 PC UEUT Oe SINK ONE 


‘ovpiiiodsox, JO 9OURpUNGe [eIoUe5) “poos AjouleI}x9 TJGT ‘(stvoA peq) OQLEGI PUe SOG 
pue (sivaXk poos) 696[ pue LOGI UWoorAjoq sooUsIEs}Ip oS1evT “WOSeaS IOMOTF AOJUIMpPIUL UL YVag : ALON 


wl 6 § i Of © OS © GS 6 Tf 8 © M& FP L LF Arp ‘poo “twat L9-XI-9T 

6 12 @ Pl 06 6 Ce © GO W LF Of © 8 @ 1 see Arp Joy “eat LO-IITA-8% 

oo SI S @6 Se Lt 8s @ 28 Of S On 6 Of 2 @ Tf came wae “Tea = TL-IILA-#1 

OLI 89 € we /alt Ab LI O 6G 6 G I 0 L G 0 I Sleateate ware “we = OL-IIIA-ST 

6. LO SO Si OG OS Ce eR 10) SO ore maa Apnop AjywWeq 69-IITA-€T ‘sdo TY 
CAS CMC IG o= Ol Oly 6 ..b-"6-0, 9 & A-- LT. Heb Apnoyo Apxeq — L9-IITA-GI poos sapnpout 
Sane not On -CheROMmIGenGl. © SC. 0 92 pe To fat wea ‘<pnoryy 9-IITA-8 ‘spvor pur sey 
Gee Oo  Y Bl Re Tt ie © GS OR G © © OY © & GER We “we —-LO-ITA-TE ‘suocueo pruny 
eo O& py Ol Sl 1 Wee OGG eG Wo i O— wy iba 8 sae joy ‘Iwas B9-ITA-6 qsor0f dooys (Aue 
QCeCOmmIGH mCi Scamce GL. 0) Ti OL € -=@l- 1- 9 GS -T. Tf -Set ULTRAM “IVITD LO-IA-61 -puooas Ap}sour) 
GCC OMC OMe IG MON GGOl US 2 fb 9 - | eo St ue “IVa =» 9-A-OI peqmysiq :(vonfry, 
C(O cn POE TON ace Oia gues EO & rs got Jooo “rea: GY-AI-0€ ep [eUoIoeN 
Coe ve © Gl am 6 Ol © Ge @ i & 1 6 4 9@ i. see ApUrA ‘ppoo “tea[ ss $9-AT-G anbivg) ozrouv| 
imc mice Ol (0 8 Ce SF £ =f 66 ° S Ff Bb ue ‘<pnojg —-9-ITI-8Z ap ory ‘greuing 

IVLOLdSaH Vd Sid OW Id AHL AIT WAN TIHH OV HII Vd Lvs Ud OW [OO sIaMO[ TOYIVI AA aVq ARI] 


‘ON 
sdnorsy) Ajfioyng 


‘porjissepo A][NF JOU [erteyeur *(%¢ |=) Joqumu 
poyeurnjss = , SGT OF LOGBT “[Izerg UtoyyNOs puK [e1]UO UI SpOYJOUL SAISUGUT Y}IM apeUt s}uUNOd AjIVp 9ULOS JO suostiedwoy ‘[ aITaV] 


* 


*jso10F SnOnploep 
Teordox1u1as 


OD 
cS ODT ec 0 ies 0 V I 6G € IT il IL 6 G 0 I F yoy “1va[D 89-XI-& oned Ov “[ossvatyy 
*“SPlely BULIOMOTFT 
YUM 4so1of 
eules JO JURUUIOY 
CAL Sve. 0 st OL Sia 6 Ol I VI G eae & I G aly per teat 89-IA-06 royuRs opsrds iy 
v8r 8g 0 €l €.f O €G O (HG LIL UL cr tf €G O 0 G afeainale ULIE AN IB9TD) LO-ILA-LG ‘orreuey) O1ped 
*JSOLOF ULSITA 
esuep AAvoT{ 
9EL OF I OL Sly VI O vo 8 0 9 0 Or 0 € G SIF 204 Tea 69-A-V royuesg opsdsy 
LET IV 0 6 VI O 8 0 GG 6 I Ol 6 SI G G G qP WHEN EMS “HEM 89-IA-&G ‘elleg Bp OBdIa0U0Z 
SIS ORI th StS OYA 0€ TI OGY 96 € OF OG 8 skep BUIPeT[OO OOT “Bore vsoroy, BUG 1O¥ STV LOL 
‘avpiiiodsayy pue (ovurpeyy, A[[eroodsa) oepruevoATT Fo Ajo1vos 
‘TUITMOOT[OFY ‘TUIjOSserg ‘soydioyfy JO soUvpUNqY ‘“1oyyVIM AEploo jsIIF 9LOFoq “1oUIUINS 9}eT UL YV9G -HLON 
6L 8I 0 6 9 0 € 0 €I GI O 6 IL L 0 I i spar Oana e921) 69-IA-ST 
€8 8I G 6 L G V 0 wl WL @ L I 8 0 I it =P WE AES 1S) 69-IA-OT 
TL eI 0 8 GI € V 0 Cie Il I G 6 0 I I st OS easels) 69-IA-6 
66 ST € OO V 0 0c 8 S| 8 0 6 € € I ala tooo Apnojo Hed 69-AL-6I1 
90I_ €I 0 6 8 0 € 0 86 8 9 L Il cil ei it SIs Uae IBQTD) 69-AI-G 
GGL VG 0 OL cr TT € 0 Ic 8 L Or O er 8 SI IL Sty 704 Apnoyo Hed 69-AI-T 
SEL &¢ T VAL © /G\ YL 9 0 S6 (SILOS S| € fa iV V G ot VE ee Cole) OL-III-6¢ 
6II VG V vil 0 9 0 €G 8 I 9 0 SI F G I ap ULIBAN Apnoye Hed 69-II-9G 
LET 86 I Gl Ol © Or O Go OL G Cw) II 9 € it =F you, pApMOTO Hed 69-II-SG 
SEI GE I iL 6G S| 0 (i OU Ih ieee il 2 17 € I as yoy INO) Hed 69-IL-VG TE rs ood, 
6II &@ € Or 9f O € 0 66 til IE IE Ul 6 SI € G “te ena Apnojo Hed OL-II-€¢G Tey U TSP I22 
VIL & I 1 vit AL € 0 O Ge Ola lh UL 6 V € I alg ULE Apno[D 69-II-GG SNC ULSILA 
CGI GG G i Sih @ 8 0 Go OL 0 6 0 Jai it I IP ULES Apnoyo Hed 69-II-0G daays *“AavoH] 
Gol 61 € Wi Gl IE L 0 86 6 i OL O iE SI il G ale NOR ChE) 69-II-GI OIE oyards ay 
SII &6 0 cI If O g 0 vA WL I lL 6 OZ € G = OMT dSEIIO) 69-II-TT ESOLS TT eIOES 
Shi? OGM tik ARS VAG 09 T SSG) Lams 96 PV ce, OL 8 skup 9G “Or wou s}so10F UleyUNOUT TOF S'TV.LOL 
‘(WOSvaS TOWIVM) OUUINSpIuUl UL YVog ‘sa1oeds Je;[euIs 9Y} UL AjJOTIvA 
yonyy “Uorser [ooo ‘YsSIy sryZ FO ‘eUNVF [e}0} YorI A[teF Ysnoy} ‘vune;z A[Iep o}e1odnedep AT[eI9uey : ALON *JSOLOF 
faa) 9uvyUOUL UT 
fae v8 LE G I 9 0 sit) 9 € 0 G 0 L I 0 G a [ooo “Apnojo Hed 69-A-T doytH fy Jo 
eal G9 ST € L G 0 V 0 lL (6) T I v G G € al a CLS) 69-I-TE 784s stpodo.ned 
q 06 8T GI GL PV I GI O 6 ©) 0 T G SI € (G G ar SARE “AMY O) 89-II-€T pau ey 
=) [ecrdor} ystoyy 
Zz OIT II 0 OT &. O G 0 Che ts} 0 V G Sil I I anal yoy “Aurvi-1vat) OQ L-IIIA-G ‘oonquieuteg “Bunt J, 
a *sdUIeMS ‘jSOLOF 
c© [OANASA ANS | [PAE PONT 
ie 6tI TV S| 91 VY € cI O (SS te) I 6 I ZL I Il I Stectnata WBA “TBOTD S9-ILA-€ VEY 3s “ose 
A IvLOLdsSaH Vd Id OW Id BHL All WAN TAH OV HII Vad Lys ud OW sip SIOMO] A IOYIVI AA 97° q AqI[eVI0'T 
| sdnois) ATFroyNgG N 
> 
(panuyuog) “T AIdV 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


190 


L6G 6S 0 Al OV 6 iAoreme (6 cr L G GI G lea it Sake SEE BUPA qe GL-IA-0G 
soe 99 I SE rae 18) E Gy 9 I 8 I 0G G Geet Ce aatacae ec 119) GLIA-LT 
soc 19 0 I he Ge 9¢ 9 0 Olea SL AS I i Ee SES) TL-A-GG 
696 9L O Ot. Fr Ir 0O Le 8 it SIE Guess G Gea [ooo “Apnoyo wed 69-A-LT 
(qnID AqyunoD 
eIIseig UL WOOUIE};e Sopny{our) *SUIv91}S 
LOG OF Ik GI OG @G 6s 0 6G 8 IT €l G SE ss G G =e EEE qea1D 69-A-ST “SSOIOF OVA 
VLG &9 © Of Ly Si IL or -8 IT GIL [l CG Of G i Uneny 2eelo 69-A-FI T]HE SL OBR) 
AI Ly 0 OF &6 & 15) 6G 9 I Ont ZL 36 it I a Ue ESS) 89-A-L fete Row CIN AST 
teow Oe Cam i 0 te Y 0 6G G@ <6 © 8 SG Fr yoy “Aurer-1B9[) 89-II-€6 Buley op enbied 
ss¢€ «OGL 2 sc ORY Of TE 09 2L v IG € 2G TP 9 SAvp SUNoaI[09 FT ‘svare O[oAIMD pue eqedovied 10F STVLOL 
‘eUNeF YSOIOF Pou, ‘dnois suo ou jo souBpUNge [euOTdeoxyY ‘uuINJNe UI yeeg HALON 
evil €€ i vik Wit On L¢ & G 9 € Or O qT @ as wu “IvaJ S9-IIX-F1 
EG e-. BR - VI ut €& Sb 1 6¢ & T L G wit (0) I G alg wuIeM ID) 89-IIX-€T HEAT) ao) 
GOI 8S G WL Nb 6 6I O We I 9 G Ol 0 Il Clea Eee 8) 69-A-GI ada} aues PE 
GIL L6G G OL OLS ZL I GG OF iE 6 G ivf — 0) I i AP WEES AEST D 89-A-F SEuTIN: Cjosin 
SEI 8t G Sil G 0 NS = 12 il OL G&G -ELbe I T a WE SESS LO-A-V 
Gol 6G G&G Slee € 9 I 6¢ & € OI G&G OES G I ar Mees AEST) 89-A-€ SSO) EET 
GET OF G lL tS) ts 8 0 Ge ¢ I GI 6G Oe IT I ae WrEaS TeeTD. LO-A-€ 43TH See) 
€Or €1 ¥ €l 9 G G it 66 F O It € Sie i I = wea ‘APNo[D 89-III-G SEEN eqodovied 
‘OpeLied “SpyPerF 
“JSOIOF 19JVM 
10G 9 OGLE EGG JS Ib Lo € I € I vo O G Ge Sk 1ou HO) 69-A-1G = Pela Resa 
00c 19 0 6 IG G Gye 66 F¥ T v IT LI O IT Cos BROT BEY) 69-A-0G ma Neg SECS i 
SRS SSI i WIC [lk Mele I OSs 2 i G 0 8 Oo G = WaEherns “Apnoyjo Wed «= SOTTX=ST : BODE ESTE 
Wal /}8) € It €1 @ eo 0 €€ 9 0 Si I Sine T G = Ee SS ESI) 69-A-ET doyttH ‘StGIaS) 
SII 66 S Si GI G 8 I LEE v, T S IT FI O 0 IL es Ts ae Ne) 89-A-S SUE SE eae 
(OL6T “Hoqg) 
(saroeds ][euls UI sSBvaIOUI 9}0U ‘UOsIIeduUIOS Fo sasodind 10F ) sAep SuUT}OeT[I0OO ye (a}UOZIIOFY O[eg PUB BUsoRqIeg 0} 
GLa sce Ol VSG ZY 1s v8 ot 9L 6 ZL Ic ¥ ve L Si reprulis) auoz pus{q ey} UT ‘sep[eD eP sodog 10} STV.LOL 
Sci OLbeGile Go Zo, OSs Sh ASL tS) GG Of ke OE 48 skep Supoeyoo QT ‘eqwozuOH Ofeg IOF STVLOL 
‘psourjeq asimioyjo ‘faeprurpory Alje1oodsa ‘oeprusvoAT Jo Ayforvog ‘uuInyne AjIva UL Yvag :ALON 
vOSC £6 € 0c & ym OF 0 So fr oT —E OF I I cleats pee. ‘Apno[y) LO-A-8T : SPIEH 
09G ¥6 si Ce Sika =e OL 30 8¢ 8 G WN tS OO I I algae EEA UTS) LO-A-ET ORE. ares 
LEG G8 La OGe GIN “Wa Ge 30) Sige m Agk 26 6 G I I Seale peer) “Apnoyo aor L9-A-OT que HUST. Ara se 
O€G £8 9 KS EL 7 te ma! wr OT € Lis 2 G I if Teale [ooo “Apnopo Hed L9-A-T SUE, SneehO Hh vied 
*‘9u0Z pue[q 
“ySo10F QUBJUOW 
Ab 09, iG [lr IS 0) Wie ok € y 0 Wye 0 G oa [ee Apnoyo ec 69-A-TT aoe SSE Shs MS | 
906 146 S Gite Ole s oI O GG 6 iy i O “Wik G I T Sieiat: [ooo “Apnoypo weg LO-A-S SEOEN CHa 
“pruny sour JN 
*[OSSeIIJY UT se 
ySo10F} snonploeqd 
SPI St € 1) G SiG Oe Ocnr A: 0 S IT He 08 = Se yoy “IwvI[D 69-A-TE Toned OBS ‘serery 
IVLOL dSdH Vd Sid OW Id AHL AIT WAN IH OV HII Vd LVS YA OW "[00 SIOAMO[ IOY}VI AA a3eq APVIO'T 
‘ON 
sdnois) Afton g 
(panuyuoy) “T ATav 


191 


VOLUME 26, NUMBER 3 


*xVEIGx008 LS O9 «OSE GS +O0SE T 00G IG & YZ, SONS KD ES} TIZVYUA NVINOZYWY-VULXH WOH S'TIVLOL 
O6L «OSG SI FF xOOT € +xO00I T OOT ZIT 2Z 9F YP OL GG 8 (oyurg ‘dsy “eryeg) yse10f prumy [e10}T]-ULIpI|W 
G86 «OSE FG OF LST FY xOST T 7) S| Ye 66 G5 AM iv (seuo0z pus[q “[oxe) oyeuUR,Tg jo uorj0d OpP1IeD 
876 «OSE E€& Fr SIT € +xO0S1 I SII 9I 8 8€ PF ey OG 8 (s[¥}0} Stoyyne) AUTOIA pue OMoURf 9p O1Y 
Gi, GSS Gl ws AL 8 LIL 1 OG SIS 96 6€ 6L 9 (HH) ((serof yur 29 MOT) AjyrUOIA pue OIoURf ep OTY 
6SZ 26 I6¢ Wr €6 € 88 SOI FI 8 OS 9F 6L LZ (H) ySo10f ureyUNOW pue puL[MOT—o[Neg OBS Uo sey 
GES F6I &£& L6 9L & 86 I S9 GI G RS €¢ L € (AH) 3Se10} [eordon puryMoy—oonquivutog U1s}sey 
(OL6L ‘Heqny Ulo1Z UoYe} S}SI]T Gay? 4S17,7) SuI~eTOO sivaA AueUu I2AO STV LOL IYVNOIOUY 
ySoLOF YSLOUL AAVOTT 
9Ice 19 V 6 9¢€ 0O 6 0 LG iL. 6I TI 6c TI G S inal AUIVI 0} IvI[D GLIA-L :eImOpuoY “OYUIZory 
‘aevul[eyy, JO A}orvos ‘ovurpeyduiANY pue oeuLtAyeg jo souvpunqe [euondeoxy :q LON 4 Sean cea 
Il€ 78 9 Sl SS G Wie i 98 IL O Gl. TL fi? {S I G ela yoy “Ive GLIIA-€ pue[qstH > OSSOIT) 
606 L8 L si Of i 6 I Z8 $1 O GI. TT Iv I G she yoy “IeVelD GLTA-G Ove “M OYfeULIoA 
org 0F nYD Op 
LOG GL g VI Iv T vI oT oe iit © ‘iL UL Gh OS I G rar yoy “Ies[D GLALATT OWES Wor [Ler], 
€eS «OLT GG OG O8 G&G os Tt 3 - iL Gc Vv Sy 6 G skep suUIde[[00 PP ‘vore THING 10} STVLOL 
‘sotoods o1vil JO ‘OU posoNnpey ‘evulTeydWANY pue ovuULIA}eg jo soUepUNnqY ‘UUIMzNe 9}3uT UL Yeog :ALON 
G16 GL &S VI GY G 9f O Lo 6 I 9I G 8E G I G + yoy “IRs GLIA-9G 
GO& OOTL LZ LI GY G G I 8S 6 € VL € 9€ € G I ai : yoy “Iva OLTA-ST 
G6G 8 9 CI OF G ES 19 OL & €l &G 1% G I = 304, ‘Apnojo Hed OL-IA-6 
GIG LS I Il 96 G&G Vv 0 VS OL G VIL G GG G G I =P 70Y4 Apnoyo Hed OL-IA-S 
€LG €6 V €l €F G 8 0 Sia ORS: GI G €€ € G I an vate rez[D ILIA-¥ 
O€G IL € VI 9F G V 0 8r 8 € GL G ve tT G I ale Auter “Apno[p) PZFASTE 
VLG 9L gi GI LS G OL O ss G GI I 66 G G G =F UBM “Te9] 69-A-LG 
I9€ GG6I & St iS Se v9 OL §& €L G Se S G :NS+ 44 ; 
LOE £6 V VI OF G yI tL 19 OL € GL G 8€ ¢G ve Set aF 704 MO) 69-A-9G : 
6SG 62 V CI GE G Ie TI ci LIE OL &G GG iL G G ain 704 REED) 69-A-GG , set Pons eta! 
IvG 7S € Sl Ge Lis 8S 6 S €l G&G ComenG G G ote 70 BEND) 69-A-VG , spoom Arp 
LIG LV € GL Gr G VI O Ww 8 G GI tT Go € G G ae yoy el) 69-A-EG SPOOM 4STOUL 
OGG €S V Sl AG I felt UI ww G&G I Il O Se FL G I =+ eee) res) 69-I-7 osreds pue 
vV6T 2S € GI L6 &G 9It 0O 96 9 I 8 I ce t. G I fe AUT Apno[) 69-1-G AAvoH CORON) 
O61T 9S € OL O€ O OL O 66 PF I OL O F6 T G I aI 104 Apnoyo Hed 89-IIX-8¢G oOveW SEMA) 
6L1 GV V €L 86 TT il O 8S '& I Ol 0 |r T G G = yoy “Apnoypo y1eg 89-IIX-GG 2p peas ey ee 
3 AAGOFI +: OSSOIH 
98IT IS I Il 66 @G VI O OS 8 I 9 G Ic O I G set yoy “IBID 69-A-86G OF “8}UBOTA OBS 
Coe OSGe Cle ce OSs 1 O€L T £6 819 GG Sh Sl wy sAvp BUT}ET[OO EF “vorIN BITISeIg 10} STVLOL 
‘Days, Ajjetoodsa ‘aeprtuevoAT pue ‘avuljeydwiANY ‘ovuliAyeg fo souepunqy -‘uunyne ur Yyweg ‘ALON *dureMs 
“ysi1eul ‘jso10F 
pruny MoT -4d 
9GI 9€ 0 VI 6 G L I GG 9 0 Si 6 0 G G Slmate uieM ‘Apnopo Ped 89-II-VG “oorsQ [007 wipe f 
IVLOLdSaH Vd Gd OW Id SHL AIT WAN ISH OV HII Vad Lys Ua OW "[00 SIOMO[Y TOYIEI NM 97e qd AY[BOO'T 
‘ON 
sdnois) Aypioyng 
(panuyuoy) ‘TT AIdvL 


192 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Regional and Seasonal Comparisons by Subgroup (Table 1) 


It is of interest to note here the seasons and areas in Brazil, and in other 
regions in the Neotropics, where, in our experience and based upon these 
methods of intensive collecting, there may be expected maximum num- 
bers of species in each of the major groups of diurnal Lepidoptera. 

Morphos in southern Brazil are most common in Santa Teresa, Espirito 
Santo; all eight species which we recognize in the extra-Amazonian area 
may be found flying together there in April. Very few other such small 
areas harbor over six species, and in more subtropical climates these rarely 
overlap during their single yearly broods in summer. If the same criteria 
of conspecificity are applied in other regions, eight species may also be 
captured (though possibly not all at the same season) in some areas of 
the eastern slopes of the Andes in Peru; and seven species may be found 
flying together at one time in many warmer parts of tropical America, 
where most species are multibrooded. 

Satyrines (including brassolines ) seem to be most diversified in areas 
of alternating dry scrubby woodland and moist riparian forest, with a 
strong additional element of open grassland and marsh, such as the 
Chapada de Guimaraes and Brasilia areas, with 54 and 58 species 
known to date. Some larger areas have the required diversity to harbor 
nearly 100 species in this subfamily. A few areas in the Andes may 
surpass 100 species, though it seems unlikely that more than fifty could 
be captured in a day. The brassolines alone seem most common in hilly 
areas where neighboring faunal regions meet; the key to their observation 
and collection is always the discovery of their preferred afternoon and 
evening flyways and the liberal use of traps with preferred local bait. 
The area of Santa Teresa has at least twenty species, of which ten have 
been observed in a single day; we know of no other area in the Neotropics 
with a similar species abundance. While satyrines and brassolines in 
general show less strict seasonal variation than other nymphalid groups, 
individual species often show marked alternations of heavy broods and 
essentially complete absences. A few temperate-zone species are single- 
or double-brooded, in summer only. 

In Brazil, the maximum number of species of Ithomiinae is reached in 
the median-littoral region, a broad area of moist coastal tablelands in 
southern Bahia and northern Espirito Santo which shelters many endemic 
species. Here, 30 species can be found in a single “pocket” in winter, and 
25 to 27 can reasonably be collected in a single day. These numbers do 
not compare with those found in the upper Amazonian and Orinocan 
tributaries on the slopes of the Andes in Bolivia, Peru, Ecuador, Colombia, 
and Venezuela, clearly the principal center of geohistorical evolution in 


VOLUME 26, NUMBER 3 193 


the subfamily. In the Chanchamayo valley (La Merced), Peru, and in 
north-central Venezuela, the author discovered nearly 50 species present, 
of which 40 could reasonably be captured in a single day, especially with 
the use of dried Heliotropium plant (Beebe, 1955; Masters, 1968) hung 
at favored locations within the “pockets.” Collection of ithomiines is 
generally best in the dry season, though a few montane species are most 
common in the warmer, moist summer season. Many species in the 
family are notable for their accentuated and unpredictable variations in 
populations and abundance from season to season and year to year. The 
localized dry season “pockets,” in which all of the species tend to fly 
together, are much diluted during the wet seasons, and occasionally switch 
their location dramatically from one year to the next. 

The maximum number of species of heliconians found in one locality 
of extra-Amazonian Brazil is 17, in Santa Teresa; 16 of these have been 
observed in a single day. Again, this compares poorly with the numbers 
present in some parts of the Andean slopes of Ecuador and Colombia; on 
the Rio Negro in Meta, Colombia, over 30 species have been found, of 
which 22 were observed by the author in two days (Brown & Mielke, 
1972). Nymphalines, many of them attracted to bait, have been found 
most commonly in extensive and very moist, slightly disturbed forests 
at the base of the mountains near Rio, and in the interior near Brasilia, 
Buriti (Chapada de Guimaraes ), and Rio Vermelho (Amazon-type high- 
land forest in western Mato Grosso), but the total lists for an area do not 
vary much from an average of 80 (locality) or 110 (larger region). The 
greatest abundance of Nymphalidae, broadly speaking, has been observed 
in Rio Vermelho (161 species in one day ); however, if total numbers ob- 
served (as a smaller number per day) over a long period is considered, 
Santa Teresa, which blends four major faunal regions, is much richer. 

Almost all tropical nymphalines, including heliconians and charaxines, 
are to be found most commonly in late summer and fall in cooler areas, 
and late winter in hotter regions. The genus Euwnica contains many 
species that are practically limited to one brood in the late winter. A few 
larger species of Charaxinae are double- or triple-brooded only, showing 
much variation in abundance and freshness of individuals caught at 
different seasons. 

For the extensive collection of Theclinae, no area can compare with 
the Brazilian cerrado in the early dry season. Although this habitat has 
much akin faunistically to an open forest, the trees rarely exceed 20 feet 
in height, and thus treetop-loving species can be netted easily, together 
with those preferring low flowers and grass. The spacing of the low trees 
assures abundant sun all day long, and the interspersal of occasional 


194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


denser growth (cerradao), more open areas (campo cerrado), and moist 
riverside and headwater woods guarantees the presence of a very wide 
variety of Theclas. The 47 species in this subfamily captured by the author 
and Karl Ebert in Itirapina, Sao Paulo, a southern outpost of the cerrado 
within the blend zone, in May 1967 (Ebert, 1970) represented an early 
and successful test of some of the methods described in this paper. In a 
good year (such as was 1972) in the Brasilia area, a day’s list of nearly 100 
Theclas should be attainable by a party of three or four collectors; the 
maximum to date, with two persons, is 84, and a total lycaenid list of 137 
species. 

The only areas we have seen to match the forests of the Chapada de 
Guimaraes for Riodininae are the lowland woods near Manaus, where 
50 species were captured in two days in January 1970, and the swampy 
woods near Belém, where 45 species were found in two days in late 
December, 1970. Presumably, an area could be found in the lower middle 
Amazon which would produce nearly a hundred species of these brightly 
colored small butterflies in a day. Diversity of flora is a good clue to 
abundance of Riodininae, and favored male patrolways (hilltops and 
open moist woods are best) must be located, and hours of flight activity 
ascertained, to give good collecting. 

In Brazil, Papilionidae are most abundant at the very beginning of the 
rainy season in September, especially in the upper Paraguay River basin, 
where over 20 species could be captured in a day; similar or greater 
numbers should be attainable in many areas of the upper Amazon basin, 
as well as in the foothills near Rio de Janeiro at this same season (24 
species recorded in Xerém, not all on one day). The fall and winter are 
very poor in Papilionidae, except for occasional Parides in warmer areas. 

Pieridae have been found most abundantly in the same areas and 
seasons as Papilionidae; the diversity is greatest in the foothills near Rio 
where mountain and lowland species mix (44 species recorded, 22 in a 
single day in Xerém). Outside Brazil, somewhat greater numbers may 
be encountered in favored seasons on the eastern slopes of the Andes. 

Hesperiidae appear commonly in almost all Neotropical areas with 
diversified ecological biotopes (though much more rarely in extensive 
virgin forests); a daily list of over 100 species should be attainable in 
any of a number of regions in the good flower seasons. Table 1 shows 
an extraordinary correlation of numbers of skippers captured with flower 
abundance, which should be taken to heart by those who specialize in this 
‘amily. Prominent in the flower groups that bring these butterflies out 
of the forest and concentrate them in accessible areas are members of the 
“upatoriae, which flower principally in fall and winter in Brazil. 


VOLUME 26, NUMBER 3 195 


SUMMARY 


A method for the conscious maximizing of selected daily butterfly lists 
in southern Brazil has permitted much new information to be obtained 
on the occurrence and biology of the small, localized, and/or high-flying 
species in the fauna, which tend to be least known. It also has provided 
a reasonably standardized basis for comparisons between different lo- 
calities and seasons, both in overall number of species present and in 
relative abundance of the various subgroups of Rhopalocera. General 
conclusions on extensive, intensive, and seasonal frequencies closely 
match those presented by Ebert (1970), though the collecting methods 
used are fundamentally different. 


ACKNOWLEDGMENTS 


The author is indebted to Dr. Heinz Ebert for the original stimulation 
of the work leading to this paper, as well as its revision at several stages 
of development. The collecting days described in Table 1 were realized 
with the assistance of a number of experienced and uncomplaining col- 
leagues, especially Dr. Ebert and his son Karl, Olaf Mielke, Claudionor 
Elias, Stan Nicolay, Nirton Tangerini, and Luis Otero; many other per- 
sons aided in the localization and exploring of the areas mentioned. Out- 
side Brazil, the author was priveleged to be introduced to the Andean 
fauna by Dr. and Mrs. L. W. Harris (Peru), Dr. E. W. Schmidt-Mumm 
(Colombia), and Drs. K. Negishi and S. S. Tillett (Venezuela). Financial 
support of the chemical study of Brazilian insects by the Conselho 
Nacional de Pesquisas (of which the author is honorary lecturing in- 
vestigator, “Pesquisador-Conferencista” ), the Banco Nacional de Desen- 
volvimento Econémico, the Conselho de Pesquisas e Ensino para Gradua- 
dos of the U.F.R.J., and the National Science Foundation (Grants GB 
5389 X and GB 5389 XI), is gratefully acknowledged. 


LITERATURE CITED 


Breese, W. 1955. Two little-known selective insect attractants. Zoologica (New 
York) 40: 27-32. 

Brown, K. S., Jr. 1970. The Heliconians of Brazil (Lepidoptera: Nymphalidae ). 
Part I. Rediscovery of Heliconius nattereri in eastern Brazil. Entomol. News. 81: 
129-140. 

1972. The Heliconians of Brazil (Lepidoptera: Nymphalidae). Part III. 
Ecology and biology of Heliconius nattereri, a key primitive species near to 
extinction, and comments on the evolutionary development of the species of 
Heliconius and Eueides. Zoologica (New York), in press. 

Brown, K. S., Jr. & O. H. H. Mierke. 1967. Lepidoptera of the Central Brazil 
Plateau. I. Preliminary list of Rhopalocera. J. Lepid. Soc. 21: 77-106, 145-168. 

1968. Lepidoptera of the Central Brazil Plateau. III. Partial list for the 

Belo Horizonte area, showing the character of the southeastern “blend zone.” 

J. Lepid. Soc. 22: 147-157. 


196 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


1972. The Heliconians of Brazil (Lepidoptera: Nymphalidae). Part II. 
Introduction and general comments, with a supplementary revision of the tribe. 
Zoologica (New York) in press. 

Brown, K. S., Jn. & R. Ferrera D’ Atmema. 1970, The Ithomiinae of Brazil 
(Lepidoptera: Nymphalidae). IJ. A new genus and species of Ithomiinae, 
with comments on the tribe Dircennini d’Almeida. Trans. Amer. Entomol. Soc. 
96: 1-17. 

Expert, H. 1970. On the frequency of butterflies in eastern Brazil, with a list of 
the butterfly fauna of Pocos de Caldas, Minas Gerais. J. Lepid. Soc. 23: supple- 
ment 3, 1—47. 

Fox, R. M., A. W. Linpsay, H. K. Crencn & L. D. Mutter. 1965. The Butterflies 
of Liberia. Mem. Amer. Entomol. Soc. 19: 1—438. 

Masters, J. H. 1968. Collecting Ithomiidae with Heliotrope. J. Lepid. Soc. 22: 
108-110. 

Mieke, O. H. H. 1967. Lepidépteros do Planalto Central Brasileiro. IV. Notas 
complementares sdbre os Hesperiidae da regiao e descricao de cinco espécies 
novas (Lepidoptera, Hesperiidae). Rev. Brasil. Biol. 27: 125-134. 

1968a. Lepidoptera of the Central Brazil Plateau. I]. New genera, species 

and subspecies of Hesperiidae (Rhopalocera). J. Lepid. Soc. 22: 1-20. 

1968b. Lepidoptera do Planalto Central Brasileiro. V. Novas espécies de 

Hesperiidae e anotacdes sdbre outras espécies conhecidas. Rev. Brasil. Biol. 

28: 447-455. 


AN UNUSUAL MOTH IN CENTRAL ILLINOIS 


After working on my car, at about 1400 on 14 November 1971, I drove it through 
a parking lot across the street from my home. I then noticed a large moth clinging to 
a building, about ten feet off the ground. A closer look revealed that it was a large 
noctuid. Immediately, I opened the back of the car, grabbed my net, made a lunge 
at the moth, and managed to capture it. 

As soon as I got home, I checked my revised edition of Holland’s Moth Book, and 
found that the moth I had just caught was Thysania zenobia Cramer. After reading 
that it was a “South American moth,” I knew that I had caught something quite un- 
usual for central Illinois. However, a moderately strong wind had been blowing from 
the south for several days preceding the capture, and as the specimen was slightly 
ven this probably explains its presence this far north, especially at such a late time 
of year. 

I am sure that this capture does not constitute a state record after checking with 
the University of Illinois and Eastern Illinois University; but it might be a new 
record at least for Coles County. If any members know of a previous capture of this 
species in this county, I would sincerely appreciate hearing from them. 

CuHartes K. Swank, 616 S. 15th St., Mattoon, Illinois 61938. 


WING-SHAPE AND ADULT RESOURCES IN LYCAENIDS 


Various small species of blues (Lycaenidae: Lycaeninae) are characterized by 
rather narrow elongated wings—a Nearctic example being Zizula cyna Edwards. 
n September of 1971 some observations were made on a related African species, 

ila hylax (F.) in the Gombe Reserve on Lake Tanganyika, Tanzania. 


VOLUME 26, NUMBER 3 197 


Fig. 1. Zizula hylax crawling into Asystasia flower. Margin of hind wing would 
be even with lip of flower when butterfly has fully entered. The length of the 
forewing in Z. hylax is about 11 mm. 


September is near the end of a long dry season at Gombe, and relatively few 
flowers were in bloom. Prominent among those blooming especially along water- 
courses, was a small herb Asystasia gangetica (L.) T. Anderson in the Acanthaceae 
(a largely tropical family related to the Scrophulariaceae). Those flowers were 
repeatedly visited by Z. hylax individuals. Their slender wings, when folded back, 
permitted the butterflies to crawl deep into the corolla to get at the nectar (Fig. 1). 
The fit was sufficiently tight to make it clear that even slightly wider wings would 
effectively prevent this behavior. 

Zizula hylax has a weak flight compared with many small relatives with broader 
wings and it does not glide. It seems highly unlikely, therefore, that the high aspect 
ratio of the long slender wings produces a significant selective advantage through its 
aerodynamic properties. On the other hand, studies by our group in California, 
Colorado and Trinidad increasingly are showing the great significance of adult re- 
sources to the dynamics of butterfly populations. It seems most likely that the 
adaptive significance of wing-shape in this case is related to access to nectar. Further 
observations on related species would be most interesting. 

We wish to thank Dr. Dennis Parnell, California State College at Hayward, for 
determining the Asystasia. This work has been supported in part by a grant GB19686 
from the National Science Foundation. 

PauL R. ERLICH AND ANNE H. Enruicu, Department of Biological Sciences, 
Stanford University, Stanford, California 94305. 


198 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


NEW DISTRIBUTION RECORDS FOR CERATOMIA HAGENI 
(SPHINGIDAE ) 


To date Ceratomia hageni Grote has been recorded from Arkansas, Kansas, 
Mississippi, Missouri, and Texas (R. W. Hodges, Sphingidae, in R. B. Dominick 
et al., The Moths of America North of Mexico, Fascicle 21, 1971). The larval food 
plant, osage orange [Maclura pomifera (Raf.) Schneid.], has a much greater range, 
having been planted extensively in the Mississippi valley and eastern United States 
including New England. The range of osage orange suggests that C. hageni should 
occur over a much greater area than previously recorded. 

Upon reviewing my collection of Indiana Sphingidae, I discovered a specimen of 
C. hageni taken 3 Sept. 1960 at lights in Indianapolis. The specimen is a male with 
a wing expanse of 89 mm and a wing length of 41 mm. Another male specimen of 
C. hageni was taken at Springfield, Ill. on 28 June 1959) (collector unknown). This 
specimen is in the Natural History Museum of Los Angeles County (J. P. Donohue, 
pers. comm. ). 

It is probable that this species has been overlooked since it is similar in macula- 
tion to C. undulosa (Walker) and C. catalpae (Boisduval), which are generally 
distributed over the eastern part of the United States. C. hageni can readily be 
distinguished from other species of the genus by the green to yellow-green shading 
of the primaries, midtibia possessing a series of apical spines (sometimes obscured by 
the vestiture), and fasciculate antennae in the female. Genitalic differences are 
detailed in Hodges (1971). 

This species should be looked for wherever osage orange occurs. The adult is a 
late flier (Howe, in Hodges, 1971). It is attracted to lights, but does not begin to 
fly until after 2200. The immature stages need to be studied more completely. 
Stallings & Turner (1944, J. Kansas Entomol. Soc. 17: 29-31) gave a brief descrip- 
tion of the egg and larva. One supposed larval specimen is in the United States 
National Museum. The pupa is unknown. Much more information concerning the 
distribution and life history of C. hageni is needed. 

CuHArLes M. Franxuin, 815 Lincoln St., Indianapolis, Indiana 46203. 


FURTHER NOTES ON W. H. EDWARDS SPECIMENS IN ILLINOIS 
MUSEUM COLLECTIONS 


In view of the taxonomic and historical importance of butterfly specimens studied 
and named by W. H. Edwards, the location of such material should be placed on 
record. These notes supplement earlier discussions of the subject by Brown (1964, 
Trans. Amer. Entomol. Soc. 90: 323-413) and Irwin (1966, J. Lepid. Soc. 20: 
156-162). The Edwards specimens in my collection listed in the latter paper 
have been permanently deposited in the Illinois Natural History Survey (Irwin, 
L971, J. Lepid. Soc. 25; 83-64), 

The Natural History Survey collection already contained eleven other Edwards 
specimens. Nine of these were in the collection of Selim H. Peabody, Regent of the 
University of Illinois from 1880 until 1891, who corresponded and exchanged butter- 
flies with Edwards. The Edwards holograph labels on the Peabody specimens consist 
only of sex signs and localities, without the names of the species. In this respect 
they are so unlike Edwards’ usual labels that I suspect that the species names were 
cut off the labels by some later worker. This may have been F. H. Benjamin, since 
most of these specimens bear Barnes and Benjamin determination labels beside the 
remaining portion of Edwards’ original labels. 

In addition to these nine specimens, there are two other Edwards specimens in 


VOLUME 26, NUMBER 3 199 


the Survey collection whose provenance is unknown. They may have originated 
from Benjamin Dann Walsh or Cyrus Thomas as well as from Peabody. Walsh and 
Thomas were state entomologists of Illinois and were correspondents of Edwards. 
Their collections were the nucleus of the present Survey collection; some of their 
material may be still extant in it, although it is no longer recognizable. One of these 
additional specimens is a male Euptychia gemma (Hibner) taken by Edwards him- 
self in West Virginia on 11 June 1878. 

I have placed identifying labels on yellow paper reading “From/W. H. Edwards” 
on all of these specimens and they, with the similarly labelled Edwards material 
that came to the Survey with my collection, make it one of the largest repositories 
of Edwards butterflies outside the Carnegie Museum, where the Edwards collection 
itself is preserved. 

The Herman Strecker collection in the Field Museum of Natural History contains 
more than 70 specimens bearing Edwards’ holograph labels. I believe that most 
of these did not originate from Edwards, but were either determined by him for 
Strecker, or before Strecker received them. One specimen bears Strecker’s label 
stating that he had sent it to Edwards for naming. A number of these specimens 
were collected by David Bruce in Colorado, and Bruce is given as their source on 
Strecker’s labels. Many of the Edwards labels in the Strecker collection contain only 
the name and sex of the butterfly, with no statement of locality or collector, and 
their pins are not the type Edwards used. All of this indicates that Edwards was not 
their original source. Two specimens bear label reading “This is the writing of/ 
W. H. Edwards. A.” “A” was Eugene Murray Aaron, who curated the Strecker col- 
lection at the Field Museum during the late 1930’s. 

The Strecker collection contains other material which does not bear Edward’s 
own labels, but which he had studied as evidenced by Strecker’s labels. Some of 
this latter group of specimens have been identified as members of the type series of 
their taxa and have been designated and labelled as holo-, lecto- or paratypes by 
Brown in his studies of the Edwards type material (loc. cit. and subsequent papers ). 

Roperick R. Irwin, Illinois Natural History Survey, Urbana, Illinois 61801. 


BOOK REVIEWS 


MICROLEPIDOPTERA PALAEARCTICA: VOLUME 1. CRAMBINAE, by Stanislaw Bleszynski. 
1965. H. G. Amsel, H. Reisser and F. Gregor, eds., in 2 parts: text (in German), 
xlvii + 553 p.; 133 col. pls., Verlag Georg Fromme & Co., Wien. Price: Austrian 
schillings 1560, Deutschmarks 240 (less if entire series is bought). 


The distinguished author specialized on the pyralid subfamily Crambinae for 15 
years, paying particular attention to the Palaearctic, Ethiopian, Indoaustralian and 
Neotropical faunas. He was the first European worker to make as complete as 
possible use in classification of the male and female genitalia, as well as of all other 
available characters. He also travelled very extensively, visiting all museums and 
collections where there might be types, as a result of which he was able to correct 
a great many errors and misconceptions that had crept into the literature since 1758. 
In all this he strictly followed the International Code of Zoological Nomenclature, 
which gave his work a sound basis that will ensure its endurance. Perhaps his most 
valuable work was breaking up the overgrown “wastebasket” genus Crambus into 
which nearly everything crambine had been thrown for over a century (it contained 
74% of all species in Staudinger and Rebel), resurrecting a number of Hibner’s 
genera and naming others himself. His most intensive work was, of course, on the 
Palaearctic fauna, and is represented by the volume being reviewed. Dr. Bleszynski’s 


200 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


untimely death in 1969 in an automobile accident was a great loss to science, as well 
as a great personal loss to many of us. This reviewer had the pleasure of meeting Dr. 
Bleszynski at various times in Vienna, London and New York, and of working closely 
with him for many years through a voluminous correspondence. He takes this op- 
portunity to acknowledge the enormous benefit which he received. Any day the mail 
might bring a letter telling of the discovery of a long-lost type in Leningrad or Berlin, 
or pointing out that the so-called type in some museum was a specimen caught 
years after the publication of the original description! 

In this volume the Palaearctic Crambinae (including what some authors separate 
as the Ancylolomiinae) are divided into 370 species in 49 genera, a far cry from the 
status of the classification in the Staudinger and Rebel Catalog, the last to cover the 
whole ground, where the totals are 165 species in 11 genera. Some genera seem 
somewhat heterogeneous, but at least the problems are clarified, and future workers 
will doubtless split still further. The type species of all generic names and the 
type specimens of all nominal species are cited, with the dates and methods of their 
designations. Generic and species synonymies and keys are given, and there is an 
excellent terminal bibliography. 

The Palaearctic fauna is, of course, extremely important for the study of the 
Nearctic, since at least 10 genera and 11 species are common to both, while other 
Nearctic species are extremely close to their Palaearctic counterparts. Obviously we 
must have full knowledge of the Palaearctic fauna in order to understand our own. 
Bleszynskis zoogeographic survey is especially interesting in this respect. All of 
this is true, of course, not merely for the Crambinae but for all of the Nearctic 
microlepidoptera that have Palaearctic relationships. 

The planning and producing of this book, and presumably also of the volumes to 
come, reflect the greatest credit possible on the editors who conceived the series. 
All sorts of unexpected things make the volume far more usable and valuable. 
These include a lexicon of terms in German, English, French and Russian, and an 
alphabetical list and index of geographic localities and another of abbreviations. The 
illustrations are divided into three groups: color paintings of adults, male genitalia 
and female genitalia. In this way illustrations that must be carefully compared with 
each other are on the same, or adjacent plates, which greatly facilitates their study. 
I was greatly intrigued by the forethought that provided three bound-in colored 
ribbons with which to mark the places of the illustrations of a species to compare 
them with each other. Another superior feature is the numbering of all the species 
in a single sequence, not in separate sequences by genera. Thus, species No. 136 in 
the text is illustrated by color painting No. 136, male genitalia no. 136 and female 
genitalia no, 136, which greatly facilitates getting the picture of the species as a 
whole. (As far as I know this idea was first used by E. P. Felt at Cornell in an 
article on North American Crambinae in 1884.) The essential data of each specimen 
illustrated are given on the legend page. The color paintings, the work of Dr. 
Gregor, are most beautifully done and printed, and are a pleasure to use. The 
whole volume sets a very high standard, hitherto unattained. 


ALEXANDER B, Kiors, The American Museum of Natural History, Central Park 
West at 79th St., New York, New York 10024. 


AUSTRALIAN Burrerriies, by Charles McCubbin. 1971. Thomas Nelson (Aus- 
tralia) Ltd., Melbourne and Sydney, Australia. vii-xxx + 206 p., 30 text figures, 
numerous colored illustrations, 1 map. Price: about $30.00 (U. S.). 


BUTTERFLIES OF THE AUSTRALIAN REGION, by Bernard D’Abrera. 1971. Lansdowne 
Press Pty. Ltd., Melbourne, Australia. 414 p., 2 text figures, numerous colored 
illustrations, 3 maps. Price: about $40.00 GUEGS. 


VOLUME 26, NUMBER 3 201 


These books are the first since “Seitz” was published about half a century ago 
which illustrate the butterflies of Australia and the Australian region, respectively, 
in color. It is somewhat unfortunate, however, that the two books were published 
in the same year, because the coverage of Australia in duplicate could affect the 
sales of both books. This overlapping is more apparent than real, though, since 
the McCubbin book deals with the butterflies of Australia, Tasmania and the off- 
shore islands only, whereas D’Abrera’s book deals with the entire Australian region 
from New Zealand to Weber's line and north to the Solomons and Fiji. 

The treatments are quite different, too. McCubbin deals with fewer than 400 
species and is able to consider these in much greater detail. D’Abrera, by contrast, 
must consider at least five times as many names, and the coverage possible under 
such conditions can be little more than skeletal, concentrating on the identification 
of the entities, not on their habits or habitats. McCubbin has been able to give 
much fuller insight on what the butterflies are like in the field, and in general, what 
they “do for a living.” These firsthand observations make the McCubbin book a 
very readable one. 

McCubbin has been able to include the Hesperioidea in his book, whereas space 
limitations have not enabled D’Abrera to do so. Perhaps because less is known about 
the habits and life histories of the Australian skippers, the section on the Hesperiidae 
in McCubbin’s book is perhaps the weakest one: if any portion of the book seems 
to have been taken from others’ observations, it would be that section, with a few 
notable exceptions where it was obvious the author had firsthand experience. Much 
of the identification and observational criteria cited seem to have been drawn from 
Evans Catalogue and from various of Waterhouse’s papers. D’Abrera did not cover 
the skippers, as stated above, but he does mention the possibility of doing this 
group in a later companion volume. I have a difficult time seeing how such a book 
could be done in the same format as the present one: there just aren’t that many 
hesperioids in the Australian region. Perhaps he would have been better advised to 
either cover the skippers in the present book, thereby adding fewer than 100 pages 
to it, or perhaps doing a second volume including the Lycaenoidea and expanding 
his discussions of all species. As it stands now, the weakest part of the D’Abrera work 
is the coverage of the Danaidae, especially the genus Danaus. The other danaids 
are at least passably covered (though Euploea could use a bit more expanded 
coverage), but the entire book could have been strengthened by adding another page 
or two of plates and discussion to Danaus, perhaps at the expense of Ornithoptera. 
Books are bought for their illustrations, however, and the bird-wings are among the 
most spectacular of the butterflies—this may have governed the decision to expand 
the Ornithoptera coverage beyond that that was necessary. 

Both books are remarkably free of disturbing typographical and other errors. The 
errata sheet that came along with our copy of D’Abrera rectifies many of these 
errors, but no such sheet accompanied the McCubbin volume. As is always the 
case, a few such errors have eluded the authors’ scrutiny: McCubbin has “Klak” 
(rather than “Kluk”) as the author of Danaus, and he has placed Pantoporia c. 
consimilis (Bdv.) in the correct genus in the text (p. 60), but on the plate the 
butterfly is listed as “N.” (presumably Neptis) c. consimilis (it is very possible that 
the plate was done before Eliot’s revision of the Neptini was done); D’Abrera 
describes as new Hypochrysops emiliae on p. 335, figures it as “emilia” on the 
preceding page (rectified in the errata sheet) and lists it as “emilia” in the index 
on p. 407 without correction. To further belabor the authors for these small errors 
would be nit-picking. In one instance, however, McCubbin has corrected a long- 
standing nomenclatorial error: Bethune Baker described Ogyris waterhouseri in 1905 
(Trans. Entomol. Soc. London: p. 273), and McCubbin has correctly cited it; 
D’Abrera has followed the logical, but incorrect emendation to “waterhousei.” None 
of these errors is serious, and they in no way detract from the usefulness of either 


book. 


202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


McCubbin’s illustration of food plants is very useful, particularly for the collector 
who might want to find the species in the field. This is an innovation which could 
be followed to advantage by other illustrators in the future. Perhaps the most 
impressive coverage of the D’Abrera book involves the Lycaenidae: he has made 
sense of a group that has stymied many of us attempting to identify these beautiful, 
but confusing, insects. The fact that many of the new taxa described by D’Abrera 
in the text are lycaenids indicates that he probably specializes in the group, and his 
careful handling of the Lycaenidae shows an expert’s touch. 

Both books owe much of their usefulness to their colored illustrations. It is difficult 
to imagine how the paintings depicted in McCubbin’s book could have been im- 
proved upon, and I only wish I had the opportunity to see the originals! The 
illustrations in D’Abrera’s book were derived from Ektachromes and are subject to 
some color shifts on some, but not all plates. The color shifting may be the fault 
of the original transparencies or they may be the fault of the printer — there’s no 
way of determining this from long distance. Comparison of the plates with actual 
specimens at hand shows that the color shift, when it occurs, involves the blue; the 
blue shows up poorly in the figures of the pierid genus Saletara, but the blue is too 
intense in several of the Satyridae. If one does not take absolute color values too 
seriously (and it would be dangerous to do so in taxonomic work, anyway), the 
D’Abrera plates lose none of their usefulness. 

From a purely personal standpoint, I wish D’Abrera had used either “upper surface” 
or “dorsal” for the “recto” he utilizes, and that he had used either “under surface” 
or “ventral” for “verso.” His terms are not in general usage, and the reason for using 
them instead of more familiar terms is obscure. 

The McCubbin book supplements the fine little Australian Butterflies by I. F. B. 
Common (1964, Brisbane, Jacaranda Press) with the excellent colored illustrations 
and in more detailed discussions of the species. D’Abrera, by contrast, has put out a 
book that has no “post-Seitz” counterpart, with the exception of Barrett and Burns’ 
Butterflies of Australia and New Guinea (1951; Melbourne, N. H. Seward), a book 
with almost nothing to recommend it scientifically and little more esthetically. 

I would very much like to see D’Abrera (or someone equally competent) do a 
companion volume of the butterflies of Indonesia, or at least from Celebes westward, 
where the book could bridge the gap between his present volume and Corbett and 
Pendlebury’s Butterflies of the Malay Peninsula. If this were done a great deal of 
the mystery surrounding the Indo-Australian fauna would be cleared up for those 
of us interested in these butterflies, but without ready access to the great European 
collections. It is very difficult to envision what McCubbin can do for an “encore” 
to produce another book of the quality of the present one, unless he undertakes 
one on the beautiful Australian diurnal moths. 

On balance both books are excellent, and if the budget can take something in 
excess of $70.00, the interested lepidopterist should have both. If, however, economics 
of this magnitude are the deciding factor, the reader must determine what his 
interests are and be guided by them. If he wants identification of Australian butter- 
flies, either book will do, or he can get by with Common’s little book; if he wants the 
finest pictures of Australian butterflies and information on their habits, he should 
buy McCubbin’s book; if his interests go beyond the confines of Australia and he 
wants more than adequate figures and updated nomenclature, he should buy 
D’Abrera’s book. As an artistic masterpiece and for workers on Australian butterflies 
in general, McCubbin’s book is the best available. The regional scope of D’Abrera’s 
book and its greater scientific “meat” make Butterflies of the Australian region the 
greater prize. Buy either, you won’t be disappointed. In any event, “Seitz” is no 
longer a “must” for the interested lepidopterist’s library. 


Lee D, Mitter, Allyn Museum of Entomology, 712 Sarasota Bank Bldg., Sarasota, 
Florida 33577, 


VOLUME 26, NUMBER 3 203 


Tue ZOOLOGICAL MISCELLANY, by John E. Gray, with an introduction by Arnold G. 
Kluge. Reprinted by the Society for the Study of Amphibians and Reptiles. 1971. 
P. viii + 86, 4 pls. Price: $8.00 paperbound, $10.00 clothbound. (Available from 
the society's publications secretary, Dr. Henri C. Seibert, Dept. of Zoology, Morton 
Hall, Ohio University, Athens, Ohio 45701. ) 


This rare natural history journal was published privately by John Edward Gray in 
London, appearing irregularly between 1831 and 1844. The reprint includes an 
editorial note about the publication dates of the three parts, and a two page introduc- 
tion giving some insight into Gray’s life and the circumstances which may have 
prompted him to publish “The Zoological Miscellany.” Most of the papers are by 
Gray himself. There are only three (of 56) papers which deal with Lepidoptera. 


GrorcE Rospert Gray. 1831. Descriptions of eight new species of Indian butter- 
flies, (Papilio, Lin.) from the collection of General Hardwicke. p. 32-33. (De- 
scriptions of Papilio glycerion, Papilio philoxenus, Papilio minereus, Parnassius 
hardwickii, Pieris horsfieldii, Pieris agathon, and Argynnis childreni, all from 
Nepaul, and of Papilio agestor, from Sumatra. ) 

JoHN Epwarp Gray. 1831. Description of a new species of Bombyx from Nepaul, 
discovered by Dr. Wallich. In the collection of Major General Hardwicke. p. 39. 
(Description of Bombyx wallichii. ) 

EDWARD DouBLEDAY. June 1842. Characters of undescribed Lepidoptera. p. 73-78. 
(Descriptions of Papilio ganesa, from Nepaul, Assam; of Papilio polyeuctes, 
Papilio xenocles, Pieris thestylis, Pieris lalage, and Rhodocera lycorias, all from 
Silhet; of Leptalis atthis from Mexico; of Leptalis cydno, locality unknown; of 
Pieris ianthe from Sierra Leone; and of Pieris anactorie from South Africa. ) 


RicHARD S. Funx, Faculty of Biological Sciences, Southern Illinois University, 
Edwardsville, Illinois 62025. 


NOTES AND NEWS 


PROPOSED AMENDMENTS TO THE CONSTITUTION 
OF THE LEPIDOPTERISTS’ SOCIETY 


The following changes in the Constitution of the Society have been proposed, in 
order to: 


1) qualify the Society as a nonprofit organization; 
2) establish the new class of Student Member; 
3) define more precisely the nature of what is now known as Honorary Member- 


4) add the Secretary-elect and Treasurer-elect to the Executive Council; 

5) provide for the nomination by the Nominating Committee of up to two candi- 
dates for each elective office, and to simplify the nominating procedure; and 

6) clarify or modernize several sections. 


Notice is hereby given, in accordance with Article XII, Section 1, that these 
proposed amendments to the Constitution will be sent to the members with the ballots 
in November, 1972. Each section containing a proposed change is reproduced below 
in full, with language to be deleted in bold-face type, and new language in italic. 

Article II, Section 1: The Lepidopterists’ Society is a non-profit educational and 
scientific organization. It shall be the purpose of the Society to promote inter- 


204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


nationally the science of lepidopterology in all its branches; to further the scientifically 
sound and progressive study of Lepidoptera; to publish periodicals and other publi- 
cations on Lepidoptera; to facilitate the exchange of specimens and ideas by both 
the professional worker and the amateur in the field; and to secure cooperation in all 
measures tending to that end. and to facilitate personal intercourse among its 
members. 

Article II, Section 2: All individual subscribers to the “Journal” and the “News 
of the Lepidopterists’ Society,’ who have paid their current annual dues, shall be 
deemed members of the Society. 

Article III, Section 3: The membership of the Society shall consist of four five 
classes—Active, Student, Sustaining, Life, and Honorary Life members. All persons 
who joined the Society before January 1, 1948, shall be designated Charter members. 

Article III, Section 4: Application for Active, Student, Sustaining, and Life 
membership in the Society, received by the Secretary or Treasurer and accompanied 
by the annual appropriate dues for the current year, shall constitute formalization 
of membership, and no nomination or election to membership shall be necessary. 
The annual and Life dues shall be fixed by the By-Laws. 

Article III, Section 6: Individuals who have made important contributions to the 
science of lepidopterology may be elected Honorary Life Members of the Society. 
There shall not be more than ten living Honorary Life Members. 

Article IV, Section 1: The officers of the Society shall consist of a President, 
President-elect, three Vice-Presidents (not more than one of whom Vice-President 
shall reside in one country), a Secretary, a Treasurer, a Secretary-elect and/or a 
Treasurer-elect, in these two offices, in years when there are incumbents. 

Article IV, Section 2: The business and affairs of the Society, not otherwise 
provided for, shall be controlled by an Executive Council, consisting of the President, 
President-elect, three Vice-Presidents, the Secretary, the Secretary-elect, the Treasurer, 
the Treasurer-elect, and nine other members of the Society. Action on all amend- 
ments to the By-Laws and all appointments and elections by the Executive Council 
shall be obtained by a canvass by the Secretary of all members of the Council. 

Article V, Section 1: The President shall before the first of July appoint a 
Nominating Committee who shall nominate one no more than two candidates for 
each elective office to be filled for the ensuing year. and a list thereof shall be 
published in one of the Society’s periodicals or mailed to the members at least 
sixty days before ballots are mailed by the Secretary. Additional candidates may 
be nominated by submission to the Secretary of written nominations signed by not 
less than ten members. Ballots containing all nominations shall be mailed in No- 
vember of each year, setting forth the officers to be elected and the names of those 
nominated for each office. If more than one person is nominated for any office, their 
names shall be arranged alphabetically on the ballot. 

Article V, Section 3: Election of Honorary Life Members. Honorary Life Members 
shall be nominated by the unanimous vote of the members of the Executive Council. 
The nominee shall be voted on by mail ballot distributed to all members of the 
Society and reported in one of the Society’s periodicals, and must receive 80% 
of all ballots cast to be elected. Not more than five Honorary Life Members may be 
elected at the first annual meeting, and not more than two in any one calendar year. 

Article XIV. General Prohibitions 
Section 1. Notwithstanding any provision of the Constitution or By-Laws which 
might be susceptible to a contrary construction: 


(a) The Society shall be organized exclusively for scientific and educational 
purposes; 

(b) the Society shall be operated exclusively for scientific and educational purposes; 

(c) no part of the net earnings of the Society shall or may under any circumstances 
inure to the benefit of any private individual; 


VOLUME 26, NUMBER 3 205 


(d) no substantial part of the activities of the Society shall consist of carrying on 
propaganda, or otherwise attempting to influence legislation; 

(e) the Society shall not participate in, or intervene in (including the publishing 
or distributing of statements), any political campaign on behalf of any 
candidate for public office; 

(f) the Society shall not be organized or operated for profit; 


(g) the 
(1) 


(2) 


(3) 
(4) 


(5) 
(6) 


Society shall not: 

lend any part of its income or corpus, without the receipt of adequate 
security and reasonable rate of interest, to; 

pay any compensation, in excess of a reasonable allowance for salaries 
or other compensation for personal services actually rendered, to; 

make any part of its services available on a preferential basis to; 

make any purchase of securities or any other property, for more than 
adequate consideration in money or money's worth, from; 

sell any securities or other property for less than adequate consideration 
in money or money's worth to; or 

engage in any other transactions which result in substantial diversions 
of its income or corpus to; any officer, member of the Governing Board, 
or substantial contributor to the Society. 


The prohibitions contained in this subsection (g) do not mean to imply that the 
Society may make such loans, payments, sales or purchases to anyone else, unless 
such authority be given or implied by other provisions of the Constitution or 


By-Laws. 


; Nag 


f 


Vie eer 


iit ¢ 


y 
“ 
= 
‘ 
: re 
»\) i 
"A . 
2 
— h 
3 
ee it 
ao } 
= } 
. 
4 
= t 
; j " 
( ’ s 
es = f i 

: = - = ‘ 

J if 

% a ta 


EDITORIAL COMMITTEE OF THE JOURNAL 


Editor: THEODORE D. SARGENT, Department of Zoology, 
University of Massachusetts, Amherst, Massachusetts 01002 


K. S. Brown, J. M. Burns, R. H. Carcasson, J. P. DonaAnuE, J. F. 
Gates CLARKE, R. O. KENDALL, J. H. MAsters, L. D. MIL er, 
A. P. Piatt, J. R. G. TURNER 


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 CrreEp, 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 814 
> 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 article’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 approx- 
imate 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. 

Page Charges: Authors with grant or institutional funds are requested to pay a 
charge of $24.00 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 
case of authors who have no grant or institutional funding, as it is not intended that 
any author should pay this charge from personal funds. However, all authors will 
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: Dr. C. V. Covell, Department of Biology, University of 
Louisville, Louisville, Kentucky 40208. 


ALLEN PRESS, INC. Lp bie: LAWRENCE, KANSAS 


Us me 


CONTENTS 


Tue Genus ZESTUSA (HESPERIDAE ) IN EL SALVADOR WITH DESCRIP- 


TION OF A New Species. Stephen R. Steinhauser 


SOME OBSERVATIONS ON THE LEPIDOPTERA OF BROMELIADS. Carlos 
R. Beutelspacher, 1000000 ee 


Tue EFFECT OF CAUTERIZING THE MNPPM oF THE PUPA OF THE 
MonarcH ButrerFLy (DANAUS P. PLEXIPPUS) (DANAIDAE). 
FOA. Urquhart 3050 so a 


CERCYONIS PEGALA BLANCA, A “Missinc TYPE” IN THE EVOLUTION 
OF THE GENUS CERCYONIS (SATYRIDAE). Thomas C. Emmel 
and Sterling O. Mattoon 00 


A New Susspecies OF LYCAEIDES ARGYROGNOMON (\LYCAENIDAE) 
FROM THE EASTERN CANADIAN Forest ZONE. John M. 
Masters 20s i 


THE ECOLOGY AND ETHOLOGY OF THE TROPICAL NYMPHALINE BUTTER- 
FLY, VICTORINA EPAPHUS. I. LIFE CYCLE AND NATURAL His- 
Tory. Allen 'M. Young 000 ee 


PIERIS NAPI L. (PIERIDAE) AND THE SUPERSPECIES CoNcEPT. S. R. 
Bowden.) sa 


Notes on UrRopus PARVULA (HENRY Epwarps) ( YPONOMEUTIDAE ). 


S.W. Frost? GE OSS AT TI 


OBSERVATIONS ON FooppLANT RECORDS FOR PAPILIO GLAUCUS 
(PApmLionwaE). Malcolm P. Levin and Mary Ann Angleberger 


THe Larva OF CHAMYRIS CERINTHA (TREITSCHKE) (NocTrumpAE). 
George L. Godfrey ei 


Maximizinc Datty ButrerFLy Counts. Keith S. Brown, Jr. —-- 


GENERAL NOTES 
An unusual moth in central Illinois. Charles K. Swank __..__.. 


Wing-shape and adult resources in lycaenids. Paul R. Ehrlich and 
Anne H, Ehalich 3.000006 ca er 


New distribution records for Ceratomia hageni (Sphingidae). Charles 
M.. Franklin (06 EA err 


Further notes on W. H. Edwards specimens in Illinois museum collections. 
Roderick R. Irwin 
vy 


Book Reviews 


127 


133 


137 


140 


150 


155 


170 


173 


177 


180 
183 


Volume 26 1972 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 


15 December 1972 


THE LEPIDOPTERISTS’ SOCIETY 
EXECUTIVE COUNCIL 


Lioyp M. Martin (Prescott, Ariz.) President 

J. F. Gates Ciarxe (Washington, D.C.) President-elect 
S. A. AE (Nagoya, Japan) Ist Vice President 

KerrH S. Brown (Rio de Janeiro, Brasil) Vice President 
H. A. FREEMAN (Garland, Texas) Vice President 

S. S. Nico.ay (Virginia Beach, Va.) Treasurer 

LEE D. MILLER (Sarasota, Fla.) Secretary 


Members at large (three year term): R. B. Dominick (McClellanville, S.C.) 

B. MATHER (Clinton, Miss.) 1972 1973 

M. Ocarta (Osaka, Japan) 1972 J. P. DonauvE (Los Angeles, Calif.) 1973 

E. C. WELLING ( Merida, Mexico ) 1972 J. M. Burns (Cambridge, Mass.) 1974 

ANDRE BLANCHARD (Houston, Texas ) R. H. Carcasson ( Vancouver, B.C.) 1974 
1973 M. C. Nievsen (Lansing, Mich.) 1974 


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 $10.00 
Student members—annual dues $5.00 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $15.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia 23455. 


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


by Cyriz F. pos Passos 


Price, postpaid: Society members—$5.00, others—$7.50; uncut, unbound signatures 
available for interleaving and private binding, same prices; hard cover bound, mem- 
bers—$8.00, others—$10.00. Revised lists of the Melitaeinae and Lycaenidae will be 
distributed to purchasers free (separately with paper covered copies and unbound 
signatures, bound in with hard covered copies ). 


The Lepidopterists’ Society is a non-profit, scientific organization. The office of 
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520. 
Second class postage paid at Lawrence, Kansas, U.S.A. 66044. 


JOURNAL OF 


Tue LEepIpoPTERISTS’ SOCIETY 


Volume 26 1972 Number 4 


THE KARL JORDAN MEDAL 


Lee D. MILLER 
Allyn Museum of Entomology, 712 Sarasota Bank Building, Sarasota, Florida 33577 


Several scientific organizations have established prizes for research 
excellence in their respective fields, such as the Brewster Medal given for 
meritorious research in ornithology. Such a prize has not been offered in 
the past for lepidopterology, but in recognition of the 25th Anniversary 
Celebration of the Lepidopterists’ Society, Mr. A. C. Allyn offered to fund 
such an award through the Allyn Museum of Entomology, Sarasota, 
Florida. This offer was accepted by the Executive Council and the 
members in attendance at the Annual Meeting of the Society in San 
Antonio, Texas. 

The award to be given by the Lepidopterists’ Society in recognition of 
outstanding original research in lepidopterology will emphasize particu- 
larly the fields of morphology, taxonomy, zoogeography and what was 
once known as “natural history.” These fields are the ones in the study 
of Lepidoptera that are currently poorly supported and not “popular’; 
it is hoped that the establishment of this award may encourage workers 
to do meaningful research along these lines. The prize will consist of 
an engraved silver medal of appropriate design, a $1,000.00 cash award 
and travel expenses for the recipient(s) to accept the award at the 
particular year’s Annual Meeting of the Society. 

This award will be known as “The Karl Jordan Medal.” There is much 
to recommend honoring this man in this way. Dr. Jordan was one of the 
original Honorary Life Members of the Lepidopterists’ Society. He was 
active in the field for over half a century, and during this period he 
produced an almost unbelievable volume of work of the highest quality. 
He was honored by The Royal Entomological Society of London with a 
Jubilee Volume prior to his death, the only man in recent history so 
honored, attesting to Karl Jordan’s preeminence among 20th Century 
lepidopterists, and entomologists in general. Dr. Jordan did not limit 


208 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


his efforts to a single fauna, but rather worked on a worldwide basis; 
neither did he restrict himself to a narrow group of insects: Jordan began 
his career at Tring as a coleopterist, later became a lepidopterist and still 
later established his credentials as the world’s leading authority on 
Siphonaptera (fleas). Finally, but by no means leastly, Karl Jordan 
excelled in those fields previously mentioned for emphasis in the awarding 
of the Medal and in general exemplifies the ideals for which The Karl 
Jordan Medal is intended. 

The procedures and criteria for awarding the Jordan Medal are as 
follows: 


1. An award committee will be established consisting of: 

A. One representative to be designated by the Allyn Museum of 
Entomology; 

B. The President of the Lepidopterists’ Society, unless he is af- 
filiated with the Allyn Museum of Entomology, in which case the 
First Vice President of the Society will be part of the award committee; 
and 

C. An elected member of the Lepidopterists’ Society to be known 
as “the Jordan Medal Representative” who will serve a one-year term 
and will not be affiliated with the Allyn Museum of Entomology. 


In practice, the President-Elect of the Society and the Allyn Museum 
representative should be in consultation on a short list of nominees for 
the succeeding year to be presented to the Jordan Medal Representative, 
who will have less than six months in which to make up his mind on 
possible candidates. 


2. The medal will be awarded only by the unanimous vote of the com- 
mittee. This, of course, will result in the medal not being awarded in 
some years, but it is felt that the requirement for unanimity of the com- 
mittee will be a safeguard in assuring that only work of the highest 
quality is awarded. 

3. The award will be for original research, not for the compilation of 
already known facts. The judgment of where the line must be drawn 
between research and compilation will be left to the discretion of the 
committee and may be expected to change from year to year as different 
people are involved. 

4. The award may be based upon a single piece of research or on a 
series of interrelated works, and the work must be at least three, but not 
more than twenty-five years old. This stipulation was suggested to assure 
that awarded work had stood the test of time and use, but it was the most 
controversial criterion for the award at the San Antonio meetings. 


VOLUME 26, NUMBER 4 209 


The Jordan Medal is not intended to be a career award; the Society 
already has such an award, Honorary Life Membership. However, this 
does not preclude awarding the Medal to an Honorary Life Member for 
a particular piece of work or for a series. 

5. Jointly awarded prizes will be allowed only in instances of co- 
authorship, and the cash prize for that year will be divided equally 
between the workers or their heirs. In instances of jointly awarded 
prizes, duplicate medals will be presented, and travel expenses for more 
than one worker will be allowed. | 

6. No member of the Jordan Medal award committee will be eligible 
to receive the prize during his tenure, but will not be ineligible in suc- 
ceeding years. 

7. The recipient of the Jordan Medal need not be a member of the 
Lepidopterists’ Society. It is hoped, however, that if the recipient is not a 
Society member that he may elect to become one. 

8. Nominations for recipients of the Jordan Medal will be accepted 
from any member of the Lepidopterists’ Society and should be sent to: 


Jordan Medal Awards Committee 
Allyn Museum of Entomology 
712 Sarasota Bank Building 
Sarasota, Florida 33577, U. S. A. 


It would be helpful to the awards committee if one or more copies of the 
nominated works were submitted to the same address for distribution to 
the award committee. 

The Karl Jordan Medal will be funded on a year-to-year basis for the 
time being. There are some problems with professional rivalry and ill- 
feelings associated with some of the similar prizes in other disciplines; it 
is hoped that lepidopterists do not become entangled in such unpleasant 
and unfortunate traps. Should this award show promise of encouraging 
workers and aiding good relationships between lepidopterists, the Karl 
Jordan Medal will receive permanent funding in the future. Cooperation, 
good relationships with fellow workers and excellence were three of the 
many fine attributes of Karl Jordan and must be fostered in the awarding 
of the prize that bears his name. 


210 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ULTRAVIOLET PHOTOGRAPHY AS AN 
ADJUNCT TO TAXONOMY? 


CLIFFORD D. FERRIS? 
College of Engineering, University of Wyoming, Laramie, Wyoming 82070 


Several papers have been published on the use of ultraviolet (uv) 
photography to visualize hidden characters on the wings of Lepidoptera 
(e.g. Nekrutenko, 1964, 1965; Eisner et al., 1969). These hidden patterns 
probably relate to differing characteristics of the scales and their arrange- 
ment on the wings. Some work has been carried out using a scanning 
electron microscope to detect differences in scale characteristics (Kolyer 
& Reimschuessel, 1969). Most of the more recently published papers 
have treated the Coliadinae, especially the genus Gonepteryx. In this 
subfamily, luminous patches appear on the dorsal surfaces of the wings 
when they are photographed under ultraviolet light. In other genera, 
Papilio for example, forms which appear quite different when viewed 
under white light appear nearly identical when photographed under uv 
light. 

Photographic techniques are necessary to visualize the ultraviolet 
evoked patterns as the human eye does not respond to that portion of the 
electromagnetic spectrum. From analysis of black light response in 
Lepidoptera, their visual perception ranges seem to peak in the range 
300-400 nanometers (millimicrons ). 

Since most of the readily available literature on ultraviolet photography 
of insects does not give details of technique, it is felt that a technique 
should be described which could be applied by anyone with a basic 
knowledge of photography. That is the purpose of this paper. 

The primary prerequisite is a camera which has a lens that will pass uv. 
Generally the better quality modern 35 mm single-lens-reflex cameras 
from Germany and Japan meet this criterion. The author uses two dif- 
ferent models of Mirandas with Soligor-Miranda lenses. A simple way to 
check a lens for uv transmission (other than by taking a photograph) is 
to use the lens to focus light from a uv source onto a fluorescent object. 
Suitable objects are the mineral Willemite (ZnSiO, which fluoresces 
bright green), if available, or various greases. Vaseline fluoresces a pale 
green. Fluorescent spray paint also may be used. 

Since only ultraviolet reflectance is of interest for distinguishing patterns 


' Published with the approval of the Director, Wyoming Agricultural Experiment Station, as 
Journal Paper no. JA 503. 


* Research Associate, Allyn Museum of Entomology, Sarasota, Florida. 


VOLUME 26, NUMBER 4 Dy 


Fig. 1. Camera support and illumination system for ultraviolet photography. 
Righthand light source swung out to expose mounting technique. 


in the Lepidoptera, it is necessary to filter out all light except that in the 
near uv spectrum (300-400 nm). This may be done with a Wratten 18A 
filter (Eastman Kodak Co.). This filter is available as a 2” x 2” square 
and appears opaque. A filter-holder adapter mount is required for the 
camera. These items can be ordered through any photographic store. 
The 18A filter transmits ultraviolet light, but blocks visible light. 

A suitable light source is necessary. Photographs can be taken in direct 
sunlight utilizing the uv content of natural light. Generally speaking, 
light which has passed through window glass should not be used, as 
most glass filters out ultraviolet rays. For an artificial source, the author 
uses two 15 watt blacklight fluorescent tubes that have a built-in filter 
glass which filters out most of the visible portion of the light spectrum. 
These are model 50058 available from Ultra-Violet Products, Inc., 5114 
Walnut Grove Ave., San Gabriel, California 91778. Similar tubes are 
manufactured by various lamp firms and can be obtained from lamp 
jobbers. 

Fig. 1 shows a simple setup for doing ultraviolet photography. The 


bo 
jamal 
bo 


JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 2. Dark @ form of Papilio g. glaucus photographed under uy illumination. 


light sources are mounted in two homemade wooden reflector housings. 
Standard 15 watt fluorescent light brackets are used (available from 
most mail-order catalog stores or from electrical suppliers). A 35 mm 
photographic enlarger easel is used to support the camera. If available, 
a Polaroid Camera Corp. copy stand makes an ideal setup as only the 15 
watt blacklight tubes are required as additional items. A conventional 
camera tripod can be used as a camera support, but is less convenient 
than an enlarger easel. In Fig. 1, one of the lamp housings has been 
swung out to illustrate the lamp mounting. The lamp appears white in 
color as the light was on when the photograph was taken. To insure 
photographing the full extent of the uv reflectance pattern with this 
setup, specimens must be mounted with the wings flat. 

Either Tri-X or Panatomic X film (Eastman Kodak Co.) is suitable for 
uv photography. The former requires much shorter exposure time than 
the latter, but tends to develop slightly more granularity. Exposure time 
can be determined by placing the 18A filter over the light entry port of a 
CdS exposure meter and measuring the reflected light from the speci- 
men. The background to which the specimen is pinned should not be 
fluorescent. The high-density polyethylene foam usually used in pinning 
trays is satisfactory. A typical exposure setting for Tri-X film (ASA 400) 


VOLUME 26, NUMBER 4 213 


Fig. 3. Normal yellow @ form of Papilio g. glaucus photographed under uv 
illumination. 


is £/16 at % second with 10 inches between the filter plane and the speci- 
men, and with the light sources approximately 6 inches from the speci- 
men. The camera lens should be stopped down to at least £/8 to produce 
sufficient depth of field to counteract the difference between focusing 
white light and near ultraviolet light. Focusing is achieved by removing 
the 18A filter. If required, a supplementary closeup lens such as a 3+ 
Portra lens (Eastman Kodak Co.) can be added. This lens does transmit 
uv light. 

Excluding the camera body, lenses, light meter, and stand or tripod, 
the present cost of setting up to do ultraviolet photography is as follows: 
18A filter and holder $25.00, two 15 watt uv tubes $14.00, fluorescent lamp 
fixtures $15.00, miscellaneous (reflector housings, etc.) $5.00. These are 
approximate prices. Special ultraviolet-transmitting lenses are available, 
but these are designed for very short wavelengths and range in cost from 
$750.00 to $1,600.00. Such lenses are not required for this type of insect 
photography. 

Figs. 2 and 3 illustrate one aspect of uv photography. The two female 
forms of Papilio glaucus glaucus Linnaeus are shown as they appear 
under uv light. Similar patterns appear when they are photographed 


214 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 4 & 5. Specimens of Colias a. alexandra from Albany Co., Wyoming: 4, 
photographed under normal illumination—male at top, yellow female in middle, white 
female form at bottom; 5, same specimens as they appear under uv illumination. 


under uv illumination, although they are quite different when viewed 
by visible light. Only a suggestion of the black bars appears in the yellow 
female form in the ultraviolet photograph. 


As a comparison, the wings of both sexes of Colias alexandra alexandra 
Edwards are shown as they appear when photographed with visible light 
(Fig. 4) and with ultraviolet light (Fig. 5). Under uv light, luminous 
patches appear on the hindwings of the male, while the female appear 
drab (a characteristic of the female sex in most North American Colias ). 
A forthcoming paper by the author on the Colias alexandra complex will 
demonstrate the utility of ultraviolet photography in taxonomic research. 

More sophisticated techniques do exist for ultraviolet photography, but 
these involve the use of special light sources and filtering techniques. 
Such matters are beyond the scope of this presentation. The intent here 


VOLUME 26, NUMBER 4 2a 


has been to describe a simple method which can be applied using a 
limited amount of equipment. 


LITERATURE CITED 


EIsNER, T., R. E. SILBERGLIED, D. ANESHANSLEY, J. E. CArrREL & H. C. HOow.anp. 
1969. Ultraviolet video-viewing: the television camera as an insect eye. Science 
166: 1472-1174. 

Konyer, J. M. & A. M. RemscuHurssEL. 1969. Scanning electron microscopy on 
wing scales of Colias eurytheme. J. Res. Lepid. (8)1: 1-15. 

NEKRUTENKO, Y. P. 1964. The hidden wing-pattern of some Palearctic species of 
Gonepteryx and its taxonomic value. J. Res. Lepid. 3(2): 65-68. 

. 1965. Three cases of gynandromorphism in Gonepteryx. J. Res. Lepid. 

4(2): 103-107. 


Additional References 


Ultraviolet and fluorescence photography. Eastman Kodak Co. Tech. Pub. M-27. 
Basic scientific photography. Eastman Kodak Co. Pub. N-9. 


A NEW SUBSPECIES OF EUMEDONIA EUMEDON 
(LYCAENIDAE) FROM CAUCASUS 


Yuri P. NEKRUTENKO 


Ukrainian Research Institute for Piant Protection, 33 Vasilkovskaya Street, Kiev 127, 
Ukraine, U.S.S.R. 


During the past few seasons I have had the opportunity to collect in 
the Western part of the Main Caucasus Ridge and to review Caucasian 
material deposited in the Lepidoptera Collection of the Zoology Museum, 
Kiev State University and in the private collection of Dr. Eugene S. 
Miljanowski (Sukhumi, Georgia ) who spent more than 30 years collecting 
in different parts of Abkhasia. From these sources I found some interest- 
ing, heretofore undescribed forms of lycaenid butterflies, one of which 
is described here with some brief remarks. 


Eumedonia eumedon modestus Nekrutenko, new subspecies 
(Figs. 1-6) 


Lycaena eumedon Esp.: Romanoff, 1884, p. 52. 
Lycaena eumedon Esp.: Wojtusiak & Niesiolowski, 1947, p. 58-59. 
Lycaena teberdina Tschetv.: Miljanowski, 1964, p. 114. 


Male. Length of the forewing (base to tip) of the holotype 14.0 mm (variation 
in type series 13.0 to 14.5 mm). Upperside of both wings of dark black-brown 
ground color, discal spots hardly recognizible. Underside ground color steel-grey, 


216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-6. Eumedonia eumedon modestus n. subsp.: 1, 2, holotype, ¢, upper and 
undersides, S.W. Caucasus, Awadhara, 1800 m, 7-12 June 1971, Y. P. Nekrutenko; 
3, 4, allotype, 2, upper and under sides, same label data; 5, paratype, ¢, ab. fylgia 
Spangb., underside, same label data; 6, paratype, 6, ab. speveri Husz., underside, 
same label data. 


discal spot and series of postdiscal spots complete, surrounded with remarkable white 
rings. The white stripe on the underside of the hindwing is well developed, but is 
twice as narrow as that in European specimens. Blue powdered area on the under- 
side of hindwing is relatively larger than in European specimens, and occupies about 
Ys of the entire wing surface. The main differential characteristic of the described 
subspecies is an extreme reduction of submarginal markings. Orange and black 
submarginal spots are almost absent on the underside of the forewing, and on the 
hindwing underside they are diffused, reduced and incomplete. 

There are no essential differences from European specimens in the male genitalic 
rmatures except for a deeper incision between valval lobes. 


emale. Length of the forewing of allotype 15.5 mm (14.9-15.9 mm). In size 


VOLUME 26, NUMBER 4 Daler, 


Fig. 7. Eumedonia eumedon modestus n. subsp., type locality. A meadow below 
upper timberline at an elevation of 1800-2000 m at Awadhara, S. W. Caucasus, 
surrounded by endemic fir-trees Abies nordmanniana (Stev.) Spach. 


and outer appearance very similar to male. This weak sexual dimorphism may often 
be confusing, especially in the field. However, in some females (including allotype) 
upperside of hindwing bears a hardly recognizible brown spot within anal angle. 
Underside pattern is quite similar to that of a male. 

Types. Holotype, male, and allotype, female, S. W. Caucasus, Abkhasian 
Autonomous Soviet Socialist Republic, Awadhara, 1800-2000 m, 7-12 June 1971, 
Y. Nekrutenko (In Zool. Mus. Kiev Univ.). Paratypes 8 ¢ 6, 2 2 2, same locality, 
dates and collector (In Zool. Mus. Kiev Univ.). Nine paratypes from Awadhara, 
June, July 1961, 1968 and 1969 (in coll. E. Miljanowski). 5 ¢ ¢ paratypes, Teberda, 
N. W. Caucasus, Mt. Chatipara, 2200-2400 m, 30 July to 6 August 1933, L. 
Sheljuzhko (Zool. Mus. Kiev Univ.). ¢ paratype, Teberda, N. W. Caucasus, Valley 
of the Teverda River, L. Sheljuzhko (Zool. Mus. Kiev Univ.), @ paratype, Teberda, 
Mt. Chatipara, 2600 m, 30 July 1933 L. Sheljuzhko (Zool. Mus. Kiev Univ.). ¢ para- 
type, Lebarde, W. Georgia, 3 July 1962 (coll. E. Miljanowski). 2 ¢ ¢, paratypes, 
Lagodekhi Reservation 25 July and 2 August 1959 (coll. E. Milianowski). 3 ¢ 6 
paratypes, Yelizavetpol (now Kirovabad), Azerbaijan, 20 and 26 June (no year), 
A. Kashtshenko (Zool. Mus. Kiev Univ.). & paratype, Bakuriani, Mt. Kochta, 31 
July 1932, B. Tkatshukov (Zool. Mus. Kiev Univ.). 

Type locality (Fig. 7). Because most specimens examined were collected in 
Awadhara, it is designated as the type locality of the described subspecies. Awadhara 
is a part of Ritsa Nature Reservation (Ritsinskiy Zapovednik) and lies 16 km up 
along the Lashipse River from Lake Ritsa, at elevation 1650-2500 m. E. eumedon 
modestus flies in rich meadows below and above upper timber line. The flight 
period extends from early June to early October (Miljanowski, personal communica- 
tion ). 


218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Subspecific characters given above are stable for all specimens of E. 
eumedon Esp. ever seen from the Caucasus range. N. M. Romanoff 
(1854) reported the occurrence of Lycaena eumedon from 8 points in 
Transcaucasia, including Bakuriani, “en général sur les lieux élevés en 
Juillet.” A male specimen from this locality, included in the type series, 
showed the appearance of the described subspecies in Caucasus Minor 
also. Wojtusiak & Niesiolowski (1947), who reported the occurrence of 
E. eumedon in the Central Caucasus (Karaugom, 1800-2500 m), have 
noted its smaller size, as compared to European specimens, and the “un- 
derside of wings not brown but grey.” E. eumedon has not been included 
in the faunistic list of E. Miljanowski (1964) as it has been confused with 
Lycaena teberdina Shel. The latter is highly local in its occurrence 
(Teberda, N. W. Caucasus ) and seems not to occur on the Southern side 
of the Great Caucasus (Sheljuzhko, 1934). 


ACKNOWLEDGMENTS 


I wish to express appreciation to Dr. Eugene S. Miljanowski for an 
interesting and useful discussion, and to Dr. Theodore D. Sargent who 
edited and corrected the manuscript. 


LITERATURE CITED 


Mitjanowski, E. S. 1964. The butterfly and moth fauna of Abkhasia. Trudy 
Sukhumskoi Opytnoi Stantsii Efiromaslichnykh Kultur 5: 91-190 (In Russian). 

Romanorr, N. M. 1884. Les Lépidoptéres de la Transcaucasie. Mém. sur les 
Lépidopteres, rédigés par N. M. Romanoff. St.-Pétersbourg. I: 1-92. 

SHELJUZHKO, L. 1934. Neue Lepidopteren aus dem Nordkaukasus.  Zeitschr. 
Oesterr. Entomol.-Ver. 19: 39-40. 

Wojrusiak, R. J. & W. Nrestocowsxt. 1947. Lepidoptera of the Central Caucasus, 
collected during the Polish Alpine Expedition in 1935, with ecological and 


zoogeographical remarks. I. Macrolepidoptera. Prace Muzeum Przyrodniczego 
PAUG62 —74e 


NOTES ON THE LIFE HISTORY OF EUGONOBAPTA NIVOSARIA 


(GEOMETRIDAE ) 

In spite of the abundance of Eugonobapta nivosaria Guenee in many localities 
in the northeastern United States and eastern Canada, there apparently has been 
no published account of the early stages. 

Success in discovering these early stages came only after several failures to 
keep the eggs alive over winter. That problem was finally solved by confining 
the females in paper lined containers, and then, after the eggs were deposited 


on the paper, placing them in a wooden box which was stored in a sheltered 


place outdoors. This box, however, was covered with snow during much of 
tne vinter, 


VOLUME 26, NUMBER 4 219 


The eggs are brick shaped and are deposited side-by-side in short, compact, 
precisely arranged rows. These eggs are similar in shape, size, and arrangement 
to those of Deuteronomos magnarius Guenee, except that they are in shorter rows. 
They are pearly white, very finely but rather sparsely pitted on the upper 
surface. They seem unusually large for so small a moth, and the number of eggs 
produced by one female is evidently small. The maximum that I have ever 
obtained from one female is about 20. 

The newly hatched larvae are translucent, almost colorless, very slender and 
very active. It is probable that they are quite general feeders on deciduous trees 
and shrubs, for I reared them on both choke cherry (Prunus virginiana) and red- 
osier dogwood (Cornus stolonifera). They soon became green, and grew very 
rapidly, the first one starting its cocoon just 15 days after hatching. 

The mature larva is light green with inconspicuous yellowish subdorsal and 
lateral lines, and somewhat wider stigmatal stripes of the same yellow color. It 
is essentially smooth, but with slightly raised flat tubercles accompanying each 
abdominal spiracle. It is easy to understand why these larvae have been completely 
overlooked, since their feeding period is quite brief, and occurs at a time when the 
superficially similar appearing larvae of Paleacrita vernata Peck are likely to be 
numerous. 

The pupa is delicate pale green, and is enclosed in a neatly woven, thin, but 
very tough cocoon of white silk spun among leaves. The most remarkable feature 
of the pupa is a complete set of conspicuous spiracular tubercles, quite unlike 
anything I have observed elsewhere. In the absence of the dorsal groove, the 
very weak development of the lateral grooves, the general arrangement of the 
cremaster hooks, the very light chitinization of the pupal skin, and the density 
of the cocoon, there are marked resemblances to pupae of the Ennomos group. 

In the past it has been a problem to determine what the closest relatives of 
Eugonobapta may be, since the adults have evidently lost most of the structural 
features that might indicate relationship, and the early stages were unknown. 
In the light of what we now know, it seems very reasonable to believe that 
Eugonobapta is a perfectly good member of the Ennomos complex. 


LAURENCE R. Rupert, Sardinia, New York 14134. 


CARDIAC GLYCOSIDES IN ASCLEPIAS SPECIES 


Since the publication of my previous note (Slansky 1971, J. Lepid. Soc. 25:294) 
Jackson Bees has brought to my attention one publication (Masler et al. 1962, 
Collection Czeckosl. Chem. Commun. 27:872—895) and others have appeared 
(Duffey 1970, Science 169:78-79; Duffey & Scudder 1972, J. Insect Physiol. 
18:63-78; Feir & Suen 1971, Ann. Entomol. Soc. Amer. 64:1173-1174; Singh 
& Rastogi 1970, Phytochem. 9:315—331) reporting the presence of cardiac glycosides 
in Asclepias syriaca and other Asclepias species indicated by Brower (1969, Sci. 
Amer. 220:22-30) to lack these compounds, as cited in my prior note. 

Perhaps, as pointed out by Duffey (op. cit.), the concentration of cardenolides 
in these Asclepias species is below a threshold level and/or these species lack 
cardenolides with strong emetic properties, such that monarchs, Danaus plexippus 
L., whose larvae feed upon these plants are palatible to predators, as found by 
Brower (op. cit.). 


FRANK SLANSKY, JR., Department of Entomology, Cornell University, Ithaca, 
New York 14850. 


220 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


THE OCCURRENCE OF CHLOROCLYSTIS RECTANGULATA (L.) 
IN NORTH AMERICA (GEOMETRIDAE) 


DoucLas C. FERGUSON 


Systematic Entomology Laboratory, U.S. Department of Agriculture, c/o National 
Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 


While collecting moths in Nova Scotia in the summer of 1970 in com- 
pany with Mr. Barry Wright of the Nova Scotia Museum, I was surprised 
to find a species with the habitus of Eupithecia Curtis but marked very 
obviously with bright green scaling on both wings and body. In more than 
20 years of intensive field work in that area I had never seen such a moth, 
yet on this occasion collected nine specimens in three localities, separated 
by as much as 175 air miles. 

A careful check on the identity of these specimens at the U.S. National 
Museum showed that they undoubtedly represent an Old World species, 
Chloroclystis rectangulata (Linnaeus ), in the subfamily Larentiinae. The 
genitalia are very distinctive in both sexes and have been figured by 
various authors, including Pierce (1914, pl. 32), Juul (1948, pl. 13), and 
Nordstrom et al. (1940, text fig. 330, female). The Nova Scotian speci- 
mens agree with European ones in every detail, and both the normal and 
dark (“nigrosericeata’) color forms are present. Colored figures of the 
adults may be found in many European works, and figures of the larvae 
are given by Juul (1948, pl. 3, fig. 27) and Nordstrom et al. (1940, pl. 40, 
fig. 211). This species will in any event be easily recognized because it is 
the only known representative of the genus Chloroclystis Hubner in the 
New World. In addition to the green markings, which do not readily fade, 
Chloroclystis differs from Eupithecia in rather obvious genital characters 
such as the obsolescence of the uncus, and the peculiar, forceps-like ter- 
minal armature of the aedoeagus. The female of rectangulata has two 
separate, crescent-shaped signa on opposite sides of the bursa copulatrix. 

It should be noted that one of the commoner species of Eupithecia 
of the eastern United States, E. miserulata Grote, also may have quite 
conspicuous green scales on the wings when very fresh. However, the 
green coloring in this species is no longer apparent in museum specimens 
more than one or two years old, having faded to yellowish brown. The 
genitalia of miserulata were figured by McDunnough (1949, fig. 4A). I 
know of no other North American species with which C. rectangulata 
is likely to be confused. 

Chloroclystis rectangulata is very widespread and common in Europe, 
occurring even north of the Gulf of Bothnia. The main host plants are 


VOLUME 26, NUMBER 4 921 


reported to be Crataegus, Prunus and Pyrus, and Prout (1915: 299) 
stated that the larva feeds in the blossoms of wild and cultivated apple 
and is often very injurious. It has also been mentioned as a pest of pear 
(Meyrick, 1927: 218) and quince (Dirimanoff et al., 1961). Harrison 
(1953) reported finding it on blackthorn (Prunus spinosa) in England, 
but noted that for the most part it seems restricted to orchards. The eggs 
hibernate and the larvae mature rapidly in the spring. 

The Nova Scotian records for this species are as follows: Smiley Brook 
Provincial Park, near Brooklyn, Hants Co., July 20, 1970 (1); Middle 
River, Victoria Co., Cape Breton Island, July 27, 1970 (6); Baddeck 
River, near Baddeck Bridge, Victoria Co., July 29, 1970 (2)!. All were 
taken at light in moist bottomland situations. Crataegus spp. and Prunus 
virginiana L., possible host plants, were common to all three localities. 
The specimens are in the collections of the Nova Scotia Museum, Halifax, 
and the United States National Museum. . 

The sudden appearance of such a conspicuously different moth, in a 
region where the Geometridae had been investigated intensively over a 
period of many years, leaves little doubt that this represents a quite 
recent introduction. However, its presence both on the mainland of 
Nova Scotia and on Cape Breton Island, in localities so far apart, indicates 
a well established population. 


LITERATURE CITED 


Dirimmanorr, M., G. SENGALEVITCH, A. CHARIZANOFF & R. ANGELOVA. 1961. Rast. 
Zasch. 9(6): 36. [in Bulgarian]. 

Harrison, J. W. Hestop. 1953. Chloroclystis rectangulata (L.) feeding on Black- 
thorn (Prunus spinosa) The Entomologist 86: 2. 

Juunt, K. 1948. Nordens Eupithecier. Gravers Andersens Forlag, Aarhus, Denmark. 
TAZ p. 

McDunnoucu, J. H. 1949. Revision of the North American Species of the Genus 
Eupithecia (Lepidoptera, Geometridae). Bull. Amer. Mus. Nat. Hist. 93(8): 
535-728, illus. 

Meyrick, E. 1927. A Revised Handbook of British Lepidoptera, 1968 reprint. 
E. W. Classey Ltd., Hampton, Middlesex. 914 p. 

NorpstROM, F., E. Wantcren & A. TULLGREN. 1935-41. Svenska Fiarilar, 
Aktiebolaget Familjeboken, Stockholm. 353 p., illus. 

Pierce, F. N. 1914. The Genitalia of the British Geometridae. F. N. Pierce, Liver- 
pool. 88 p., 48 pls. 

Prout, L. B. 1912-21. The Palaearctic Geometrae, vol. 4 in Seitz, The Macro- 
lepidoptera of the World. Alfred Kernen, Stuttgart. 479 p., 29 pls. 


1 Another specimen, a fresh female, was collected at the Hants Co. locality on 23 July 1972, 


indicating the continued presence of the species. 


292, JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


NEW RECORDS OF LEPIDOPTERA FROM THE UNITED STATES 
(ARCTIIDAE, GEOMETRIDAE, EPIPLEMIDAE) 


DoucLas C. FERGUSON 


Systematic Entomology Laboratory, Agricultural Research Service, USDA, 
c/o U.S. National Museum, Smithsonian Institution, Washington, D.C. 20560 


Collections of moths that I have recently examined from various sources 
included five species hitherto unreported from the continental United 
States, or indeed from anywhere in America north of Mexico. One of these 
is an especially interesting species described from Siberia; the others are 
of neotropical distribution. Their occurrence in this country is believed 
to be natural, not a result of introduction by man. They are as follows. 


Arctiidae 
Hyperborea czekanowskii Grum-Grshimailo 


Hyperborea czekanowskii Grum-Grshimailo, 1900: 464. 
Phragmatobia czecanousci Hampson, 1920: 349-350. 

One male specimen of this species (Fig. 1) was collected on the 
Seward Peninsula of Alaska and sent to me for identification by Dr. 
Kenelm W. Philip of Fairbanks, who recognized it as something unusual. 
It bears the following data: Grassy delta near mouth of Serpentine River 
20 mi. SE of Shishmaref, Seward Peninsula, Alaska, 25 June 1970, W. 
Foster. I had expected that this would prove to be an undescribed 
species, but found that it so closely matches the description and figure 
of H. czekanowskii in Seitz (1910: 103, pl. 18h) that there can be little 
doubt as to its identity. Seitz gave for its distribution only the Tunguska 
River [in central Siberia], but I found on consulting the original descrip- 
tion that the type series consisted of 17 males and one female for which 
were given the following data: Valley of the lower Tunguska River, 
1873; valley of the Olenek River between the lower Tomba and the Alakit, 
July, 1874; valley of the Adytscha [Adycha] River at its confluence with 
the Jana [Yana], July 3-4, 1885. The spellings in brackets are as given 
on recent National Geographic Society maps. The Adycha River locality 
is closest to Alaska, being about 1,700 miles west of the Seward Peninsula 
and slightly farther north. 

Hf. czekanowskii is the monobasic type-species of the genus Hyperborea, 
which Grum-Grshimailo proposed for his new species in the same paper. 
Although it has very much the appearance of an Apantesis species, the 
antennae are peculiar, appearing nearly simple on comparison with the 
oipectinate male antennae of all known species of the genus Apantesis 


VoLUME 26, NUMBER 4 223 


Fic. 1-5. 1, Hyperborea czekanowskii 6, Seward Peninsula, Alaska (full data in 
text); 2, Triphosa affirmata 2, Guatemala; 3, Semaeopus cantona, holotype ¢, 
Orizaba, Mexico; 4, Erosia incendiata 2, Kingsville, Texas, 15 November 1968, J. E. 
Gillaspy; 5, Antiplecta triangularis 2, Brownsville, Texas (full data in text). (Figs. 
1-4 natural size; Fig. 5 twice natural size.) Photographs made by photographic 
laboratory of National Museum of Natural History, Smithsonian Institution. 


Walker. The male antennae of czekanowskii are slender but biserrate, 
with a large bristle arising from the end of each process, and with the 
shaft setose ventrally and quite heavily scaled dorsally. The eyes are re- 
duced, as would be expected of an arctic species with diurnal flight 
habits. The body is more slender than is usual in Apantesis, although 
not as slender as it appears in Seitz’s figure, and the hindwings are some- 
what subhyaline. The moth is rather colorless compared to most species 
of this group; the forewing is blackish, traversed with pale lines as in 
Apantesis; the hindwing is dull whitish with dark gray-brown spots, and 
the abdomen is banded with the same shades except for a small caudal 
tuft of yellow hair. In the region where it occurs, czekanowskii is likely 
to be confused only with Apantesis quenseli (Paykull), or possibly with 
A. turbans (Christof), both however stouter-bodied, differently marked 
species with pectinate male antennae. Also, czekanowskii is unlike any 
species of Apantesis in our fauna in having two strongly oblique pale 
lines meeting the inner margin of the forewing. Although I suspect that 
this species may fit just as well in the genus Apantesis, I leave it for the 
present as originally proposed. As the Alaskan specimen is the only one 


224 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


available, I prefer not to remove the abdomen for genital dissection until 
it has been figured in color for The Moths of America North of Mexico. 

Hampson (1920: 349) placed this species in the genus Phragmatobia, 
but it does not appear at all closely related to the group that includes 
the type-species, Phragmatobia fuliginosa (L.). Hampson also changed 
the spelling of the species name to czecanousci, this being an unjustified 
emendation which is a junior objective synonym of the name in its 
original form. 


Geometridae 
Semaeopus cantona (Schaus ) 


Cnemodes cantona Schaus, 1901: 192. 
Semaeopus cantona Prout, 1936: 88, pl. Ile. 

Several specimens of this species were collected in the Santa Ana Wild- 
life Refuge, Hidalgo, Texas by Mr. and Mrs. André Blanchard. The 
specimen that I examined was a male, taken 20 October 1970, and it 
appeared to agree exactly with the type in the collection of the U.S 
National Museum from Orizaba, Mexico (Fig. 3). The Texas specimens 
are all in the Blanchard collection. 


Triphosa affirmata (Guenée), new combination. 
Scotosia affirmata Guenée, 1857: 447, pl. 9, fig. 2 (not affirmaria Walker, 1860). 


I identified and returned to Mr. Charles P. Kimball a very rubbed 
female of this species taken at Homestead, Florida, 29 March 1969 (C. E. 
Hallas collection). It is a neotropical species described from Brazil but 
known from Mexico to Argentina. The specimen illustrated (Fig. 2) is 
from Guatemala. 


Epiplemidae 
Erosia incendiata Guenée 
Erosia incendiata Guenée, 1857: 35, pl. 8, fig. 4. 


A female of this species in very nearly perfect condition was found 
resting on the brick wall of the Department of Biology building at Texas 
A & I University, Kingsville, Texas, on 15 November 1968, and kindly 
donated to the U.S. National Museum by the collector, Dr. J. E. Gillaspy. 
The species is widespread in the neotropics from southern Mexico, in- 
cluding Yucatan, to French Guiana and Brazil. There is a good series in 
the U.S. National Museum mostly from Jalapa, Orizaba and Cordova, 
N . ico, Costa Rica and French Guiana. It was described from Brazil. 


pecies is sexually dimorphic, which will help to explain the dif- 


VOLUME 26, NUMBER 4 DONS) 


ference between the Texas specimen (Fig. 4) and the male figured by 
Guenée. The red coloring on the hindwing in Guenée’s illustration is 
greatly exaggerated. 


Antiplecta triangularis Warren 
Antiplecta triangularis Warren, 1906: 401. 


A single poor specimen in the collection of the United States National 
Museum taken at Brownsville, Texas, “March 27: 28,” at light, F. H. 
Benjamin (Fig. 5), appears to agree exactly with the type material of 
this species from Orizaba, Mexico, also in the U.S. National Museum. 
Benjamin had tentatively identified it, but as far as I know the record 
was never published. The specimen is a female. A closely related but 
obviously different species of Antiplecta occurs in southern Florida, and 
this one is probably undescribed. An example loaned to me for study 
by Mr. Charles P. Kimball was taken on Key Largo, 20 July 1962. 

After this paper was submitted, I found additional material of Anti- 
plecta triangularis in the collection of Mr. André Blanchard at Houston. 
A series of fresh specimens of both sexes was taken at the Santa Ana 
Refuge, Hidalgo Co., Texas, 14 November 1971. 


LITERATURE CITED 


GruM-GrsHmaiLo, G. 1900. Lepidoptera nova vei parum cognita _ regionis 
palaearcticae. Annuaire du Musée Zoologique de |’Académie impériale des 
Sciences de St. Pétersbourg, 4(1899): 455-472. 

GuENEE, A. 1857. Histoire Naturelle des Insectes, Species Général des Lépidoptéres, 
10: 1-584, illus. Paris. 

Hampson, G. F. 1920. Catalogue of the Lepidoptera Phalaenae in the British 
Museum, Suppl. 2, xxiii + 619 p., 30 pl. 

Prout, L. B. 1932-38. Die Amerikanischen Spanner (incomplete). In Seitz, Die 
Gross-Schmetterlinge der Erde, Band 8: 1-144, 17 pls. Alfred Kernen, Stuttgart. 

ScHAus, W. 1901. New Species of Geometridae from Tropical America. Trans. 
American Entomol. Soc. 27: 165-194. 

SEITZ, A. 1910-12. In Seitz, Macrolepidoptera of the World, 2, Palearctic Bombyces 
and Sphinges, 479 p., 56 pls. Alfred Kernen, Stuttgart. 

WARREN, WiLL1AM. 1906. Descriptions of new genera and species of South 
American geometrid moths. Proc. U.S. National Museum. 30: 399-557. 


bo 
bo 
(ep) 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


POPULATION EXPANSIONS AND MASS MOVEMENTS OF 
NYMPHALIS CALIFORNICA (NYMPHALIDAE ) 


Jerry A. POWELL 


Division of Entomology, University of California 
Berkeley, California 94720 


Mass movements by California tortoise shell butterflies have been 
sporadically recorded during the past 115 years in widespread parts of 
the Pacific Coast states. The flights, which follow massive population 
increases in relatively localized areas, have been called migrations in the 
literature (e.g. Williams, 1930, 1938). 

After about a decade of comparative stability, N. californica has again 
disrupted its equilibrium, and its swarms became conspicuous during 
1971. According to an Associated Press release dated 11 August, which 
was published in various parts of the country, butterflies “by the millions” 
were flying in the vicinity of Mt. Shasta, Siskiyou County, California, 
“slickening the highways” in the towns of Mt. Shasta, Dunsmuir, and 
McCloud (see Lepid. News, 1971, No. 5). 

Truck drivers were quoted as saying they could go only 25 miles per 
hour owing to the dense clouds ot buttterflies which crossed the highways. 
The report stated that the butterflies began appearing in hordes about 
five weeks previously [in early July]. Local naturalists reported that the 
flights consisted of six nymphalid and one lycaenid species (three of 
which do not occur in northern California ), but the roster did not include 
N. californica. That swarms did consist of the latter species, however, is 
indicated by several lines of evidence: a) correspondence to the editor 
of Lepidopterists’ News by five observers during the fall of 1971 agreed 
that this was the dominant species (Lepid. News, 1971 No. 6:1); b) R. 
Lyon (Pacific Southwest Forest and Range Experiment Station, Berkeley ), 
who witnessed the flight on 6 August, reported (in litt.) that the butter- 
flies appeared to be all one species and that it was identified as Nymphalis 
californica by R. Hill (formerly of the Pacific SW Forest and Range Expt. 
Sta., now retired). Hill made an on-site detection report to the Forest 
Service; c) J. Helfer (1971, The Mendocino Beacon, 20 Aug. 1971, p. 4) 
reported that the flights were observed by J. Myers who had just returned 
from Mt. Shasta, bringing a specimen which Helfer identified as N. 
californica. 

rom all reports the flight structure and timing are strikingly similar 
‘0 those which I observed in 1958 when I lived at Mt. Shasta all summer. 

iat season the butterflies appeared in large numbers on the mountain 


VOLUME 26, NUMBER 4 DONT 


slopes at 4,000-6,000 feet during 22-26 June, but were not seen to emigrate 
(snowline was then below 7,000 feet). Then during 3-7 August, a brood 
of fresh individuals appeared in tremendous numbers at 6,000 to 8,000 
feet and moved outward to subtending areas. I observed them in ag- 
gregations at damp soil in the town of Mt. Shasta and as far as three 
miles northeast of Weed (6 SW and 6 NW airline miles from the colonies 
observed on the mountain). The three towns mentioned in the AP 
release above are approximately 6 SW, 11 SSW, and 8 SSE airline miles 
from timberline, respectively, so that it seems probable that dispersal 
occurs in all directions into the 2,000-3,000 ft. elevation valleys which 
surround the 14,000 ft. mountain. 

How distant these flights extend is unknown, but in October 1971, 
mass flights were noted in two other parts of California where N. 
californica does not commonly occur in high numerical density, east of 
the Sierra Nevada in Mono County and in the San Francisco Bay area. 
These places are approximately 300 SE and 250 SW airline miles from 
Mt. Shasta. At Highway 395 near Mammoth Lakes, E. Kane (State De- 
partment of Agriculture, Sacramento ) (in litt.) counted 173 N. californica 
flying easterly during a five minute period at about 1530 on 8 October. 
In the Berkeley Hills near San Francisco Bay the butterflies were flying 
southeastward, along the axis of the hill ridge, at rates of 1.5 to 11/minute 
on a 50-foot sighting line during midday on 5, 7, and 12 October and 
were absent on 14 and 28 October (Powell, 1972). Moreover, circum- 
stances indicate that sightings reported by Arnaud (1972, in: Proc. Pacific 
Coast Entomol. Soc. 341st meeting, Pan-Pacific Entomol. 48: 72) at a 
height of 600 feet in downtown San Francisco on 4-5 October, also in- 
volved N. californica. 

The last time that an outbreak occurred in the San Francisco Bay area 
was in 1959-1960, when moderate aggregations were followed in the 
second season by tremendous numbers of the adults in June in Marin 
County. Various personal communication and literature reports which I 
have accumulated in addition to those summarized by Williams (1930, 
1938) show that the outbreaks are recurrent, especially at Mt. Shasta, 
where there are reports of the swarms for 1889, 1902, 1911, 1926, 1931, 
1932, 1952, 1958, and 1971. Most likely the phenomenon occurred in other 
years as well but was not reported, such as during the 1918-1922 and 
1941-1944 eras, judging from records at other localities. In any case, it 
appears that localized defoliation of Ceanothus, the preferred host plant, 
and mass movements of the butterflies occur at intervals of around 5-13 
years but too irregularly to allow precise prediction of any cyclic 
periodicity. 


228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Recorded outbreaks at secondary areas, such as Marin County in 1959- 
1960, apparently have resulted from preceding emigration from Mt. 
Shasta or other epidemic centers and have masked generalizations con- 
cerning possible periodic phenomena which may have obtained. 

The records suggest that this species periodically develops an im- 
balance with factors in its population equilibrium at isolated sites, fol- 
lowed by mass emigration of adults in various directions, and sometimes 
subsequent colonization and population explosions in secondary districts 
in the first or second following year. The populations then return to low 
density numbers or may disappear altogether in outlying colonies. Thus, 
Nymphalis californica should not be considered a migratory species 
except in the broadest sense. 


LITERATURE CITED 


Powe LL, J. A. 1972. Mass movements of Nymphalis californica (Boisduval) in 
the San Francisco Bay area during 1971 (Lepidoptera: Nymphalidae). Pan- 
Pacific Entomol. 48(2): 144. 

WiuiaMs, C. B. 1930. The Migration of Butterflies. Oliver and Boyd, London. 
473 p. 

1938. Recent progress in the study of some North American migrant 

butterflies. Ann. Entomol. Soc. Amer. 38: 211-239. 


TWO NAME CHANGES FOR SUBTROPICAL AMERICAN PIERIDAE 


On a recent visit to Oxford I had an opportunity to examine the collection 
of unpublished drawings known to early authors as “Jones's Icones.” Many of 
these drawings, and probably the specimens upon which they were based, were 
used by Fabricius. Some time ago it had been suggested to me that the current 
interpretation of castalia Fabricius, 1793, is grossly incorrect. This is one of the 
names based upon a Jones figure. Upon examining the plate in question at Oxford 
I found that it is a clear representation of what today is called Appias drusilla 
(Cramer), [1777]. Therefore, Papilio castalia Fabricius, 1793, is a synonym of 
Papilio drusilla Cramer, 1777. 

There are several recognized subspecies of drusilla. Jones’s figure is based upon 
a Jamaican specimen and represents the subspecies named jacksoni Kaye, 1920. 
This name must yield to the earlier Fabrician name. The Jamaican subspecies 
must be called Appias drusilla castalia (Fabricius), 1793. 

Transfer of the name castalia from Kricogonia to Appias requires recognition of 
lyside Godart, 1819, as the specific name for West Indian Kricogonia. Although 
dos Passos in his Synonymic List . . . recognized two species in the genus, castalia 
and lyside, Mr. Thomas Turner has demonstrated through breeding experiments 
that the two develop from eggs laid by a single female. Therefore only one 
species is involved. Its name is Kricogonia lyside (Godart). Haiti probably is the 
type locality for lyside. 


NorMAN D. Ritey, British Museum (N. H.), London, England. 


VOLUME 26, NUMBER 4 229 


OBSERVATIONS AND NEW RECORDS 
OF IOWA RHOPALOCERA 


STEPHEN MILLER 


12585 Jones Bar Road 
Nevada City, California 95959 


Our knowledge of the Lepidoptera of Iowa has suffered in recent 
years from minimal collecting and a resultant scarcity of published in- 
formation. With the exception of a short paper by Miller (1961), 
Christenson’s (1971) recent work is the first significant study of the 
state’s butterfly fauna to appear in nearly fifty years. Encouraged 
through communication with Christenson and by the accessibility of 
available data, I decided to make a personal survey of Iowa butterfly 
populations in the vicinity of Iowa City (Johnson County) during a short 
period of residence in the state in 1971. 

The more fragile elements of Iowa's native flora and fauna have suf- 
fered profoundly from the extensive commercial modification the state 
has experienced during the last fifty years. Thus I was interested in ex- 
amining the few areas which had escaped the plow and were relictual 
associations of flora which predominated in the state prior to the advent 
of cultivation. 

Christenson (1971) mentions 22 species of butterflies that have not 
been collected in lowa since 1920, but were recorded prior to that time. 
I was eager to investigate the possibility that some of those species sur- 
vived the drastic modification of the Iowa landscape and continued to 
maintain populations in the biotic refugia of the area. These areas might 
be expected to support populations of species of limited distribution in 
Iowa, owing to plant associations and other biotic requirements which 
only such isolated refugia could provide. Of the 130 species of butter- 
flies known to occur in Iowa, 19 had been recorded from one county 
only (Christenson, 1971). Though 10 species appear to reach their dis- 
tributional limits within the state (op. cit.), which might partially account 
for some of the single-county records, this fact also suggested the im- 
portance of examining the fauna of plant communities which had pre- 
viously been overlooked. The recent state records of Miller (1961) also 
seemed to support this notion. 

Several localities in Johnson County were selected which appeared to 
meet these considerations and, due to their proximity to my home, could 
be visited on a regular basis throughout the season: Williams Prairie, 
a small private preserve about 4 miles north of Oxford; the vicinity of 


230 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Cou Falls (the northeast corner of the 13,000-acre Hawkeye Wildlife Area 
which surrounds the Coralville Reservoir west of Highway 218); Mac- 
Bride Field Campus of the University of Iowa, about 4 miles north of 
North Liberty; and the area to the southeast of the Field Campus and 
north of the Coralville Reservoir known as Sugar Bottoms. These localities 
were visited at least bi-weekly for the three-month period 20 April—20 
July, following which other considerations were responsible for terminat- 
ing my investigations. In addition, Muskrat Slough, a public hunting 
access owned and maintained by the State Conservation Commission and 
located approximately 7 miles north-northeast of Mechanicsville in Jones 
County, was visited on two occasions in July, and one trip was made 
to the Paint Creek Unit of the Yellow River State Forest, near Waukon 
Junction in Allamakee County, on 10-11 July. 

Of the areas selected, Muskrat Slough is a typical marshland situation. 
Cou Falls, MacBride Field Campus, Sugar Bottoms and Yellow River 
State Forest are, in general, representative of the deciduous oak-hickory 
woodland bordering most of the state's major rivers. Virgin prairie, 
once the characteristic feature of the Iowa landscape, has succumbed 
nearly in toto to extensive cultivation and exists today essentially as four 
state-owned preserves and a few private holdings. Williams Prairie is 
representative of what little remains of these fascinating ecological com- 
munities. The flora of these areas has been described by Conard (1958). 

Following is a descriptive list of the more significant records and ob- 
servations. Included are 35 new county records representing 31 species, 
5 of which had previously been reported from one or two counties only. 
In addition there are new records of all four species recently reported 
as new to the state by Miller (1961). Though I failed to discover popu- 
lations of any of the 22 species which have not been collected in the 
state since 1920, 7 species were taken for which only one or two recent 
records exist. It is my hope that some of the observations which follow 
will serve to stimulate future investigations of Iowa’s butterfly fauna, 


especially in view of the continuing threat of additional commercial ex- 
ploitation. 


HESPERIIDAE 


Euphyes dion (Edwards). 5, 13 July 1971, Muskrat Slough, Jones Co. (5 $64). 
This species was first credited to the Iowa fauna by Miller (1961), who took speci- 
mens at Pilot Knob State Park and a swamp near Klemme, both Hancock County, 
on 22 July 1960. With the exception of a few specimens in a display collection 
at the University of Iowa labelled “Banner area, Warren County” and bearing dates 
in ne mid-1960s, the present series is the first taken since that time. New county 
record, 


Euphyes conspicua (Edwards). 28 June, 1, 20 July 1971, Williams Prairie, Johnson 


VOLUME 26, NUMBER 4 Dx) 


Co. (13 64,4 22); 5, 13 July 1971, Muskrat Slough, Jones Co. (4 64,1 @). 
New county records. 

Euphyes bimacula (Grote & Robinson). 19, 21, 23, 28 June, 1 July 1971, Williams 
Prairie (10 6 6,6 2@@). New county record. 

It should be noted that E. dion appears to be absent from the Williams Prairie 
fauna. On several occasions during July, when the species was flying at Muskrat 
Slough, an intensive search of the area failed to produce specimens. In conjunction 
with the need to determine whether E. bimacula occurs at Muskrat Slough (my first 
visit to the area was made at the terminus of the species flight), this observation 
provides excellent opportunity for subsequent investigations of host-plant specificity 
and biotic studies in general of these poorly-known hesperiids. 

Poanes viator (Edwards). 5 July 1971, Muskrat Slough, Jones Co. (1 @). All 
previous records of this species have been from Pilot Knob State Park, Hancock 
County, where Miller (1961) first discovered it in Iowa on 22 July 1960. New 
county record. 

Polites origines (Fabricius). 23 June 1971, MacBride Field Campus, Johnson Co. 
(1 6). Christenson (1971) mentions the rarity of this species in lowa. New county 
record. 

Polites mystic (Edwards). 19, 21 June 1971, Williams Prairie, Johnson Co. 
(7 68,6 22). P. mystic seems to be very locally distributed throughout Iowa. 
This previously unrecorded population is of some interest due to the extent of varia- 
tion apparent in the series. Specimens run as dark in the ground color of the ventral 
hindwing as typical eastern mystic to even lighter than the norm of the prairie sub- 
species dacotah (Edwards). Specimens collected farther west in Dallas County by 
Miller (in litt.) seem referable to dacotah. Other Iowa populations should be closely 
examined to determine the extent of this blend zone. New county record. 


LYCAENIDAE 


Harkenclenus titus (Fabricius). 28 June, 1 July 1971, Williams Prairie, Johnson 
Co. (6 6 6,1 2); 3 July 1971, Cou Falls, Johnson Co. (1 ¢ ); 11 July 1971, Yellow 
River State Forest, Allamakee Co. (2 92 ¢@). An unusually abundant species in 1971. 
New county records. 

Satyrium liparops strigosa (Harris). 11 July 1971, Yellow River State Forest, 
Allamakee Co. (1 @). One of three recent captures. New county record. 

Satyrium acadica (Edwards). 1 July 1971, Williams Prairie, Johnson Co. (1 ¢). 
With the exception of three specimens in a display collection at the University of 
Iowa taken at Sheeder Prairie, Guthrie County, on 23 June 1965, the present record 
is the first for the species in recent years. New county record. 

Callophrys (Incisalia) henrici (Grote & Robinson). 22, 23, 24 April 1971, Sugar 
Bottoms, Johnson Co. (2 6 6,1 @). Prior to my discovery of this species in Johnson 
County it had been recorded only from Fremont and Pottawattamie Counties in the 
extreme southwestern corner of the state. Christenson (1971) implies that the 
paucity of records in Iowa may be due to the absence of redbud (Cercis canadensis 
Linnaeus ), a commonly-mentioned host, from most areas of the state. This plant does 
not occur in the Sugar Bottoms area. Prunus, however, also mentioned as a host, is 
abundant there. New county record. 

Callophrys (Mitoura) gryneus (Hiibner). 22 April 1971, Sugar Bottoms, Johnson 
Co. (1 6, 1 2); 23, 25 April, 8 May, 14 July 1971, 2 miles west of Cou Falls, 
Johnson Co. (14 66, 6 9@). Previously recorded only from Henry and Linn 
Counties, I found C. gryneus to be fairly abundant at both locales where I discovered 
it in Johnson County, though restricted to the immediate vicinity of its foodplant, 
Juniperus virginiana Linnaeus (red cedar). It was also observed, though not col- 
lected, at Effigy Mounds National Monument, Allamakee County, on 11 July 1971. 
Further collecting should prove the species to be much more widely distributed in 
Iowa than the records indicate. New county record. 


232 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Panthiades m-album (Boisduval & LeConte). 3 May 1971, Cou Falls, Johnson 
Co. (1 @). Miller (1961) took a single female of this species at Waubonsie State 
Park, Fremont County (extreme southwest Iowa), on 22 May 1960, thus discovering 
it for the first time in Iowa. The fact that the present specimen, the second recorded 
from the state, was taken in fresh condition at the same time of year seems to indicate 
that P. m-album very likely breeds in Iowa, if only sporadically. Miller (in litt.) 
is of the opinion that the species probably occurs in small, very local populations 
throughout at least the southern half of the state. New county record. 

Lycaena thoe Guérin-Ménéville. 28 May, 5, 12, 19, 21, 23 June 1971, Williams 
Prairie, Johnson Co. (11 66,5 2@). 

Lycaena xanthoides dione Scudder. 19, 21, 23, 28 June, 1 July 1971, Williams 
Prairie, Johnson Co. (11 66,9 9 @). 

Lycaena helloides (Boisduval). 28, 29 May, 5 June, 1 July 1971, Williams Prairie, 
Johnson'Co, (15) GG ores 

Lycaena phlaeas americana Harris. 28 May, 23 June 1971, Williams Prairie, Johnson 
Co. (11 24, 7 2@). Two of the 11 males collected at this locality are of the 
aberrant phenotype “fasciata” (Strecker). 

Mention is made of the occurrence at a single locale of all four species of Lycaena 
native to the state in light of the unusual nature of this observation. There are 
probably very few spots in Iowa where this situation exists. There are records for 
each of the four species from Story County (Christenson, 1971), though it is not 
known whether there is any area in the county where the populations occur sym- 
patrically. Data on host-plant specificity for each of the species at Williams Prairie 
would be of great interest, as well as observations on the territorial aspects of court- 
ship and mating. 


SATYRIDAE 


Lethe eurydice fumosa (Leussler). 19, 21, 23 June 1971, Williams Prairie, Johnson 
Co. (20 6 6,3 22); 5 July 1971, Muskrat Slough, Jones Co. (4 6 6,1 2). Special 
notice is given the present records of L. e. fumosa in light of the recent treatment 
of the Lethe eurydice complex by Cardé et al. (1970). This subspecies appears to 
be restricted to “small, isolated colonies (many now extinct) in sedgy, permanent 
marshes in the prairie regions from Minnesota and South Dakota to Indiana, Ne- 
braska and Colorado” (op. cit.). The early stages of fumosa are unknown, and the 
discovery of two large populations of the subspecies in eastern Iowa should increase 
the feasibility of subsequent investigations of the biology of this insect. Miller (in 
litt.) has seen typical eurydice from northern Iowa, but does not have the exact 
locality. Collectors should also be aware of the possible occurrence of Lethe appalachia 
R. bs Chermock within at least the eastern third of the state. New Jones County 
record. 

Cercyonis pegala (Fabricius). 28 June, 1 July 1971, Williams Prairie, Johnson 
Co. (15 @ @). Emmel (1969) states in his discussion of C. pegala that the alope 
(Fabricius ) phenotype “ranges from Virginia and New Jersey north to eastern Quebec 
and Maine,” and indicates that to the north and west of this area it intergrades with 
the nephele (Kirby) phenotype, which evidences none of the yellow forewing 
patch of alope. One would expect Iowa populations to be predominantly of the 
nephele phenotype and, indeed, most records from the state refer to this morph. It 
was with some surprise, then, that of the 15 males taken at Williams Prairie two 
display the conspicuously yellow-patched forewing of alope. All 13 additional 
specimens taken at Williams Prairie and those collected at other localities in Iowa 
during 1971 are typical nephele. Miller (in litt.) has occasionally taken alope-like 
emales farther west in Polk and Dallas Counties, but has seen no males from those 

3 with a yellow-patched forewing. Further collecting may uncover other 
orphic populations of C. pegala in Iowa. 


VOLUME 26, NUMBER 4 rors) 


Additional New County Records 
HESPERIIDAE 


Euphyes vestris metacomet (Harris). 10, 16 June, 3 July 1971, Cou Falls, Johnson 
Co. (3 6 ¢@); 11 July 1971, Yellow River State Forest, Allamakee Co. (2 9 @ ). 

Poanes hobomok (Harris). 6, 9, 10, 15, 16 June 1971, Cou Falls, MacBride Field 
Campus, Johnson Co. (7 646,322). 

Pompeius verna (Edwards). 3 July 1971, Cou Falls, Johnson Co. (2 ¢@ @ ). 

Wallengrenia otho egeremet (Scudder). 14 July 1971, Cou Falls, Johnson Co. 
(1 6); 11 July 1971, Yellow River State Forest, Allamakee Co. (1 ¢). 

Polites themistocles (Latreille). 28 May, 6, 9, 15 June 1971, Cou Falls, MacBride 
Field Campus, Williams Prairie, Johnson Co. (6 ¢ 6,1 @). 

Ancyloxypha numitor (Fabricius). 9, 12, 19, 21 Jume 1971, MacBride Field 
Campus, Williams Prairie, Johnson Co. (4 66,1 @). 

Erynnis brizo (Boisduval & LeConte). 22, 23, 24 April, 3, 9 May 1971, Sugar 
Bottoms, Cou Falls, Johnson Co. (7 646, 6 @¢@). 

Erynnis horatius (Scudder & Burgess). 3 May 1971, Cou Falls, Johnson Co. (1 ¢ ). 
(det. H. A. Freeman). 

Epargyreus clarus (Cramer). 9 May, 6, 9, 15, 16 June 1971, Cou Falls, MacBride 
Field Campus, Johnson Co. (5 ¢ 6,1 @). 


LYCAENIDAE 


Satyrium calanus falacer (Godart). 11 July 1971, Yellow River State Forest, 
Allamakee Co. (1 ¢ ). 

Strymon melinus humuli (Harris). 3 July 1971, Cou Falls, Johnson Co. (1 ¢). 

Everes comyntas (Godart). 23 April, 15 May, 9, 12 June 1971, Sugar Bottoms, 
MacBride Field Campus, Williams Prairie, Johnson Co. (4 66,3 @@). 

Celastrina argiolus pseudargiolus (Biosduval & LeConte). 11 July 1971, Yellow 
River State Forest, Allamakee Co. (1 @ ). 


NYMPHALIDAE 


Chlosyne gorgone carlota (Reakirt). 20, 27, 28, 29 May 1971, Cou Falls, Williams 
Prairie, Johnson Co. (5 ¢ 46,2 2@). 

Boloria toddi ammiralis (Hemming). 11 July 1971, Yellow River State Forest, 
Allamakee Co. (1 ¢ ). 

ae eee (Fabricius). 11 July 1971, Yellow River State Forest, Allamakee 
Con(2 $s ); 

Speyeria aphrodite alcestis (Edwards). 11 July 1971, Yellow River State Forest, 
Allamakee Co. (1 ¢). 


SATYRIDAE 
Lethe anthedon (Clark). 11 July 1971, Yellow River State Forest, Allamakee 
Co. Gi®,): 


ACKNOWLEDGMENTS 


My thanks to Lee D. Miller for reading the manuscript and providing 
additional data from his personal collecting in Iowa, and to C. Don Mac- 
Neill and Michael M. Collins for reviewing the manuscript and making 
several pertinent suggestions. 


234 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


LITERATURE CITED 


Carpr, R. T., A. M. SHaprro & H. K. Crencn. 1970. Sibling species in the 
eurydice group of Lethe (Lepidoptera: Satyridae). Psyche 77(1): 70-103. 
CuristENson, M. C. 1971. An annotated checklist of the butterflies of Iowa. M.A. 

Thesis, University of Northern Iowa, Cedar Falls (unpublished). 

Conarp, H.S. 1958. Plants of Iowa. 7th edition (published by the author). 90 p. 
EMMEL, T. C. 1969. Taxonomy, distribution and biology of the genus Cercyonis 
(Satyridae). I. Characteristics of the genus. J. Lepid. Soc. 23(3): 165-175. 
Miter, L. D. 1961. Notes on nine Iowa butterfly species, including four new 

to the state. J. Lepid. Soc. 15(2): 97-98. 


SOME NOTES ON THE SPHINGIDAE 


Since the appearance over a year ago of the first published part of the continuing 
work on the Moths of America (Hodges, R. W. in Dominick, R. B. et al., 1971, 
Moths of America North of Mexico, fasc. 21, Sphingoidea), some additional informa- 
tion has been gathered by the author on this group in the area of McClellanville, 
South Carolina. 

Darapsa myron (Cramer) is taken frequently at bait. We use fermented peaches 
or bananas, no extras added, with good results. It should be added that myron 
and D. pholus (Cramer) generally are not easy to differentiate in the traps, if only 
to emphasize the difficulties inherent in field identification. 

Darapsa versicolor (Harris) occurs here somewhat later than stated in the reference, 
being taken at light in latter July and August, though we have only half a dozen 
specimens in the Wedge Plantation collection (WPC). 

Paonias astylus (Drury) in this locality has two definite broods, the first in April, 
and the second in late July to early August. All WPC specimens to date have been 
taken at light. 

Sphinx franckii Neumoegen. Four specimens have been taken. One on 8 June 
1968, and three in 1971, dated 23 June, 14 August, and 1 September. All were ¢ 2, 
and all came to light. This sudden 1971 take of three induced us to look for the 
foodplant, ash, so far without success. Either there is undiscovered ash nearby, or 
in this locality franckii has fixed on some other foodplant. 

Erinnyis obscura (Fabricius), one 6, 25 October 1968, to light. 

Deidamia inscripta (Harris). Hodges notes that it flies just before sunrise. This 
information was taken from a note in the WPC collection which related to only 
two specimens that were actually seen by me to fly into one of our traps at that time. 
The collection, however contains not only several specimens flying at this time of 
day, but also a goodly number flying from roughly midnight on. It would therefore 
seem more accurate to say that it is a late flier. Both sexes have been taken at light 
(none at bait), and the larva has been reared on Vitis sp. 

I should like to add at this point that the editors of Moths of America North of 
Mexico hope that such additional information as this will be published as available 
by various workers, for we fully realize that there is much still to be learned and 
much that collectors and institutions have already available which has not been 
assembled. Many life histories, distribution records, habits and other information 
of interest have been studied and recorded by individuals who have not published, 
with resulting gaps in the literature. 


RicHarp B, Dominick, The Charleston Museum, Charleston, South Carolina 29401. 


VOLUME 26, NUMBER 4 U5 


CONFIRMATION OF A DISPUTED FOODPLANT OF 
PAPILIO GLAUCUS (PAPILIONIDAE) 


J. Mark SCRIBER 
Department of Entomology, Cornell University, Ithaca, New York 14850 


Although reported as a polyphagous species, feeding on more than 13 
families, 21 genera, and 34 species of plants, Papilio glaucus L. does 
appear to have different foodplant preferences in different regions over 
its range (Brower, 1958; Remington, 1968). 

Because foodplant records in the literature are frequently suspect 
(Brower, 1958; Shields, Emmel, & Breedlove, 1970), I feel it important 
to verify a single, very early observation by Sir John Abbot, who recorded 
P. glaucus feeding on hop tree, Ptelea trifoliata (Rutaceae). This 
observation was made in Georgia and was reported in his manuscripts 
(Abbot, 1792-1804), which have been cited by Boisduval & LeConte 
(1833), D’'Urban (1857) and Scudder (1889). I have been unable to 
find any other recorded observations of P. glaucus on Ptelea. 

My observation was made on 5 July 1971, in the Cornell Plantations 
near Ithaca, New York. A very early second instar larva was found 
resting on a leaflet approximately six feet off the ground. This larva was 
taken back to the laboratory and reared for positive identification through 
all of its stadia on leaves from this plant. This wafer ash (hop tree) 
is in the open and is maintained as part of the Plantation. 

Within the rest of the P. glaucus group, Kendall (1957, 1964) reported 
finding Papilio multicaudatus Kirby larvae on Ptelea trifoliata in Texas, 
confirming Behrs (1884) observation in California. Comstock (1927) 
reported Ptelea baldwinii as a foodplant for Papilio rutulus Boisduval 
in California. This hop tree has been incorrectly interpreted as ‘hop’ 
(Humulus) since that time (see Brower, 1958), and perhaps such con- 
fusion is responsible also for the rather unlikely records of P. glaucus 
feeding on Humulus lupulus (e.g. Scudder, 1889; Teitz, 1952). Possibly 
the ‘ash, Fraxinus trifoliata, referred to by Couper (1874) as a foodplant 
of P. glaucus, was also in reality the wafer ash, Ptelea trifoliata. 

I feel that my observation of P. glaucus on Rutaceae is interesting from 
the standpoint of the ‘synergistic co-evolution’ of the Papilionidae and 
their foodplants (Slansky, in press). Unlike related smooth, green, eye- 
spotted larvae in Asia (Papilio bianor Cram. group) which feed on 
Rutaceae (Jordan, 1908), the P. glaucus group and the Papilio troilus L. 
group are both believed to have arisen in the New World and to have 


236 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


fed originally on Lauraceae and Magnoliaceae before expanding their 
foodplant diets (Forbes, 1932, 1958; Munroe, 1960). 

Having both the essential oils of the Umbelliferae and the alkaloids of 
the Aristolochiaceae and Magnoliales (including Magnoliaceae, Lauraceae, 
and Annonaceae ), the Rutaceae seem to have played a key role in much 
of the co-evolution of the Papilionidae and their foodplants (Dethier, 
1941: Ehrlich & Raven, 1965). Whether or not this P. glaucus-Rutaceae 
interaction is some relic from the phylogenetic past of P. glaucus, or a 
secondary consequence of its polyphagous habit, is unknown. 


LITERATURE CITED 


Azssor, J. 1792-1804. Drawings of the Insects of Georgia. 17 vols. Library of 
the British Museum. Also Boston Society of Natural History: Oemler and Gray 
Collections of Harvard University Library. 

Beur, H.H. 1884. Biological synopses of California Lepidoptera. Bull. Calif. Acad. 
Sci. 1: 63-65. 

BorspuvAL, J. A. & M. J. LeConre. 1833. Histoire générale et iconographie des 
Lépidopteres des Chenilles de |’ Amerique septentrionale. Librarie Encyclopedique 
de Roret. Paris. 228 p. 

Brower, L. P. 1958. Larval foodplant specificity in butterflies of the P. glaucus 
group. Lepid. News 12: 103-114. 

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

Couper, W. 1874. A dissertation on northern butterflies. Can. Entomol. 6: 91-96. 

Detuier, V. G. 1941. Chemical factors determining the choice of foodplants by 
Papilio larvae. Amer. Nat. 75: 61-73. 

D’Ursan, W. S. M. 1857. On the order Lepidoptera, with the description of two 
species of Canadian butterflies. Can. Nat. & Geol. 2: 223-236. 

Eneuicu, P. R. & P. H. Raven. 1965. Butterflies and plants: a study in co-evolution. 
Evolution 18: 586-608. 

Forses, W. T. M. 1932. How old are the Lepidoptera? Amer. Nat. 98: 452-460. 

. 1958. Caterpillars as botanists. Proc. 10th Int. Cong. Entomol. 1: 313-317. 

Jorpan, K. 1908. The Indo-Australian Rhopalocera, in A. Seitz, Macrolepidoptera 
of the World. Vol. 9. Stuttgart. 

KENDALL, R. O. 1957. New foodplants in Texas for Papilio multicaudatus. Lepid. 
News 11: 224. 

. 1964. Larval foodplants of twenty-six species of Rhopalocera (Papilionidae ) 
from Texas. J. Lepid. Soc. 18: 129-157. 

Mounrog, E. 1960. The generic classification of the Papilionidae. Can. Entomol., 
Suppl. 17. 5 p. 

Remincron, C. L. 1968. Suture zones of hybrid interaction between recently 
joined biotas, in T. Dobzhansky, ed., Evolutionary Biology 2: 321-428. 

ScuppER, S. H. 1889. The Butterflies of the Eastern U. S. and Canada. 3 vols. 
Publ. by the author. Cambridge, Mass. 

SuieLps, O., J. M. EMmMen & D. E. BreEpLove. 1970. Butterfly larval foodplant 
records and a procedure for reporting foodplants. J. Res. Lepid. 8: 20-36. 

SLANSKY, F, 1973. Latitudinal gradients in species diversity of the New World 
swallowtail butterflies (Papilionidae). J. Res. Lepid. (in press). 

Tmtz, H. M. 1952. The Lepidoptera of Pennsylvania, a Manual. Pennsylvania 
State College, Pa. 194 p. 


VOLUME 26, NUMBER 4 Doi 


THE BIOLOGY OF CALLOPHRYS (INCISALIA) 
FOTIS BAYENSIS (LYCAENIDAE)! 


JoHN F. EMMEL 
1808 Eighth Avenue, San Francisco, California 94122 


AND 


CLIFFORD D. FERRIS? 
College of Engineering, University of Wyoming, Laramie, Wyoming 82070 


Despite its occurrence in a region which has been explored by lepi- 
dopterists for over a century, Callophrys (Incisalia) fotis bayensis Brown 
was not discovered until 1962 (Brown, 1969a). At the present writing, 
the type locality of this unique butterfly, the San Bruno Mountains in 
San Mateo County, California, is about to be “developed” for homesites 
and commercial properties. Although efforts are being made by con- 
servationists to halt this destruction of the natural habitat, it appears 
unlikely that the San Bruno Mountains will remain unaltered. For this 
reason, numerous visits have been made to this area over the past four 
years to record as much of the biology of the butterfly fauna as possible 
before it is permanently lost to science. Many of the data in this paper 
were gleaned during these investigations. 

Brown (1969b) published a brief description of the larva and habitat 
of C. f. bayensis. Our purpose is to provide additional detailed informa- 
tion on the biology of the insect. Emmel carried out intensive field work 
in the San Bruno Mountains to document preferred habitat, flight span, 
and behavior of adults and immatures. Ova were sent to Ferris for rearing 
and describing of the immature stages. To determine the regional dis- 
tributions of bayensis, Emmel surveyed the San Francisco Bay area from 
1968 to 1971 to locate populations of Callophrys fotis outside of the type 
locality. 


Distribution 


The San Bruno Mountains are located on the San Francisco Peninsula 
at the northern end of San Mateo County, California. The range is ap- 
proximately four miles long and one to two miles wide, and runs from 
northwest to southeast. Elevation ranges from sea level at the eastern 
end on San Francisco Bay to 1314 feet at the highest peak. Callophrys 


1 Published with the approval of the Director, Wyoming Agricultural Experiment Station as 
Journal Article no. 513. 
2 Research Associate, Allyn Museum of Entomology, Sarasota, Florida. 


238 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


fotis bayensis has been collected in at least six localities in the San Brunos. 
These localities are generally situated on north-facing slopes where 
moisture is conserved and the foodplant grows in abundance. 

C. f. bayensis has been found in two localities outside of the San Bruno 
Mountains, both in San Mateo County. One locality is on a north-facing 
slope near the south end of Milagra Ridge, just north of Sharp Park Road 
at 400-550 feet elevation. This is about four airline miles southwest of the 
San Bruno Mountains. Immatures were taken here in April and May 
1969, and in May 1970. Only about 50% of the adults from this population 
are typical bayensis; the remainder superficially resemble C. f. schryveri 
Cross, or C. f. mossii (H. Edwards ). 

The other locality, outside of the type locality, is on the northwest flank 
of Montara Mountain at 900-950 feet elevation, 1.1 air miles south of the 
Linda Mar School in Pacifica. This area, in the northernmost extension 
of the Santa Cruz Mountains, is about nine miles from the San Bruno 
Mountains. Immatures were collected here in April 1969, and adults were 
taken in March 1970. All specimens from this locality appear to represent 
typical bayensis. 

Suitable habitats have not been found south of the Montara Mountain 
area. To the north of the San Bruno Mountains, two localities which have 
habitats similar to the type locality have been discovered. These localities, 
however, apparently do not support populations of bayensis, presumably 
because of limited foodplant biomass. One locality is on the north slope 
of Twin Peaks in San Francisco at 700-800 feet elevation where a fairly 
extensive Sedum spathulifolium colony grows. In the past, this area may 
have supported a bayensis colony which was exterminated through the 
gradual destruction of the habitat by housing developments. 

The other locality is on a north-facing slope 1.0 air mile west of Yellow 
Bluff in the Fort Baker Military Reservation, south of Sausalito in Marin 
County. This area is about seven airline miles from the San Bruno 
Mountains. While the foodplant is locally abundant here, repeated visits 
have failed to produce evidence that bayensis occurs in the area. Further 
north in Marin County, a distinct new subspecies of Callophrys fotis has 
been found in a habitat markedly different from that which supports 
bayensis. This new subspecies is being described in a separate paper. 

Fig. 1 shows the known distribution of bayensis; records of adult and 
larval collections are given below. 


Habitat and Foodplant 


An excellent description of the climate, geology, and flora of the San 
bruno Mountains is provided by McClintock & Knight (1965). The 


VOLUME 26, NUMBER 4 239 


SS OS 
_— 


O san 


8SRUNO 


San Andreas 


Lake 


Fic. 1. Map showing locations where C. fotis bayensis occurs. The stippled circles 
represent the known colonies. SBM = San Bruno Mountains; SP = Sharp Park; 
M = Montara Mountain. 


Milagra Ridge and Montara Mountain localities have a climate and 
habitat which are very similar to those of the localities in the San Bruno 
Mountains. 

As noted above, bayensis is typically found on steep north-facing slopes 
where its foodplant, Sedum spathulifolium Hooker (Crassulaceae ), grows 
abundantly. Representative vegetation on these slopes includes Rhus 
diversiloba Torr. and Gray (Poison Oak), Berberis pinnata Lag. (Coast 
Barberry), Baccharis pilularis DC. (Coyote Brush), Anaphalis mar- 
garitacea (L.) Gray (Pearly Everlasting), Erigeron glaucus Ker. (Seaside 
Daisy), Dudleya farinosa (Lindl.) Br. and R., Arabis blepharophylla H. 
and A. (Coast Rock Cress), Eriogonum latifolium Sm. (Coast Buck- 
wheat), Ranunculus californicus Benth. (California Buttercup), and 
Lomatium utriculatum ( Nutt.) C. and R. (Bladder Parsnip ). 

Sedum spathulifolium blooms from April to June. In areas where the 


YAO JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


plant is abundant, the bright yellow flowers and red stems form a virtual 
carpet of color over the rocky ground. These colors are reflected in the 
cryptic patterns on the third and fourth instar larvae (see larval descrip- 
tion below). 


Field Observations of Adults and Immatures 


Emergence of adults takes place primarily during the month of March. 
A series of adults taken on 15 March 1970 exhibits a complete spectrum 
between worn and fresh specimens. In some years, a few adults probably 
emerge as early as late February. 

No visits to the habitat of bayensis were made earlier than 1030 PST, 
at which time flight activity appeared to be maximal. It would appear 
that flight begins at least an hour earlier in the day. No data were ob- 
tained on flight activity during the afternoon hours. Adults of both sexes 
remained in close proximity to the foodplant. Males were much more 
active than females, often landing repeatedly on small shrubs, and they 
appeared to exhibit territorial behavior. Several males were noted nectar- 
ing at flowers of Ranunculus californicus, Arabis blepharophylla, and 
Lomatium utriculatum. Females tended to settle on the foodplant where 
they remained unless disturbed, and then they flew only short distances. 

Oviposition takes place in March and early April. In one of the San 
Bruno Mountains localities, a total of five ova were located on leaf upper- 
sides of the foodplant. Confined females oviposited freely on both upper- 
sides and undersides of Sedum leaves. 

The first and second instar larvae bore into the succulent leaves. By the 
time the third instar is reached in nature, the Sedum plants are beginning 
to bloom and the larvae generally move up to the flowerheads to feed. 
In many cases, the larvae fail to locate flowering stalks, and feed through 
to maturity on the centers of leaf rosettes. Brown (1969b) states that the 
color of the mature larva depends upon the color of the part of the food- 
plant on which it feeds; we noted no such relationship. Larvae reared 
by Ferris on leaves alone displayed three distinct color morphs. In the 
field, Emmel observed mature larvae of the various color morphs to be 
randomly distributed with regard to the color of the foodplant parts on 
which they were feeding. Myrmecophily was never observed in the field 
studies. 

Pupation was not observed in the field, but probably occurs in ground 
litter under or near the Sedum plants. Eclosion of adults in the laboratory 
invariably occurred during the early morning hours, usually within an 
hour of being exposed to the first light of day. 


VOLUME 26, NUMBER 4 241 


Fic. 2. Larva, ovum, and pupa of Callophrys fotis bayensis: a. first instar larva, 
dorsal view; b. first instar larva, cross-section through middle segment; c. first instar 
larva, lateral view; d. second instar larva, lateral view; e. second instar larva, cross- 
section through middle segment; f. third instar larva (mature), dorsal view in normal 
feeding position; g. third instar larva, cross-section through middle segment; h. ovum, 
dorsal and lateral views and enlargement of hexagonal cell pattern; i. pupa, lateral 
view. 


YAP, JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Description of Immature Stages 


In March 1970, several females collected by Emmel in the San Bruno 
Mountains were confined over Sedum spathulifolium plants to obtain ova 
for life history studies. These females were placed in small jars containing 
several leaf rosettes of the foodplant. The jars were covered with netting 
and placed under a goose-neck lamp using a 75 watt bulb. The females 
oviposited freely on the Sedum. The ova obtained were sent to Ferris, who 
reared them on transplanted specimens of Sedum spathulifolium and 
described the immature stages. 


Ovum: The eggs were pale green oblate spheroids approximately 0.8 mm in 
diameter. The eggs hatched in five to six days, and turned opaque white prior to 
emergence of the larvae. 

First instar: The first instar larvae were pale yellow-green with clear hairs upon 
emergence from the eggs. They turned somewhat darker prior to moulting. The 
newly emerged larvae did not eat the egg shells. Stadium one lasted two days and 
the larvae increased in size from 1 mm to 2 mm. 

Second instar: In this instar, the larvae were pink with pale hairs. Stadium two 
lasted three days with growth increase from 2 mm to 3 mm. The mature second 
stadium larvae varied in color from pale to dark pink. 

Third instar: Initially, the larvae were a dark cream or straw color with con- 
siderable pink mottling, and were covered with short black spines. The larvae re- 
mained in the third instar from 9-12 days. Larval length reached 1 cm. In this 
stage, various color morphs developed. The mature stadium three larvae varied from 
yellow to cherry red, with greenish mottling. The caterpillars spent 24 to 48 hours 
in the transformation to the fourth instar. They remained motionless and contracted 
in length to about 7 to 8 mm before moulting. 

Fourth instar: The fourth instar larvae exhibited three color morphs from the 
beginning. Some were yellow, some pale orange, and others cherry red, all with short 
dark bristles. There were pronounced chevron markings dorsally on the segments, 
at first in a similar, but darker color than the background. Later, the chevrons were 
dark cherry red. Some of the yellow larvae had very light or absent chevron markings. 
This instar lasted about 15 days with growth from 1 cm to 2 cm. A sample of 89 
larvae collected in the San Bruno Mountains on 23 May 1971, showed the following 
numbers of the three principal color morphs: “red,” 63 (71%); “yellow,” 5 (6%); 
“light orange” or “intermediate,” 21 (24%). 

Pupa: The fourth stadium larvae spent three days in the prepupal stage. They 
were motionless during this period and contracted considerably in length. Initially 
the pupae were pale pink, but turned quickly to pale brown. They exhibited short 
hairs dorsally. The pupae measured from 0.7 to 0.9 cm in length. Pupation occurred 
on the earth in the debris at the bottom of the rearing jars. 

The transformation from ovum to pupa required an average of 34 days under 
laboratory conditions of 20° C. and 12 hours of artificial sunlight per day. 


The larvae were voracious feeders and were peripatetic. While feeding, 
they remained relatively motionless and burrowed into the leaves of the 
hostplant, leaving a large pile of frass behind them. This created some 
problem with rearing, as the excreta molded quickly, and tended to 
produce mold on the anal end of the larvae. The first instar larvae fed 
on the small leaves in the center of the Sedum rosette. They appeared in- 


VOLUME 26, NUMBER 4 243 


Fic. 3. Callophrys fotis bayensis: a. habitat in the San Bruno Mountains, San 
Mateo Co., California; b. second instar larva preparing to moult; c. new third instar 
larva, note cast-off skin and black head capsule; d. mature fourth stadium larva; 
e. pupa; f. young hostplant Sedum spathulifolium, the stage when oviposition occurs; 
g. ovum. 


244 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


quisitive or pugnacious and reared up their heads whenever an object 
came near. The third and fourth instar larvae totally devoured the food- 
plant, both leaves and stems. Prior to each moult, the larvae spun a light 
silk mat to which they anchored themselves. The cast-off skins were not 
eaten. 


Distribution Records 


CALIFORNIA. San Mateo Co.: slope 2 air miles W of Sierra Point, 1000’, San 
Bruno Mts., 1 2,1 9, 21-III-69; ravine 1.7 air miles WNW of Sierra Point, 700-800’, 
San Bruno Mts., 5 ova on leaf uppersides of Sedum spathulifolium, 13-IV-69; at S 
end of Milagra Ridge, along and N of Sharp Park Road, 4 second-third instar larvae on 
flower buds and leaf rosettes of S. spathulifolium, 29-IV-69; same locality, 24 third- 
fourth instar larvae on flowers of S. spathulifolium, 14-V-69; same locality, 22 first- 
third instar larvae on flower buds and leaf rosettes of S. spathulifolium, 29-IV-70; on 
NW flank of Montara Mountain, 1.1 air miles due S of Linda Mar School, 925’, 7 
second-third instar larvae in centers of leaf rosettes of S. spathulifolium, 29-IV-69; 
same locality, 5 6, 1 @, 16-III-70; ravine 1.6 air miles WNW of Sierra Point, 800-— 
1000’, 15 third-fourth instar larvae on flowers of S. spathulifolium, 29-V-69; slope 1.2 
air miles ESE of 1314’ summit, on NE slope of San Bruno Mts., 850-1000’, 24 ¢, 
20 ¢, 15-III-70; same locality, ca. 40 third-fourth instar larvae on flowers and leaf 
rosettes of S. spathulifolium, 17-V-70; same locality, 27 ¢, 16 92, 28-III-71; same 
locality, 89 second-fourth instar larvae on flower buds and leaf rosettes of S. 
spathulifolium, 23-V-71; along crest of range 0.7 air mile SE of 1314’ summit, San 
Bruno Mts., 1000-1100’, 1 &, 28-III-71; same locality, 1 4, 2 9, 3-IV-71; same 
locality, 16 second-fourth instar larvae on flower buds and leaves of S. spathulifolium, 
14-V-71; east-facing slope 0.6 air mile NNE of 1314’ summit, 600-650’, San Bruno 
Mts., 15 second-third instar larvae on flower buds and leaves of S. spathulifolium, 
7-V-71; same locality, 29 third-fourth instar larvae on flowers of S. spathulifolium, 
31-V-71. 

All of the above specimens were collected by J. F. Emmel. The type locality cited 
by Brown (1969a) is a northwest-facing slope 0.2 air mile west and slightly south 
of the 1314’ summit, 1000-1100’, San Bruno Mts. 


LITERATURE CITED 


Brown, R. M. 1969a. A new subspecies of Callophrys fotis from the San Francisco 
Bay area (Lycaenidae). J. Lepid. Soc. 23(2): 95-96. 

1969b. Larva and habitat of Callophrys fotis bayensis (Lycaenidae). 
J. Res. Lepid. 8(2): 49-50. 

McCuintock, E. & W. Knicur. 1968. A flora of the San Bruno Mountains, San 
Mateo County, California. Proc. Calif. Acad. Sci. 32(20): 587-677. 


VOLUME 26, NUMBER 4 245 


THE TYPE LOCALITY FOR TWO MOTHS (PYRALIDIDAE, 
SATURNIIDAE) COLLECTED BY LT. W. L. CARPENTER, 
U.S.A., IN COLORADO, 1873 


F. Martin BROWN 
Fountain Valley School, Colorado Springs, Colorado 80911 


In 1874 A. S. Packard described Crambus carpenterellus and Hemileuca 
diana from material collected by Carpenter while on the Hayden Survey 
in Colorado. Since Dr. A. B. Klots has need for a reasonably precise 
type locality for the crambid, I have investigated for him and find that 
one can be selected. The basic data are found in F. V. Hayden’s “Annual 
Report .. . for the year 1873,” published in 1874. 

The solution to the problem for Hemileuca diana is easily found. 
Packard wrote (1874:557) of the source: “Plum Creek, September 12 
(Lieutenant Carpenter).” There are a number of Plum Creeks in Colo- 
rado. The one involved (see Peale, p. 199) lies in Douglas County and 
drains the foothills from Palmer Lake northward almost to Littleton where 
it enters the South Platte River. In 1873, as today, two roads traveled 
along the principal branches of Plum Creek. The older road follows 
Plum Creek southward from the junction with the South Platte and gains 
the Platte-Arkansas Divide via the west branch. The railroads and the 
main highway, Interstate 25, follow the east branch to Larkspur and then 
strike due south, or continue to Palmer Lake. 

The easternmost tributary of East Plum Creek heads at Palmer Lake 
and is the stream farthest south in the system. This stream was named 
Carpenter Creek by the Hayden Survey. A good type locality for 
Hemileuca diana would be headwaters of East Plum Creek between 
Larkspur and Palmer Lake. The extent of Plum Creek is found on the 
U.S.G.S. 7% min. quadrangles Dawson Butte, Kassler, Larkspur, Littleton 
and Sedalia. 

Packard’s statement about the source of his series of carpenterellus 
is vague. He wrote (p. 548) “Mountains of Colorado, July 19, August 12, 
September 8 (Lieutenant Carpenter).” The September date is the earliest 
one mentioned in the report for return to the eastern foothills of the 
mountains after a summer near and about the continental divide. The 
published evidence (p. 556) is that Carpenter was still on the “Pacific 
slope” as late as 6 September. Other evidence in the report places the 
party at the head of Eagle River on the north slope of Tennessee Pass at 
this time. The shortest possible horseback route from the camps on the 


246 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


upper Eagle River to the foothills around Turkey Creek is 120 miles. Even 
a hardened cavalryman of the 1870’s would find that ride a noteworthy 
one to accomplish and collect specimens on both 6 and 8 September. 
There are three references (p. 548, 549 and 556) to material collected in 
the foothills on 8 September by Carpenter. Perhaps the 6 September date 
is wrongly placed on the Eagle River. 

The date 12 August is referred to several times. On p. 551, 553 and 554 
it is associated with “in the mountains.” On p. 567 Osten-Sacken wrote of 
willow galls collected by Carpenter: “those taken on East River, August 
12, still contained the caterpillars... .” East River is a tributary of the 
Gunnison River in western Colorado. It has its headwaters in a lake on 
the south side of Schofield Pass above Gothic in Gunnison County. The 
river flows southward and joins the Gunnison at Almont. Willows (Salix) 
are common along the banks of the East River throughout its course. Since 
the Survey spent considerable time mapping the Elk Mountains, and 
Schofield Pass is in those mountains, I suspect that Carpenter collected 
somewhere near Gothic on the East River, rather than in the Atriplex- 
grasslands nearer the Gunnison River. The entire East River lies in 
Gunnison County and is shown on U.S.G.S. 7% min. quadrangles Almont, 
Cement, Crested Butte, Gothic, Oh-be-joyful and Snowmass Mountain. 

“In the mountains” is associated with the date 19 July on a number of 
pages—548, 549, 551, 556, etc. The nearest dates associated with definite 
localities are 16 July at Fairplay (p. 548) and 21 July at Twin Lakes (p. 
962). In 1873 there were three routes by which Carpenter might have 
traveled from Fairplay to Twin Lakes. Gannett (p. 675) described each 
of them as follows: 


“In the Park Range the most northerly pass is Mosquito Pass at the head of 
Mosquito Gulch. Its elevation is 13,438. The ascent is steep, and difficult for pack 
ae on both sides; except in mid-summer, there is a great deal of snow on the 
trail.” 

“Weston’s Pass, Park Range, at the head of the Little Platte. Elevation 11,676. 
A good wagon-road crosses this pass. The ascent on the South Park side is by easy 
grades, but on the Arkansas side it is much steeper.” 

“Trout Creek Pass, Park Range. Elevation 9,346 feet. This pass is through the 
low rugged hills south of Buffalo Peaks, and near the salt works. The stage-road to 
Arkansas Valley crosses this pass. It is an extremely easy one.” 


Today, 100 years later, the last is the only one of the three passes in 
regular use, being the pass by which U.S. Highway 24 crosses the “Park 
Range,” now called the Mosquito Range. 

Mayden himself gives us the clue to which pass was used. On p. 49 he 
wrote of the work in the Park Range “our last move was along the divide 
‘rom Weston’s Pass to the base of Buffalo Peaks.” This move was made 


VOLUME 26, NUMBER 4 247 


on 22 July. Apparently while Hayden and the surveyors went to occupy 
their station on Buffalo Peaks, Carpenter and an advance party moved 
to Twin Lakes. Thus we can be quite sure that the 19 July specimen(s ) 
of Crambus carpenterellus were collected in the vicinity of Weston Pass 
on the boundary between Park and Lake counties. Good camp could 
easily be made on the Park County side of the pass. In fact, I camped 
there myself in the 1930's. It is an area of typical Hudsonian forest, grass- 
land and bog. The region is well shown on U.S.G.S. 7% min. quadrangles 
Mount Sherman and South Peak. 

Faced with selecting one of these three diverse localities as the type 
locality for carpenterellus, I reneged and passed the problem to Dr. Klots. 
He wrote to me “July 19 is the most logical of the three dates . . . for this 
species to be flying. August 12 is possible, but it would be pretty well 
gone by then. I think September 8 would be much too late.” Thus Dr. 
Klots settled upon Weston Pass, Park County, Colorado to be the type 
locality for Crambus carpenterellus Packard. 


LITERATURE CITED 


GANNETT, H. 1874. Geographical Report of Henry Gannett, M.E., in Hayden, 
1874, p. 670-681. 

Haypen, F. V. 1874. Annual Report of the United States Geological and Geo- 
graphical Survey of the Territories, embracing Colorado, being a report of 
progress of the exploration for the year 1873. Government Printing Office, 
Washington, D.C. 718 p. illus. 

OsTEN SAcKEN, C. R. 1874. Notice on the galls collected by Lieutenant W. L. 
Carpenter, in Hayden, 1874, p. 567. 

PackarD, A. S., JR. 1874. On the geographical distribution of moths in Colorado, 
in Hayden, 1874, p. 543-560, 15 figs. 

PEALE, A. C. 1874. Report of A. C. Peale, M.D., geologist of the South Park 
Division, in Hayden, 1874, p. 193~273, illus. 


THE MATURE LARVA OF SPHINX VASHTI (SPHINGIDAE) 


Sphinx vashti Strecker is widely distributed in the western half of North 
America (Hodges 1971, in Dominick et al., The Moths of America North of 
Mexico, Fascicle 21, Sphingoidea: p. 59-61). The egg, larva, and pupa were 
first described by Dyar (1894, Psyche 7:177), who reared it on Snowberry 
(Symphoricarpos albus). Recently, Comstock (1966, J. Res. Lepid. 5:218—-219) 
described and figured the egg and first instar larva. The mature larva is depicted 
here for the first time. 

On 14 July 1958 I found a larva feeding on Coralberry (Symphoricarpos 
orbiculatus) in the front yard of my home in Ottawa, Kansas. The mature 
larva is pale apple green with blue-green granulations on the dorsum. The lateral 
oblique lines on the abdomen are lavender or purplish-red. The caudal horn is 
dark red to deep blue at the tip. In Dyar’s specimen the lateral lines were 


48 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Mature larva of Sphinx vashti. 


white, edged anteriorly with a narrow black line, and the caudal horn was 
greenish white; black above, below, and at the tip. 

The larva pupated in the soil on 22 July 1958, spent two winters as a pupa 
(the soil was never moistened), and produced an adult female on 6 May 1960. 

This species, single-brooded and always rare in Kansas, has occasionally been 
taken visiting honeysuckle blossoms in late May and June. I have taken several 
adults visiting columbines (Aquilegia) in the early evening in Gunnison National 
Forest, Ohio City, Colorado in mid-July. Fleming (1970, Mich. Entomol. 3:17- 
23) did not include any feeding records for the adults of Sphinx vashti, but his 
review was primarily of eastern species. 


Witit1amM H. Howe, 822 East Eleventh St., Ottawa, Kansas 66067. 


VOLUME 26, NUMBER 4 249 


A PROPOSAL FOR THE UNIFORM TREATMENT OF 
INFRASUBSPECIFIC VARIATION BY LEPIDOPTERISTS 


Joun H. Masters? 
Lemon Street North, North Hudson, Wisconsin 


The binomial system of zoological nomenclature dates back to Lin- 
naeus 10th edition of Systema Naturae in 1758. The trinomial was not 
conceived by Linnaeus, however, and did not come into extensive use 
until the last half of the 19th century. While Linnaeus created the 
binomial system, he did not propose any sort of rules for the naming of 
animals. A great deal of confusion resulted and in the early 19th century 
a number of codes were proposed, mostly imposing a basic philosophy for 
priority of names, in attempts at solution. At the First International 
Congress of Zoology, held in Paris in 1889, Raphael Blanchard submitted 
a proposed set of international rules for naming animals. Blanchard’s 
rules were formally adopted at the Second Congress in Moscow in 1892 
and have been subsequently revised until the present International Code 
of Zoological Nomenclature was adopted by the Fifteenth Congress in 
London in 1958 and was officially published in 1961. 

Under the present Code (1961) the trinomial is restricted in usage to 
geographical subspecies and all other types of infraspecific variation are 
considered as infrasubspecific and are removed and excluded from the 
provisions of the Code. This decision, by the International Commission, 
was not meant to imply that the study of infraspecific categories other 
than the subspecies is unimportant, but to emphasize the fact that sub- 
specific variation is essentially different from any of the others. Sub- 
specific variation is generally considered to be the first stage of the 
speciation process and those populations which are currently treated as 
subspecies are so treated subjectively and may be, in any later revision, 
elevated to the species level. Because the subspecies names is subject to 
elevation to the species level (and conversely, species names are subject 
to reduction to the subspecies level), it is essential to retain it in the 
species-group where it is liable to those rules and criteria, including 
priority, which apply to the species name. 

The geneticist, and many others, may regard other types of infra- 
specific variation as more important than subspecies; however infrasub- 
specific variants are not subject to elevation to the species category and 
there was deemed to be no need to conserve priorities or other protection 
under the provisions of the Code. 


1 Research Associate, Carnegie Museum, Pittsburgh, Pennsylvania. 


250 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


INFRASUBSPECIFIC VARIATION 


Other than a general agreement that infrasubspecific names should not 
be placed in italics, as are the species-group names, lepidopterists have 
not given them anything approaching a standard treatment in the last 
decade. There has been, however, a very sharp decrease in the publica- 
tion of formal names to apply to infrasubspecific variation during the last 
twenty years. While a few names still appear, most authors are content 
to describe examples of infrasubspecific variation without attempting to 
formally name them. 

Designations for many types of infrasubspecific variation seem at least 
useful, if not necessary, and if they are to be designated, it is highly de- 
sirable that a consistent and uniform method be used for citing them. It 
is not necessary that priorities or rules of Latin word formation be fol- 
lowed, nor is it essential that usage in Lepidoptera be consistent with 
that in other orders of insects or in other classes of animals. The op- 
portunity is clearly present for lepidopterists to develop a system of 
designation suitable for present day needs without concern for the 
stigmas of priorities and validities that have made this impossible in the 
past. With the vacilation of the I.C.Z.N. in the area of infrasubspecific 
designation, it is unlikely that mandatory rules will ever be adopted. 
Consistent usage can only come through common acceptance of the 
majority of lepidopterists. It is my hope that this proposal for a rational, 
uniform treatment of infrasubspecific categories will eventually lead to 
a standard and uniform treatment by lepidopterists around the world. 

Infrasubspecific designations in Lepidoptera have been employed for 
an assortment of variations, all of which involve maculation (phenotype 
expression), some, but not all, of which involve genotype and none of 
which, at least directly, involve physiotype. For practical purposes, they 
all can be grouped into four distinct categories: (1) polychromatic or 
polymorphic forms, (2) mutant or aberrational forms, (3) seasonal or 
brood forms, and (4) hybrids. Each of these presents an entirely different 
set of circumstances and the criteria for designation must be dealt with 
individually. 


Polychromatic or Polymorphic Forms 


Ford (1940) defined the condition of polymorphism as “the occurrence 
together in the same habitat of two or more discontinuous forms, or 
phases, of a species in such proportions that the rarest of them cannot 
be maintained merely by recurrent mutation.” Polymorphic forms are 
perhaps best known to North American lepidopterists in the yellow or 
vhite color phases of female Colias species. They are most pronounced 


VOLUME 26, NUMBER 4 PAD | 


in the various mimetic female forms of certain African and Indo- 
Australian Papilio species, e.g. Papilio dardanus Brown or Papilio polytes 
Linnaeus. These are clearly genetic situations and, in fact, the genetics of 
many of them have been carefully studied and worked out by breeding in 
the laboratory. The discontinuous factor is important for considerations 
of polymorphism, for continuous variation that could be plotted on a 
curve of normal distribution, such as the length of forewings, is excluded. 
(For detailed discourse on the genetic aspects of polymorphism, see 
Ford (1965). ) 

Nearly all of the North American Colias species exhibit dimorphic 
yellow vs. white female color phases (for detailed information, see 
Hovanitz, 1950), and many names have been proposed to refer to the 
white form: “alba” Strecker, “albida’” Chermock, “canescens” Comstock, 
“flavocincta” Cockerell, “hatui” Barnes & Benjamin, “neri’ Barnes & 
Benjamin, “medi” Gunder, “lambillioni” Dufrane, “martini” Gunder, 
“pallida” Cockerell, “pallida” Skinner, “pallidice” Scudder, “pallidissima” 
Bowman, and “shastae” Barnes & Benjamin. While roughly 50% of these 
names are more or less descriptive of the color condition, the genetic 
factors creating the white or yellow phases are identical, or nearly so, 
for all of the species in the genus and there is no reason why a single de- 
scriptive name should not be employed as a nomen collectivum to apply 
to the equivalent forms in each species. However, since both white and 
vellow forms are normal genetic components of the population, it would 
not be proper to apply a designation to the white form without an 
equivalent designation for the yellow form. It is my proposal that the 
name “alba” be employed as a descriptive and collective name for the 
white color phase in Colias species, and that “flava” be similarly em- 
ployed to designate the yellow phase. If desired, “chrysa” could be 
added to distinguish those populations with an orange phase from those 
with a yellow phase. 

The use of a collective-descriptive designation, as cited in the ex- 
ample of Colias, seems to be the most practical way to deal with poly- 
morphic variation. To avoid confusion with species-group names, these 
names should be enclosed in quotation marks but not italicized. Since 
they are not subject to the laws of priority and since they are descriptive. 
there is no need to append an author’s name. (Author’s names are ap- 
pended to species-group names to facilitate the reference to an original 
description, not to honor the author.) The use of Latin to derive the 
collective-descriptive names seems preferable to a contemporary language 
because it will have equal meaning in international usage and will dis- 
courage translation into vernacular vocabulary. There is, of course, 


252 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


some loss of descriptive value in the Latin derived names, as most 
biologists in the present day are not Latin scholars. With slight modifica- 
tion, collective-descriptive designations can be adapted to polymorphic 
as well as polychromatic situations. 

Additional examples of polychromatic situations are: Heliconius doris 
(Linnaeus) which is sympatrically trichromatic in both sexes with red, 
green and blue color phases which could be referred to, respectively, 
as ‘erythro,’ “chloro” and “cyano.” Similarly, the dimorphic female 
color phases of Papilio glaucus Linnaeus could be treated as “flava” for 
the yellow examples and “atrata” for the dark ones. The blue and brown 
color phases of certain female Lycaenidae, e.g. Plebejus saepiolus 
(Boisduval), could be collectively indicated as “cyana” and “atrata.” 
Similarly, “cyana,” “atrata” and “bicolor” could be used in reference to 
the brown, blue and mixed females of Morpho aega Hubner. The silvered 
and unsilvered varieties of Speyeria and related genera might be re- 
ferred to as “argentamaculosus” and “flavomaculosus” respectively. 

For non-chromatic polymorphic forms, the system is not as simple 
for there may be no simple descriptive term that could be used to 
designate the various forms. In many of these cases, the various poly- 
morphic forms are apparent mimics of other species and it is appealing 
to compose a compound name of the prefix “pseudo-” along with the 
specific name of the model species.2, As an example of non-chromatic 
polymorphism consider the various polymorphic females of Papilio 
dardanus cenea Stoll in South Africa. Van Son (1949) lists eleven dis- 
tinct named forms of the female of Papilio dardanus cenea all of which 
show a remarkable phenotype expression and all of which are strikingly 
distinct from the male phenotype. All of these forms are apparent mimics 
of distasteful or protected species of Danaidae or Craeidae, which allows 
us to coin a collective-descriptive name based on the model-mimic re- 
lationship. Papilio d. cenea form “hippocoonides” Haase, which mimics 
Amauris niavius dominicanus Trimen, would be designated as Papilio d. 
cenea “pseudodominicanus” which is both simpler and more meaningful; 
similarly, form “trophonius” Westwood, which is a mimic of Danaus 
chrysippus Linnaeus, would become “pseudochrysippus”; and the other 
forms could be similarly named for the species they mimic. 

For situations where males and females are dimorphic with respect to 
each other but constant within the same sex, e.g. Neophasia terlootii 
Behr, with white males and brick orange females, no infrasubspecific 
designation is required or desirable. For species which display a great 


The Code (recommendation D13) advises against the use of the prefix pseudo- with non- 


K nouns or idjectives; however this is the simplest procedure to use in collective-descriptive 
yn and infrasubspecific usages are clearly not governed by the Code in any case. 


VOLUME 26, NUMBER 4 aye 


deal of random variation, e.g. Parnassius phoebis Fabricius, collective- 
descriptive names could be utilized to refer to the various individual 
variants. Eisner (1955) (see also Brown, 1956) proposed a total of 66 
collective-descriptive names for application to variation that he had ob- 
served in the genus Parnassius. Many of the variants referred to by 
Eisner represent aberrations rather than polymorphisms and are treated 
in the following section. Unless a variant is fairly regular in occurrence 
and there is good reason to suspect a genetic cause for it, I see no reason 
or need for an infrasubspecific designation. 

In using collective-descriptive names for polymorphic forms, setting 
the names in another typeface (e.g. boldface), but not italics, could 
be considered as an alternative to enclosing them in quotation marks. 
Intervening qualifying phrases (e.g. form as in Colias gigantea 2 form 
“alba” ) would be optional usage. 


Mutant or Aberrational Forms 


Aberrations, mutants or “sports” are encountered with fair frequency 
among Lepidoptera. Many of these forms (mutants) have genetic cause 
but, unlike polymorphic forms, they are extremely rare in occurrence 
and not a normal part of the population. If the same sort of mutant re- 
appears from time to time, it is assumed to be maintained by recurrent 
mutation rather than by selection. Other aberrational forms are produced 
by environmental causes. For example, it is well known that aberrant 
specimens of Euphydryas phaeton (Drury) can be artificially produced 
by exposing pupae to near freezing temperatures at a critical time in 
their development. As a general rule, these forms are much rarer in 
occurrence than are polymorphic forms; in the majority of cases their 
actual percentage of occurrence in a population would be less than 
0.01% (one in 10,000). A polymorphic form may be this rare in a local 
population, but not throughout its entire range and, in some cases, an 
environmentally induced aberrant may be considerably more common 
than this during a single brood, but not on a continuing basis. 

In the not too distant past, there was a strong tendency to adorn each 
mutant or aberrational form with a formal name. At present, they are 
rarely named, but are frequently described and reported in the literature. 
Whether genetic or non-genetic in cause, aberrants normally are not an 
integral part of any population; each specimen is an individual without 
direct continuity with any succeeding individuals which may resemble 
it. Putting a name, formally or informally, on aberrant specimens serves 
no useful purpose, and might serve to confuse them with polymorphic 
forms. It should be kept in mind, however, that mutant forms are the 


254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


raw material for evolution and that they may become established as 
polymorphic forms through selection. 

Sexual mosaics and gynandromorphs are considered by me to fall into 
this category of aberrational forms. 


Seasonal Forms 


Seasonal forms are most pronounced in the areas of Africa where there 
is a considerable seasonal variation in rainfall. The “wet” and “dry” 
season forms of some species, particularly of the genus Precis (Nym- 
phalidae), are so completely distinct in appearance that they were de- 
scribed as distinct species. In North America, seasonal forms are best 
known in the distinct brood forms of Eurytides marcellus (Cramer) 
(Papilionidae), Celastrina argiolus (Linnaeus) (Lycaenidae), various 
Pieris species (Pieridae) and various Polygonia species ( Nymphalidae ). 

Seasonal forms may be considered environmental in nature, as the 
changes in appearance are brought about in response to environmental 
conditions, in most cases differences in solar radiation or differences in 
humidity, and not by genetic change. Of course genetic factors control 
the seasonal changes, since some species have seasonal forms while 
closely related ones may not. In some cases, changes in active genes 
may be responsible for the phenetic differences, but both forms are 
identical in terms of total genome, one form flowing from the other via 
direct inheritance. 

A great deal of nomenclature has been expended, in the past, in treat- 
ment of seasonal variation where the variation involved is clearly the 
result of common factors. For instance, Pieris sisymbrii Boisduval, Pieris 
protodice Boisduval & LeConte, Pieris occidentalis Reakirt, Pieris napi 
(Linnaeus) and Pieris rapae (Linnaeus) have, in common, a situation 
in which the early spring brood is heavily marked on the ventral hind- 
wing while summer or fall broods are relatively immaculate. Many 
names have been formally proposed to cover this situation in Pieris 
(including “transversa” Barnes & Benjamin, “vernalis’” Edwards, and 
“nasturtii” Edwards for the spring forms; and “acadica” Edwards, 
“cruciferarum” Boisduval, “aestiva” Harris, “castoria” Reakirt, “iberidis” 
Boisduval, “pallida” Scudder, “pallidissima” Bames & Benjamin, and 
“yreka” Reakirt for the summer forms). All of these could readily be 
eliminated by using “vernalis” as a descriptive-collective name for all 
of the spring broods and by using “aestivalis” as an equal descriptive- 
collective name for the summer broods. This same system could be used 
in the case of all species having seasonal or brood forms; the descriptive 


VOLUME 26, NUMBER 4 255 


names “autumnalis” and “hyemalis” could be added to cover fall or 
winter forms as required. 

However, I feel that a preferable treatment for seasonal or brood forms 
would be to assign the successive generations a Greek letter designation. 
Thus the spring broods of the Pieris species cited above, could be given 
the designation a (alpha), and the summer broods could be designated 
by B (beta). 

Compare the following methods of designating the two distinct spring 
and summer broods of Eurytides marcellus. 


Method I, using commemorative names with priorities: 


Eurytides marcellus (Cramer) form “walshii” (Edwards ) 
; [early spring brood] 
Eurytides marcellus (Cramer) form “telamonides” (Felder & Felder) 
Eurytides marcellus (Cramer) form “lecontei” (Rothschild & Jordan) 
[summer brood] 


Method II, using collective-descriptive names: 


Eurytides marcellus (Cramer) “monovernalis” [early spring brood] 
Eurytides marcellus (Cramer) “bivernalis” [spring brood] 
Eurytides marcellus (Cramer) “aestivalis” [summer brood] 


Method III, using Greek-letter designations: 


Eurytides marcellus (Cramer) a brood [early spring brood] 
Eurytides marcellus (Cramer) 8 brood [spring brood] 
Eurytides marcellus (Cramer) y» brood [summer brood] 


It is my feeling that method III is preferable, as there would be no 
ambiguity such as might result if collective-descriptive names, albeit 
different ones, were used for both polymorphic and seasonal form 
designations. 


Hybrids 

In Lepidoptera, hybrids are not excessively rare in nature and, in some 
cases, can be produced with a degree of efficiency in the laboratory. We 
must, however, recognize four distinct classifications of hybrids: (1) 
hybrids between two subspecies of the same species, (2) hybrids between 
two distinct species of the same genus, (3) hybrids between two 
species of different genera, and (4) hybrid populations that are viable 
and breeding, although resulting from the hybridization of two distinct 
species. 


256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Where both parents are known, a hybrid can be readily designated by 
joining the two species-group names with a multiplication sign (x). 

(1) For hybrids between subspecies of the same species, such as 
between Limenitis arthemis arthemis (Drury) and Limenitis arthemis 
astyanax (Fabricius), the designation would be Limenitis arthemis 
(arthemis xX astyanax) or Limenitis arthemis (astyanax X arthemis). 
The name of the male parent, if known, should precede that of the female 
parent. 

(2) For hybrids between two species of the same genus, such as 
Limenitis arthemis and Limenitis archippus (Cramer), the combination 
would be Limenitis arthemis < archippus or Limenitis archippus x 
arthemis. 

(3) For the much rarer situation of hybrids between two species of 
distinct genera, e.g. hybrids between Phyciodes tharos (Drury) and 
Chlosyne nycteis (Doubleday), the designation would be simply 
Phyciodes tharos x Chlosyne nycteis, or Chlosyne nycteis x Phyciodes 
tharos. 

(4) In Lepidoptera there are, albeit very rarely, viable, breeding popu- 
lations resulting from hybridization between two closely related species. 
Papilio kahli Chermock & Chermock is one of the better known examples 
of these, and has been treated in detail by Remington (1958), who con- 
cluded: 


“1) Riding Mountain is the locus of origin of P. kahli, an isolated, distinctive off- 
shoot of P. polyxenes with black wings, spotted abdomen, and large acentric ‘pupil.’ 
2) In relatively recent times P. machaon arrived on the plateau, perhaps carried from 
Alberta in hay or straw during the development of the National Park or of highways 
or railroads. 3) These two Papilio at first lacking behavioral and other isolating 
mechanisms, hybridized rather freely; the distinctive genotype of P. kahli allows 
the F, heterozygotes to show some yellow-wing characters never seen in laboratory 
crosses of machaon with true polyxenes. 4) Since P. kahli and P. avinoffi belong to 
separate species, one expects that isolating mechanisms are evolving in Riding 
Mountain populations and that eventually natural hybrids will no longer be produced. 
Meanwhile, each species may be incorporating into its genotype new adaptive alleles 
from the other species (introgression). There is little basis for regarding kahli as 
dimorphic in the sense of P. glaucus females. For the present, these Riding Mountain 
file Sr may be called P. kahli (or P. polyxenes kahli), P. machaon avinoffi and 
their hybrids.” 


Papilio nitra Edwards is another North American Papilio that repre- 
sents a situation similar to that of Papilio kahli. Warren (1969) cited 
four populations of Old World Pieris (dubiosa Rober, balcarica W. & N., 
pseudorapae Vty. and meridionalis Heyne) which he considered hybrid 
TAaCes. 

(lybrid races such as these may be referred to by specific names of the 
species group type—subject to priorities and other provisions of the 


VOLUME 26, NUMBER 4 Dion 


Code—as though they were true species; however the generic name 
should be preceded by the sign of multiplication, e.g. x Papilio kahli, 
or X Pieris balcarica. 


HOW TO DETERMINE WHETHER A NAME IS SUBSPECIFIC 
OR INFRASUBSPECIFIC 


It is necessary to recognize subspecific names and infrasubspecific names and to 
distinguish between them. The provisions of the Code (my treatment here is adapted 
from Field, 1971) are summarized here. 


Subspecific Names 


Article 45 (d) of the Code dictates three situations under which we are to accept 
a proposed name as a subspecific name. 

1. The original status of any name of a taxon of lower rank than species is de- 
termined as subspecific if the author, when originally establishing the name, clearly 
stated it to apply to a subspecies. Obviously the best way to propose a subspecific 
name is to state that it is a subspecies. 

2. The original status of any name of a taxon of lower rank than a species is de- 
termined as subspecific if the author, when originally establishing the name, did not 
state its rank. This clearly means that if an author proposed a trinomial name without 
explaining the trinomen in any way, we are to accept it as a proposal of a subspecies. 

3. The original status of any name of a taxon of lower rank than a species is 
determined as subspecific if the author, when originally establishing the name, stated 
the taxon to be characteristic of a particular geographical area (or geological 
horizon) and did not expressly refer it to any infrasubspecific category. This clearly 
covers all names proposed in the past as races, local forms, altitude forms, and the 
like, provided they were proposed as trinomial names. 


Infrasubspecific Names 


Article 45 (d) (iii) gives two ways of recognizing when a taxon is of infra- 
subspecific status. 

1. The original status of any name of a taxon of lower rank than species is de- 
termined as infrasubspecific if the author, when originally establishing the name, ex- 
pressly referred the taxon to an infrasubspecific rank. This necessarily includes 
names given to all categories lower in rank than the subspecies and includes all 
names given to individual specimens and segments of populations such as aberrations, 
transition forms, seasonal forms, wet and dry forms, cold forms, color forms, sexual 
forms, and the names given to the separate generations of the same population. 

2. The original status of any name of a taxon of lower rank than species is de- 
termined as infrasubspecific if the author, when originally establishing the name, after 
1960, did not clearly state that it was a subspecies. 


“Varieties” and “Forms” 


Paragraph (e) of article 45, interprets the usage of the terms “variety” and “form” 
as follows: (i) before 1961, the use of either of the terms ‘variety’ or ‘form’ is 
not to be interpreted as an express statement of either subspecific or infrasubspecific 
rank; (ii) after 1960, a new name published as that of a ‘variety or ‘form’ is to be 
regarded as of infrasubspecific rank. (This is also stated in article 15.) 

For publications dated before 1961, we must study the author's text to determine 
what he meant by his use of the terms “variety” and “form.” If the author clearly 


258 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


indicates in his text that he is writing about an individual variant (such as an aberra- 
tion, color form, or the like) that is part of a population, then we have to reject 
the name. On the other hand, if the original author in no way shows what he meant 
by the term “variety” or “form,” or if it is not clear that he is naming either sub- 
species or individual varients of such population, or if his text clearly indicates that 
he is describing geographical variation of the modern subspecies concept, then we 
have to accept any such proposed trinomial name as an acceptable trinomial under 


article 45 (d) (i). 


Acceptable Subspecific Names 


In summary, there are five types of trinomial names that should be acceptable 
as subspecific names under the Code. 

1. Names given as trinomials (article 5), as subspecies (article 45 (d) (i)). 

2. Names given as trinomials (article 5), before 1961, where no expressed category 
or rank is indicated (article 45 (d) (i)). 

3. Names given as trinomials (article 5), before 1961, as races, local forms, altitude 
forms, or given as trinomials, to any other geographically based population (article 
ANS: (cl) (Git). 

4. Names given as trinomials (article 5), before 1961, as “varieties” and “forms” 
where the author indicates or even hints that they represent geographically based 
populations (article 45 (d) (ii) and (e) (i)). 

5. Names given as trinomials (article 5), before 1961, as “varieties” and “forms” 
where the author in no way indicates what he meant by the use of these terms (article 
45 (d) (i) and (e) (i)). Many authors used these terms for subspecies. Article 45 
(e) (i) allows us to accept these terms as subspecific unless it is apparent that an 
infrasubspecific category is intended. 


Names Unacceptable under the Code 


There are seven types of names that are excluded as subspecific names by the Code. 

1. All names proposed as quadrinomials (article 5 by recognizing only the generic 
name, the specific name, and, when applicable, the subspecific name). 

2. All names given to aberrations as such, transitional forms as such, seasonal forms, 
wet and dry forms, color forms, sexual forms, generation forms as such, and similar 
forms (article 1; article 45 (d) (iii) and glossary of the Code: definition of the 
term infrasubspecific ). 

3. All names given to “varieties” and “forms” before 1961, where the author 
clearly indicates that he is dealing with an individual variant such as one of those 
mentioned aboye under number 2 (article 45 (e) (i) ). 

4. All names proposed as trinomials after 1960, where it is not clearly stated that 
such names are subspecific names (article 45 (d) (iii) ). 
rnin proposed for “varieties” or “forms” after 1960 (article 15, article 45 

e) (ii) ). 

6. All names proposed for races, local forms, altitudinal forms, or any geo- 
graphically based populations, after 1960, where they are not expressly called sub- 
species (article 45 (a) and article 45 (d) (iii) ). 

7. All names given to hybrids (article 1). 


Excluded Names Becoming Available 


Names rejected or excluded under the Code may later become available, for 
article 10 (b) states that “a name first established with infrasubspecific rank becomes 
available if the taxon in question is elevated to a rank of the species-group, and takes 
the date and authorship of its elevation.” 


VOLUME 26, NUMBER 4 259 


SUMMARY 


1. Taxonomic categories of lower rank than subspecies (infrasub- 
species ) have been removed from the protection of the “Code” (Inter- 
national Code of Zoological Nomenclature, 1961). This means that names 
proposed for infrasubspecific variation (including those proposed for 
polymorphic forms, aberrations, seasonal forms, sexual forms, color forms, 
altitudinal forms, etc.) do not have the regulation and protection of the 
Code under the laws of priority and uniform usage. 

2. If names are to be used to designate infrasubspecific variation, they 
should be used in such a way as to avoid confusion with the subspecies 
or trinomial usage. 

a. Names should be placed in quotation marks or some other type- 
face (such as boldface) and not in italics as are used for the species- 
group names. 

b. These names should not take an author's name. 

3. Since infrasubspecific names are not subject to the laws of priority 
or other provisions of the Code, the opportunity is clearly present for 
lepidopterists to develop a uniform system of designation suitable for 
present day needs without concern for the stigmas of priorities and 
validities that, ostensibly, have made this impossible in the past. 

4. The following proposals are advanced to cover four major categories 
of infrasubspecific variation. 

a. POLYCHROMATIC OR POLYMORPHIC FORMS should be described by 
collective-descriptive names. The use of Latin to derive the collective- 
descriptive names is preferred because it will have equal meaning in 
international usage and will discourage translation into vernacular 
vocabulary. The name “alba” as used to describe the white color phase 
of female Colias butterflies is an example of a descriptive name. Since 
a similar white color phase occurs by a similar genetic mechanism in 
nearly all species of Colias, the same name should be applied to similar 
color phases in all of them—thus it is a collective name. 

b. MUTANT OR ABERRATIONAL FORMS are not an integral part of the 
population; each specimen is an individual and does not have any 
direct continuity with any succeeding specimen which may resemble it. 
Placing a name, formally or informally, on aberrant specimens serves no 
useful purpose and is to be discouraged entirely. 

C. SEASONAL FORMS are environmental and not genetic in nature be- 
cause the differing broods involved are genetically identical. Collective- 
descriptive names, such as are suggested for use with polymorphic 
variation, could be applied in this case; however, since polymorphic 
variation is quite distinct from seasonal variation and since a given 


260 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


specimen may exhibit both polymorphic and seasonal variation, it is 
recommended that Greek-letter designations be employed to indicate 
seasonal variation. Starting with the first spring or first wet season 
generation, broods could be designated, successively, as a (alpha), 
B (beta), y (gamma), etc. broods. 

d. HYBRID INDIVIDUALS can be indicated by joining the two species- 
group names with a multiplication sign. In those rare situations where 
an actual breeding population of hybrid origin exists, a species group 
name may be applied, but only if the entire name is preceded by a 
multiplication sign. 


LITERATURE CITED 


Brown, F. M. 1956. Infra-subspecific names among Parnassius. J. Lepid. Soc. 
10: 140-142. 

“Cope.” 1961. International code of zoological nomenclature adopted by the XV 
international congress of zoology. International Trust for Zoological Nomen- 
clature, London. 176 p. 

Eisner, C. 1955. Parnassiana Nova. V, Nomina _ Collectiva. Zoologische 
Mededlingen, Rijksmuseum van Natuurlijke Histoire te Leiden 33: 177-179. 

Fretp, W. D. 1971. Butterflies of the genus Vanessa and of the resurrected 
genera Bassaris and Cynthia (Lepidoptera: Nymphalidae). Smithsonian Contr. 
Zool., No. 84. 105 p. 

Forp, E. B. 1940. Polymorphism and Taxonomy, in, J. Huxley, ed., The New 
Systematics. Clarendon Press, Oxford, p. 493-513. 

. 1965. Genetic Polymorphism. All Souls Studies No. V., Faber & Faber, 
London. 101 p. 

Hovanirz, W. 1950. The biology of Colias butterflies. II. Parallel geographic 
variation of dimorphic color phases in North American species. Wassman J. 
Biol. 8: 197-219. 

REMINGTON, C. L. 1958. Genetics of populations of Lepidoptera. Proc. 10th 
Intern. Congr. Entomol. 2: 787-805. 

VAN Son, G. 1949. The butterflies of southern Africa. Part I. Papilionidae and 
Pieridae. Mem. 3, Transvaal Museum, Pretoria. 237 p. 41 pl. 

Warren, B. C. S. 1970. Some aspects of hybridisation and their significance. 
Entomol. Rec. 82: 305-313. 


VOLUME 26, NUMBER 4 261 


LIMENITIS LORQUINI (NYMPHALIDAE) ATTACKING A 
GLAUCOUS-WINGED GULL 


Readers of the encounter between a Monarch and a Red-winged Blackbird 
reported by Slansky (1971, J. Lepid. Soc. 25:294) may be interested in a 
similar pugnacious interlude. This occurred on the warm and sunny morning of 
4 July 1970, in Beacon Hill Park, Victoria, British Columbia. On that day, along 
with lepidopterists JoAnne Pyle, H. Whetstone Pyle and Chuck Dudley, I was 
photographing butterflies for a work on the fauna of Washington. We were 
drawn into an ornamental rose garden by a superb, fresh Limenitis lorquini 
burrisonii Maynard. The Admiral, a male, was sunning on the roses and soaring 
slowly around the garden. There were no nectar sources, no female lorquini, 
nor any other attractants in view. 

The butterfly was exceptionally approachable, more so than any others of its 
species I had encountered before. Photographing it presented little difficulty. 
Indeed, the “friendly” creature crawled onto my fingers and landed upon many 
bemused visitors to the garden. Despite considerable human activity in the area, 
the lorquini maintained this “tame” behavior. Then a sound, which had been 
constantly in the background, came nearer. A shadow passed over the garden 
as a Glaucous-winged Gull appeared from behind the crowns of a Douglas Fir 
wood. Suddenly, as the gull came directly overhead at a height of 20 to 30 feet, 
the Admiral darted up from his rose-blossom perch and accosted the gull. Again 
and again lorquini darted at the huge bird, never descending until the gull quit 
the area. 

This remarkable spectacle occurred several times in a_ half-hour period. On 
each occasion, the pale seabird entered the arena of action and began to circle, 
only to be immediately enjoined by the Lorquin’s Admiral. The height of engage- 
ment was consistently the same as Slansky reported for the Monarch and _ the 
blackbird—from 20 to 30 feet. Unlike the Red-winged Blackbird, however, this 
gull did not seem to react to what I interpreted as the butterfly’s aggression. 
It called raucously, but no more than before or after the encounter; and it seemed 
to depart volitionally and not under stress. The gull made no attempt to deter 
or eat the Admiral. 

Why was the lorquini indifferent to other movement through the garden, yet 
so demonstrative toward the seagullP One may speculate that the bird was 
recognized as a potential foe, rival male, or mate, and that the appropriate 
behavioral response was elicited. Attacking a potential predator would not seem 
very adaptive for a butterfly, and where I have seen male lorquini together 
before, they have passively coexisted. Therefore, the possibility of a courtship 
chase would seem most likely. Butterfly males have been known to pursue experi- 
mental super-female models: perhaps the same sort of perception and reaction 
was seen here. 

It would be pleasing to see more ethological notes in this journal. I feel 
that, in regarding butterflies strictly as specimens for acquisition or research, we 
often ignore events which render these organisms fascinating in a living context: 
events such as this encounter between an Admiral and a seagull, which was 
just slightly less intriguing than a well known episode involving another kind of 
seabird, an albatross, and an aged seafarer. 


Ropert MicuaEL Pyie, Monks Wood Experimental Station, Abbots Ripton, 
Huntingdonshire, England. 


bo 
op) 
bo 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


BOOK REVIEWS 


MICROLEPIDOPTERA PALEARCTICA; VOLUME 2. ErumupAr, by Klaus Sattler. George 
Fromme & Co., Vienna. In two vols., text xvi + 185 p.; plates, 9 color 97 black & 
white, each with unnumbered exp]. page facing. Dec., 1967. Subscription price: 
1,160 Austrian schilling; full price: 1,365 Austrian shilling (about $58.00). [text in 
German]. 


This, the second contribution of the ambitious project which plans to systematically 
treat and illustrate in color all microlepidoptera of the Palearctic Region, follows the 
same format set forth in Volume 1, the Crambidae, by the late Stanislaw Blezynski, 
in 1965. Principles of the project are explained in the forward to Volume 1 (see J. 
Lepid. Soc., 19: 117-125, 1965). 

The treatment of the Gelechioid family Ethmiidae is taxonomically and mechanically 
accurate and concise throughout. It includes introductory parts dealing with histori- 
cal development and associated problems in handling material and literature, technical 
aspects of the work, explanation of taxonomic characters, early stages and bionomics, 
especially foodplants, and the systematic placement of the family. The systematic 
portion encompasses 72 species in the genus Ethmia in addition to 4 species of 
uncertain placement and 4 species transferred to other taxa. In synonymizing other 
previously described genera, Sattler displays a sensible respect for the appreciation 
of problems of higher category taxonomy on a world basis. He issues a warning 
that questions of genera should not be answered on the basis of limited knowledge of 
only the European or other fauna; and he points out that introduction of new generic 
names inevitably implies distinctions which may not in reality exist. Among 135 
species level names, type specimens of 95 were examined during this study; the 76 
species which precipitated include 12 previously undescribed, 28 new synonyms, 
and 5 entities treated as new status at the subspecific level. 

Each species is characterized by original citation, type locality, a brief diagnosis 
of external features and of male and female genitalia, summary of biology and early 
stages where known, geographical distribution, and other pertinent remarks. Mor- 
phological characters are illustrated by excellent drawings, including genital features 
of both sexes for all species where both are known, and by the incredibly good color 
paintings done by F. Gregor. 

As a matter of format the literature cited is fairly brief. Synonymies consist of 
original citations only and do not include generic name combination changes. Refer- 
ences to subsequent papers are in short form (author-date-page). Not all references 
are included in the terminal bibliography (e.g., those on p. 19), or else the system 
is not explained clearly. On the other hand, citations in the terminal bibliography are 
given in complete form and include reference to the individual species treated. 

Geographical distribution summaries are based on specimens examined, with 
additional information from the literature clearly specified. Moreover, improbable 
reports are discussed and reports considered clearly false by Sattler are characterized 
as such, Faunistic compilations are mentioned, but not every local list is recorded. 

Similarly, biological information is well documented. Life cycle and hostplant 
data are summarized (larvae will not be described in any of the volumes according 
to the preface). In an extensive tabular record of foodplants by Ethmia species, 
Sattler has attempted to verify all doubtful records and has weeded out errors. He 
emphasizes the need for certainty in identifications of both moth and host in listing 
foodplant records or other biological features. The widespread practice of uncritical 
copying of host lists results in a misleading picture when compilations are attempted. 
ile gives the example of 12 citations of Cerinthe by French, German, and Italian 
authors causing the impression of widespread use of this plant by Ethmia pusiella. 
‘n reality the records all trace back to one 1868 observation, and it actually refers 

o another Ethmia. Without Sattler’s critical eye the host list would have been 


VOLUME 26, NUMBER 4 263 


impossible to interpret in assessing biological characteristics of the American fauna. 
Lepidopterists everywhere should take note of this critical screening of the distribu- 
tional and biological records, which characteristically are passed down from genera- 
tion to generation like folklore. 

It is surprising that Palearctic ethmiids are so poorly known. One would not have 
expected a dozen undescribed species of moderate sized moths in a group as colorful 
as this. The biological scene is very incomplete, and Sattler points out the need for 
additional knowledge. For example, foodplants are known for only about 23 species 
(jess than 33% of fauna). This is about the same as the state of knowledge in 
Nearctic and West Indian species, a comparable sized fauna, despite a century or so 
headstart by European lepidopterists. For many Palearctic species even the genera- 
tion sequence is not well-known. Ethmia lybiella is said to lack the peculiar “anal 
legs” of the pupa which are characteristic of all Holarctic species groups in the genus, 
but the 1915 record has not been confirmed. Two species, E. rothschildi and E. 
pseudoscythrella, which are exceedingly rare and each known only from one sex, are 
likely to be diurnal moths which fly in early spring, judging from appearances and 
what we know of similar species in the western United States. It seems possible 
that they represent a diversity of overlooked species in arid parts of the Palearctic. 

It was a disappointment to this reviewer that the work includes no discussion of 
possible phylogenetic associations and only cursory treatment of systematic relation- 
ships of the ethmiids, a matter which is by no means well defined when one looks 
at the world fauna of Gelechioidea. However, it can be argued that a faunal study 
of this nature is not the place for such speculation, and Sattler points out in the 
introduction that this could not be a detailed monograph owing to its forced 
scheduling. 

An innovation in format which is bound to be well received is the cross-indexing of 
species references. Eiach species is assigned a number; there is a systematic checklist 
in numerical order and an alphabetical list which doubles as an index to species 
number, page, genitalia figures, and plate numbers. The numbers, given in brackets, 
also are used to identify foodplant and literature references. There is also an index 
to geographical places. 

A major drawback to the format is the resultant cost. No effort has been made to 
economize. The quality of materials is excellent, the artwork and reproduction 
superb, and the high quality binding includes such luxuries as tricolor ribbon-markers. 
The work is presented in two volumes, which may be advisable for larger groups 
but was unnecessary for the ethmiids. Evidently in order to justify a separate plates 
volume, the black and white illustrations are reproduced at a much larger scale than 
was needed, with much wasted space (sometimes only one genitalia figure per 
10.5 x 7.5 inch page), with explanations on a blank opposing page rather than at 
the bottom of the same page. The 190 pages thus consumed could easily have 
been reduced to 30-40% that total without any loss of accuracy or clarity to the 
figures. 

Provided that the authorities of Microlepidoptera Palearctica can solicit contribu- 
tors capable of executing with preciseness comparable to that of Klaus Sattler, they 
will indeed produce the truly monumental work promised in the forward. Too often 
in such faunal works temptation to quickly encompass all taxa dictates lax editorial 
control and selection of specialists of varying ability, resulting in uneven quality 
from one volume to another. It will be interesting to see if the standard of discrimina- 
tion and accuracy established in this treatment of Ethmiidae can be maintained. 


MICROLEPIDOPTERA PALEARCTICA. VOLUME 3. CocHyLiDAE, by Josef Razowski. G. 
Fromme & Co., Vienna. In two bound vols.: text, xiv + 525 p.; plates, 27 color, 
134 black & white, each with unnumbered expl. page facing. Sept., 1970 (Full 
price about $108, subscription price data not available). [text in German] 


264 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


This volume treats 291 species in 21 genera and 5 species of uncertain placement 
in the tortricoid family known in the past as Phaloniidae. Although Razowski has 
published extensively on the group previously and has catalogued the Palearctic 
species separately in 1970 (Acta Zool. Cracov., 15: 341-399), the present work is 
said to contain the following nomenclatural additions and changes: 21 new species, 
1 new subspecies, 14 new synonyms of genera or new status as subgenera (including 
11 Razowski names), 59 new synonyms of species, 7 new status as subspecies, and 
39 new combinations. 


J. A. PowELi, Department of Entomology, University of California, Berkeley 94720. 


JAMAICA AND ITs BUTTERFLIES, by F. Martin Brown and Bernard Heineman, illus- 
trated by Marjorie Statham Favreau and others. 1972. xv + 478 p., 11 colored plates. 
The Curwen Press and E. W. Classey Ltd., London, England. Price: $44.00 U.S. 


This long-awaited work will be a “must” for the amateur of tropical butterflies, for 
the professional systematist, for those interested in the biogeography of the Americas 
and for all who delight in a sumptuous natural history, placed in a tropical island 
setting and illuminated with a wealth of human anecdotes and detailed classificatory 
and biological information. The book consists of a general introduction, containing 
historical, environmental and biological background, supplemented by personal 
reminiscences in Barney Heineman’s inimitable style; then there is an extensive ac- 
count of the biogeography of West Indian butterflies, prepared by Martin Brown, 
long a student of the area; the body of the book follows, consisting of detailed 
treatment of the 120 species, with their classification, probable history and biology; 
the whole is complemented by the set of beautiful and extraordinarily accurate 
colored plates prepared by Marjorie Statham Favreau and finely reproduced by the 
Curwen Press. The book as a whole reflects the brilliance and erudition of Martin 
Brown, the energy and detailed local knowledge of Barney Heineman and the 
warmly human personalities of both authors. 

It would be hard to write a work of this scope without giving some grounds for 
criticism. To expect an individual with as many competing and compelling interests 
as Martin Brown, working against a deadline, to write a completely fault-free account 
of the classification and geography of the Antillean butterflies in the framework of 
the whole neotropical fauna would be asking a good deal of human nature. A num- 
ber of errors or questionable interpretations are accordingly evident in the text, 
though they are far outweighed by the immense value and interest of the material 
as a whole. 

I know less about West Indian butterflies than I did 25 years ago and at the time 
of writing I lack ready access to a good deal of the literature. Consequently I have 
not attempted to review the taxonomy and distribution completely, but I feel obliged 
to pick up a few points. 

In the zoogeographic section there are a number of questionable items. The sub- 
species of Calisto smintheus are listed under the Bahaman C. sibyla, whose range 
is given as Cuba. The recent paper by de la Torre y Callejas, who has revised the 
Cuban populations on the basis of extensive collecting and field work, has been 
passed over without comment. The statements on the sedentary and forest-loving 
habits of Satyridae are exaggerated. Several species of Calisto, including the 
Jamaican C. zangis, are common in open country and some have been recorded as 
pests of sugar cane. The continental species of Hermeuptychia include forms that 
are ubiquitous in a wide variety of habitats, and confinement to forests is not a 
sufficient explanation for their failure to reach the Antilles. Though some of the 
Hispaniolan species and subspecies of Calisto are, as Brown says, closely similar, many 
of them are very distinct. Most of them are local and as the island has been very 


VOLUME 26, NUMBER 4 265 


superficially investigated I expect more, rather than less, species to be recognized when 
the fauna is fully known. 

In the Lycaenidae, Hemiargus ceraunus is not mentioned in the table on p. 60; on 
p. 252-255 it is treated as a subspecies of H. hanno, but on p. 61 we read, “From 
Hispaniola and Costa Rica northward H. hanno is replaced by H. ceraunus, a good 
species related to H. hanno.” Nabokov’s view that the two species have separate 
clines in the West Indies and a zone of overlap in Hispaniola is not mentioned. 
In the Pieridae I consider Ascia menciaee to be probably no more than the Cuban 
subspecies of A. josephina. Eurema messalina is shown in the table as occurring 
in Hispaniola and Puerto Rico, whereas in the text it is correctly cited as occurring 
in Jamaica, Cuba and the Bahamas. Eurema larae, E. lathyi, E. nicippiformis, E. 
euterpiformis, E. pyro and E. chamberlaini are omitted without comment. The 
view taken by d Almeida and by myself that E. dina and E. leuce are distinct species 
separable on genital characters is not mentioned, nor is the sympatric occurrence of 
two very distinct populations of this complex in Hispaniola. The Hispaniolan 
Phoebis editha is omitted from the table, though Brown himself first recognized and 
described the strikingly dimorphic, superficially sennae-like male. P. avellaneda [sic] 
certainly occurs in Hispaniola, as does P. philea thalestris. I agree with the authors 
that this last may well be a good species rather than only a subspecies. In the 
Papilionidae Battus devilliers [sic] is listed as occurring in Hispaniola, where it is 
in fact replaced by the related but very distinct B. zetides. Papilio polyxenes is 
listed under the suppressed name P. ajax. P. machaonides is omitted. 

This is not the place for detailed discussion of biogeography. I think Brown would 
be the first to say that many of his speculations as to dates of arrival should be taken 
with a grain of salt. The role of the probably emergent Honduras Banks in pop- 
ulating the West Indies in the Pleistocene and Tertiary seems to me to be probably 
underestimated, whereas the relatively recent Yucatan-Cuba immigration route is 
possibly given undue prominence, though it has certainly been important for the 
recent fauna. The existence of special Jamaican elements, consisting first of species 
not found elsewhere in the Antilles, but closely allied to Central American forms 
(e.g. Papilio thersites, Anaea johnsoni and Mestra dorcas) and second of segregates 
or duplications of widespread Antillean forms with considerable endemic differentia- 
tion (e.g. Hemiargus dominica, Electrostrynon pan and Pyciodes proclea) is not 
mentioned explicitly or specifically explained. The role of geological rifting and 
drift in the Antilles also deserves attention and assessment. In the comparison of 
total faunas, the method adopted underestimates the degree of differentiation from 
large faunas and overestimates that from small faunas. The fauna of Hispaniola will 
certainly prove to be much underestimated in Table 2, both because of the omission 
of a number of known species and because Hispaniola has been far less thoroughly 
explored entomologically than the other Greater Antilles. 

The taxonomic section, with its thorough treatment of the classification and dis- 
tribution of the species and groups, is of course the part of the book that provides 
the largest amount of detailed and often new scientific information. The extensive 
distributional record is a tribute to the industry both of the authors and of the other 
collectors whose work they so carefully summarize and document. The biogeographic 
and evolutionary interpretations of the various taxa are of course more speculative, 
and opinions on them will vary. 

A rather superficial scrutiny reveals some doubtful points. I won't debate the 
division into families and other higher groups, as the relative ranking is really in 
large part a matter of personal preference. I do think the separation of Heliconiinae 
from Argynninae at the family level is rather hard to justify. I would rather have 
seen Battus and Parides separated in the Papilionidae, for these two genera differ 
in really striking morphological characters. I myself also consider the New World 
Eurytides significantly different from the Old World Graphium; the structural dif- 
ferences are for instance substantially more important than those that separate 


266 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Libytheana from Libythea or Aphrissa from Phoebis. On the whole, however, I con- 
sider the generic classification moderate and well balanced. 

The discussion of the Danaidae is taxonomically sound, but the biogeography 
suffers here as elsewhere for making dispersal contingent on mass migration. 
Actually the latter is often confined to males (though not so in the Danaidae), and 
there is abundant indirect evidence that waif dispersal of non-migratory species 
is a significant factor in populating islands even when they are much more strongly 
isolated than the Greater Antilles. In the present context, oversea dispersal of D. 
eresimus is excluded because this species is not known to be migratory, though Bates 
has recorded an obvious waif from the Florida Keys and the occurrence of the 
species is in general rather erratic and local. The detailed status of D. plexippus 
in the West Indies is still confused, though the general picture of resident populations 
contaminated in varying degree by migrants from the north seems clear enough. 
We badly need a statistical study comparable to that which Brown so competently did 
on Heliconius charitonius. The supposed subspecies of Lycorea cleobaea are based, 
to the best of my belief, on very scanty material; in my opinion they will require 
review. Anetia jaegeri and A. cubana are treated as distinct in the text, though 
united in the geographic table. I agree with the former interpretation. The actual 
capture of an Anetia of this group in Jamaica is to be awaited with the keenest interest. 

Mr. Gerardo Lamas M. has kindly provided me with some notes on the Ithomiidae 
(or Ithomiinae, as he prefers to call them). He points out that hair tufts are 
found on the hindwings of females as well as males, at least in some species of 
Thyridia. The type-specimen of Papilio irene Drury apparently is not at the Uni- 
versity Museum, Oxford, where a search by Mr. E. Taylor has failed to reveal it. 
Although the general image of ithomiids as local and shade-dwelling is correct, many 
of the species are widely distributed, some live in open places rather than in the 
shade, and at least one, Mechanitis lysimnia nessaea (Hiibner), has been recorded 
as a migrant (May 1924, cited by C. B. Williams, The Migration of Butterflies, 1930: 
341). Most or all the species are highly resistant to mechanical injury, and thus 
might survive transport by storms. Greta was placed by Fox (1968) in the tribe 
Thyridiini. Fox did not divide Greta into two subgenera. Godman and Salvin in 
1879 erected the new genus Hypoleria with two sections, A and B. These correspond 
today to Hypoleria s.s. (Section A) and Greta (Section B), as was stated by Fox. 
Fox (1968) lists seven species from Central America. The proper name of the 
Jamaican ithomiine appears to be Greta diaphana (originally Papilio diaphanus); 
diaphane is an unnecessary nomen novum given by Hibner, who thought that 
diaphanus was preoccupied. 

The treatment of Junonia (“Precis”) reflects continuing uncertainty as to the 
status of the forms. Thorough biological and statistical study using modern methods 
will be needed to resolve the problems of this group. Such a study will make a 
rewarding project for some future student. Ideally it will extend to the members 
of the Old World orithyia complex as well, whose relationships to evarete and 
allies were debated by Eliot and myself. The larval differences observed by Turner 
between genoverva-like and zonalis-like stocks in Jamaica are mentioned, but Brown 
and Heineman wisely await more comprehensive evidence before assessing these 
differences. 

In the Lycaenidae the species pan (Drury) is placed in Strymon in the text but 
in Electrostrymon in the zoogeographic table. 

The authors confess themselves puzzled by Avinoff’s record of Anteos lacordairei 
Boisduval. I can shed some light on this from personal recollection. Avinoff con- 
sidered lacordairei and maerula to be sibling species analogous to Kricogonia lyside 
and castalia, which he also considered distinct. He thought lacordairei was a rela- 
tively scarce species which differed in the blackish not reddish antennae, shorter 
wing and more strongly striated underside from the more abundant maerula. When 
I was more familiar with the problem I thought this view had some merit, but 


VoLUME 26, NUMBER 4 2.67 


looking back I don’t think the evidence was very strong. However the idea should 
perhaps be checked out by someone who has good material at hand. 

On the Papilionidae, Papilio troilus has turned up once or twice as a stray in 
Cuba. As several sight records and speculative occurrences are discussed, I was 
disappointed to see no reference to Avinoffs “Papilio arawak” a name he used 
hypothetically and in conversation to refer to several sightings of a large white 
butterfly, not an Anteos, which he thought might be an unknown swallowtail. It 
was one of his lifelong ambitions to find and capture this butterfly and the hope 
should not be lost to the present generation of Jamaican lepidopterists. 

There are a number of spelling lapses in both the general and the taxonomic 
sections of the book, for example Sideronia for Siderone (p. 52), teleboa for teleboas 
(p. 54, 55), avellanada for avellaneda (p. 62), thetys for tethys (p. 85, 86), 
Glossypteris for Glossopteris (p. 98), exotische for exotischer (p. 103, et al.), 
bekannte for bekannter (p. 124 et al.), eleucha for eleuchea (p. 137), dorcus for 
dorcas (p. 156), Suite for Suites (p. 197), Ithoballus for Ithobalus (p. 321) and 
André for Andrey (p. 325). The capitalization and punctuation of bibliographic 
references in the text are erratic. A number of relevant references are omitted from 
the bibliography. These include papers by Wm. P. Comstock, Eliot, Ford, Hemming, 
Martorell, de la Torre y Callejas and a couple by myself, among others. 

These imperfections, though numerous, are in general minor. They will not 
seriously detract from the standing of this book as the most thorough treatment ever 
given to any Antillean lepidopterous fauna. Above and beyond this the book is 
produced in such a way that it is a collector's piece worthy of a place on any book- 


shelf. 


EUGENE Monroe, Entomology Research Institute, Canada Department of Agricul- 
ture, Ottawa. 


NOTES AND NEWS 


I wish to thank the many persons who provided assistance to me during this first 
year of my editorship. 

The members of the Editorial Committee of the Journal were most helpful as 
primary reviewers of submitted manuscripts. In addition, the following individuals 
reviewed one or more manuscripts upon request: D. E. Berube, H. K. Clench, C. V. 
Covell, C. D. Ferris, D. J. Klingener, A. B. Klots, A. P. Mange, W. B. Nutting, 
C. L. Remington, D. F. Schweitzer, A. M. Stuart, and O. R. Taylor. All of these 
persons performed a valuable service to the Society, and thanks are warmly extended 
to them. 

My wife, Katherine, provided the cover drawing (Catocala flebilis Grote) for this 
volume, and aided in many other ways. Nancy Wells served as my editorial as- 
sistant throughout the year, and her invaluable help is most gratefully acknowledged. 
Finally, from among the many, I especially wish to thank D. F. Hardwick and S. S. 
Nicolay for their particular kindnesses. 


THEODORE D. SARGENT 


268 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


OBITUARY 


HUBERT DE LESSE (1914-1972) 


One of the best-known of European lepidopterists, and a former vice- 
president of the Lepidopterists’ Society, Hubert de Lesse, was found dead 
in his house in mid-March, following a period of two years of progres- 
sively failing health, with recurrent illnesses and hospitalizations. 

Chief Research Scientist of the Centre national de la Recherche 
scientifique, and Corresponding Member of the Muséum national 
d Histoire naturelle in Paris, Dr. de Lesse was well-known to professional 
lepidopterists through his many publications on Rhopaloceran chromo- 
somes, as well as his systematic studies on Satyrinae and Lycaenidae. 
Fewer of his scientific colleagues knew of his devotion to alpinism 
(mountain climbing), or were aware of his extensive travels to tropical 
lands; his intense interest in field studies of Lepidoptera literally led him 
to visit the climatological “ends of the earth.” These travels resulted in the 
gathering and publication of an uncountable number of significant ob- 
servations on the zoogeography, behavior, sympatry, and interaction of 
butterfly species, which will modify and perfect presently accepted con- 
cepts in these areas for several generations to come. 

Born in Paris on April 7, 1914, and an active member of the Société 
entomologique de France since 1932, Dr. de Lesse spent most of his 
research career successfully applying to butterfly systematics the methods 
of cytological analysis which he learned during two years of botanical 
research (1938-1939). At the end of World War II, he dedicated his 


VOLUME 26, NUMBER 4 269 


talents to a study of the satyrine genus Erebia, combining his love for 
Lepidoptera with that for mountain heights. After a trip to Greenland 
with the Second French Polar Expedition (1949), which resulted in a 
series of publications on the botany and entomology of this frozen land, 
he applied morphological studies of both male and female genitalia to the 
revision of the satyrine genera Satyrus, Pararge, Maniola, and Lethe. 
Continuing studies on various nymphalid groups and Lycaenidae, espe- 
cially alpine forms, led to a variety of publications on these and related 
subjects; in the two years 1951-1952, over thirty papers appeared from his 
hand. 

Starting in 1950, a new field—that of chromosome study—came to be 
more significant in the work and publications of Hubert de Lesse. During 
the twenty years that he devoted himself to this line of research, he 
established his name as one of the pioneers and principal workers in the 
field, publishing nearly sixty papers on the subject and its direct systematic 
implications. Many changes in specific and subspecific designations and 
relationships were introduced by his studies of chromosome numbers in 
various Lepidopteran groups. However, the real value of this work lay 
in the derived understandings of the fundamental biological processes 
of speciation, despeciation, hybridization, sympatric isolation and the 
formation of sibling species, and—in fact—evolution itself, observable 
as though it were a contemporary phenomenon, through examinations 
of cytological preparations and systematic field studies in carefully chosen 
localities. 

His doctoral thesis, on chromosome variation and speciation in Rhopa- 
locera, was successfully presented and defended in the Sorbonne on 
November 16, 1960. In the following years, Dr. de Lesse undertook ex- 
tensive travels to all parts of Europe, Turkey, Iran, Libya, eastern and 
southern Africa, and Central and South America. He also received ma- 
terial fixed for chromosome study from colleagues and correspondents 
in North America, northern and equatorial Africa, and Madagascar. 

This author had the privilege of working with Hubert de Lesse over 
the last two years of his life, following his visit to Brazil at the end of 
1969. In joint and individual excursions during his visit, we fixed for 
chromosome examination and studied in the field a large number of 
species of Neotropical Lepidoptera, representing the families and groups 
already known to be most interesting for further study. Although already 
suffering from ill health, Dr. de Lesse showed himself to be a persistent, 
observant, and effective field worker, dedicating himself many hours 
each day to the task of studying and fixing Brazilian Rhopalocera, during 
a period of highly unfavorable weather conditions. 


270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The continued deterioration of his health prevented Dr. de Lesse from 
bringing to fruition the large-scale project developed during this briet 
period of our collaboration. However, a short joint paper restricted to the 
examination of the material fixed during the 1969 visit has been pub- 
lished, and laid the basis for a forthcoming series of studies on the evolu- 
tionary significance of chromosome numbers and patterns in Neotropical 
Lepidoptera. The father and originator of this series unfortunately has 
not survived to see it come to maturity, but his careful instructions, time- 
tempered methods, and wise orientation will continue in the work now 
being carried to its conclusion in a variety of laboratories in Europe and 
the Americas. 

Hubert de Lesse was, above all, an individualist and field naturalist, 
happiest in the isolation of alpine meadows or tropical forests, far from 
the urban noise and population pressure of his home city. However, he 
constrained himself to spending long hours in his flower-surrounded Paris 
house on rue de l'Est (Reuil-Malmaison), searching for, drawing, and 
studying, under a microscope at over a thousand magnifications, tiny 
bits of concentrated biological and evolutionary information. The result 
of this painstaking sacrifice, which took a progressive toll on his vision, 
nerves, and general health, is a legacy of suggestive and authoritative 
publications, which will serve the world community of lepidopterists 
and biologists in general for many years in the future. 

Dr. de Lesse’s extensive collections are in the Entomological Laboratory 
of the Muséum national, in Paris. A selected list, including the more 
significant of his 137 publications, conserving original numbers in the 
complete list, follows: 


2. Contribution a l’étude du genre Erebia. Rev. Fr. de Lépid. 11: 97-118 (1947). 
4. meee fagi Scop. et H. aelia Hffmsg. Bull. Soc. Lin. Lyon n° 7: 123-129 
1948). 

5. Contribution a létude des Rhopalocéres du départment de la Dréme. Lambil- 
lionea 48: 59-64 (1948); 49: 8-30 (1949). 

8. Contribution a l’étude du genre Coenonympha. Lambillionea 49: 68-80 (1949). 

10. Contribution a étude du genre Erebia: armures génitales femelles. Rev. Fr. 
d’Entomol. 16: 165-198 (1949). 

11. Expéditions polaires francaises. Zoologie. Premiére note: Microlepidoptera 
(in collaboration with P. Viette). Ann. Soc. Entomol. France 115 (1946) 
81-92 (1949). 

13. Observations sur les Lépidoptéres du Groenland, et remarques sur leur homo- 
chromie. Rev. Fr. de Lépid. 12: 163-169 (1949). 

14. Quelques indications sur Melitaea britomartis Assm., espéce a rechercher en 
France. Bull. Soc. Lin. Lyon n’ 2: 38-41 (1950). 

17. Notes zoologiques et botaniques sur l'Ouest du Groenland. La Terre et la 
Vie n’ 4: 175-201 (1950). 

18. Expéditions polaires frangaises. Zoologie. 4° note: Macrolepidoptera. Ann. Soc. 
Entomol. France 118 (1948), 51-78 (1951) 


? 


VOLUME 26, NUMBER 4 Dial 


Mlle 
23. 
24. 


25. 


27. 


28. 


ol. 
32. 


33. 


34. 


36. 


37. 


Al. 


43. 
44, 
AT. 


48. 


50. 


53. 


56. 


Sur une espéce de Satyridae mal connue: Hipparchia (Pseudotergumia) wyssii 
Christ. Bull. Soc. Entomol. France n°’ 4: 50-53 (1951). 

Divisions génériques et subgénériques des anciens genres Satyrus et Eumenis 
(s.l.). Rev. Fr. de Lépid. 13: 39-42 (1951). 

Quelques Lépidoptéres de Besse en Chandesse (in collaboration with P. Viette ). 
Rev. Fr. de Lépid. 13: 78-83 (1951). 

Contribution a l’étude du genre Erebia (3° note). Répartition dans les Pyrénées 
de E. tyndarus Esp. et E. cassioides Rein. et Hohenw. Vie et Mileau 2: 95-123 
(1951). 

Contribution a l'étude du genre Erebia (4° note). Répartition de E. pandrose 
(Borkh.) et de sa sous-espéce sthennyo Grasl. dans les Pyrénées. Vie et Mileau 
2: 267-277 (1951). 

Les types de Nymphalidae paléarctiques du Laboratoire d’Entomologie du 
Muséum d’Histoire Naturelle de Paris (in collaboration with G. Bernardi). 
Bull. Soc. Entomol. France n’ 9: 136-143 (1951). 

Révision de l’ancien genre Satyrus. Ann. Soc. Entomol. France 120: 77-101 
(1951). 

Expéditions polaires francaises. Zoologie (6° note). Biogéographie des 
Lépidoptéres du Groenland. Ann. Soc. Entomol. France 119: 97-116 (1951). 
Contribution a létude du genre Erebia (6° note). Notes de répartition et 
nouvelles indications sur E. eriphyle (Frr.) et E. stirius (Godt.) récemment 
signalés de France. Rev. Fr. de Lépid. 13: 130-137 (1951). 

Un Rhopalocére de plus a inscrire au Catalogue des Lépidoptéres de France— 
Euphydryas ichnea Bdv. (= intermedia Mén.) (Nymphalidae). Rev. Fr. de 
Lépid. 13: 1438-152 (1951). 

Note sur les genres Precis Hb. et Junonia Hb. (Lep. Nymphalidae). Bull. Soc. 
Entomol. France 57: 74-77 (1952). 

La variabilité géographique de Lycaena helle Denis et Schiff. (Lep. Ly- 
caenidae ) (in collaboration with G. Bernardi). Rev. Fr. de Lépid. 13: 203-213 
(952): 

Liste des Grypocéres et Rhopalocéres de la Faune francaise conforme aux 
Régles internationales de la Nomenclature. Satyridae (Rev. Fr. de Lépid. 13: 
241-245 (1952). 

Révision des anciens genres Pararge (s.l.) et Maniola (= Epinephele auct.). 
Ann. Soc. Entomol. France 121: 61-76 (1952). 

Cytologie—Quelques formules chromosomiques chez les Lycaenidae (Lépi- 
doptéres Rhopalocéres ). Comptes Rendus Acad. Sci. 235: 1692-1694 (1952). 
Révision des Neohipparchia (Lep. Satyridae) d’Afrique du Nord. Bull. Soc. 
Sci. Nat. Maroc. 32: 91-105 (1952). 

Formules chromosomiques nouvelles du genre Erebia (Lepid. Rhopal.) et 
séparation d’une espéce méconnue. Comptes Rendus Acad. Sci. 236: 630-632 
(1953). 

Formules chromosomiques de Boloria aquilonaris Stichel, B. pales D. et Schiff., 
B. napaea Hoffmsg. et quelques autres Lépidoptéres Rhopalocéres. Rev. Fr. de 
Lépid. 14: 24-26 (1953). 

Formules chromosomiques nouvelles chez les Lycaenidae (Lepid. Rhopal.). 
Comptes Rendus Acad. Sci. 237: 1781-1783 (1953). 

Les types de Nymphalidae paléarctiques du Laboratoire d’Entomologie du 
Muséum d'Histoire Naturelle de Paris (2° note). (in collaboration with G. 
Bernardi). Bull. Soc. Entomol. France 58: 154-160 (1953). 

Recherches cytologiques et biogéographiques sur quelques Lépidoptéres. Bull. 
Soc. Zool. France 78: 287-291 (1953). 

Découverte d'un nouvel Erebia dans les Hautes-Pyrénées (in collaboration with 
H. Descimon). Rev. Fr. de Lépid. 14: 119-122 (1953). 


86. 


90, 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Formules chromosomiques nouvelles chez les Lycaenidae (Lepid. Rhopal. ). 
Comptes Rendus Acad. Sci. 238: 514-516 (1954). 


- Discussion de certains caractéres morphologiques d’Arethusana arethusa Schiff. 


et des formes boabdil Rambur et dentata Stgr. Rev. Fr. de Lépid. 14: 140-142 
(1954). 


- Comments on two recent papers on the genus Erebia: a reply. The Entomologist 


87: 91-96 (1954). 


_ Expériences de croisements dans le genre Erebia (Lépidoptéres Satyridae) (in 


collaboration with Z. Lorkovic). Bull. Soc. Zool. France 79: 32-39 (1954). 
Contribution 4 l’étude du genre Erebia (12° note). Description des premiers 
états. Rev. Fr. de Lépid. 14: 167-179 (1954). 

Nouvelles découvertes concernant le degré de parenté dErebia tyndarus 
Esp. et E. cassioides Hohenw. (in collaboration with Z. Lorkovic). Lambillionea 
54: 58-68 and 78-86 (1954). 

Etat actuel de la systématique du groupe d’Erebia tyndarus Esp. (s.l.). Etude 
dun travail récent. Rev. Fr. de Lépid. 14: 228-236 (1954). 

Contribution A étude du genre Erebia (13° note). Description des premiers 
états. Rev. Fr. de Lépid. 14: 251-257 (1954). 

Recherches de formules chromosomiques chez les Zygaena. Bull. Soc. Lin. 
Lyon 24: 142-144 (1955). 

Distribution holarctique d’un groupe d’espéces du genre Erebia (Lépidopteres ) 
récemment séparées d’aprés leurs formules chromosomiques. Comptes Rendus 
Soc. Biogéogr. 276: 12-18 (1955). 

Note supplémentaire sur le groupe d'Erebia tyndarus Esp. (in collaboration 
with Z. Lorkovic). Lambillionea 55: 55-58 (1955). 

Etude cytologique des Lysandra fixés par M. H. Beuret. Mitt. Entomol. Ges. 
Basel, 77-80 (1956). 

Quinze jours de récoltes en Italie centrale et découverte d’Erebia pandrose Bkh. 
aux Monti della Laga. Lambillionea 56: 9-16 (1956). 


. Notes on the species groups in the genus Erebia: a reply. The Entomologist 


89: 278-282 (1956). 

Fixation de lectotypes et description d’une nouvelle sous-espéce dans le groupe 

d’Erebia tyndarus Esp. (Lepid. Nymphalidae Satyrinae). Rev. Fr. de Lépid. 
15: 147-150 (1956). 

Révision du genre Lethe (s.l.) (Lep. Nymphalidae Satyrinae). Ann. Soc. 

Entomol. France 125: 75-95 (1957). 

Liste des Lépidoptéres Rhopalocéres récoltés en 1955 au Liban par H. de 

Lesse. Introduction, Hesperiidae, Nymphalidae. Bull. Soc. Lin. Lyon 26: 

238-241 (1957). 

Une nouvelle forme de Boloria aux Alpes Pontiques (in collaboration with A. 

Crosson du Cormier). Lambillionea 57; 34-57 (1957). 

Description de deux nouvelles espéces d’Agrodiaetus (Lep. Lycaenidae) séparées 

a la suite de la découverte de leurs formules chromosomiques. Lambillionea 57: 

65-71 (1957). 

(acess Rhopalocéres récoltés en Iran. Introduction. Alexanor 1: 39-46 
1959 ). 

Séparation spécifique d’un Lysandra d’Afrique du Nord 4a la suite de la 

découverte de sa formule chromosomique. Alexanor 1: 61-64 (1959). 

Caracteres et répartitions en France d’Erebia aethiopellus Hoffmsg. et E. 

mnestra Hb. Alexanor 1: 71-81 (1959). 

Caracteres externes et formule chromosomique d’Agrodiaetus baytopi n. sp. 
(Lep. Lycaenidae). Bull. Soc. Ent. Mulhouse, 45-48 (1959). 

Note sur deux sous-espéces d’Agrodiaetus (Lep. Lycaenidae) récemment 

séparées d’aprés leurs formules chromosomiques. Lambillionea 59: 5-10 (1959). 


VOLUME 26, NUMBER 4 2h 


Oike 


92. 


93. 


95. 


98. 


99: 


100. 
OL: 


102. 


103. 
104. 
107. 
109. 


110. 
WI. 


WAL), 


114. 


ILS), 
BLE 
118. 
I). 


120. 
123. 


124. 


125. 


Nouvelle étude chorologique et cytologique conduisant 4 changer l’application 
du nom d’Agrodiaetus iphigenia H.S. (Lep. Lycaenidae). Lambillionea 59: 
57-61 (1959). 

Les nombres de chromosomes dans la classification du groupe d’Agrodiaetus 
ripartii Freyer (Lepidoptera, Lycaenidae). Rev. Fr. d’Entomol. 27: 240-264 
(1960). 

Recherches sur la distribution des représentants du groupe d’Erebia tyndarus 
Esper. Erebia calcarius au Monte Cavallo au nord de Venise (in collaboration 
with Z. Lorkovic). Bollet. Soc. Entomol. Ital. 90: 123-129 (1960). 

Spéciation et variation chromosomique chez les Lépidoptéres Rhopalocéres. 
Ann. Sci. Nat., Zool. 1: 1-223 (1960). Doctoral thesis defended in the Sorbonne 
on November 16, 1960. 

Les hybrides naturels entre Lysandra coridon Poda et L. bellargus Rott. 
Alexanor 2: 22-30 (1961). 

Variations géographiques des caractéres externes chez les espéces autrefois 
réunies sous le nom d’Agrodiaetus ripartii Frey (Lep. Lycaenidae). Rev. Fr. 
d’Entomol. 28: 93-100 (1961). 

Les nombres de chromosomes chez Agrodiaetus dolus Hb. et les espéces voisines. 
Alexanor 2: 57-63 (1961). 

Signification supraspécifique des formules chromosomiques chez les Lépi- 
doptéres. Bull. Soc. Entomol. France 66: 71-83 (1961). 

Deux nouvelles formules chromosomiques ot n dépasse 100 chez les Ly- 
caenidae (Lépidoptéres Rhopalocéres). Comptes Rendus Acad. Sci. 253: 1982— 
1984 (1961). 

Variation chromosomique chez Agrodiaetus actis H.S. et A. altivagans Forst. 
Rev. Fr. d’Entomol. 29: 66-75 (1962). 

Formules chromosomiques de quelques Lépidoptéres Rhopalocéres du Sénégal 
(in collaboration with M. Condamin). Bull. I.F.A.N. 24 A: 464-473 (1962). 
Lépidoptéres Lycaenidae récoltés en Iran en 1961. Alexanor 2: 305-312 
(1962); 3, 33-38 (1963). 

Variation chromosomique chez les Agrodiaetus (Lep. Lycaenidae). Rev. Fr. 
d’Entomol. 30: 182-189 (1963). 

Nomenclature des Erebia francais. Alexanor 3: 127-136 (1963). 

Les nombres de chromosomes chez quelques Erebia femelles (Lep. Satyrinae). 
Rev. Fr. d’Entomol. 31: 112-115 (1964). 

Formules chromosomiques de quelques Lépidoptéres Rhopalocéres du Gabon 
(in collaboration with G. Bernardi). Biologica Gabonica 1: 65-71 (1964). 
Formules chromosomiques de quelques Lépidoptéres Rhopalocéres du Sénégal 
et de Céte dIvoire (in collaboration with M. Condamin). Bull. I.F.A.N. 27: 
1089-1094 (1965). 

Impressions d’un Lépidoptériste en Amérique du Sud. Alexanor 4: 171-178 
(1965). 

Formules chromosomiques de quelques Lépidoptéres Rhopalocéres d Afrique 
Centrale. Ann. Soc. Entomol France, N.S. 2: 97-101 (1966). 

Les nombres de chromosomes chez les Lépidoptéres Rhopaloceéres néotropicaux. 
Ann. Soc. Entomol. France, N.S. 3: 67-136 (1967). 

Formules chromosomiques des Rhopalocéres d'Afrique du Nord. Bull. Soc. 
Entomol. France 72: 20-25 (1967). 

Note sur le genre Euptychia. Lambillionea 67: 34-39 (1967). 

Formules chromosomiques de Lépidoptéres Rhopalocéres d’Uganda et du Kenya. 
Ann. Soc. Entomol. France, N.S. 4: 581-599 (1968). 

Les hybrides naturels de Lysandra coridon Poda et Lysandra bellargus Rott. 
(2° note). Alexanor 6: 73-82 (1969). 

Les nombres de chromosomes dans de groupe de Lysandra coridon Poda. Ann. 
Soc. Entomol. France, N.S. 5: 469-522 (1969). 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


. Quelques nombres de choromosomes de Lépidoptéres Rhopaloceres d’Amerique 


du Nord. Bull. Soc. Entomol. France 74: 57-58 (1969). 


. Les nombres de chromosomes chez les Lépidoptéres Rhopaloceres em Amérique 


centrale et Colombie. Ann. Soc. Entomol. France, N.S. 6: 347-356 (1970). 


. Systématique du groupe de Lysandra coridon (Lep. Lycaenidae). Alexanor 


6: 203-224 (1970). 


. Formules chromosomiques de quelques Lépidoptéres Rhopalocéres de Guyane. 


Ann. Soc. Entomol. France, N.S. 6: 849-855 (1970). 


. Une nouvelle sous-espéce de Melanargia galathea L. (Lep. Nymphalidae 


Satyrinae). Alexanor 7: 156-157 (1971). 


. Formules chromosomiques de Lépidoptéres Rhopalocéres du Brésil (in collabora- 


tion with K. S. Brown, Jr.). Bull. Soc. Entomol. France 76: 131-137 (1971). 


. Variations géographiques des nombres chromosomiques chez les Lycaenidae. 


Comptes Rendus Soc. Biogéogr. 48 (407-421): 33-38 (1972). 


. Formules chromosomiques de Lépidoptéres Rhopalocéres de Madagascar. Bull. 


Soc. Entomol. France 77, in press (1972). 


ACKNOWLEDGMENTS 


This obituary notice was prepared with information received from, and 
with the help and correction of, Dr. Pierre Viette of the Muséum national 
d Histoire naturelle in Paris, a long-time friend and colleague of Hubert 
de Lesse. 

The author acknowledges financial assistance for the study of the 
chemistry of Brazilian insects, from the National Science Foundation 
(Grant N? GB 5389 XI), and the Brazilian Conselho Nacional de Pesquisas 
and Banco Nacional do Desenvolvimento Econémico. 


KeirH S. Brown, Jr., Centro de Pesquisas de Produtos Naturais, U.F.R.J., Rio de 
Janeiro, Brazil. 


VOLUME 26, NUMBER 4 


275 


INDEX TO VOLUME 26 


(New names in boldface) 


Acrolophus vigia, 136 
Aglais urticae, 116 
Angleberger, M. A., 177 
Antiplecta triangularis, 225 
Arctiidae, 222 
Austin, G. T., 63 
Badger, F. S., 13 
behavior, 24, 28, 29, 33, 65, 82, 89, 97, 
Me TIG 131, 155, 161, 996. 261 
Beutelspacher, C. R., 133 
Blanchard, A., 56, 79 
book reviews, 123, 124, 125, 199, 200, 
203, 262, 263, 264 
Bowden, S. R., 170 
Bowe, J. J., 122 
Brown, F. M., 111, 245 
Brown, K. S., 183, 268 
Callophrys fotis bayensis, 237 
gryneus, 112 
siva, 112 
Callosamia securifera, 86 
Cashatt, E. D., 1 
Castilonia, R., 64 
Catocala relicta, 94 
spp., 35, 105 
Ceratomia hageni, 198 
Cercyonis pegala blanea, 141 
Chamyris cerintha, 180 
Chloroclystis rectangulata, 220 
Chlosyne rosita browni, 51 
Colias alexandra, 214 
collecting procedures, 35, 64, 183 
collections, 108, 198 
Crambus carpenterellus, 245 
Danaidae, 21, 137 
Danaus plexippus, 137, 219 
Darapsa myron, 234 
versicolor, 234 
Deidamia inscripta, 234 
Dominick, R. B., 69, 234 
Donahue, J. P., 108 
ecology, 105, 112, 133, 155, 196, 226 
Ehrlich, A. H., 196 
Ehrlich, P. R., 196 
Emmel, T. C., 140, 237 
Epiplemidae, 222 
Erosia incendiata, 224 
Erinnyis obscura, 234 
Eueides cleobaea zorcaon, 55 
Eugonobapta nivosaria, 218 
Eumedonia eumedon modestus, 215 


Euphydryas phaeton, 122 

evolution, 112, 140 

Ferguson, D. C., 220, 222 

errs, GD. 210 237 

Franklin, C. M., 198 

Eros: S) We E73 

Funk, R. S., 203 

genetics, 100, 109, 250, 253 

Senitalia, 1) 58, 61, 117, 129. 130, 154. 
220 

Geometridae, 218, 220, 222 

Glaucopsyche lygdamus, 63. 

Godfrey, G. L., 180 

Hardwick, D. F., 24, 29, 82, 89 

Heliconiidae, 49 

Hemileuca diana, 245 

Hesperiidae, 15, 127, 230, 233 

Hogue, C. L., 33 

Howe, W. H., 247 

hybridization, 28, 104, 255 

Hylesia sp., 33 

Hyperborea czekamowoskii, 222 

Irwin, R. R., 198 

Ithomiidae, 123 

Jae, R. J., 28 

Johnson, K., 112 

Jordan medal, 207 

Kellogg, C. G., 35 

Kendall, R. O., 49 

Klots, A. B., 199 

larval foodplants, 112, 122, 155, 177, 
219, 235 

Kevin, MOP: £77 

Libytheidae, 19 

life histories, 24, 29, 82, 86, 89, 143, 155, 
180, 218, 237, 242, 247 

Limenitis lorquini, 261 

list, state, 13 

Lycaeides argyrognomon nabokovi, 151 

Lycaena heteronea, 63 

Lycaenidae, 49, 63, 112, 150, 196, 215, 
ISN 233, 237 

Makielski, S. K., 109 

Masters, J. H., 123, 125, 150, 249 

Mattoon, S. O., 140 

Miller, L. D., 200, 207 

Miller, S., 229 

mimicry, 63 

Munroe, E., 123, 264 

name changes, 111, 228 

Napaea eucharilla picina, 136 


276 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Nekrutenko, Y. P., 215 Riodinidae, 18 
new records, 49, 116, 196, 198, 220, 222, Rupert, L. R., 218 
229, 234 Sargent, T. D., 35, 94 
new species, 11, 56, 59, 61, 79, 128, 150 Saturniidae, 33, 86, 245 
new subspecies, 141, 151, 215 Satyridae, 21, 140, 232, 233 
Noctuidae, 24, 29, 35, 56, 82, 89,94, 105, Schinia citrinellus, 82 
180, 196 intrabilis, 24 
notes and news, 65, 126, 203, 267 jaegeri, 89 
Nymphalidae, 19, 49, 116, 122, 155, 226, pallicincta, 29 
233, 261 Scott, F. W., 116 
Nymphalis californica, 226 Scriber, J. M., 236 
obituaries, 66, 268 Semaeopus cantona, 224 
Oidaematophorus balanotes, 1 sex ratios, 102, 176 
glenni, 11 Shields, O., 116 
grandis, 5 Shull, E. M., 13 
kellicottii, 7 Sibatani, A., 117 
lacteodactylus, 9 Slansky, F., Jr., 219 
Oncocnemis eottami, 56 Sphingidae, 198, 234, 247 
heterogena, 59 Sphinx franckii, 234 
Owen, D. F., 105 vashti, 247 
Paonias astylus, 234 Steinhauser, S. R., 127 
Papilionidae, 17, 109, 177, 235 Strymon albata sedecia, 51 
Papilio glaucus, 109, 177, 212, 213, 235 Swank, C. K., 196 
Paramiana eallaisata, 61 taxonomy and systematics, 117, 170, 249 
Pieridae, 17, 104, 170, 228 techniques, 64, 69, 137, 184, 210 
Pieris napi, 170 Thecla basalides, 135 
protodice, 104 Thysania zenobia, 196 
rapae, 104 Triphosa affirmata, 224 
spp., 170 type localities, 245 
Platte Ae ee Urodus parvula, 173 
polymorphism, 94, 109, 250 Urquhart, F. A., 137 
Powell, J. A., 226, 262, 263 Van Buskirk, M. D., 86 
Priestaf, R. C., 104 Victorina epaphus, 155 
Pseudophilotes, 111 Wirth, W. W., 65 
Pterophoridae, 1 Wright, B., 116 
Pyle, R. M., 261 Young, A. M., 155 
Pyralididae, 245 Yponomeutidae, 173 
Pyromorpha eaelebs, 79 Zestusa levona, 128 
Pyromorphidae, 79 spp., 127 


Riley, N. D., 228 Zizula hylax, 196 


EDITORIAL COMMITTEE OF THE JOURNAL 
Editor: THEODORE D. SarcENT, Department of Zoology, 

University of Massachusetts, Amherst, Massachusetts 01002 
K. S. Brown, J. M. Burns, R. H. Carcasson, J. P. Donanue, J. F. 


Gates Criarke, R. O. KENDALL, J. H. Masters, L. D. MILLER 
A. P. Piatt, J. R. G. TuRNER 


? 


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 LirERATuURE 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 414 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 article’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 approx- 
imate 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. 

Page Charges: Authors with grant or institutional funds are requested to pay a 
charge of $24.00 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 
case of authors who have no grant or institutional funding, as it is not intended that 
any author should pay this charge from personal funds. However, all authors will 
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: Dr. C. V. Covell, Department of Biology, University of 
Louisville, Louisville, Kentucky 40208. 


ALLEN PRESS, INC. erInreo LAWRENCE, KANSAS 
usm 


CONTENTS 
THe Karu JorpAN Mena. Lee D. Miller _.__..__._ 207 


ULTRAVIOLET PHOTOGRAPHY AS AN ADJUNCT TO Taxonomy. Clifford 
D. Fertig: (eS IN AN OE ee a 210 


_A New Svusspecies OF EUMEDONIA EUMEDON (LYCAENIDAE) FROM 
.Caucasus,, Yuri P. Nekrutenko)_.2...__ 215 


THE OCCURRENCE OF CHLOROCLYSTIS RECTANGULATA (L.) IN NORTH 
_AMERICA (GEOMETRIDAE). Douglas C. Ferguson _..._. 220 


New Recorps oF LEPIDOPTERA FROM THE UNITED STATES ( ARCTIIDAE, 
GEOMETRIDAE, EPIPLEMIDAE ). Douglas C. Ferguson ________. 222 


POPULATION EXPANSIONS AND Mass MOVEMENTS OF NYMPHALIS CALI- 
FORNICA (NYMPHALIDAE ). Jerry A. Powell __--------_-_- 226 


OBSERVATIONS AND NEw Recorps or Iowa RHoOPALOCERA. Stephen 
Miller) il 0 a 229 


CONFIRMATION OF A DisPUTED FOODPLANT OF PAPILIO GLAUCUS ( Pa- 
PILIONIDAE ). J. Mark Scriber 0s. 935 


THE Brotocy oF C ALLOPHRYS (INCISALIA) FOTIS BAYENSIS (LYCAE- 
NIDAE).. John F. Emmel and Clifford D. Ferris 237 


Tur Type Locaurry FOR Two Motus (PyYRALIDAE, SATURNIIDAE) — 
Couiectep By Lr. W. L. Carpenter, U.S.A., Iv COLORADO, 
1873. F. Martin Brown 0 245 


A PROPOSAL FOR THE UNIFORM TREATMENT OF INFRASUBSPECIFIC 
VARIATION BY LEPIDOPTERISTS. John H. Masters ___....-- 249 


GENERAL NOTES 


Notes on the life history of Eugonobapta nivosaria (Geometridae). 


Lddtence: Ri. Ruperies i Os Re 218 
Cardiac glycosides in Asclepias species. Frank Slansky, Jr. __...-.- 219 
Two name. changes for subtropical American Pieridae. Norman D. Riley _. 228 
Some notes‘on the Sphingidae. Richard B. Dominick __..... 234 
The mature larva of Sphinx vashti (Sphingidae). William H. Howe ____. QA7 


Limenitis lorquini attacking a Glaucous-winged Gull. Robert Michael Pyle 261 


Book REVIEWS ie MUNN N ANEMIC RINDMAMAN ND CM 262 
NOTES (AND! NEw ida OS aOR Ae) TSN Oar a Se er 267 
OBITUARIES I UN i aaa oN SN GAN le et 268 


ee 


ee 


Volume 27 1973 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 


23 February 1973 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


J. F. Gates Crarxe (Washington, D.C.) President 

Harry K. Criencu (Pittsburgh, Penn.) President-elect 
ALEXANDER B. Kiots (New York, N.Y.) Ist Vice President 
C. F. Cowan (Berkhamsted, England) Vice President 

E. G. Munroe (Ottawa, Ontario) Vice President 

S. S. Nicoxay (Virginia Beach, Va.) Treasurer 

LEE D. Mixzer (Sarasota, Fla.) Secretary 


Members at large (three year term) : M. C. Nietsen (Lansing, Mich.) 1974 
A. BLancHARD (Houston, Texas) 1973 D. C. Frercuson (Washington, D.C.) 
R. B. Dominick (McClellanville, S.C.) 1975 

1973 R. O. Kenpatxi (San Antonio, Texas) 
J. P. Donanve (Los Angeles, Calif.) 1973 1975 
J. M. Burns (Cambridge, Mass.) 1974 J. A. Powext (Berkeley, Calif.) 1975 
R. H. Carcasson (Vancouver, B.C.) 1974 


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 $10.00 
Student members—annual dues $5.00 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $15.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia 23455. 


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


by Cyr. F. pos Passos 


Price, postpaid: Society members—$5.00, others—$7.50; uncut, unbound signatures 
available for interleaving and private binding, same prices; hard cover bound, mem- 
bers—$8.00, others—$10.00. Revised lists of the Melitaeinae and Lycaenidae will be 
distributed to purchasers free (separately with paper covered copies and unbound 
signatures, bound in with hard covered copies). 


The Lepidopterists’ Society is a non-profit, scientific organization. The office of 
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520. 
Second class postage paid at Lawrence, Kansas, U.S.A. 66044. 


el 


Se 0 cite ih Di in tiie 


JOURNAL OF 


Tur LEepipopTreERiIsts’ SOCIETY 


Volume 27 | 1973 Number 1 


LIFE HISTORY OF ISOPARCE CUPRESSI (SPHINGIDAE) 


RicHARD B. DoMINICcK : 
The Charleston Museum, Charleston, South Carolina 29401 


References to the life history of Isoparce cupressi (Boisduval), the 
Cypress Sphinx, are scarce in the literature. The only description of the 
larva known to the author is that by Bates (1928) referred to by Hodges 
(1971). The adult moth has been taken with regularity in coastal South 
Carolina at light in the neighborhood of its food plant, Bald Cypress 
(Taxodium distichum (L)). Two main broods occur in this area, the 
spring brood from mid-March through mid-May with a high peak in 
April, and the second brood predominately during August and September. 
A scattering of specimens is also taken during all the summer months. 
Hodges (1971) has given an excellent description of the adult moth, but 
one further distinctive feature should be mentioned. Just to the costal 
side of the heavier broken black dash on the forewing, at about the center 
of the wing surface, is a reddish brown streak running parallel with and 
adjacent to the black dash. This marking, together with the black dash 
(there are often two or three of the latter), is quite conspicuous and 
diagnostic in the fresh specimen. 

On 13 August 1971 at McClellanville, South Carolina a female Isoparce 
cupressi was captured at a mercury vapor light, labelled 2 A’71, and per- 
mitted to lay eggs in captivity. She did so without hesitation, laying ap- 
proximately one hundred eggs, first on paper in a plastic box and later 


< 

Color Plate. Isoparce cupressi (Bdv.) and its foodplant (Taxodium distichum) 
(natural size). 1. Adult 6, caught at light 13 Sept. 1968; 2. 2nd instar larva, 5 days 
old; 3. 3rd instar larva, 11 days old; 4. 3rd instar larva, 10 days old; 5. 4th instar 
larva, 13 days old; 6. 4th instar larva, 14 days old; 7. 5th instar larva, 3 weeks old, 
full grown; 8. pupa, 1 week old, ex larva placed on wild cypress tree; 9. and 10. 5th 
instar larvae, 3 weeks old, full grown; 11. 5th instar larva, eeks old, about to 


pupate. (All larvae are freeze-dried specimens. ) 


2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


on sprigs of cypress introduced for her benefit. Being confined to a small 
area, of course, a natural pattern of egg-laying could not be definitely 
determined; however the eggs were laid singly rather than in clusters, 
and when the fresh cypress was presented she chose to lay singly on the 
plant. The eggs are slightly flattened spheres 1 mm in diameter, grossly 
smooth, microscopically very slightly and evenly roughened, and of a 
pearly green color. The micropyle appears to lie dorsolaterally. The larva 
hatches in seven days, leaving a transparent uneaten eggshell. We killed 
our female after three days. No food was presented as it is a superfluous 
commodity for a moth with rudimentary mouthparts. 


Larval Instars 


Coloration and markings of the larvae through the successive instars 
progress as a model of camouflage on their food plant. The first instar 
larva is green and at one day old is about 3 to 4 mm in length, with a black 
dorsal horn about 3 mm long. The head is round, green and immaculate. 
A dorsal and two lateral stripes, one on each side, extend the length of 
the caterpillar except for the prothoracic segment where they fade toward 
the head. These three stripes are of a slightly darker green. Very thin 
paler bands run transversely over the dorsal surface from one lateral 
stripe to the other, marking off the segments. The result is a tiny larva just 
about the size of one of the pinnae of the young shoots of the cypress, and 
the larval variation of greens blends beautifully with a lengthwise posi- 
tion on one of the leaflets. 

The second instar occurs about four days later and the larva grows to 
about 10 mm in length. The basic body color is a forest green, while the 
head acquires a pair of yellowish-green vertical stripes down each side 
and a similar but thinner pair down the center line. These stripes diverge 
somewhat from the top of the head where they commence at the base of 
a pair of tiny hornlike protuberances. On all segments of the body a 
dorsolateral and a vertrolateral stripe of a yellowish cast develop, these 
stripes becoming slightly constricted at the juncture of the segments and 
just beginning to show signs of segmental angling posteriorly toward 
the median. The first thoracic segment resembles the top of the head in 
tending to define a rather sharp angle dorsally. At this stage the first 
whitish lateral lunules appear, running obliquely posterodorsally from the 
mesothoracic segment to and including the seventh abdominal segment. 
On the prothoracic segment, the lunule is replaced by a horizontal white 
dash which continues posteriorly, fading and connecting the tops of the 

e lunules. On the first thoracic segment it remains a distinctive 
ighout the succeeding instars. The hor, on the eighth ab- 


VOLUME 27, NUMBER 1 g 


dominal segment, has a roughened surface, a touch of red to its pre- 
dominately black color, and is about 4 mm in length. 

The third instar occurs in about another four days. For that matter, 
four or five days proved to be the average for each of the stadia. The 
pattern of the head shows no significant change. The stripes on the body 
remain in the same general pattern, but the longitudinal yellowish stripes 
are now more broken with each segment; the dorsolateral pair assume a 
slightly oblique aspect on each segment, slanting even more toward the 
median posteriorly. At this time a reddish spot appears at the site of the 
spiracle on each segment except for the second and third thoracic seg- 
ments. The second thoracic spiracle shows instead a small black spot 
with one or two tiny white punctae, and that of the metathoracic segment 
tends to be featureless. The junction of the lateral oblique lunules (which 
are now acquiring the yellowish tinge of the other markings) and the 
ventrolateral broken stripes is now forming the shape of an arrowhead 
pointing cephalad on each segment, with the colored spiracular dot at the 
point of the arrow. It is during this third instar that the forest green of 
the dorsal surface begins to change to a red-brown (later darkening to a 
purple-brown) dorsal stripe that is a conspicuous character of the larva 
from this period on. It characterizes a camouflage consistent with the 
growth stage when the larva travels by way of the brown twigs of the 
cypress, and this stripe down the back with the oblique yellow-white 
markings on the green base color assumes the pattern of light through 
the foliage. This dark dorsal stripe begins on about the mesothorax and 
extends posteriorly onto the horn. The larva is also just beginning to show 
signs of developing the tiny yellow punctae that will appear more 
markedly later on, adding to the broken-light pattern. These spots first 
appear anteriorly. At the end of this instar the larva is about 18 mm long, 
and the posterior horn recurves slightly. The thoracic legs also gradually 
change color from green to reddish brown as the larva grows. 

The fourth instar begins roughly two weeks after hatching. The head 
acquires a pair of black lines diverging downward from the upper pointed 
“horns” and ending about two-thirds of the way down the face. All the 
prolegs, including the anal, now acquire the red-brown color of the 
spiracles. The metathoracic spiracle may or may not have a reddish 
punctum; if so, it remains less conspicuous than the others. The small 
yellowish punctate excrescences now appear on all body surfaces. The 


yellowish ventrolateral segmental markings are by now wel! broken up by 
segments, and tend to disappear from the second theracic to a variable 
number of segments between the mesothoracic and anterior abdominal 


segments. The dorsal markings are now more purplish-brown and are 


4 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


punctuated by rows of minute whitish dots, while the spiracular color 
remains reddish as does the base and dorsum of the horn. The horn is 
now beset with spicules. The dorsal brown stripe takes on a newly de- 
veloping shape. Beginning with the mesothorax the purplish brown of 
each segment has the shape of a half hourglass with the thin end behind. 
This is laterally edged by a touch of black and in turn with the remains 
of the broken yellowish dorsolateral stripe. The first thoracic segment 
follows the general pattern but presents a different appearance because 
the dorsal brown, having disappeared here, is replaced by a thin black 
mark flanked by the yellow and in turn by a pair of new black spots, 
flanked yet again by the yellow line first seen in the second instar which 
is in reality the lateral lunule pointing anteroposteriorly. By the end of 
the fourth instar the larva has reached a length of about 3 cm or slightly 
less. One freeze-dried specimen (not illustrated) just shedding into the 
fifth instar is marked as being 16 days old. 

The fifth instar larva continues rapid growth to a length of about 5 
cm, with the posterior horn measuring slightly under 1 cm. The abdominal 
spiracles and the dorsolateral surface of the prolegs become partially dark 
brown to blackish; the second thoracic spiracle tends to maintain its dark 
color with white punctae; and the rest of the spiracular coloration as- 
sumes some black along with the red-brown, that of the metathorax 
tending to remain relatively modest. The yellow punctate excrescences 
have become more numerous, following to some extent the direction of the 
yellow lines and lunules. These latter acquire traces of brown or black 
edgings. Posterior to the eighth abdominal segment (i.e., abaft the 
horn), the lateral spiracular brown ceases and the yellow lateral lunule 
straightens out to run posteriorly down and across the anal plate where 
it meets its mate. Some of these lines and lunules may be more white 
than yellow. The anal plate is greyish green dotted with tiny black spots. 
All in all we have an extremely handsome caterpillar, camouflaged in such 
a way as to be very difficult to detect on a wild cypress. 

According to Bates (1928) the larvae feed at night. I was able 
neither to confirm nor deny this observation since this brood from 2 A’71 
was reared in confined and therefore artificial quarters and demonstrated 
no meal-time preferences. 


Pupating Habits 
At the end of approximately three weeks the larvae turn a purplish 
brown color all over and begin to wander in preparation for pupation. 
This color suits them well for travelling more openly on the larger 
branches and trunk of the tree. Their pupating habits, being for all 
practical purposes unknown, were of especial interest, and I endeavored 


VOLUME 27, NUMBER 1 5 


to simulate natural conditions as well as possible with some interesting, 
and at times perplexing and amusing results. 

Three sets of conditions seemed indicated in order to discover the ac- 
tions of a cupressi larva seeking to pupate. First was the simulation of a 
cypress tree standing in the water, described herein as the “water cage.” 
Second, the simulation of a cypress growing on dry land, described as the 
“ground cage. Third, taking a larva directly out to a wild cypress pond 
and hoping for the best. 

The “water cage” was made by cutting a cypress branch about two 
feet long and 2’2 inches in diameter and suspending it by a thread from 
the top of a cage with its bottom end submerged in a can of water. Such 
a branch is quite smooth, and in order to create the likeness of a crevice 
or two, a couple of branching twigs were tied to the side of the limb. 
A smear of vaseline was applied to the suspending thread to discourage 
the caterpillar from climbing off the wood. The “ground cage” was made 
by filling a large coffee can with fine sand covered with a couple of 
inches of peat moss, and a similar-sized cypress branch with twigs was 
stuck upright into the center of the can. Vaseline was smeared around 
the edges of the can to discourage wandering out of the allotted area. 
In all, seven pupating cupressi larvae were used, six in the cages and one 
reserved for the wild. 

Larva #1, on 12 September, was placed on the bark of the “water cage.” 
It climbed up and down the eighteen inches or so of available dry trunk 
several times, being turned at the top by the vaseline on the thread and 
at the bottom by the water. It was not averse, however, to dunking its 
head quite thoroughly into the water before turning back. After a while 
it began to settle into one of the slight crevices provided by a twig, but 
after an hour or so of trying to snuggle in was evidently not satisfied and 
wandered off again, whereupon it fell into the water and drowned. 

Larva #2 on the same date was put on the cypress in the “ground cage.” 
This one was observed for twenty minutes or so crawling all about the 
“tree trunk” and then curled up on the top of the stump and to all ap- 
pearances took a nap. At this point I was called away for about ten 
minutes, and upon my return there was no sign of the larva. There was 
no possible route of escape from the cage, and subsequent investigation 
proved that it had descended the “tree” and burrowed underground where 
it proceeded to pupate. 


The next day, 13 September, three more larvae were ready to pupate. 

Larva #3 was put in the “water cage” and it wasted very little time indeed 
e e . e a “% > > | | 7 »* 

before either crawling or falling into the water when I had my head 


. ° . E 7 ~ | +7 ] ’ .Ofar— 
turned for just a moment. This time rescue was p it, and the cater 


6 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


‘ ’ a: Ws 4 
if 4, ‘ £ 
F i % i, id 
A ae, fee ys Ls 
7 . , 


Fig. 1. The author points to spot where larva #7 pupated. Photo by C. R. 
Edwards, 26 Sept. 1971. 


pillar was put into the “ground cage,” where it immediately crawled 
down the trunk and burrowed straight into the peat moss. Larvae #s 4 
and 5 were then put directly into the “ground cage,’ and neither spent 
more than fifteen or twenty minutes before burrowing underground. The 
sand of course in this cage was packed tightly enough to support the 
cypress limb firmly, and the peat was fairly loose on top, but an average 
of only thirty seconds was required for each larva to disappear com- 
pletely. 

Since the “water cage” seemed a fatal hazard, I then compromised 
and replaced the water with sand and peat, inserting the trunk in a man- 
ner similar to the “ground cage,” but in this case greasing the bottom two 
inches of exposed trunk with vaseline, hoping to discourage headlong 
downward progress but providing a safe landing just in case. Caterpillar 
#6 was then put on the trunk and was observed to head downward, hardly 
hesitating at the greased bottom. Crawling right over the vaseline, it 
too disappeared rapidly underground. One week later I carefully spooned 
out the peat and sand and found #6 had pupated in a slightly packed 
chamber about three inches below the surface, in the sand. There was 
no evident sign of silk spinning. 

Larva #7 was ready to pupate on 15 September. This was the: one 


VOLUME 27, NUMBER 1 


~| 


reserved for the wild, and was taken to a nearby cypress pond. A smallish 
cypress about a foot or two in diameter that was standing in the water 
having been selected, the larva was placed on the trunk about breast high 
and left to its own devices. A smallish tree such as this has relatively 
smooth bark, and this particular tree was chosen for a relative paucity of 
crannies, moss or other hiding places. This larva crawled all about and 
twice tried to dig into some small patches of moss, but it was evident 
they did not offer the protection it desired. After an hour or so of being 
left to explore this tree, it was transferred to a neighboring tree which 
offered greater chances of concealment by way of crevices and heavier 
patches of moss. It was interesting to note a determined attempt on the 
part of the larva to burrow into one mossy niche which was almost but 
not quite deep enough to hide it completely. The power of forcing its 
way into tight corners and under quite thickly rooted moss, and its evident 
strong desire to burrow, were noteworthy. After a half hour's attempt 
the caterpillar gave up and went off in search of a better spot. It found 
one to its satisfaction a short distance away in a patch of moss thick and 
deep enough to permit its burying itself completely out of sight. I 
marked the spot and returned a week later, and then carefully peeling 
back the moss, found the pupa deep in a mossy cell, smoothly shaped but 
again with little or no appreciable sign of spinning (Fig. 1). 

These observations would seem to indicate a strong burrowing instinct 
on the part of the pupating cupressi larvae, and would explain the great 
difficulty of successfully searching for pupae in the wild, but will per- 
haps give an indication of where and how to look. The two cases of larvae 
falling into the water reported here can possibly be best explained by the 
artificial conditions imposed with insufficient room for manoeuver. 


Hodges (1971) mentioned two cupressi pupae found by this author the previous 
year. These were in fact found in a remarkable and unique tract of very old virgin 
cypress, of the type known to the lumber industry as “Black Cypress” and noted for 
its fine quality. It is a tract of swampland known as Four Holes Swamp, consisting 
of less than 3000 acres of uncut timber near Summerville, South Carolina. It is one 
of the last two tracts of such trees left in the Southeast, and an urgent conservation 
measure is under way in an attempt to save this remnant of the past where the trees 
are several hundred years old and some measure over ten feet in diameter above the 
massive buttress. Great shards of bark hang loosely from the towering trunks, and a 
day’s canoeing through this magnificent primeval swampland forest is an experience to 
stir the blood. Brown water snakes and alligators abound, or an otter may be seen 


gazing with inquisitive whiskers aslant before gliding silently under the dark waters. 
A full day of examination of every cypress within reach produced the two pupae, 
mentioned by Hodges, tucked under the protection of loosely hanging bark. As with 
those reared, neither pupa of that day showed any indication of having spun. 
? i |) | Lar vara °O.. 
Of the reared brood of larvae ex 2 A771, a goodly number were pre 


served in the various instars by the technique of freeze-drying, some of 


8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


which are shown in the illustration. Two of the six pupae (remember 
one larva drowned) were likewise preserved whole, and four adult 
cupressi moths emerged under indoor conditions in early October. The 
pupal cases of these latter were unearthed from the sand in the cages, 
and were all found at approximately the same depth and in the same type 
of cells as mentioned above. 

The pupa is dark brown, about 3.5 cm in length and on the moderately 
slender side. It lacks a free tongue case. Its surface is punctate on all 
sides, though ventrally the abdominal segments are smooth posteriorly. 
It should be noted in passing that while our pupae agree in description 
with that given by Bates (1928) except for the placement of the punctae, 
none of our reared larvae reached the length quoted by that authority of 
65 mm, though the four emerging adults were of normal size. 


ACKNOWLEDGMENT 


I wish to extend my thanks to Dr. Ronald W. Hodges, who has read the 
manuscript, offered helpful suggestions, and shuffled many commas. 


LITERATURE CITED 


Bates, M. 1928. Notes on the Cypress Sphinx (Isoparce cupressi). Florida 
Entomol. 12: 20-21. 

Hopnces, R. W. in Dominick, R. B. et al., 1971, The Moths of America North of 
Mexico, fasc. 21, Sphingoidea, p. 48. 


BEHAVIORAL ADAPTATIONS OF CRYPTIC MOTHS. VI. FURTHER 
EXPERIMENTAL STUDIES ON BARK-LIKE SPECIES 


THEODORE D. SARGENT 
Department of Zoology, University of Massachusetts, Amherst, Massachusetts 01002 


Prior studies have demonstrated that a number of bark-like moths will 
select appropriate backgrounds in various experimental apparatuses which 
provide a choice of backgrounds differing in reflectance (Kettlewell, 
1955; Sargent, 1966; Sargent & Keiper, 1969). Several experiments indi- 
cate that these selections are based on innate preferences of the moths 
for certain background reflectances (Sargent, 1968, 1969a, b). 

The present paper summarizes additional experiments which shed light 
on (1) some factors promoting “errors” in the background selections of 


VOLUME 27, NUMBER 1 9 


Fig. 1. The experimental apparatus used in the present studies. 


moths in experimental situations, and (2) the nature of the background 
selection process itself. The experiments were carried out during the 
summers of 1970 and 1971 in Leverett, Franklin County, Massachusetts. 


General Methods and Materials 


The basic apparatus in all of the experiments consisted of a cylinder 
(19 in. high x 44 in. circumference ) made up of blotting paper sections, 
set into a plywood box (19 in. high X 15 in. square), and covered with a 
pane of window glass (Fig. 1). The blotting paper sections were painted 
black, gray, or white, and so provided a choice of backgrounds differing 
in reflectance. (The percentage-reflectance values of the backgrounds, 
as measured with a General Electric recording spectrophotometer against 
a white standard of pressed BaSO;, were: black 4.30, gray 39.50, and 
white 85.61.) The apparatus was placed in a wooded area, and moths 
were introduced into the cylinder at night by sliding the glass top to one 
side. Background selections of the moths were noted shortly after dawn 
on the following morning. 


The Experiments and Results 


EXPERIMENT 1. Early observations suggested that some species which 
are known to select appropriate backgrounds in nature (e.g. Catocala 
relicta Walker (Noctuidae) which prefers to rest on white birch trees; 
Keiper, 1968; Sargent & Keiper, 1969) did not exhibit background prefer- 
ences in the present experimental apparatus. It was noted, however, that 


the background selections of these moths might vary according to their 
position in the apparatus, as moths resting at the very top of the cylinder, 
i.e. immediately adjacent to the pane of glass, were very often over an 
inappropriate background. Therefore an experiment was conducted, 


using reared Catocala relicta (Sargent, 1972), in which the precise posi- 


10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TasLe 1. Experiment 1: Background selections of Catocala relicta. 


Number of Moths 


Position 

within White Black 
Cylinder Background Background Pt 
At Top a 8 > 090 
Below Top 10 0 < 0.01 


* Based on chi-square tests of goodness-of-fit to a 1:1 distribution. 


tions of the moths, as well as the background selections, were recorded. 
The experimental cylinder for this experiment consisted of alternating 
black and white sections (each 11 x 19 in.). 

The results of this experiment (Table 1) clearly revealed that moths 
resting at the top of the cylinder did not exhibit a background preference, 
while moths resting at lower levels preferred the white backgrounds. 
It would appear that some behavior, perhaps a phototactic or geotactic 
escape response, interfered with the background selections of certain 
moths in this experimental situation. 

EXPERIMENT 2. Another factor which seemed to influence background 
selections in the experimental apparatus was the number of moths tested 
on any given night. Accordingly, a simple experiment was conducted 
with Phigalia titea (Cramer) (Geometridae), a species known to prefer 
pale backgrounds (Sargent, 1969b). In this experiment, tests of back- 
ground selections were carried out using samples of from 1-30 individuals 
in the experimental apparatus. The moths were collected at lights, and 
the experimental cylinder in this case consisted of a white, gray, and 
black section (each 14.7 x 19 in.). 

The results of this experiment (Table 2) showed that there was a 
critical sample size of about 20 individuals, below which a background 
preference was clearly exhibited, and above which no background prefer- 
ence was shown. This finding suggests that some behavioral interaction, 


Taste 2. Experiment 2: Background selections of Phigalia titea. 


Number of Moths 
Sn, eee eee 


Sample No. White Gray Black 
Size Tests Background Background Background Pe 
1-10 9 18 q 8 < 0.05 
11-20 3 21 10 7 < 0.02 
21-30 z; 19 19 IES) > 0.70 


* Based on chi-square tests of goodness-of-fit to a 1:1:1 distribution. 


VoLUME 27, NUMBER 1 ek 


TaBLE 3. Experiment 3: Background selections of Cosymbia pendulinaria. 


Number of Moths 


Experimental White Black 
Condition Background Background ee 
Without Acetate Cylinder 55 6 < 0.001 
With Acetate Cylinder 30 2 < 0.001 


* Based on chi-square tests of goodness-ot-fit to a 1:1 distribution. 


presumably one resulting from the mutual intolerance of individuals and 
functioning to disperse the moths, interfered with background selections 
at high densities in this experimental apparatus. 

EXPERIMENT 3. Some question has been raised about the ability of 
bark-like moths to select appropriate backgrounds from a distance, ie. 
when not in direct physical contact with a substrate (e.g. Kettlewell, 1955; 
Ford, 1964). If contact were required, then it might be argued that 
thermal, rather than visual, cues were playing the important role in 
background selections. In that event, moths would be detecting and 
responding to small, surface temperature differences associated with 
backgrounds of different reflectances. 

In order to test this possibility, the experimental apparatus was equipped 
with a cyclinder of clear cellulose acetate which could be set within the 
cylinder of blotting paper sections, and which separated experimental 
moths from the painted backgrounds by approximately two inches. The 
species tested was Cosymbia pendulinaria Gueneé (Geometridae ), which 
prefers white backgrounds (Sargent, 1968). The painted cylinder in 
this experiment consisted of alternating black and white sections (each 
11 x 19 in.), and moths were tested for background preferences when 
the acetate cylinder was absent (controls) and present (experimentals ). 
It was assumed that the separation of moths and backgrounds by the 
acetate cylinder was sufficient to prevent detection of any surface tem- 
perature differences. 

The results of this experiment (Table 3) showed that direct physical 
contact with the backgrounds was not required for the moths to exhibit a 
background reflectance preference. 


DISCUSSION 


The results of Experiments 1 and 2 indicate that considerable care 
must be taken in the design and execution of experiments which are 
intended as tests of background preferences in cryptic moths. Certain 
behaviors, particularly escape and avoidance responses, may be elicited 


12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in some individuals and under some circumstances in an experimental 
apparatus, and these behaviors may interfere with background prefer- 
ences. Thus, a failure to detect background preferences may result from 
inadequacies in an experimental test, rather than from inabilities of the 
moths themselves. 

The results of Experiment 3 tend to corroborate the prevailing view 
that background reflectance preferences of bark-like moths are based 
on responses to visual stimuli. This conclusion is strengthened by field 
observations of released individuals of Cosymbia pendulinaria (and 
Catocala relicta; Sargent & Keiper, 1969). In these field tests, the erratic 
escape flight of certain individuals would become directed after some 
seconds, and this directed flight (sometimes up to 50 feet in length) 
usually led to an appropriate background (white birch tree). This 
observation, the results of Experiment 3, and numerous other experi- 
mental results (Sargent, 1968, 1969a, b) fail to support the reflectance- 
matching mechanism proposed by Kettlewell (1955) and Ford (1964) to 
explain the selection of appropriate backgrounds by bark-like moths. 
On the contrary, all of the evidence to date supports the view that these 
background selections are genetically fixed, or innate, responses. 


ACKNOWLEDGMENTS 


I thank my wife, Katherine, for the drawing of the experimental 
apparatus, and Dr. F. J. Francis of the Department of Food Science and 
Technology, University of Massachusetts, for the determinations of the 
background reflectances. 


LITERATURE CITED 


Forp, E. B. 1964. Ecological Genetics. Methuen, London. p. 256. 
Kerper, R. R. 1968. Field studies of Catocala behavior. J. Res. Lepid. 7: 113-121. 
KETTLEWELL, H. B. D. 1955. Recognition of appropriate backgrounds by the pale 
and black phases of Lepidoptera. Nature 175: 943-944. 
SARGENT, T. D. 1966. Background selections of geometrid and noctuid moths. 
Science 154: 1674-1675. 
1968. Cryptic moths: effects on background selections of painting the 
circumocular scales. Science 159: 100-101. 
1969a. Behavioral adaptations of cryptic moths. II. Experimental studies 
on bark-like species. J. N. Y. Entomol. Soc. 77: 75-79. 
1969b. Background selections of the pale and melanic forms of the 
cryptic moth, Phigalia titea (Cramer). Nature 222: 585-586. 
. 1972. Studies on the Catocala (Noctuidae) of southern New England. 
II. Mating results with C. relicta Walker. J. Lepid. Soc. 26: 94-104. 
& R. R. Kerper. 1969. Behavioral adaptations of cryptic moths. I. Pre- 
liminary studies on bark-like species. J. Lepid. Soc. 23: 1-9. 


VoLUME 27, NUMBER 1 13 


NOTES ON LIFE HISTORIES AND HABITS OF SOME 
WESTERN THECLINAE 


Ki. J. N&wcoMer 
1738 N.E. Naomi Place, Seattle, Washington 98115 


Realizing that very little was known about life histories of several 
Theclinae, I made some studies of some of them. It was not difficult to 
get adults to oviposit in small (1.50 X 3.75 in.) “season salt” bottles which 
have plastic shaker tops. Eggs would be deposited on a supposed food- 
plant or on the sides of the bottles. Larvae were reared 'in metal pill 
boxes with fresh food supplied usually twice daily. 

Callipsyche behrii (Edwards). This species occurs throughout most of 
the area from the Rocky Mountains to the Pacific Coast. It has been 
recorded as feeding on Lupinus ssp., Astragalus ssp., and Lotus ssp. 
(Brown, 1957; Jones, 1951; Comstock, 1927). But in central Washington 
the foodplant is primarily something quite different, antelope bush 
(Purshia tridentata (Pursh) DC, which is in the Rosaceae. This plant 
is quite common in the arid transition zone and it occurs from British 
Columbia and Montana to central California and New Mexico. The eggs 
are deposited on the stems and occasionally on the leaves of this shrub. 


Egg: typical theclid form, diameter 0.8 mm, very light greenish, does not appear 
to be denticulated but is covered with small nodules. First instar: 2 mm, head 
black, body pinkish with many smali dark dots and some setae. Second instar: 
3.5-4 mm, light green with many dark green dots and small setae. Later instars: 
light green with diagonal lighter green markings laterally on each segment and a 
lateral light stripe. This coloring blends very well with the small leaves of the 
foodplant. Pupa: not seen. 


Incisalia fotis (Strecker) is also found throughout the West but rather 
localized. It flies early in the spring and hence may be missed by some 
collectors. The foodplant is sedum. 


Egg: diameter 0.6 mm, height 0.4 mm, very finely reticulated, light blue green. 
First instar: light tan with double pink stripe dorsally and single pink stripe 
laterally; two tubercles on each segment lateral of the pink stripes, each with two 
setae: head dark brown. Mature larva: light greenish, pronounced double pink 
stripe dorsally, lateral pink stripes and a ventral white stripe; dark red diagonal 
“hash” marks on each segment; hairs numerous, golden brown; head, light brown, 
sunken into the first segment. Pupa: length 10.5 mm, width 5.25 mm, purplish 
brown, dorsum lighter with many fine darker reddish lines; two rows of dark spots 
on each side of dorsum; dorsal line dark, many short, light brown hairs scattered 
over body except on wings. 


Incisalia eryphon (Boisduval) is well distributed over the area west of 
the Rocky Mountains. In May 1964, I was able to get 30 or 40 eggs of 


14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


this species but have not succeeded in getting any since that time. 
Holland (1930) stated that the early stages were unknown. He reported 
that Scudder had “elaborately described” the early stages of the related 
niphon, and that the caterpillars fed on pine. Scudder (1889) did not 
elaborate on what part of the pine tree the larvae fed upon. The only 
published information I could find on eryphon was a brief paper by Hardy 
(1959). Hardy reared larvae on lodgepole pine (Pinus contorta Loudon ) 
and found that they fed “in the soft and succulent base” of the needles. 
He described the various stages. Llewelyn Jones (1951) also reported 
that the larvae fed on the western white pine (P. monticola Douglas ). 

When the eggs started to hatch, I assumed the larvae would feed at the 
base of the needles of the yellow pine (P. ponderosa Lawson) that I had 
them on, but they died without feeding. So I tried the staminate catkins, 
which were ripening in May, and found that the larvae fed on them 
readily. They had some difficulty in boring into the catkins, so I scraped 
away a bit of the outer coating, after which they had no difficulty and 
soon had disappeared within. As the larvae grew, they were often 
partially outside of the catkins, but usually had the head and fore part 
of the body buried. One would think that, with such food, their color 
would be different than if they were feeding on the needles, but this was 
not the case. In the first instar they were honey colored and I would have 
expected them to remain that color in the later stages. But when they 
reached the third instar they became bright green. They were usually 
well powdered with pollen, which constituted a very rich food for them. 
Under laboratory conditions eggs hatched in 5 or 6 days. Because of the 
manner of feeding, it was not feasible to keep track of the various instars, 
but the total feeding time was about two weeks. 


Egg: light green, the usual lycaenid form, diameter 1 mm, height 0.6 mm. 
Newly hatched Jarva: length 1.5 mm; head brown, body honey colored, a whitish 
line on each side of the dorsum with very short setae on each segment; anal plate 
small, round. Third (?) instar: length 11 mm; light green with a lighter green 
line on each side of the dorsum; many very small dark points on the abdomen from 
which grow short, light-brown hairs of various lengths; cervical shield greenish 
brown; head light brown; anal plate not evident. Full-grown larva: length 18 mm; 
ground color green; head and thoracic shield light brown; a latero-dorsal white 
stripe on each side of dorsum. Pupa: length 7.5-8 mm, width 3.5-4 mm; at first 
green, turning brown; dark area on each segment dorsally and a smaller round, dark 
spot subdorsally; a few irregular dark brown spots on wings; very fine setae scattered 
over body but not on wings. 


Strymon saepium ( Boisduval) occurs from the Rocky Mountains to the 
Pacific Coast and from southern British Columbia to southern California. 
In this region there are some 50 species of Ceanothus which is evidently 
the preferred foodplant. It undoubtedly feeds on more than one species, 


VOLUME 27, NUMBER 1 15 


possibly even in one area, but its preferences among the many species are 
not known. 

Eggs were deposited on the stems and leaves, mostly on the under sur- 
face, of Ceanothus velutinus Douglas, and a few on C. sanguineus Pursh.; 
and on oak, in this case between the buds. Larvae would not feed on oak 
but fed readily on both species of Ceanothus, starting on the under sur- 
face. The upper surface of C. velutinus is somewhat sticky and feeding 
there is difficult. The larger larvae ate holes through the leaves and they 
also fed on the flower buds. 


Eggs: the usual flattened shape, reticulated over the entire surface except the 
base; area about the micropyle depressed but also reticulated; light greenish; diam- 
eter 0.8 mm. Newly hatched larva: length 1 mm, sluglike; cream colored with many 
long hairs; a darker mid stripe and a lateral stripe; head light brown. Second instar: 
length 2.5 mm; light green, latero-dorsal ridges whitish; lateral ridge whitish; head 
black. Third instar: length 4-5 mm; same color and stripes; dorsal area dark 
green; many short spines all over body. Fourth instar: length 10 mm, width 3.5 mm; 
uniform light green except for narrow dorso-lateral whitish line. Full-grown larva: 
length 15 mm, width 5 mm; light green, two whitish dorsal stripes separated by a 
narrow green area; lateral stripe whitish; body covered with short golden hairs; 
ventral side lighter green; head small, dark brown, posterior part lighter brown. 
Pupa: length 11 mm, width 5 mm; chestnut brown with many irregular darker 
flecks on dorsum; head and wing covers with few hairs, many light brown ones 
dorsally. 


LITERATURE CITED 


Brown, F. Martin. 1957. Colorado Butterflies. Proceedings Denver Museum of 
Natural History. Nos. 3-7. Denver, Colo. 

Comstock, J. A. 1927. Butterflies of California. Los Angeles. 

Harpy, A. 1959. On the life history of Incisalia eryphon (Lycaenidae) on south- 
ern Vancouver Island. J. Lepid. Soc. 13(2): 70. 

Hoxuanp, W. J. 1930. The Butterfly Book. Doubleday, New York. 

Jones, J. R. J. LuEwettyn. 1951. An Annotated Check List of the Macrolepidoptera 
of British Columbia. The Entomological Society of British Columbia. 

ScuppER, S. H. 1889. The Butterflies of the Eastern United States and Canada. 
Cambridge. 


PRELIMINARY REPORT ON COMMUNAL RESTING OF 
SMYRNA KARWINSKII ADULTS (NYMPHALIDAE ) 


Since 1962 we have observed adults of Smyrna karwinskii (Geyer), resting in 
groups of ten individuals to more than 100, at all times of the day, either in small 
caves on a lava wall, or under concrete slabs covering alleys between some cabins 
on Cerro Verde. Cerro Verde, a mountain ca. 2000 m. eley., overlooks Izalco 
Volcano, about 50 km. WNW of San Salvador, capital of El Salvador. This phe- 
nomenon has been observed at different times of the year: March-April, August, and 
November-—January, during both the dry and rainy season. 


16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


It had been our assumption that the groups were drinking filtration water, but 
this year we had a chance on 31 March to observe five groups more closely: one of 
about 15 individuals, one of about 20, two of about 50 and one of more than 100. 
All were in places protected against the wind. None of the individuals had the 
proboscis uncoiled, and there was no moisture present, yet the groups stayed there 
from the time first detected, 1015, to the time we left, 1700. From time to time 
individuals would fly away, and individuals would join the group. Males and females 
were mingled in each group, but no sexual activity was noticed. 

All groups followed a certain pattern: a nucleus of about eight individuals had 
the heads together, so that the antennae, upraised, formed a tight fascia. Then a 
row of individuals formed a tight circle around this nucleus, with the antennae 
touching the wings of the inner group. In this way row after row were formed. 
Some rows did not form a complete circle, and eventually there was a line of four 
to five individuals breaking the regularity of the pattern, but they were encircled 
by the next row. All heads were pointing inward. The groups looked very much 
like dried moss growths. 

When at the resting places the butterflies are very tame, so that specimens can be 
captured by hand from any place in the group, without disturbing the rest. If 
handled roughly the whole group will take flight, fully alert, and disperse among 
neighboring trees and rocks. In the surrounding woods individuals were observed 
while feeding on sap from wounds caused by other insects or woodpeckers high up 
in tree trunks. 

We have not investigated yet if the individuals keep to their own groups after their 
occasional flights, or if they exchange locations with others; nor have we investigated 
if this species has acquired the behavior of forming groups just to protect themselves 
from the wind or predation. This behavior has not been observed in Smyrna 
blomfildia datis Fruhstorfer, a closely related species abundant in the lowerlands. 

We intend to present the full results of our investigation on this phenomenon in a 
future article. 


ALBERTO MuysHonpT, 101 Ave. N., 322, Lomas Verdes, San Salvador, El Salvador. 


BIRD PREDATION ON PAPILIO POLYXENES F. (PAPILIONIDAE) 


Birds have been shown to exert substantial predation pressure on many butterfly 
species (Carpenter 1940, Entomol. Mon. Mag. 76: 224-229: Rawson & Bellinger 
1953, Lepid. News 7: 27; Betts 1956, Entomol. Mon. Mag. 92: 68-71; Gibb 1958, 
J. Anim. Ecol. 27: 375-396). However, a shortage of field observations exist in the 
literature dealing with bird predation on larvae and adult butterflies. 

During the summer of 1971, while carrying out field studies in Ithaca, New York, 
on the eastern black swallowtail (Papilio polyxenes), several observations on bird 
predation were made. On 3 June, a female butterfly was taken by a _ bluejay 
(Cyanocitta cristata LL.) as she was ovipositing on carrot (Daucus carota L.). In 
the same area on 1 July, another bluejay was seen attacking a 5th instar larva on a 
carrot plant. Near Homer, New York, on 31 July, a male swallowtail was taken on 
the wing by a catbird (Dumetella carolinensis L.) just after the butterfly flew 
from the common burdock (Arctium minus (Hill) Bernh.) flower upon which it had 


nee teeding 
| >) | reeqaing. 


‘es M. Erickson, Department of Entomology, Cornell University, Ithaca, New 


VOLUME 27, NUMBER 1 1G 


HABITAT SELECTION AND POPULATION STRUCTURE IN 
PLEBEJUS SAEPIOLUS BOISDUVAL (LYCAENIDAE) 


MARGARET A. SHARP 
Missouri Botanical Garden, St. Louis, Missouri 63110 


AND 


Davin R. Parks 
Department of Physics, Stanford University, Stanford, California 94305 


The importance of resource distribution has been demonstrated for the 
survival of both the larvae and adults of butterflies (Dethier, 1959; Gil- 
bert, 1971; Singer, 1971). Nearly all butterfly larvae feed on the leaves 
and flowers of angiosperms, and most temperate zone adults depend on 
nectar for food. Furthermore, the relationships of butterflies and their 
larval food-plants may be quite specific and complex (Breedlove & 
Ehrlich, 1968, 1972; Downey & Fuller, 1961; Ehrlich & Raven, 1964). It 
is therefore reasonable to suppose that the distribution of specific plant 
resources may have great influence on the habitat selection of the mobile 
adult butterflies. 

During the summer of 1971, a study was made of the structure of a 
population of the lycaenid butterfly Plebejus saepiolus Boisduval and the 
distribution of its resources in a subalpine meadow in Gunnison Co., 
Colorado. The females of this species oviposit singly on the flowers and 
developing fruits of the alsike clover, Trifolium hybridum L., and prob- 
ably also on other species of Trifolium. It is noteworthy that Trifolium 
was found to be by far the most important adult nectar source for P. 
saepiolus. 

The objectives of the study were to investigate how P. saepiolus 
distribute themselves with respect to the Trifolium resource and to dis- 
cover how frequently individuals move within and between areas of 
favorable habitat. A capture-recapture technique involving a number ot 
discrete areas was chosen in order to obtain this information. 

The study site was a gently sloping meadow at 2708 m. elevation, just 
east of the Crested Butte, Colorado, Town Cemetery. Located in a valley, 
the site was not noticeably affected by any constant prevailing winds. 
The vegetation of the meadow was dominated by Artemisia tridentata 
Nutt. and grasses, with other plant associations occurring locally, particu- 
larly in wetter regions. The distribution of Trifolium was mapped over 
the site, and on this basis six 30 X 60 meter areas, designated “a” through 
“f” were selected for the study (Fig. 1). Three of these (a, e, and f) 


18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Plebejus saepiolus study areas. 


were in relatively wet regions of high Trifolium density, while the others 
(b,c, and d) were drier and contained little or no Trifolium. 

On each day of the study, two people spent ten minutes in each of the 
six areas collecting all P. saepiolus found. At the end of ten minutes, all 
individual butterflies were sexed, marked with an individual number 
(except for recaptures for which the number was recorded ), and released 
from the center of the area. Less than 1% of the insects were unable to 
‘ly when released. The marks were not conspicuous and thus were un- 


VOLUME 27, NUMBER | 19 


TasBLeE 1. Total captures of Plebejus saepiolus in study areas (1971). 


Study Areas 


Date* a b Cc d e if 
6/24 10 0 iL i 17/ 23 
6/25 15 3 0 2 2, 24 
6/26 12 1 I 3 22, 28 
6/27 12 0 0 IL 22) 19 
6/28 1) 2 2 1 28 28 
6/29 10 0 0 0 19 20 
6/30 15 i! 0 ll 2, 21 
UW = 15 0 0 0 26 20 
T/4 10 0 0 0 20 w) 
Mean 12.33 78 44 1.67 2.2.00 22.78 
Variance 4.75 1.19 D3 5.00 11.25 11.19 
Std. Dev. 2.18 1.09 Wo DDA! B85) BoD 
Std. Dev. of Mean se 36 24 WD LIP TL Je 


* 7/1 and 7/5 were omitted due to cloudy weather conditions which caused unusually low 
butterfly captures. 


likely to affect either the survival of marked individuals or their prob- 
ability of being captured. 


RESULTS 


Before considering in detail the results of this experiment, the sex ratio 
of the captured butterflies should be examined. The female to male 
ratio was 0.19 whereas the ratio for laboratory-reared butterflies of various 
species is normally about 1.0 (Brussard & Ehrlich, 1970). The hypothesis 
that males are much more likely to be captured is supported by the recap- 
ture data in which 13% of the males marked were recaptured at least once, 
while only 4% of the females were recaptured. Since the two sexes are 
quite similar in appearance, the disparity should be due largely to be- 
havioral differences, including greater flight activity by males. The recap- 
ture data therefore refer principally to males; females are expected to be 
more sedentary than the data for males would suggest. If this were true, 
we might expect males to be found more often in less favorable areas 
because of their greater mobility. The sex ratio for areas a, e, and f 
(0.20) versus that for b, c, and d (0.12) support this, but the data are 
based upon too few captures to be relied upon. 

Table 1 presents information on total captures of P. saepiolus in the 
study areas. The difference between the numbers captured in the Tri- 
folium areas (a, e, and f) and the non-Trifolium areas (b,c, end..d).is 
highly significant (p < 001; modified T-test). The scarcity of captures 
in non-Trifolium areas indicates that individuals are seldom found more 


20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Area of Recapture 
a b C d e { - Por 


Area of 
Capture 


Fig. 2. Matrix of capture-recapture data arranged by location. 


than a few meters from Trifolium. This is emphasized by the low yield 
from area d, whose east and west edges follow the borders of Trifolium 
regions containing many Plebejus saepiolus. The data strongly suggest 
that habitat selection involving the oviposition plant is occurring for P. 
saepiolus. 

One valuable feature of mark-release-recapture techniques is that they 
provide information about the movements of individuals. Fig. 2 presents 
in matrix form the locations of all recapture events; entries on the main 
diagonal represent recaptures in the same areas as the original capture 
and off-diagonal entries represent transfer recaptures. It is clear that 
most recaptures occurred in the original capture area and that most of 
the transfers were between areas e and f which are separated by 60 m. 
of favorable habitat. The ratio of transfers to same-area recaptures is 
0.27 for all areas together and 0.27 for areas e and f alone. If the positions 
of individuals were randomized while retaining the observed number of 
individuals in each area, these ratios would be 2.12 and 1.00, respectively. 
Thus, even male Plebejus saepiolus do not travel freely over the 60 m. 
between areas e and f. An additional indication of the sedentary nature 
of these butterflies comes from observations of P. saepiolus concurrent 
with this study in which only 2 of the 37 individuals observed in the 30 X 
60 meter region just east of area d were marked, while 7 of the 27 in 
area @ were marked. These favorable areas are separated by 30-40 meters 
of drier ground not supporting Trifolium. 


VOLUME 27, NUMBER 1 21 


In the course of this study, male-male encounters were frequently ob- 
served. A possible effect of such encounters would be to encourage the 
even distribution of males over favorable habitat. The mean and variance 
data of Table 1 indicate that this is occurring. If the captures were totally 
independent events and the average population constant through time, 
the number of captures would follow a Poisson distribution (variance 
equal to the mean). Butterfly responses to weather or changes in popula- 
tion would increase the variance. For areas a, e, and f, however, the 
variances are about half the means. The captures are therefore not in- 
dependent events, and the number of butterflies in an area is more 
uniform than would be expected if they did not interact. 


CONCLUSIONS 


Plebejus saepiolus, in contrast to other species of butterflies found in 
subalpine Colorado (Sharp, Parks & Ehrlich, MS in prep.), shows a 
striking degree of correlation with the micro-distribution of its oviposition 
plant Trifolium, which itself has a patchy distribution. P. saepiolus ap- 
pears to be quite sedentary and, as with its relative Plebejus icarioides 
Boisduval, its populations are localized. Other butterfly species in the 
area, notably Erebia epipsodea Butler (Brussard & Ehrlich, 1970) and 
Colias alexandra Edwards (Ward B. Watt, pers. comm.), range widely 
and have large populations. The distributions of individuals in these 
populations do not correlate strongly with that of their oviposition plants, 
and the plants themselves are widely distributed. 

It seems likely that P. saepiolus distributions represent one strategy 
available to a small, weakly-flying butterfly in a seasonally unpredictable 
environment such as subalpine Colorado. By maintaining sedentary pop- 
ulations closely associated with the perennial plant which provides both 
larval food and nectar for the adults, Plebejus saepiolus can minimize 
uncertainties in finding a source of food. 


ACKNOWLEDGMENTS 


Support for this work was provided by the National Science Founda- 
tion Graduate Fellowship Program and by NSF Grants #GB 19686 and 
#GB 22853X. The authors are grateful to the Rocky Mountain Biological 
Laboratory for the use of their facilities, and to M. C. Singer and Ward 
B. Watt for much valuable information. Thanks are due to Paul R. Ehrlich 
and Richard W. Holm, who read and criticized the manuscript. 


LITERATURE CITED 


BREEDLOVE, D. E. & P. R. Enruicu. 1968. Plant-herbivore coe olution: Lupinus 
and lycaenids. Science 162: 671-672. 


22, JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


1972. Coevolution: patterns of legume predation by a lycaenid butterfly. 
Oecologia, in press. 

Brussarp, P. F. & P. R. Enrutcn. 1970. Contrasting population biology of two 
species of butterflies. Nature 227: 91-92. 

Derumr, V. G. 1959. Food-plant distribution and density and larval dispersal as 
factors affecting insect populations. Can. Entomol. 91: 581-596. 

Downey, J. C. & W. C. Futter. 1961. Variation in Plebejus icarioides (Lycaeni- 
dae) I. Food-plant specificity. J. Lepid. Soc. 15(1): 34-52. 

Enruicu, P. R. & P. H. Raven. 1964. Butterflies and plants: a study in coevolu- 
tion. Evolution 18: 586-608. 

Gupert, L. E. 1971. The effect of resource distribution on population structure in 
the butterfly Euphydryas editha: Jasper Ridge vs. Del Puerto Canyon colonies. 
Ph.D. dissertation, Stanford University. 

Sincer, M. C. 1971. Evolution of food-plant preference in the butterfly Euphydryas 
editha. Evolution 25: 383-389. 


FOODPLANT ECOLOGY OF THE BUTTERFLY 
CHLOSYNE LACINIA (GEYER) (NYMPHALIDAE). 
I. LARVAL FOODPLANTS 


RayMonp W. NECK 
Department of Zoology, University of Texas at Austin, Austin, Texas 78712 


For several years I have studied field populations of Chlosyne lacinia 
(Geyer) (Nymphalidae: Melitaeini) in central and south Texas for 
genetic (Neck et al., 1971) and ecological genetic data. A considerable 
amount of information concerning foodplants of this species has been 
collected. Foodplant utilization information is an important base from 
which ecological studies may emerge. Such information is also invaluable 
in evaluating the significance of tested foodplant preferences of larvae 
and adults. Such studies have been under way by other investigators 
and will be available for comparison with natural population observa- 
tions. 

In addition to personal observations (which cover a four-year period 
encompassing some 20 generations ), an extensive search of the literature 
reveals numerous, though scattered, previous reports of foodplants. Litera- 
ture references to populations in central and south Texas are integrated 
into ‘Table 1 with personal observations. All reports of foodplants outside 


tha ct ara: ara cM 10 

the study area are discussed separately at the end of the study area 

foodplant discussion. 
| Wey, 


odplants (see Table 1) are arranged into three basic categories ac- 


VOLUME 27, NUMBER 1 23 


cording to their frequency of utilization by C. lacinia: 1) major food- 
plants, Il) occasional foodplants, and III) rarely utilized foodplants. 
Obviously these three groups are arranged in a descending order of im- 
portance. Within each group, however, foodplants are noi arranged in 
any particular order of importance. 

The first three plants on the list are by far the most important foodplants 
of C. lacinia in central and south Texas. These three plants are the most 
prone of all foodplants (except #7 and #10) to form large, nearly mono- 
specific stands. Larval populations of C. harrisii have greater develop- 
mental success in dense fields of the foodplant, Aster umbellatus, as a 
result of smaller inter-plant distances which allow more successful larval 
movement to fresh plants (Dethier, 1959), rather than as a result of 
greater foodplant biomass which would increase the number of oviposi- 
tion sites. 

The last group of foodplants is almost negligible in importance and 
some records should be considered anomalous. These foodplants are 
included because some “anomalous” foodplants may be utilized at a high 
level under certain conditions (see foodplant #4). This latter group of 
plants might not fit the definition of a suitable foodplant given by 
Remington & Pease (1955) which is as follows: “A full test of the 
suitability of a plant requires that the larva be reared solely on that plant 
and that the adults be induced to mate and lay eggs which then hatch.” 
In actuality, however, one plant species need not be the sole foodplant 
utilized in order to be considered a “full” foodplant (see Discussion ). 

All foodplant records are the result of adult female discrimination 
which resulted in oviposition of an egg mass (marked by “O” in Table 1) 
unless a particular record is cited as being larvae that have switched 
from another foodplant. Skeletonized leaf damage is typical of the 
gregarious early-instar larvae. Post-dispersal larvae of the fourth and 
fifth (final) instars chew holes through all leaf layers. Thus, it is quite 
simple to determine whether leaf damage typical of immature larvae is 
present. If none is present, the larvae are assumed to have switched from 
some other plant. In several instances larvae were actually observed 
while they were switching. This phenomenon of larval foodplant switch- 
ing will be discussed in more detail elsewhere (Neck, in prep. )- 

All observations within the study area involve the taxon of C. lacinia 
known as adjutrix Scudder. Some references to populations in the western 
United States may refer to either adjutrix or crocale ( Edwards) or mixed 
populations of these two phenotypes. Records from Latin America refer 
to saundersi (Doubleday), a term used to describe a form close to or in- 
cluding adjutrix (Higgins, 1960). 


JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


ASH UI TS hs 
-usy ut AvoeT ‘*D ‘ 


ouON 


6S6I ‘Tlepuey 


ouUON 


OLOI “Te 9 puouumAg 


6S61 ‘TIPPUex 


6S61 ‘TIPPUEx 


OL6I “|e 32 puout 
“UNI “PPE, “Wey yULy, 


OL6T “[B@  puouw 
“und, +661 ‘TT2Puex 


Sp10d9yY SNOTAIIg 


s[IOs sno 
-LIBA ‘So}IS paq.in}stp 
epeys [ented 10 
uns |[Nf UL [IOS 9}seA\ 


sjtos Apues 


S[IOS peALlop 9U0}SOUMT] 

[fos Fo 
a1eq ysoul[e syors1o Ul 
ssurddo.190}no suo0saUIT] 


syoyoryy 
pue spue[pooa papeys 


sule[dpooyF 10 spoq 
yeotd yoM AT[eUOSBeS 
syueq 

ulvoljs pur SspyjelfF Fo 
sjios Apues poqinjsip 


sAejo siofoid ynq 
s[Ios snore ‘soztd j4.1p 
pue punois poieeyo 


yequqey 


}SVI ING svate [TB 


sooog 
-suvI[, pue o[puey 
-ueg ynq sroie |e 

svare 
jequeo pur Ulo}sva 


SUIv[ dg BpurI 
Oly ‘Nevoye[q spireapy 

nvoye[q spiear 
-py Aljetoedsa ‘svore 


UId}SoM puP [e1}UI 
SvaIv 
[equeo = pue Ute}sva 


9}e}s 9.11}7U9 


vole 4svad 
UL O1BI ‘9}e}S FO SOUL 


Bore seo UL UOU 
-U09 ssa] “eye}S 9.11} 


(sexoy,) osuey 


PIOMBRI IS[R 


(y) svoyjurleyE = O-VW “T snioydosajshy wmuay.iwg °6 
PIeMseI WOUUIOT) 
(Vy ) oveyjuerpoyT -V TT DYOfusiuayD DISOLqUY °3 
sjuv[dpooy paziyim Ajorwey *yT] 
IOAMOTFUNS 
Jeoploquinony “5 YL 
(A) svoyyuRloyT = O-V snyofiwaunonas snyjyupyar °L 
Asiep quays “Avig 
(A) svomuelpeH = O-d (MAH) opidsiy pwuawxaZ ‘9 
aAouep[Os AIMOTFUNG 
‘sua1ds 
(A) aveyjuele_ §=O-d (‘AB)) DIDJUap DIaINZIA °C 
poaA\}so.Ly 
(A) svoyeley, O-d ‘TT DIWID.MA DUISAGIAA “Ff 
Sjuv[dpooy Jeuoise.g [I] 
PIOMSRI JULI) 
(vy) evoyquelpey, =O-V "T ppifily Diso1quiy *¢ 
Asivp uadavorn 
(A) svoyeley, O-V ‘ABT SAPlONaIUa DIsdUdUL KY “FZ 
IIMOTFUNS UOWIUOD 
(A) svoyqUeley] §=O-V ‘TT snnuup snyjupyay Ty 
sjuvjdpooy sofeyy *y 
oq, PPp0D soroads jury ae 
‘T q1avL 


‘gore Apnjs oY} UIYYWM DIWMID) aufisojYyD JO syuL[dpooF [eare'T 


25 


VoLUME 27, NUMBER 1 


ouUON 
“ULUTOD 

Suscl TER Ol 
OLET [Be 39 puotaumMadg 
ur qepuey ‘oO ‘Y 
OUuON 

OLOI “[e 9 puowumnidg 


sUON 


ouON 


sjios Apues 


S[Ios sno 
“IVA JO Spolf AOT[eF 
‘soinjsed  pouopueqe 

sotireid 


pue s[jouy ouo}soUtI] 
sjios AAvoy 
sjios Arp 


SoPIS poqinjsip }srout 


sjios Apuvs asooT “deep 


‘SUOIPPUOD Pel} J9puN poepi0de1 UOI}sOdIAo—Q ‘[eruuer1ed = g ‘[enuue = V _; 
psemioyduiey “yong 
SBOIB |e IvIAIOISV -V DYOf14D] DIAYIOLALA FY 


yseo ynq swore {Pe 
10}0aS . [e.1yU9O 
sevoiev [e1ju90 pue YNOsS 
SBAIB UId}SOM 
pue = [e.jue9 “Yynos 


9721s o11}US 


sured 
[eqysvoo pur [e.QU90 3svO 


‘IBULISOIqMIy oqingns (Vy) favurtseqi9aA sqragqns = (A), 


avIue[oH 


ovoywelloy 


ovoy uelloH 
(A) oveyueleH 


Cy) eeoutet or 


(A) evoyrurreH 


d-V 


O-d 


mv 


[eeyMoiy “sn 
Dyayand vypiojyjwey 
peeMUISOY 

‘ds wmydjis 
JOMOTJUMBT 978.14S01g 


‘ssa'T_ sypia sndipoojdhypoy 


JamMoyjunsysng “Avis 
(D9 °H) vaypo piswig 
LIMqe]J20r) 

"T WNLpWNs4s WNYy.UdDX 
IaMoyzuns 


JeIPIOATIS “D ¥ WL 
snoyhydosin snyyupya H 


et 


VI 


agli 


‘OL 


ie ee ee ee ee eS SS SS SS ee ae 


Splod9y SNOTASIg 


wyUqey 


(sexoy,) odsuvy 


ePQLtL 


IO) 


soroods yueTg 


ee ee eee ee ee——e——EeEEEE—E—— 
a 


(panuyuoy) “T AIAv], 


26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Larval Foodplants Within Study Area 


The botanical nomenclature of Gould (1969) was followed. All lo- 
calities, unless otherwise noted, are in central and south Texas. Obviously 
some plants not on this list may serve as foodplants, but, due to the time 
spent observing C. lacinia, all major and occasional foodplants utilized 
in this area are believed to be known. Various sources (Heiser, 1947; 
Shinners, 1958; Lynch, 1968; Gould, 1969; Correll & Johnston, 1970) were 
utilized to obtain the information on geographical range (within Texas ) 
and preferred habitats of the various foodplants. This information is 
summarized in Table 1. 

Helianthus annuus L. serves as the major foodplant from spring (first 
adults are normally seen around April 1 in Austin) until August. By 
August the great majority of these plants have become senescent due to 
hot, dry summer weather. The few sunflower plants that survive through 
summer until September rains occur are often able to survive until early 
winter. Larval broods have been found on this species as late as November 
(1971) in central Texas in favorable sites and years. Infestations have 
been found on H. annuus in deep south Texas (Santa Ana Wildlife Refuge) 
as late as the last week of December (1970). 

Although egg masses have been found on cultivated monocephalic 
varieties of H. annuus, larval development does not appear to be as 
successful as on the wild form. Populations would not likely occur in 
cultivated fields of this species because of insecticidal treatments used 
against the sunflower moth, Homeosoma electellum (Phycitidae) (Teetes 
& Randolph, 1968). 

Ximenesia encelioides Cav. is by far the major foodplant of C. lacinia 
from August (Drummond et al., 1970) until the end of the larval feeding 
period in November or early December. Although this annual germinates 
in the spring (February), most seedlings grow rather slowly until late 
summer rains occur (normally in September). These plants then grow 
rapidly and are able to support huge larval populations. 

Ambrosia trifida L. serves as a foodplant only when there is a nearby 
population of C. lacinia on another foodplant. A. trifida is often found in 
the moister parts of a field of H. annuus. I have never found a pure, 
isolated stand of A. trifida to be infested with C. lacinia. Possibly adult 
females are not particularly attracted to this plant unless they have 
already been partially stimulated by one of the other two plants listed 
previously. 

\lthough I have always found limited numbers of larvae on A. trifida, 
<endall (1959) reported thousands of larvae in Medina Co. on a mixed 
stand of Hf. annuus and A. trifida. He also located another large popula- 


VoLUME 27, NUMBER 1 27 


tion which was feeding solely on A. trifida in Kendall Co. This exception 
may have been the result of 1957 being a particularly good year for C. 
lacinia (Kendall, 1959). The large populations that year were prob- 
ably the result of a wet year following the longest Texas drought (1950- 
56) on record. 

Until 1971 the sole record for Verbesina virginica L., which is a central 
Texas foodplant for Chlosyne nycteis, was Kendall’s (1959) report of 
a single larva. In autumn 1971 I located three different infestations on 
this species including one large larval population. Possible reasons for 
utilization of this plant by C. lacinia in 1971 will be discussed elsewhere 
(Neck, in prep.). , 

Xanthium strumarium L. was not recorded as a foodplant for C. lacinia, 
despite continual checking, until autumn 1971. W. H. Calvert (pers. 
comm.) has reported difficulty in rearing C. lacinia larvae on this species 
(i.e., near complete arrest of development). Gaillardia pulchella Foug. 
was reported (R. O. Kendall, pers. comm.) as a foodplant in the autumn 
of 1970 during a severe drought when other foodplants may not have been 
suitable. Some foodplant records (species #’s 8, 10, 12 & 16) consist 
solely of instances where larvae have switched foodplant species. One 
of the aforementioned larval switch foodplants, Helianthus argophyllous 
T. & G. (#10), is known as a foodplant only in the artificial situation of a 
transplant garden at a field laboratory. No records, as a result of either 
oviposition or larval switching, are known under natural conditions, de- 
spite examination of numerous large stands of this species. 


Larval Foodplants Outside Study Area 


Cockerell (Edwards, 1893) reported larvae on what “appears to be the 
common H. annuus” near Las Cruces, New Mexico. Later Cockerell 
(1900) reported them to be very common on H. annuus in the Mesilla 
Valley, New Mexico. He also found them on Helianthus ciliaris while a 
few larvae were found on Xanthium canadense and Palafoxia hookeriana. 
In southern California, larvae have been commonly found on H. annuus 
but a few have been found on Viguiera deltoidea var. parishii (Thorne, 
1962). Comstock (1927) reports the foodplant merely as “sunflower.” 
In Arizona it feeds on H. annuus and Xanthium saccharatum (Gorodenski, 
1969). H. annuus is by far the most important foodplant in Arizona. 
Generally, Xanthium is utilized only in association with the former 
species, although one large population has been reported on the latter 
plant (S. A. Gorodenski, pers. comm.). According to Love & Dansereau 
(1959), the two taxa of Xanthium mentioned above are taxonomic entities 
of the highly variable species, X. strumarium (foodplant #11). 


De) JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The three above states, in addition to Texas, encompass the entire range 
of C. lacinia in the United States except for occasional strays as far north 
as Kansas and Nebraska (Klots, 1951). Its range extends southward 
to Argentina (Bauer, 1951) where it feeds on a composite which Koehler 
(1927) believed to be a Helianthus. L. E. Gilbert (pers. comm.) has 
observed larvae on Eupatorium sp. in Trinidad and Baltimora sp. in 
Guanacaste Province near Canas in Costa Rica. 


Ovipositional Mistakes 


Ovipositional mistakes by adult females of various lepidoptera have 
been previously reported (Remington, 1952; Dethier, 1959a). At times 
mistakes are on exotic toxic species which have native palatable con- 
geners (Straatmen, 1962; Kendall, 1964). Only one mistake has been 
observed for C. lacinia. A normal-sized egg mass was found on a leaf of 
Solanum pseudocapsicum (Solanaceae) growing in the transplant garden 
of the Brackenridge Field Laboratory in the summer of 1971 (A. Hart- 
gerink, pers. comm.). Unfortunately the egg mass was not discovered 
until the leaf had been kept in a refrigerator for several days, thus pre- 
venting the eggs from hatching. Normally utilized foodplants (H. annuus, 
A. trifida and Z. hispida) were present nearby in the same garden. Al- 
though observing such mistakes is largely a matter of chance, one would 
not expect a species laying large egg masses, e.g. C. lacinia, to make as 
many mistakes as a species which lays smaller masses or single eggs as 
the selective disadvantage would be much greater. Dethier (1959b) re- 
corded no such errors for C. harrisii. 


Larval Foodplant Taxonomy 


The above larval foodplants, all members of the family Compositae, 
belong to the tribe Heliantheae (Correll & Johnston, 1970) except 
Gaillardia and Palafoxia (both Heleniae), Heterotheca (Astereae) and 
Eupatorium (Eupatoriae). Cronquist (1955) independently derived 
these last two tribes from the Heliantheae, considered to be the most 
primitive tribe of the Compositae. He acknowledges, however, the pos- 
sibility that the Eupatoriae may be related to the Heliantheae through 
primitive members of the Senecioneae. 

Fle puts the tribe Heleniae into the Heliantheae claiming that the 
former tribe is a catch-all group for several groups which independently 
evolved from the basic Heliantheae stock. The phylogenetic arrangement 


1 CATT vl - . « . ° ° ° 
described above would indicate a tight taxonomic relationship among 


the various foodplants of C. lacinia. It should be mentioned, however, 
that Cronquist derived all but one of the tribes of the Compositae from 


imitive tribe Heliantheae. 


VOLUME 27, NUMBER 1 29 


Of the fourteen foodplants which belong to the Heliantheae, all but two 
belong to the subtribes Ambrosiinae (Ambrosia, Xanthium, and Par- 
thenium) and Verbesiinae (Helianthus, Viguiera, Simsia, Zexmenia, 
Ximenesia and Verbesina). Quite far removed from these two groups, 
and from each other, are Silphium and Calyptocarpus (B. L. Turner, pers. 
comm. ). 

The genus Viguiera (see foodplant #5) is very closely related to 
Helianthus (Heiser, 1969) with generic distinctions breaking down in a 
few species. Blake (1918) believed that Helianthus evolved from the 
ancestral stock of Viguiera, an idea which has received recent support 
from Heiser (1957). Also noteworthy is the fact that some authorities 
(Correll & Johnston, 1970) place Ximenesia (see foodplant #2) within 
the genus Verbesina (see foodplant #4). 

The three species of Helianthus utilized as foodplants by C. lacinia 
in central Texas are all members of the section Annui of Helianthus 
(Heiser, 1969). H. argophyllous is considered to be extremely closely 
related to H. annuus (Heiser, 1951). The only other species of this genus 
which is commonly encountered in central Texas is H. maximiliani, a 
perennial which is not known to be a foodplant. It belongs to the section 
Divaricati of Helianthus. 


DISCUSSION 


It appears that although C. lacinia feeds on a relatively large number 
of plant species, little taxonomic diversity is involved. C. lacinia may, 
therefore, be considered oligophagous, i.e., feeding on only a few species 
of many present throughout its geographical range. 

Actually, since there has never been a definition of just how many 
foodplants are a “few,” application of this term is somewhat the result of 
personal discretion. Dethier (1947) argued that the terms monophagous, 
oligophagous and polyphagous should be based on the number of chemi- 
cals which are perceived as attractants. Using this criterion, none of the 
above terms could be applied to C. lacinia until the chemistry of the 
foodplant-butterfly relationship has been analyzed. C. lacinia might be 
considered monophagous if all of the above foodplants were perceived 
via one chemical or a “group of closely related chemicals confused as one” 
(Dethier, 1947). Thorsteinson (1960) has presented a more diversified 
model involving differential relative importances of stimulatory and 
inhibitory phytochemicals. Plants related to foodplants, but not utilized 
by C. lacinia, might be distinguished by repellant or inhibitory factors 
(chemical or physical) not present in foodplants. For the purposes of this 
discussion, however, C. lacinia will be designated as oligophagous. 


30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Previously we saw that C. lacinia is oligophagous as a species, i.e., it 
occurs on a number of foodplants over its geographic range. A single 
larval population of C. lacinia is most often found infesting several 
foodplants unless only one is available. This may be due to variance in 
adult ovipositional behavior or foodplant switching by larvae. Thus, one 
can say that C. lacinia may be oligophagous as a population and also as 
an individual. A single colony usually feeds on more than one plant and 
the same may be true for an individual larva. 

This situation in C. lacinia may be contrasted with examples, from the 
literature, of other butterflies. Plebejus icarioides (Lycaenidae) can be 
found on numerous species of Lupinus (Leguminosae) although an in- 
dividual population feeds on only one species, even if several species of 
Lupinus are available (Downey & Fuller, 1961). Thus, P. icarioides is 
oligophagous as a species but monophagous as a population and as an 
individual except in rare cases where ants, which may take the larvae into 
their nests for the winter, place them on a second Lupinus in the following 
spring (Downey, 1962). 

Singer (1971) reports larvae of Euphydryas editha (which is in the 
same tribe as C. lacinia) on plants of several families, but many colonies 
are found on practically only one foodplant species. However, at least 
one population of E. editha regularly feeds on two different plant species 
(of two different plant families) in the pre-diapause and post-diapause 
periods (Singer, 1971). Using the same criteria utilized above, E. editha 
can be classified as being oligophagous as a species but monophagous or 
oligophagous as a population. One colony has been found to be regularly 
oligophagous as individuals. A similar situation occurs in E. maturna 
in Europe (Higgins, 1950; Higgins & Riley, 1970) and E. phaeton in the 
eastern United States (Klots, 1951). All of the species discussed in this 
section are oligophagous, but this oligophagy can be exhibited in various 
ways. 

Singer (1971) reported inter-populational differences in foodplant 
specificity in E. editha. This phenomenon has not been observed in C. 
lacinia. Populations of E. editha are genetically isolated from one an- 
other, primarily due to the sedentary habits of the adults (Ehrlich, 1961, 
1965). C. lacinia adults, while most commonly found in the vicinity of 
foodplant stands, are quite capable of flying the distance between popula- 
tions. Such flights probably occur more frequently as a result of nectar 
source shortages which arise when adult populations become very dense. 
Dethier & MacArthur (1964), working with C. harrisii, reported an adult 
emigration which occurred following an increase in the larval population 
due to artificial stocking. This same phenomenon, i.e., density-dependent 
dispersal, is believed to occur in C. lacinia. 


VoLUME 27, NUMBER 1 31 


There is no significant spatial foodplant polymorphism in C. lacinia 
(differential foodplant preferences in spatially separated populations ) as 
discussed for E. editha. Rather, C. lacinia exhibits a temporal foodplant 
polymorphism, i.e., it feeds on H. annuus (spring and early summer) and 
X. encelioides (late summer and fail) during two roughly separable time 
periods. Temporal foodplant polymorphism is known for Papilio machaon 
in desert areas where it feeds on a succession of different foodplants 
(Buxton, 1923). Tephroclystis virgaureata, a European geometrid moth, 
has two generations annually, the first of which reportedly feeds on 
Senecio and Solidago (both Compositae) while the second feeds on 
Prunus and Crataegus (both Rosaceae) (Klos, 1901). | 

It is interesting to note in Table 1| that field oviposition has been ob- 
served for all major and occasional foodplants (groups I and II). How- 
ever, field oviposition is not known for some of the rarely utilized food- 
plants of group III. Force (1966) reported that adults of the three-lined 
potato beetle, Lema trilineata, were “more fastidious in selecting hosts for 
feeding and oviposition” than were larvae which attempted to feed on 
many unsuitable plant species. This is not really unexpected since the 
more mobile adults can search for other food items whereas the relatively 
sedentary larvae are normally forced to eat what the adult selects for its 
young. If the adult selected suitable foodplants, there will be no selec- 
tion for larval discrimination, because the correct foodplant is always 
available. 


SUMMARY 


Sixteen plant species are known as foodplants of Chlosyne lacinia 
(Geyer) in central and south Texas. All of these plants are in the family 
Compositae and all except two are in the tribe Heliantheae. What is 
known of the foodplants outside the study area is reported. The type of 
oligophagy exhibited by C. lacinia is discussed in relation to oligophagy 
as exhibited by other butterfly species. A spatial foodplant polymorphism 
has not been observed, but a temporal foodplant polymorphism is re- 
ported. 


ACKNOWLEDGMENTS 


For encouragement, discussion and comments on previous drafts of 
this paper, I wish to thank G. L. Bush, W. H. Calvert, L. E. Gilbert, 
F. E. Hanson and R. O. Kendall. This research was supported in part by 
PHS-Training Grant GM-00337 and a grant from the National Institute of 
General Medical Sciences GM-15769. 


32 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


LITERATURE CITED 


Bauer, D. L. 1961. Tribe Melitaeini, in P. R. and A. H. Ehrlich, How to Know 
the Butterflies. Wm. C. Brown, Dubuque, Iowa. 

Buake, S. F. 1918. A revision of the genus Viguiera. Contr. Gray Herb, 54: 1- 
205. 

Bruges, C. T. 1924. The specificity of foodplants in the evolution of phtyophagous 
insects. Amer. Nat. 58: 127-144. 

Buxton, P. A. 1923. Animal Life in the Deserts. Edward Arnold, London. 

CockERELL, T. D. A. 1900. Synchloe lacinia. Entomol. News. 11: 503. 

Comstock, J. A. 1927. Butterflies of California. Publ. by author, Los Angeles. 

CorreELL, D. S. & M. Jonnston. 1970. Manual of the Vascular Plants of Texas. 
Texas Research Foundation, Renner, Texas. 

Croneuist, A. 1955. Phylogeny and taxonomy of the Compositae. Amer. Mid. 
Nat. 53: 478-511. 

Deruter, V. G. 1947. Chemical Insect Attractants and Repellants. Blakiston, 
Philadelphia. 

1959. Foodplant distribution and density and larval dispersal as factors 

affecting insect populations. Can. Entomol. 91(9): 581-596. 

. & R. H. MacArruur. 1964. A field’s capacity to support a butterfly 
population. Nature 201: 728-729. 

Downey, J. C. 1962. Myrmecophily in Plebejus (Icaria) icarioides (Lycaenidae). 
Entomol. News. 73: 57-66. 

. & W. E. Furtuter. 1961. Variation in Plebejus icarioides (Lycaenidae). 
I. Foodplant specificity. J. Lepid. Soc. 15(1): 34-52. 

Drummonp, B. A. III, G. L. BusH & T. C. Emmet. 1970. The biology and lab- 
oratory culture of Chlosyne lacinia Geyer (Nymphalidae). J. Lepid. Soc. 24(2): 
135-142. 

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

EnruicH, P. R. 1961. Intrinsic barriers to dispersal in checkerspot butterfly. 
Science 134: 108-109. 

. 1965. The population biology of the butterfly, Ewphydryas editha. II. The 
structure of the Jasper Ridge Colony. Evolution 19: 327-336. 

Force, D. C. 1966. Reactions of the three-lined potato beetle, Lema trilineata 
(Coleoptera:Chrysomelidae), to its host and certain nonhost plants. Ann. 
Entomol. Soc. Amer. 59(6): 1112-1119. 

GoropEnskI, S. A. 1969. The genetics of three polymorphic larval color forms of 
Chlosyne lacinia (Lepidoptera, Nymphalidae). Genet. Res. 14: 333-336. 
GouLp, F. W. 1969. Texas plants—A checklist and ecological survey. Rev. Ed. 

Texas Agri. Exper. Sta. MP-585. 

Heiser, C. B. 1947. Hybridization between the sunflower species Helianthus 
annuus and H. petiolaris. Evolution 1(4): 249-262. 

1951. Hybridization in the annual sunflowers: Helianthus annuus « H. 

argophyllous. Amer. Nat. 85(820): 65-72. 

. 1957. A revision of the South American species of Helianthus. Brittonia 

8(4): 283-295. 

1969. The North American sunflowers (Helianthus). Mem. Torrey Bot. 
Chip 223) l= 28! 

liccins, L.G. 1950. A descriptive catalog of the Palearctic Euphydryas (Lepidop- 
tera: Rhopalocera). Trans. Roy. Soc. Entomol. Soc. London 101: 435-489. 

———. 1960. A revision of the Melitaeine genus Chlosyne and allied species 
( Lepidoptera: Nymphalinae ). Trans. Roy. Entomol. Soc. London 112: 381- 


VOLUME 27, NUMBER 1 33 


.& N. D. Ruzy. 1970. A Field Guide to the Butterflies of Britain and 
Europe. Houghton Mifflin, Boston. 

KENDALL, R. O. 1959. More larval foodplants from Texas. J. Lepid. Soc. 13(4): 
221-228. 

. & C. A. Kenpatt. 1971. Lepidoptera in the unpublished field notes of 
Howard George Lacey, naturalist. (1856-1929). J. Lepid. Soc. 25(1): 29-44. 

Kios, R. 1901. Zur lebensgechichte von Tephrocylstis virgaureata Dbld. Verh. 
K. K. Zool. Bot. Ges. Wien 51: 785 (referred to in Brues, 1927). 

Kors, A. B. 1951. A Field Guide to the Butterflies. Riverside, Cambridge, Mass. 

Korner, I. P. 1927. Biologia de Chlosyne saundersi Dbl. & Hew. Rev. Soc. 
Entomol. Argentina. 1-2: 3-4. 

Love, D. & P. DANsEREAU. 1959. Biosystematic studies on Xanthium: taxonomic 
appraisal and ecological status. Can. J. Bot. 37: 173-208. 

Lyncu, D. 1968. Plants of Austin, Texas. St. Edward’s, Austin. 

Neck, R. W., G. L. BusH & B. A. Drummonp III. 1971. Epistdsis, associated 
lethals and brood effect in larval color polymorphism of the patch butterfly, 
Chlosyne lacinia. Heredity 26(1): 73-84. 

SHINNER, L. H. 1958. Spring Flora of the Dallas-Ft. Worth Area Texas. Publ. 
by author, Dallas. 

SincER, M.C. 1971. Evolution of food-plant preference in the butterfly Euphydryas 
editha. Evolution 25(2): 383-389. 

Teetes, G. L. & N. M. RanpoueH. 1968. Chemical control of the sunflower moth 
on sunflowers. J. Eco. Entomol. 61(5): 1344-1347. 

TuornE, F. 1962. Larval notes on Chlosyne lacinia and C. californica. J. Lepid. 
soe: 16: 61. 

THORSTEINSON, A. J. 1960. Host selection in phytophagous insects. Ann. Rev. 
Entomol. 5: 193-218. 

TinxuaM, E. R. 1944. Faunistic notes on the diurnal lepidoptera of the Big Bend 
region of Trans-Pecos, Texas, with the description of a new Melitaea. Can. 
Entomol. 76: 11-18. 


THE COLLECTION OF BUTTERFLIES MADE BY JACK DENNIS 
AT BEULAH, MANITOBA 


Joun H. Masters 
Lemon Street North, North Hudson, Wisconsin 54016 


During a period of almost 50 years A. J. Dennis, better known as Jack, 
resided and collected Lepidoptera at Beulah, Manitoba. He engaged in 
extensive exchange and sale of specimens and, as a result, his materials 
are distributed throughout the world. His personal coliection, now in the 
possession of the Manitoba Museum of Man and Nature in Winnipeg, 
is the basis of this paper. 

1 have not been able to ascertain a great deal concerning Jack Dennis 
himself. Only three lepidopterists had direct contact with him, Jack May, 


34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Hugh Gibbon and Shirley Brooks, all of whom are now decreased. Most 
of what I have been able to learn is from the G. Shirley Brooks cor- 
respondence file in the Manitoba Museum, from Don Eff of Boulder, 
Colorado who visited him at Beulah in 1941, and from L. P. Grey of 
Lincoln, Maine who had extensive correspondence with him. 

Dennis started collecting Lepidoptera in the 1890s (specimens in his 
collection date from 1898) at Beulah. He saw action during the first 
world war and was disabled. After the war he returned to Beulah, where 
the rest of his life was a rather impoverished existence on an army pension. 
Of necessity, he raised vegetables and fished, but this left him with 
considerable time for butterfly collecting. His collecting was confined, 
however, to the immediate vicinity of Beulah and the nearby town of 
Birtle, since he had no means to travel and was apparently not physically 
able to do much hiking. To supplement his pension he sold specimens 
and also exchanged them. Among those to whom he sold specimens was 
Jean Gunder, who named two aberrational forms after him. Judging 
from the mounted specimens in his collection, his primary exchange 
partners were G. S. Brooks, J. A. Comstock, D. Eff, L. P. Grey, V. Harper, 
W. Hovanitz, H. A. Howland, J. C. Hopfinger, and J. May. After an 
extended illness, Jack Dennis died of cancer in Winnipeg on 12 July 1946. 
G. S. Brooks, at that time curator of entomology for the Manitoba 
Museum, purchased his collection shortly afterwards for the Museum. 

The Dennis collection is contained in home-made plaques of the Riker 
Mount type. Specimens were carefully mounted, depinned and placed 
in these mounts over cotton. In spite of the fact that the collection has 
had virtually no care for 25 years, it is in excellent shape with no dermestid 
infestation. The remaining collection, in the Manitoba Museum of Man 
and Nature, consists of about 400 of these plaques. At one time there 
were considerably more. Some of the details of the original collection 
can be ascertained from a letter which Shirley Brooks wrote to Don Eff, 
26 September 1946: “The collection proper is coming here. About half 
of the stuff is going to Hugh Gibbon. Well I never saw such a mess of 
butterflies as I found there. The collection, as you may call [sic! recall] 
it is all in Riker mounts, and has full data, but that consists of about a 
third of the total, all the remainder lacks any data, much of it beautiful 
stuff. Quite useless from a collectors point. Then there were about 70 
boxes of papered stuff, probably 10% with data, and some of it many 
years old. I looked through most of it, and after selecting about 4 cigar 
boxes full, had a bonfire with the remainder.” 

The fact that Brooks selected what he wanted from the collection to 


tola han VA; o A 
ake back to Winnipeg probably accounts for a number of species, known 


VOLUME 27, NUMBER | 35 


to be taken at Beulah, being absent from the collection and the fact that 
there are only one or two plaques of some species but many many of 
others. Some of the gaps can be filled in by the fact that Dennis submitted 
all of his records from Beulah and Birtle to Brooks for inclusion in his 
(1942) list of Manitoba butterflies. All of the Beulah and Birtle records 
cited there are from Dennis. 

Brooks had intended to organize and catalogue the Dennis collection 
that he took back to Winnipeg. He died in 1947 before having a chance 
to do this and the collection was kept in storage in the Manitoba Museum 
until Harvey Beck, then Keeper of Collections for the Museum, invited 
me to use the collection in 1968. I have ignored the non-Manitoban ex- 
change materials in the collection, but have catalogued the remainder, 
mostly specimens that Jack Dennis collected himself. 

Beulah and the nearby town of Birtle are located on the prairie in 
southwest Manitoba, just south of the Manitoba Escarpment. The 
lepidopterous fauna here shows predominately eastern species, but with 
notable incursions from the west. A few forest species stray into the area 
from Riding Mountain, which is part of the Manitoba Escarpment just 
to the north. All in all it is an interesting collecting locality and with 50 
years of collecting here by Dennis, the species list should be fairly 
complete. 


Species Collected at Beulah and Birtle by Jack Dennis 


HESPERIIDAE 


Amblyscirtes samoset (Scudder). Three specimens, 14 May 1901 and 14 June 1906. 

Amblyscirtes vialis (Edwards). Two specimens, 16 June 1920 and 24 June 1925. 

Euphyes vestris metacomet (Harris). Ten specimens, 7 to 18 July, 1904 to 1943. 

Poanes hobomok (Harris). Nine specimens (one plaque), 6 to 18 July, 1940 and 
1941. Additional plaques are undoubtedly missing. 

Polites coras (Cramer). Twelve specimens, 1 to 24 July, 1906 to 1941. 

Polites themistocles (Latreille). Four specimens, 11 July 1901. 

Polites mystic (Scudder). Eight specimens, 10 to 20 July, 1921 to 1940. 

[Hesperia uncas uncas Edwards]. Not present in the Dennis collection, however 
MacNeill (1964) records a male from Beulah (presumably of Dennis origin) in the 
Los Angeles County Museum, 22 August 1901. 

Hesperia manitoba laurentina (Lyman). Well represented, 41 rather variable speci- 
mens, all in August, 1920 to 1940. 

Hesperia nevada (Scudder). Seven specimens, labeled by Dennis as H. comma 
colorado, 18 to 24 June, 1940 and 1941. Also recorded from Beulah by MacNeill 
(1964). 

Hesperia ottoe Edwards. One specimen, 18 August 1943, Miniota, Manitoba (possibly 
collected by Hugh Gibbon). 

Oarisma powesheik (Parker). Nine specimens, 10 to 20 July, 1902 to 1920. These 
were labeled O. garita by Dennis and recorded as that species by Brooks (1942). 

Ancyloxypha numitor (Fabricius). Four specimens, one dated 28 June 1940 noted 
“first ever caught,” other three 4 and 10 July 1941. 


36 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Carterocephalus palaemon mandan (Edwards). Ten specimens, 16 to 24 June, 1901 
to 1905. 

Pyrgus communis (Grote) ssp. 18 specimens, June to August, 1912 to 1936. 

Erynnis icelus (Scudder & Burgess). Nine specimens in three plaques, three of them 
labeled E. brizo; 14 May to 18 June, 1906 to 1938. 

[Erynnis brizo (Boisduval & LeConte)]. Three specimens of E. icelus were labeled 
as this species by Dennis. Brooks (1942) recorded it from Beulah and Birtle on 
Dennis’ word. 

Erynnis persius (Scudder) ssp. One female, 18 June 1940, labeled E. icelus. 

Erynnis lucilius (Scudder & Burgess). One female, 18 June 1940, labeled E. icelus. 
This constitutes the only known record for Manitoba. 

[Erynnis juvenalis (Fabricius) ssp.]. Brooks (1942) records this species from 
Beulah and Birtle on Dennis’ word, but there are no specimens in the collection 
nor are there any labeled as such. 


PAPILIONIDAE 


x Papilio kahli Chermock & Chermock. I consider Papilio kahli to be a viable breed- 
ing population of hybrid origin. The species is well represented in the Dennis 
collection with over 50 specimens, mostly reared from larvae. The specimens are 
quite variable, ranging in appearance from that of Papilio asterius to Papilio 
hudsonianus, which are the presumed parental stocks. They are variously labeled 
Papilio polyxenes, Papilio zelicaon, Papilio bairdii, Papilio nitra and Papilio 
oregonia by Dennis, which accounts for these names on Brooks’ (1942) checklist. 
A detailed study of the variation in this species is in preparation. 

[Papilio glaucus canadensis Rothschild & Jordan]. Not represented in the collection, 
but Dennis is known to have taken it at Beulah and Birtle, as would be expected. 


PIERIDAE 


Pieris protodice protodice Boisduval & LeConte. A dozen specimens, mixed with the 
following species, May, July and August, 1920 to 1937. 

Pieris occidentalis Reakirt ssp. Over 30 specimens, mixed with P. protodice, May, 
July and August, 1920 to 1937. Variable and very darkly marked specimens in 
both spring and summer broods, obviously distinct from P. protodice in the 
summer brood. These were labeled by Dennis as var. calyce Edwards, hence this 
name in Brooks’ checklist. 

Pieris napi oleracea Harris. Six specimens, 15 June to 26 July, 1904 to 1922. 

Pieris rapae rapae (Linnaeus). Only one specimen, undated, in the collection. This 
species is very abundant at Beulah now, and must have been during Dennis’ later 
years there. 

Colias eurytheme eurytheme Boisduval. Represented by 95 specimens, 1 July to 
6 September, 1924 to 1941. Of special interest are eight miniature specimens 
(forewing lengths 17 to 19 mm.) taken during July 1934 and labeled “caught 
at Beulah, Manitoba, the year of the drought.” 

Colias eriphyle Edwards or Colias philodice Godart ssp. Represented by 89 specimens, 
24 May to 18 October, 1910 to 1944. 

Colias alexandra christina Edwards. This is the best represented species in the 
Dennis collection. There are over 400 examples, including at least one plaque 
for nearly every year between 1902 and 1944. Male capture dates range from 12 
June to 5 July, female from 16 June to 14 July. The variation in these specimens 
is the subject of another paper (Masters, in press ) 


. 


DANAIDAE 


[Danaus plexippus plexippus (Linnaeus)]. Not represented in the collection, but 
known to have been taken by Dennis at Beulah. 


VOLUME 27, NUMBER | 37 


SATYRIDAE 


[Lethe anthedon borealis (Clark)]. Not represented in the collection, but recorded 
for Beulah and Birtle by Brooks (1942). The species is abundant at Riding Moun- 
tain, a short distance to the north. 

Lethe eurydice fumosus (Leussler). Thirty specimens, 1 to 24 July, 1903 to 1944. 
Series of six specimens 12 July 1903, are labeled, “Uno near Beulah, first time 
caught.” 

Coenonympha tullia nr. benjamini McDunnough. Represented by 25 specimens 
divided by Dennis into groups with ocelli, which he labeled benjamini, and groups 
without ocelli, which he labeled insulana. Dates include 16 June to 10 July, 1898 
to 1936. 

Cercyonis pegala nr. ino (Hall). Ten specimens in one plaque, 10 to 20 July, 1906 to 
1922. 

Erebia discoidalis discoidalis (Kirby). Four specimens, undated, possibly from 
Riding Mountain or some other point in Manitoba; however Beulah and Birtle were 
included by Brooks (1942) as localities for this species. 

Erebia epipsodea freemani Ehrlich. One specimen, undated. Beulah is given as 
a locale for this species by Brooks, and in correspondence to L. P. Grey, Dennis 
lamented on the fact that epipsodea is no longer seen although it was once common 
before extensive agriculture. 

Oeneis uhleri varuna (Edwards). Jean Gunder named an aberration of this species, 
taken by Dennis at Beulah, O. uhleri varuna trans. form dennisi. There are 25 
specimens in the collection, 1 to 14 June, 1934 to 1938. 

Oeneis alberta alberta Elwes. Represented by 29 specimens, 16 to 26 May, 1899 to 
1938. 


NYMPHALIDAE 


Vanessa atalanta rubria (Fruhstorfer). Represented in the collection by one plaque 
of ten specimens, 28 July to 28 August, 1908 to 1912. 

[Cynthia virginiensis Drury )]. Not represented in the collection, but listed for Beulah 
and Birtle by Brooks (1942). 

[Cynthia cardui (Linnaeus)]. Not represented in the collection, but assumed to have 
been taken at Beulah by Dennis. 

[Precis coenia coenia (Huebner)]. Listed by Brooks (1942) for Birtle, however not 
represented in the collection. 

Nymphalis j-album j-album (Boisduval & LeConte). Nine specimens, 24 July to 
4 August, 1900 to 1912. 

Nymphalis milberti milberti (Godart). Five specimens, 24 July to 1 September, 1910 
to 1926. 

[Nymphalis antiopa antiopa (Linnaeus)]. Not represented in the collection, but 
known to have been taken by Dennis. 

Polygonia interrogationis (Fabricius). Six specimens in the collection, all 24 June 
1896. 

Polygonia comma (Harris). Two specimens in the collection, undated. Recorded 
from Beulah by Brooks (1942). 

Polygonia satyrus neomarsyas dos Passos. This is the commonest Polygonia on the 
western Manitoba prairies; it is represented in the collection by 17 specimens, 17 
May to 25 July, 1903 to 1942. 

[Polygonia faunus (Edwards)]. Recorded from Beulah and Birtle by Brooks (1942), 
but not represented in the collection. This species is fairly abundant at Riding 
Mountain. 

[Polygonia zephyrus (Edwards)]. The record from Beulah that Dennis gave to 
Brooks for his Manitoba list (1942) for this species, turns out to be erroneous. 
The specimen is a rather darkly marked P. progne. 


38 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Polygonia progne (Cramer). Nine specimens, 9 to 24 June, 21 July and 10 September, 
1936 to 1941. 

Phyciodes tharos (Drury) ssp. There are 30 specimens of Phyciodes in the collection, 
all labeled Phyciodes tharos pascoensis by Dennis, but only a very few of them 
are actually tharos; most are P. campestris. I suspect that tharos is at least as com- 
mon, if not more so, than campestris at Beulah but that Dennis primarily mounted 
campestris because they are slightly larger. 

Phyciodes campestris camillus Edwards. See comments on preceding species; 2 to 
20 July, 1906 to 1938. This is the first published record for P. campestris in 
Manitoba, although I have found it moderately abundant in recent years at the 
Lake Audy Prairie area in Riding Mountain National Park. 

Chlosyne gorgone carlota (Reakirt). Represented by 19 mounted and many, many 
papered specimens, this is evidently one of the most abundant species at Beulah; 
8 to 20 June, 1936 to 1944. The Beulah population seems to be distinct from 
more typical carlota. The ground color of the wings is more “pinkish” and the 
black markings on the dorsal surfaces are reduced in much the same manner as in 
C. nycteis reversa. The southwest Manitoba population of this species could be 
given subspecific recognition. 

Chlosyne nycteis reversa (Chermock & Chermock). One specimen only, a female, 
28 June 1900, included in a plaque with six specimens from Sylvania, Ohio with 
1939 and 1940 dates. The inference is that the species is very rare here and the 
one specimen more than likely the only one caught. 

[Chlosyne harrisii (Scudder) and Chlosyne hanhami (Fletcher)]. Neither of these 
are represented in the collection, but Dennis apparently captured both at Beulah 
or Birtle as recorded by Brooks (1942). In a letter to G. S. Brooks, dated 9 May 
1943, Dennis states that he takes C. harrisii in green spots near springs and C. 
hanhami in more open terrain. The two are sympatric over much of southern 
Manitoba and remain distinct; harrisii is smaller and prefers wooded habitats while 
the much larger hanhami is found on the prairies. 

Boloria selene (Denis & Schiffermiiller) ssp. One very lightly marked male, 28 July 
1944 at Birtle; presumably the only example of this species taken by Dennis. 

Boloria bellona (Fabricius) ssp. Only one plaque of nine specimens in the collection, 
18 June to 14 July, 1900 to 1940, would indicate a rarity at Beulah. Hugh Gibbon 
took these in numbers at Miniota during the 40’s and 50’s. 

Speyeria callippe calgariana (McDunnough). Four specimens; female 10 July 1900, 
male 16 July 1902, male 14 July 1909, and female 20 July 1909. 

Speyeria edwardsii (Reakirt). Four specimens; male and female 17 July 1898, male 
14 July 1928, and female 24 July 1936. 

Speyeria atlantis dennisi dos Passos & Grey. This was originally described by Jean 
Gunder as “Argynnis lais tr. f. dennisi.” This name, as proposed by Gunder, is not 
available; transitional form was his term for designating aberrant specimens. The 
name dennisi was elevated to the subspecies rank by dos Passos & Grey (1947); 
this is its first availability. The species is better known as Argynnis or Speyeria 
lais Edwards in the literature, but unfortunately this name is not available, the 
Argynnis lais of Scudder having priority. The species is common at Beulah in late 
summer. 

Speyeria mormonia eurynome (Edwards). Two specimens at Beulah, a female 23 
July 1937, and a male 28 July 1938. There is another Manitoba specimen in the 
Dennis collection, 25 July 1935, from Riding Mountain. 

Speyeria cybele pseudocarpenteri (Chermock & Chermock). Represented by 18 
specimens, 24 July to 18 August, 1920 to 1934. These are relatively small for 
cybele. Speyeria aphrodite manitoba (Chermock & Chermock) would be expected 
to occur here with cybele, but it is not represented in the collection, nor is it 

ecorded from Beulah by Brooks (1942). 

“uptoieia claudia claudia (Cramer). Six specimens, 10 to 26 July, 1920 to 1938. 


| 
| 


VoLUME 27, NUMBER 1 39 


LYCAENIDAE 


Satyrium liparops fletcheri (Michener & dos Passos). One plaque with three specimens 
from Birtle, 15-16 July 1940, plus four more specimens taken by Hugh Gibbon at 
Miniota, 18 July 1942. 

Satyrium acadica watrini (Dufrane). Two specimens, only one dated, 12 July 1941. 

[Harkenclenus titus (Fabricius) ssp.]. Not represented in the collection, however 
recorded from both Beulah and Birtle by Brooks (1942). 

Lycaena thoe Guérin-Méneville. Eight specimens, only six dated, 10 to 17 July, 
1906 and 1907. 

Lycaena xanthoides dione Scudder. Nine specimens, 16 to 20 July, 1904 to 1940. 

Lycaena helloides (Boisduval). Nineteen examples, 8 to 24 July, 1900 to 1930. 

Lycaeides melissa melissa (Edwards). Four specimens, all females, 1 to 16 July, 1924 
to 1938. 

Plebejus saepiolus (Boisduval) ssp. Eleven examples, 8 to 20 July, 1900 to 1914. 

Agriades rustica manuscript ssp. Brown. Represented by 24 specimens, 10 to 20 June, 
1900 to 1936. 

Everes comyntas comyntas (Godart). There is one plaque of five specimens, un- 
dated, that were labeled as comyntas by Dennis. Of these only one male is 
actually comyntas, the others are amyntula. Southern Manitoba is part of a very 
limited area where these two species are sympatric. 

Everes amyntula albrighti Clench. Two males and two females, undated; see com- 
ments under E. comyntas. 

Glaucopsyche lygdamus afra Edwards. Eight specimens, only six of them dated, 
14 June to 17 July, 1907 to 1910. 

Celastrina argiolus argentata Fletcher. Twelve specimens, all from the spring brood, 
20 to 25 May, 1906 to 1920. 


ACKNOWLEDGMENTS 


I would like to acknowledge the assistance of J. Richard Heitzman of 
Independence, Missouri who made the determinations of Hesperiidae for 
me; and H. Harvey Beck and Robert W. Nero, both formerly of the 
Manitoba Museum of Man and Nature, for making the arrangements 
that allowed me to make this intensive study of the Dennis collection. 


LITERATURE CITED 


Brooxs, G. S. 1942. A checklist of the butterflies of Manitoba. Can. Entomol. 
74: 31-36. 

MacNertt, C. D. 1964. The skippers of the genus Hesperia in western North 
America with special reference to California (Lepidoptera: Hesperiidae). U. 
Calif. Publ. Entomol. 35: 1-230. 

Masters, J. H. Variation in Colias alexandra christina Edwards (Pieridae) in 
southwest, Manitoba. J. Res. Lepid. (in press ). 

pos Passos, C. F. & L. P. Grey. 1947. Systematic catalogue of Speyeria ( Lepidop- 
tera, Nymphalidae) with designations of types and fixations of type localities. 
Amer. Mus. Novit. 1370: 1-30. 


40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


A REVIEW OF THE AMBLYSCIRTES WITH THE DESCRIPTION 
OF A NEW SPECIES FROM MEXICO (HESPERITDAE) 


Hucu AverRY FREEMAN? 
1605 Lewis Drive, Garland, Texas 75040 


Genus Amblyscirtes Scudder 1872 
Generotype Hesperia vialis Edwards 


Amblyscirtes Scudder, Rept. Peabody Acad. 1871, 75 (54), 1972. 
Stomyles Id. op. cit. 76 (55). Generotype Pyrgus textor Hubner. 
Mastor Godman, Biol. Cent.-Am., Rhop. II. 567, 1900. Generotype Mastor anubis 


Godman. 
Epiphyes Dyar, Jn. N. Y. Entomol. Soc. XIII, 132, 1905. Generotype Pamphila 


carolina Skinner. 


Antennae short, approximately % costa; shaft checkered, white under the club; 
club stout, %4 shaft; apiculus obtuse from thickest part of club, short equal width 
of club; nudum 4/6. Palpi cylindrical, upturned; second segment with shaggy 
vestiture, third segment slender, smooth, and vertical, almost as long as second in 
most species. Mid and hind tibia spined. Males with either brands or a stigma on 
the upper side of the primaries. Primaries in most species similar in both sexes. 
Costa flattened; apex rounded-rectangular; outer margin strongly rounded except 
toward anal angle; cell about two-fifths as long as wing. Secondaries rounded, in 
most of the species longer through cell than in most of the related genera. Genitalia 
rather peculiar in that the saccus and aedoeagus, in most species, are very long, ex- 
tending nearly to the thorax. 

There are 31 species in the genus Amblyscirtes, and for the present 
I do not recognize any subspecies. Biological and distributional data as 
well as basic morphological characteristics indicate to me that several 
previously recorded subspecies actually represent valid species and are 
thus treated in this review. The members of this genus are placed into 
four groups by Evans and I follow his arrangement. The first group is 
the Exoteria Group which is characterized by the presence of more or 
less ochreous scaling on the upper surface of the wings and by the presence 
of at least grey scaling on the lower surface of the secondaries. A cell spot 
or spots may or may not be present. The second group is the Aesculapius 
Group which is characterized by being greyish-brown to black, with a 
well marked single or double cell spot on either the upper or lower surface 
of the primaries, and discal spots or grey scaling on the lower surface of 
the secondaries. The third group is the Vialis Group which is characterized 
by being greyish-brown to black, with no cell spot on the primaries, and 
by having discal spots or greyish scaling on the lower surface of the 


secondaries. The fourth group is the Phylace Group which is character- 


As - woes like to express my appreciation to the American Philosophical Society for a research 
nt which made it possible for me to conduct this work on the Amblyscirtes. 


VOLUME 27, NUMBER 1 4l 


ized by having no markings on the lower surface of the secondaries. 
There is considerable diversity superficially among members of this 
genus, however with the exception of simius Edwards there is a re- 
markable similarity in the male genitalia of all of the species. Often 
worn specimens are very difficult to identify even after an examination of 
the genitalia due to this great consistency in basic form. 

The genus Amblyscirtes reaches its greatest development in the south- 
western part of the United States with Texas being the metropolis as 13 
species have been recorded from that state. There have been two species 
recorded from Canada, 22 from the United States, 19 from Mexico, and 
one from Cuba. Records from Central and South America are lacking 
indicating that this is strictly a North American genus. 


Key to the Species 


la. Under surface of secondaries marked with at least grey scaling; usually 


QpLcell HOU. OREN TT e  eeeeeee e 2 
lb. Under surface of secondaries unmarked; apical spots absent 27 
Zas Wipper surface with more or less ochreous scaling —..--...- 3 
2b. Upper surface without ochreous scaling, greyish-brown or brownish-black _ 12 
SMeoncs man a pale cell spot; 15-20 mm wide <2. 4 
Bhwenimantes wiathouta_cell spot 13-4 mm wide 2. 9 
4a. Under surface of primaries cell brown; cilia checkered; a distinct broken 

SAGITD, OUS STE eee D 


4b. Under surface of primaries cell orange; cilia checkered or uncheckered; an 
MIeISMMCEDLOkem stigima OG brands present —-.. 2. 8 
5a. Under surface of primaries have a suffused white discal area in space lb. 6 
Sb. Under surface of primaries lacks suffused white discal area in space 1b —- 7 
6a. Cilia white, faintly checkered; primaries 15-18 mm wide; distinct white 
discal and cell spots on lower surface of secondaries ~~ folia 
6b. Cilia sordid white, uncheckered; primaries 20 mm wide; indistinct yellow- 
ish spots in spaces 2 and 3; lower surface of secondaries uniform chocolate 


brown with faint indication of yellowish discal spots _ raphaeli 
6c. Cilia brown; primaries 16 mm wide; white discal spots present on lower 
UGE CMMO RE SCCOMG AICS) seve ns insulae-pinorum 


7a. Cilia distinctly checkered brown and white; distinct white spots present 
on primaries and lower surface of secondaries; primaries 15-18 mm 


Cl a LN a eS ee easel 5. = /CKOCETIG 
7b. Cilia indistinctly checkered light brown and white; all spots on ais wings 
indistinct or absent; primaries 18-20 mm wide — _.. immaculatus 


8a. Cilia white, uncheckered; apex on lower surface of primaries and of lower 
surface of secondaries grey, overscaling the usual white markings; indistinct 
broken stigma present; primaries 13 mm wide — simius 

8b. Cilia checkered light and dark brown; grey overscaling on apex and lower 
surface of secondaries sparse; distinct brand present against cubitus and 
between origins of veins 3 and 2 and under vein 2; primaries 13 mm 


wide Ook. < ae Os i ee eee : Cassus 
9a. Maculation on primaries and lower surface of second: rries usually prominent, 

especially they apical spots 2 Ae. LO 
9b. Maculation on primaries and lower surface of secondaries reduced and in 


Some: cases: mearlysalsemt ss s*\ eens 1] 


10a. 


10b. 


10c. 


lelrar 


LIU. 


an 
12b. 
lear 
13b. 


14a. 


14b. 


tear 
15b. 
16a. 


16b. 


lear 


17b. 


18a. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Maculation on primaries and lower surface of secondaries prominent, with 
an ochreous spot or spots in space la and sometimes 1b on the upper sur- 
face of primaries; maculation on lower surface of secondaries usually clear 
white; brands between origins of veins 3 and 2 and under vein 2 in- 
CONSPICUOUS 22-21 aenus 
Maculation on primaries and lower surface of secondaries somewhat re- 
duced with no spot in space la or 1b; maculation on lower surface of 
secondaries never clear white, usually dusky; brands prominent on upper 
surface of -primaries 2_..0.0.00 ol erna 
Maculation on primaries reduced, prominent on lower surface of secondaries, 
often clear white; generally ground color darker than in above two species; 
brands prominent on upper surface of primaries —_______-___________ linda 
Lower surface grey; heavy grey overscaling on lower surface of secondaries; 
cilia sordid white; coloration on upper surface pale brown; brands on pri- 
maries conspiciious: 2.22.2 Eee oslari 
Lower surface dark brown; grey overscaling very sparse on lower surface 
of secondaries; cilia dusky; maculation ochreous on upper surface of pri- 


maries;) brands) inconspicuous —aes sme nese es ee fluonia 
A distinct single or double cell spot on upper or lower surface of primaries 13 
No cell spot on upper or lower surface of primaries 20 
Brands present on upper surtace of primaries 2 14 


Grey stigma present on upper surface of primaries; primaries 11 mm wide; 
cilia usually plain brown; maculation on primaries faint or absent; lower 
surface of secondaries dark brown, overscaling sparse, with distinct tiny 


white diseal and cell spots present 222 = ae elissa 
On the upper or lower surface of the primaries there is a distinct white spot 
in space Ibe 22382) ee ee 15) 


On the upper or lower surface of the primaries there is an obsolete olive 
spot in space lb, or it may be absent; cilia checkered; maculation on pri- 
maries above olive scaled; brands under cubitus and vein 2 inconspicuous; 
on the lower surface of the secondaries there is a heavy grey overscaling 


causing the white markings to be blurred; primaries 13 mm wide ____ samoset 
Cilia checkered 2.2..:.0000 a ee 16 
Cilia not checkéred 2.00 Oe ee 19 
On the lower surface of the secondaries there is distinct grey scaling and 

small white spots; veins are concolorous with rest of wing lef 


No distinct spots on upper surface of secondaries; on the lower surface of 
the secondaries grey scaling absent; veins more or less white on the lower 
surface of the secondaries, with the discal spots united into a band; pri- 
maries 14 mm wide; brands inconspicuous aesculapius 
There is a small white cell spot and discal spots on the upper surface of 
the secondaries; on the upper surface of the primaries there are spots in 
spaces 4 and 5; on lower surface of secondaries there is an extra spot in 1c 
under origin of véin 2 0 ee 18 
No spots on the upper surface of secondaries; on the lower surface of the 
secondaries there is grey overscaling with more or less distinct white spots 
present; there may or may not be an ochreous spot in space lb on the 
primaries; brands inconspicuous; primaries 13 mm wide __________.____ texanae 
Small, primaries 13 mm wide, pale in coloration; upper and lower cell 
spots large and usually fused on primaries; brands narrow under cubitus 
and vein 2: on the under surface of the primaries the costa and apex, as well 
as the entire under surface of the secondaries, densely overscaled with 
violet-grey scalés: (2) Eg ee coc prenda 
Larger, primaries 14 mm wide, darker in coloration; wpper and lower cell 
spots small, seldom fused; narrow brands under cubitus and vein 2 in- 


VOLUME 27, NUMBER 1 43 


19a. 


Lobe 


20a. 


20b. 


Dia: 


23b. 


Aa 
QAb. 
25a. 


25b. 


26a. 


26b. 


distinct; violet-grey overscaling greatly reduced on lower surface of wings _ 
UE Un ADDN Es) es oe ee tolteca 
Primaries 13 mm wide; brands distinct under cubitus and vein 2: on the 
lower surface of the primaries the costa and apex is heavily overscaled with 
dull yellow as is the entire lower surface of the secondaries; indistinct rusty 
brown maculation on the lower surface of the secondaries == carolina 
Primaries 13 mm wide; brands distinct under cubitus and vein 2; dull 
yellow overscaling on lower surface of wings sparse or absent, with the 
ground color rusty brown; maculation on lower surface of secondaries dull 
SPSL iy. GHSUTA GT Ee Cn reversa 
Cilia white, not checkered; primaries 13 mm wide; brands short and fairly 
broad under cubitus and vein 2; maculation on upper surface of primaries 
SUEDE, extending into space lb; there are discal spots on the upper 
surface of the secondaries; on lower surface of secondaries there is heavy 


greenish overscaling with he usual whitish discal spots present nereus 
Cilia more or less checkered; no spot in space 1b on the primaries; no spots 
present on the upper surface of the secondaries _......-- 21 


Lower surface of secondaries abnormal, variegated with large white, dark 
brown and greenish patches; primaries 12 mm wide; brands long and nar- 
row under cubitus and vein 2; apical spots prominent on primaries, other 


PEED “SO TS ee ee ee nysa 
Lower surface of secondaries normal with grey scaling and white spots 
MaweeMcesent On at least mdicated, 2. 22 


On the lower surface of the secondaries the grey overscaling is heavy and 
the white discal spots are conspicuous and dark edged; primaries 12 mm 


wide; brands short and fairly broad under cubitus and vein 2 __......_-_ €0S8 
On lower surface of secondaries the grey overscaling is sparse and the white 
Gusedlaspors are poorly Gefined or absent 2. 23 


Apical spots on primaries abnormal, increasing in size from space 6 to space 
8, an additional spot in space 9, and white streaks on the costa at the ends 
of spaces 8-11; primaries 13 mm wide; very short brand present under vein 
2; spots in spaces 2 and 3 on primaries indistinct or absent; on lower surface 
of secondaries grey overscaling indistinct, with only the slightest indication 


Bis GTS@aM, Symi U: eet eee vialis 
Apical spots on primaries normal, dots in spaces 6, 7, and 8, and no streaks 

ED, See OO, a a ee ee PP 24 
Usually prominent spots in spaces 2 and 3 on the primaries —--------_ 25 
Wsuallyano spots in spaces 2 and 3 on the primaries. 26 


Spot in space 2 rounded on the primaries; upper cell spot on primaries 
often present; primaries 12-14 mm wide, not produced; brands well de- 
veloped, against cubitus and a long streak under vein 2; on lower surface 
of secondaries grey overscaling fairly heavy, with the discal spots sordid 
wilhs to ¢u@er 2 ee ee ee ek at CCN 
Spot in space 2 V-shaped on the primaries; no upper cell spot on primaries 
present; primaries 12-14 mm wide, produced; brands well developed 
under cubitus and vein 2; on lower surface of the secondaries the grey 
overscaling is sparse, with the discal spots grey and often rather in- 
Aegis: Se SS  _ ee ee ee ee ; = _ belli 
Primaries 11 mm wide; brands reduced to a tiny eon aoe vein 2 against 
the cubitus; apical spots present on primaries; grey overscaling fairly sparse 
on lower surface of secondaries, with greyish, indistinct discal spots 


present Bene Be Be eS oa see Se alte rnata 
Primaries 12-14 mm wide; long brand present under vein 2; no maculation 
present on the upper Smnnee of primaries; on lower surface of primaries 


indistinct apical spots are present; on lower surface of secondaries there 


44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


I 2 


Figs. 1, 2. Amblyscirtes raphaeli n. sp. Holotype male. Candelaria Loxicha, 
Oaxaca, Mexico, 7 August 1969 (E. C. Welling; A. M. N. H.) 


may be present some scattered white scales and the slightest indication of 

of discal spots( 022. tL ee florus 
27a. Palpi below orange or yellowish white = = ee 28 
27b. Palpi below black with ochreous hairs intermixed; cilia on primaries 

yellowish-white, on secondaries dark brown; narrow stigma broken at vein 

2 and extending just below that yeim 2 =) anubis 
28a. Cilia sordid white in the males, sordid white to grey in the females; palpi 

below orange in the males, yellowish-white in the females; males have a 

broad and short brand covering vein 2 near its origin on the primaries _.. phylace 
28b. Cilia orange in both sexes; palpi below orange in both sexes; males have a 

narrow broken grey stigma from origin of vein 3 to vein 1 _._- fimbriata 


Exoteria Group 
1. Amblyscirtes folia Godman 1900 


Synonym: tutolia Dyar 1913: Mexico. 

Type locality: Mexico. 

Distribution: Mexico. Colima: Salada, VI-67. Guerrero: Acahuizotla, IX-60; 
Chilpancingo; Mexcala, VII-5-56; Tierra Colorado, IX-61. Jalisco: Ajijic, IX, X-65, 
66; Lake Chapala, VI. 


This species is readily recognized by the well marked discal and apical 
spots on the primaries, the whitish streak in space 1b on the lower surface 
of the primaries, and the clear white discal and cell spots on the lower 
surface of the secondaries. It occurs in semi-tropical and tropical areas 
of western Mexico. 


2. Amblyscirtes raphaeli Freeman, new species 


MALE (Upper Side). Primaries dark brown, with an indistinct apical spot in space 
6. A distinct yellowish spot in space 2, and another similar spot in space 3 situated 
outward from the spot in space 2. A fairly broad, slightly broken, black stigma ex- 
tending from space 1 to the end of the cell. Fringes yellowish-white, not checkered. 
Secondaries dark brown, immaculate. Fringes yellowish-white, not checkered. 

MALE (Under Side). Primaries, brown, with the discal spots distinct. Apical spot 
in space 6 present but very indistinct. A broad suffused yellowish area in space 1b 


VoLUME 27, NUMBER | 45 


Fig. 3. Male genitalia. Amblyscirtes raphaeli n. sp. Paratype. Candelama Loxicha, 
Oaxaca, Mexico, 21 July 1969 (E. C. Welling; H. A. F.). 


extending from the middle of the space to the outer margin. Secondaries dark 
chocolate brown, with practically no overscaling present. The slightest indication 
of three yellowish discal dots, otherwise immaculate. 

Thorax dark brown, both above and below. Abdomen dark brown above, some- 
what lighter below. Head brown. Palpi brown with yellowish intermixed scales 
present. Antennae, shaft brown above, below indistinctly ringed with narrow yellow 
lines; club, basal half greyish above, yellow below, terminal end and apiculus black 
above, lighter below with the apiculus being tan. 

Wing Measurements: Primaries: base to apex, 20 mm; apex to outer angle, 
14 mm; outer angle to base, 15 mm. Secondaries: base to end of vein 3, 15 mm; 
center of costa to anal angle, 14 mm. Total expanse: 39 mm. 

Type Material: Holotype. Male, Candelaria Loxicha, Oaxaca, Mexico, 7 August 
1969, obtained from Eduardo C. Welling, will be placed in the American Museum 
of Natural History, New York. There are nine male paratypes, all obtained from 
Eduardo C. Welling, from the same location, 21 July 1969, 6 August 1971, 11 
August 1971, 14 August 1971, 20 August 1971, and 25 August 1971, which will remain 
for the present in the H. A. Freeman collection. 


This new species is the largest known Amblyscirtes. Previously im- 
maculatus Freeman represented the one with the greatest expanse of wings 
as the primaries from the base to the apex were 18 mm wide, while in 
raphaeli that distance is 20 mm. Raphaeli slightly resembles immaculatus, 
however there are four basic differences: (1) raphaeli has discal spots 
in spaces 2 and 3 on the primaries which are absent in immaculatus; 
(2) raphaeli has a yellow apical spot in space 6 which is absent in im- 
maculatus; (3) raphaeli has on the lower surface of the primaries a broad 
suffused yellowish area in space lb which is absent in immaculatus; 
and (4) there are basic differences in the genitalia. 

This new species is named in honor of my good friend Senor Raphael 
Aguirre, manager of Hotel Covadonga, south of Ciudad Valles, S. L. P., 
who has given me much assistance in my collecting in Mexico. 

This distinctive species can be readily recognized by its large size and 
the chocolate coloration on the lower surface of the secondaries. Ap- 


46 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


parently this is a jungle species as the known specimens were collected 
in such a habitat in Oaxaca. 


3. Amblyscirtes immaculatus Freeman 1970 


Type locality: Salada, Colima, Mexico. 

Distribution: Mexico. Colima: Salada, VI-67. Guerrero: Acahuizotla, VII-65. 

This species is conspicuous by its large size and the reduction of all 
maculation. It flies in company with folia and seems to prefer semi- 
jungle areas as its natural habitat. 


4. Amblyscirtes nisulae-pinorum Holland 1916 


Type locality: Isles of Pines, Cuba. 
Distribution: cusa. Isles of Pines. 


Evans considered this species to be a subspecies of folia, however 
morphological and biological characteristics indicate to me the distinct- 
ness of the two species. Data available indicate this species to be endemic 
to the Isles of Pines, Cuba. 


5. Amblyscirtes exoteria (Herrich-Schaffer) 1869 


Synonyms: nanno Edwards 1882: Arizona. marcus Strand 1907: “Delagoa Bay.” 

Type locality: not known. 

Distribution: UNITED strates. Arizona. Apache County: White Mountains, 
VII-4-51. Cochise County: Carr Canyon; Chiricahua Mountains, VI-28-42; Huachuca 
Mountains, VIII; Onion Saddle Pass, Chiricahua Mountains, VII-11-60; Paradise, 
VI-10-60. Graham County: Fort Grant; Mount Graham. Pima County: Madera 
Canyon, VII-6-10-60, VII-10-15-64; Mount Lemmon, VII-60. mexico. Jalisco. 
Veracruz: Orizaba, VI. Sonora: northern section, VI, VII. 


This species can easily be separated from folia by the absence of the 
whitish streak in space 1b on the lower surface of the primaries. Personal 
observations and recorded data reveal that this species is usually found 
in semi-arid mountainous regions of Arizona and northern Mexico, and 
rarely in the mountains of Veracruz. 


6. Amblyscirtes simius Edwards 1881 


Type locality: Oak Creek and Pueblo, Colorado, 

Distribution: UNITED staTEs. Arizona, VI. Colorado. Oak Creek Canyon. Baca 
County: Regnier, VI. El Paso County: Fountain Valley School, V. Larimer County: 
Ft. Collins, V-18-69. Pueblo County: Pueblo, VII. Saguache County: Saguache, 
VIII-2-67. Nebraska. Sioux County: VII. New Mexico. Colfax County: Raton, 
VII-15-45. Grant County: Silver City, VIII. Texas. Armstrong County: Palo Duro 
Canyon, IV, V-42. Jeff Davis County: Davis Mountains, VII-30-53. 


Superticially simius resembles other members in_ its group of 
Amblyscirtes, however its genitalia do not approach the basic genitalic 


- Eee EES ee Ee 


VoLUME 27, NUMBER 1 AT 


pattern of other members of this group. This is a species confined to more 
or less semi-arid mountainous terrain in the western section of the United 
States. 


7. Amblyscirtes cassus Edwards 1883 


Type locality: Mount Graham, Arizona. 

Distribution: UNITED sTaTEs. Arizona. Hannegan, VII-6-51. Cochise County: 
Chiricahua Mountains, VI-28-42, VII-1-51; Ramsey Canyon, VI-28. Pima County: 
Madera Canyon, VII-9-60, VII-10-64. New Mexico. Sandoval County: Jemez 
Springs, V. Texas. Jeff Davis County: Davis Mountains, VI-13-60. mexico. Baja 
California north, VI. Jalisco: Ajijic, X-7-65. Sonora, VI. 


Cassus prefers mountainous canyons for its normal habitat, with south- 
ern Arizona being the center of distribution. Rarely is it found in north- 
western Mexico. The orange cell area on the lower surface of the pri- 
maries as well as the double cell spot on the primaries makes this small 
species easy to recognize. 


8. Amblyscirtes aenus Edwards 1878 


Type locality: Southern Colorado. 

Distribution: UNITED sTATES. Arizona. Cochise County: Chiricahua Mountains, 
VII-10-60; Paraside, VII-10-60. Pima County: Madera Canyon, VII-6-60, VII-12-64; 
Mount Lemmon, VII-15-64. Colorado. Baca County: Regnier, VI. Boulder County: 
Boulder, V—V1; Bounder Canyon, VI; gluch south of Jamestown junction, V-31-54; 
Lefthand Canyon, VI-12-54; Mesa Trail, V-16-54; Six Mile Canyon, V-12-54; north 
Soda Springs, VI-8-53; Sugar Loaf Mountain, V-29-55. El Paso Canyon: Rock Creek, 
V-21-32. Jefferson County: Chimney Gulch, VII; Clear Creek Canyon, V-17-29; 
Coal Creek, VI-14-38; Plainview, 4-6, VII. New Mexico. Quay County: Tucumcari, 
VI-1-42. Sandoval County: Jemez Springs, VI. Texas. Armstrong County: Palo 
Duro Canyon, V-9-42. Brewster County: Alpine, IV-20-62, VI-16-60; Chisos Moun- 
tains, VIII-9-61, VII-10-62. Jeff Davis County: Davis Mountains, VI-10-60; Fort 
Davis, VI-9-42. Palo Pinto County: Palo Pinto, II-31-50. Mexico. Durango: Rio 
Nazas Valley near El Rodeo, VIII-18-22-68 (Peter Hubbell), AMNH, (new record 
for Mexico). 


This is a rather variable species as to the extent of the white discal 
spots on the lower surface of the secondaries. Some specimens have 
these spots clear white and very prominent, while others may have them 
dusky and almost obsolete. It occurs rather commonly over the south- 
western and midwestem parts of the United States, usually in the moun- 
tains. 


9. Amblyscirtes erna Freeman 1943 


Type locality: Palo Duro Canyon, Texas. 


Distribution: UNITED sraTEs. Kansas. Barber County: \-5-45, Oklahoma. 
Cimarron County: Black Mesa, V-3-47, V-21-49. Comanch ounty: Cache, VI- 
: >, ‘ ) : 

27-42. Woods County: Freedom, IV-21-46. Texas. Armstrong County: Palo Duro 


Canyon, IV-30-44, V-10-42, VII-28-42. Brewster Coun’ pine, VI-10-42. Gray 


48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


County: Lake McClellan, V-14-44. Roberts County: Miami, VIII-6-42. Mexico. 
Tamaulipas: Ciudad Victoria, VIII-15-16-62. 

Erna was placed by Evans as a synonym of fluonia Godman based on 
a specimen that I sent him plus one in his collection labelled Texas, ex 
coll Fruhstorfer. This was done without any knowledge of the biology 
of the two species and insufficient morphological data—actually the two 
are very distinct. Both fluonia and erna fiy in the mountains just west of 
Ciudad Victoria, Tamaulipas, Mexico, with erna occurring at slightly 
lower elevation than fluonia normally; however, they have been collected 
together in several locations. Erna prefers rocky canyon areas, whereas 
fluonia is usually located in shaded spots on the mountain sides. There 
are differences in the genitalia; however the easiest point of distinction 
between the two species is found on the lower surface of the secondaries, 
as fluonia has rather heavy overscaling giving the wing a mottled appear- 
ance, while erna has very light overscaling giving a uniform appearance 
to the wing. Usually the discal spots are better defined on this surface 
of the wing in erna as rarely are they discernable in fluonia. 


10. Amblyscirtes linda Freeman 1943 


Type locality: Hope Hill Farm, Faulkner County, Arkansas. 

Distribution: UNITED states. Arkansas. Carroll County: Eureka Springs, V-3-64. 
Faulkner County: Hope Hill Farm, VI, VII; Pinnacle Springs, VII-2-42. Missouri. 
Barry County: Cassville, V, VI. Oklahoma. Comanche County: Cache, VII-20-42, 
VIII-16-42. Washington County: Blue Springs, IV-67. 


This species was treated as a subspecies of aenus by Evans from which 
it is very distinct both morphologically and biologically. Morphologically 
the maculation is very different and the brands on the primaries are more 
prominent in linda than they are in aenus. Biologically linda is a woods 
species, whereas aenus is confined to more semi-arid, mountainous terrain. 


In its western range linda overlaps erna in the Wichita Mountain section 
of Oklahoma. 


11. Amblyscirtes oslari Skinner 1899 


Type locality: Chimney Gulch, Colorado. 

Distribution: UNITED states. Arizona. Pima County: Tucson, VII-6-60. Colorado. 
Archuleta County: Arboles, VI-25-55; Juanita, V-13-36. Boulder County: Boulder, 
VI; Boulder Canyon, VI-13-53; Four Mile Canyon, VI-7-53; gulch south of James- 
town Junction, V-31-54; Lefthand Canyon, VI-10-53; Six Mile Canyon, V-29-54. 
El Paso County: Williams Canyon, VII-7-31. Jefferson County: Chimney Gulch, 
V-28-18; Coal Creek; Golden, VI. Park County: Mill Gulch, V. Kansas. Barber 
County: IV-28-46, V-27-45. New Mexico. Sandoval County: Jemez Springs, VI. 
North Dakota. Slope County: Bad Lands, VI-11-61. Oklahoma. Woods County: 
Freedom, IV-21-46, VI-9-45. Texas. Armstrong County: Palo Duro Canyon, IV, V, 
19 Bay ‘or County: IV-30-70. Brewster County: Alpine, VI-5-42. Carson County: 

ite Deer, V-20-43. Jeff Davis County: Fort Davis, VI-3-40. 


VoLUME 27, NUMBER 1 49 


This species has a rather wide range over the midwestern section of 
the United States. It is readily recognized by the reduced maculation 
on the primaries and on the lower surface of the secondaries. It seems 
to prefer semi-arid, rather mountainous terrain for its natural habitat. 


12. Amblyscirtes fluonia Godman 1900 


Type locality: Mexico. 

Distribution: MExIco. Federal District: Zoquiapan, VIII-6-56. Guerrero: Amula; 
Chilpancingo; Xucumanatlan. Hidalgo: Jacala, VIII-1-63. Jalisco: Ajijic, VIII, IX, 
65; Lake Chapala. Michoacan: San Juan Pura, VI-27-47. Morelos: Cuernavaca, 
VII-28-61. Oaxaca: Oaxaca, VI-22-64. Puebla: Acatlan, VIII-20-64. Tamaulipas: 
Ciudad Victoria, VIII-62. 


Fluonia is readily separated from erna by its darker coloration and by 
the heavy, mottled, overscaling on the lower surface of the secondaries. 
Fluonia usually flies in areas of fairly high elevation. In the mountains 
west of Ciudad Victoria, Tamaulipas, erna usually occurs at an elevation 
of 2850 feet, whereas fluonia occurs from 3600 to 7000 feet. Fluonia is 
fairly common in the state of Jalisco as well as in the general vicinity of 
Oaxaca, Oaxaca. 


Aesculapinus Group 
13. Amblyscirtes elissa Godman 1900 


Type locality: Guerrero, Mexico. 

Distribution: Mexico. Chiapas: Acapetahua, III-60. Guerrero: Acahuizotla, 
VII-60; Dos Arroyos; Iguala, VIII-51; Rincon; Tierra Colorada. Morelos: Jantepec, 
VI-42. 


This small, dark greyish-brown, species can be readily recognized by 
the reduced maculation and grey stigma on the primaries, and by the 
lower surface of the secondaries being dark brown with tiny distinct 
white discal and cell spots present. Apparently elissa is a rather rare 
species confined mainly to Guerrero and Chiapas. 


14. Amblyscirtes samoset (Scudder) 1863 


Synonyms: hegon Scudder 1863: White Mts., New Hampshire. nemoris Edwards 
1864: Portsmouth, Ohio. argina Plotz 1884: “Brisbane.” 

Type locality: Massachusetts. 

Distribution: CANADA. Manitoba: Stone Mountain, VI. New Brunswick. Quebec. 
UNITED STATES. Arkansas. Faulkner County: Hope Hill Farm, [V-13-68. Connecticut, 
Avon, VI. Georgia. Fulton County: Indian Trail, Atlanta, [V-14-16-50. lowa. 


Grinnell. Maine. Mt. Desert Island, VI-5-31. Cumberland County: Portland, VI-38 
Kennebec County: Augusta, VI, VII. Penobscot County: Enfield, VI-15-37; Pas- 
sadumkeag Bog, VI-16-40. Piscataquis County: Baxter Park, Mt. Katahdin, VII- 
2-40. Massachusetts. Boston, VI. Minnesota. Aitkin, Carlton, Roseau, Lake of the 


Woods Counties, VI-67. New Jersey. Sussex County: Ogdensburg, VI. New York. 


50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Tompkins County: Oneonta, VIII-2-68. Ohio. Hamilton County: Cincinnati, V- 
15-38. 

This is a rather common and widespread species over the eastern part 
of the United States and southern Canada. It can be recognized by the 
general characteristics given in the key to the various species. The only 
specimens that I have collected came from Arkansas in a wooded area 
during the spring of the year. 


15. Amblyscirtes texanae Bell 1927 


Type locality: Sunny Glen Ranch, Alpine, Texas. 

Distribution: UNITED staTEs. New Mexico. Quay County: Tucumcari, VIII-25- 
41. Texas. Armstrong County: Palo Duro Canyon, IV-26-43. Brewster County: 
Alpine, V-31-42, VI-13-60; VI-2-42, VIII-7-61. Jeff Davis County: Davis Moun- 
tains, VI-13-60; Fort Davis, VI-11-49. 

This species is common in the Alpine and Davis Mountain sections 
of southwestern Texas. It is most often found in rocky ravines where it 
flies rapidly for short intervals then abruptly comes to rest on the rocks 
where it will remain for a short time then repeat the same procedure 
again. 


16. Amblyscirtes tolteca Scudder 1872 


Type locality: Tehuantepec, Oaxaca, Mexico. 

Distribution: Mexico. Guerrero: Acapulco, VII-36; Acuitlapan, VII-21-56; Iguala, 
VII; Rio Balsas, VI. Jalisco: Guadalajara, VII. Oaxaca: Candelaria Loxicha, VII-14-71; 
Tehuantepec, VIII-64. San Luis Potosi: Hotel Covadonga, 6 miles south Ciudad 
Valles, VI, VII, VIII; Tamazunchale, VII-63, VIII-24-67. Tamaulipas: Ciudad 
Mante, VI-9-41; Ciudad Victoria, VIII-16-62; El Solto, VIII-19-62, VIII-24-67. 
Veracruz: Jalapa, VI-64; Orizaba, VIII-67; Presidio, VIII. Yucatan: Valladolid. 


Tolteca is primarily a jungle species, however I have found specimens 
in brush environments in the state of Tamaulipas. In areas of dense 
vegetation most specimens will be found feeding on flowers that are 
usually shaded from the sun or else resting in the jungle shade. 


17. Amblyscirtes prenda Evans 1955 


Type locality: Tucson, Arizona. 

Distribution: UNITED sTATES. Arizona. Pima County: Tucson, VI-22-55. MEXICO. 
Chiapas: Comapapa, VII-24-69. Guerrero: Taxco, VII-1-36. Nayarit: Tepic, IX-64. 
Sonora: 7 miles southeast of Alamos, VIII-67; Guaymas. 

vans considered prenda to be a subspecies of tolteca, however from 
available information prenda appears to be a distinct species. Morphologi- 
cally prenda differs from tolteca in the maculation, and by being smaller 
size and lighter in coloration. There are slight differences in the 

| Biologically the two are very different as prenda occurs in arid 


VOLUME 27, NUMBER 1 51 


or semi-arid terrain very often flying in the heat of the day, whereas 
tolteca is a jungle species nearly always found in shaded areas. 


18. Amblyscirtes aesculapius (Fabricius ) 1793 


Synonyms: ftextor Geyer 1831: U. S. A. oneko Scudder 1863: Connecticut. 
wakulla Edwards 1869; Apalachicola, Florida. 

Type locality: North America. 

Distribution: UNITED staTEs. Alabama. Mobile County: Mobile. Florida. Orange 
County: Rock Springs, III-30-38. Georgia. Chatham County: Savannah, V-24-51. 
Kentucky. Jefferson County: VIII-9-69. Mississippi. Hinds County: Brownsville, 
IX-2-57; Clinton, VIII-25-56. Tennessee. Davidson County: Nashville, VIII. Texas. 
Harris County: Sam Houston National Forest, VI-6-71. Harrison County: Caddo 
Lake, VIII. Virginia. Nansemond County: Dismal Swamps, near Suffolk, VI-8-41; 
Jerico Ditch, near Suffolk, [IX-7-59; Magnolia, VII-17-59; Suffolk, VI-21-40. 

This very distinctive species is basically confined to the southern and 
southeastern part of the United States. It prefers wooded areas for its 


normal habitat. 


19. Amblyscirtes carolina Skinner 1892 


Type locality: South of Hamlet, Richmond County, North Carolina. 

Distribution: UNITED STATES. North Carolina. Gaston County: Gastonia, VIII- 
27-38. Richmond County: Hamlet, VIII. Virginia. Nansemond County: Great 
Dismal Swamps, VII; Suffolk, VII-1-40. 

Carolina seems to be confined to the North Carolina-Virginia area. 
This species is readily recognized by the heavy overscaling of dull yellow 
on the costa and apex of the lower surface of the primaries and the entire 
lower surface of the secondaries. The lower surface of the secondaries 
is indistinct rusty brown. This species is usually associated with swampy 
areas. 


20. Amblyscirtes reversa Jones 1926 


Type locality: Suffolk, Virginia. 

Distribution: UNITED sTATES. Georgia. Fulton County: Atlanta, V-12-56; Harris 
Trails, Atlanta, VII-17-55. Rabun County: V-4-55. North Carolina. Leland, VI- 
28-44, Virginia. Nansemond County: Sutfolk, V-28-45, VI-9-41, VII-20-59, VII-9-44. 

Reversa has long been considered to be a synonym of carolina or 
at most a form. I believe that actually it is a distinct species due to 
morphological differences such as the absence or obselete dull yellow 
overscaling on the lower surface of the wings, and the differences on the 
lower surface of the secondaries where the ground color is rusty brown 
and the maculation is dull yellow and distinct. There are also slight 
differences in the genitalia. This species ranges farther south than 
carolina being found in Georgia as well as in the same areas as carolina. 
I have both species from Suffolk, Virginia. 


52: JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Vialis Group 
21. Amblyscirtes nereus Edwards 1876 


Type locality: South Apache, Arizona. 

Distribution: UNITED sTATES. Arizona. Graham County: Mount Graham, VI. 
South Apache. New Mexico. Texas. Brewster County: Alpine, II-27-61, V1-2-42, 
VII-19-51. Jeff Davis County: Davis Mountains, VI-30-60, VII-27-53, VII-30-63, 
VIII-19-51; Fort Davis, VI-9-11-49. mexico. Chihuahua. Sonora. 


This is a distinctive species from the southwestern part of the United 
States and Chihuahua, Mexico. It is found in arid or semi-arid terrain 
and often in rocky ravines. It will often rest on greyish soil where its 
coloration blends well with its surroundings. 


22. Amblyscirtes nysa Edwards 1877 


Synonym: similis Strecker 1878: New Braunfels, Texas. 

Type locality: Texas. 

Distribution: UNITED STATES. Arizona. Pima County: Baboquivari Mountains, 
IX-50; Tucson, VII-10-60. Arkansas. Carroll County: Beaver, VI. Kansas. Douglas 
County: V-X. Greenwood County: Eureka, IX-1-40. Harper County: X. Mont- 
gomery County: V-X. Scott County: X. Shawnee County: VII. Summer County: 
Caldwell, X-2-41. New Mexico. Eddy County: Carlsbad Caverns, VI-10-58. Quay 
County: Tucumcari, VIJ-11-42. Texas. Armstrong County: Palo Duro Canyon, 
IV-26-43, V-16-42, IX-2-43. Bexar County: San Antonio, VI, VII, VIII, X. 
Brewster County: Alpine, VI-5-42; Sunny Glen Ranch, Alpine, VI-2-42. Cameron 
County: Brownsville, VI-6-71. Carson County: White Deer, IV-VI. Childress 
County: Childress, VIII-7-41. Comal County: New Braunfels, VI, VIII. Dallas 
County: Dallas, IV-3-38; Garland, VIII-7-71; Lancaster, VIII-1-41; Vickery, VI-16- 
40. Gray County: Lake McClellan, V-14-44. Hidalgo County: Pharr, V-30-47. 
Jeff Davis County: Fort Davis, VII-10-49. Terrell County: Sanderson, VII-12-49. 
Uvalde County: Uvalde, V-31-42. Val Verde County: Del Rio, VI-5-49. mexico. 
Northern Mesa. Nuevo Leon: Monterrey, VI-10-12-35. 


This small species is common over a large section of the midwest and 
southwestern sections of the United States and on down to the state 
of Nuevo Leon in Mexico. It is readily recognized by the variegated 
lower surface of the secondaries. Nysa is a familiar visitor to city flower 
gardens as well as rocky ravines in arid parts of the southwest. 


23. Amblyscirtes eos (Edwards) 1871 


Synonyms: comus Edwards 1876: Texas. nilus Edwards 1878: Texas. quinque- 
macula Skinner 1911: Las Cruces, New Mexico. 

Type locality: Dallas, Texas. 

Distribution: UNITED sTATEs. Arizona. Cochise County: Portal, VI-20-63. Santa 
Cruz County: Nogales, VII-11-64. New Mexico. Dona Ana County: Las Cruces, 
VI. Eddy County: Carlsbad, VII-9-49. Texas. Armstrong County: Palo Duro 
Canyon, IV-17-43, V-1-43. Brewster County: Alpine, VIII-19-57; Marathon, VI-3-40. 
Carson County: White Deer, VIII-23-41. Culberson County: Guadalupe Pass, VII- 
9-49. Dallas County: Dallas, VIII-7-50; Garland, IV-10-49; Lancaster, III-21-54. Jeff 


VoLUME 27, NUMBER 1 53 


Davis County: Davis Mountains, VII-27-53; Fort Davis, VI-11-49. Kerr County: Kerr- 
ville, VI-4-49. Palo Pinto County: Palo Pinto, VIII-13-55. Potter County: 15 miles north 
of Amarillo, VIII-10-41. Val Verde County: Del Rio, VII-12-49. mexico. Northern 
Sonora. Northern Mesa. 


Eos is associated with arid or semi-arid regions of the southwest. It can 
readily be recognized by the clear white, distinctive maculation on the 
lower surface of the secondaries. 


24. Amblyscirtes vialis (Edwards) 1862 


Synonym: dsella Herrich-Schaffer 1869: locality not known. 

Type locality: Rock Island, Illinois. 

Distribution: CANADA. British Colombia, Corfield, Vancouver, VI. Manitoba, 
Miniota, VI-16-37. Ontario. UNITED sTATES. Arkansas. Cleburne County: Quitman, 
VII-12-40. Faulkner County: Enders, VII-12-43; Hope Hill Farm, IV-20-45, IV- 
26-33, VI-12-40, VI-26-43, VI-27-44, VII-2-42; Pinnacle Springs, VII-1-43. Pulaski 
County: North Little Rock, VI-2-32. California. Sierra County: w. of Downieville, 
VI-17-67. Trinity County: Yosemite, VI. Colorado. Boulder County: Boulder 
Canyon, VI-19-41; Eldora, VI-24-33; Spring Gulch, VII-1-55. El Paso County: Bear 
Creek, V-31-32; Broadmoor, V-30-49; North Cheyenne Canyon, V-31-32; Rock 
Creek, VI-11-30, VII; William’s Canyon, VII-7-31. Grand County: Muddy Pass, 
VII-5-41. Jefferson County: Clear Creek Canyon, V-17. La Plata County: Junction 
Creek, VI-17-37; La Plata Mountains, VII-6-38. Larimer County: Rocky Mountain 
National Park, VII-5-35. Park County: Tappan Creek 6 mi. NW of Lake George, 
V-30-49. Florida. Georgia. Fulton County: Indian Creek Road, Atlanta, VI-6-57. 
Idaho. Priest Lake. Illinois. Mercer County: Perryton Township, V, VI, VII, 67. 
Scott County: Rock Island. Kansas. Douglas County: Lawrence, IV-6-67. Franklin 
County: VII-5-54. Greenwood County: Eureka, VIII-28-40. Pottawatomie County: 
VI. Scott County: VI. Maine. Penobscot County: Enfield, VI-11-39, VI-4-40: 
Passadumkeag, VI-1-36, VI-12-39. Caratumb, VI-2-41. Minnesota. Nicollet County: 
VIII-12-67. Mississippi. Tishomingo County: Tish State Park, IV-20-57. Missouri. 
Greene County: Willard, VII-13-38. New Hampshire. Randolph. Franconia, White 
Mountains. New Jersey. Woodbury; Elizabeth. New Mexico. Sandoval County: 
Jemez Springs, V, VI. New York. Rensselaer County: Berlin, V-30-41. Tompkins 
County: VI-40. North Carolina. Cranberry, VII. Ohio. Oklahoma. McIntosh 
County: Checotah, VII-25-41. Pennsylvania. Texas. Dallas County: Cedar Hill, 
IV-5-42, IV-7-44; Dallas, V-13-37; Garland, VIII-20-71; Lancaster, V-16-41; Vickery, 
VI-16-40. Vermont. Mt. Equinox. Virginia. Wisconsin. 


This is the most common and widespread species in the genus. In 
most areas where it is abundant it is usually associated with wooded 
areas. I have failed to locate specimens in arid or semi-arid habitats. 
Vialis is readily recognized by the distinct apical spots and absence or 
reduction of other spots on the primaries and by the uniform coloration 
on the lower surface of the secondaries. 


25. Amblyscirtes celia (Skinner) 1895 


Type locality: New Braunfels, Texas. 

Distribution: UNITED sTATEs. Texas. Bexar County: San Antonio, VI-5-96, 
VII-30-42. Cameron County: Brownsville, VI-8-40, VI-5-7! mal County: New 
Braunfels, VI-30-40, VII-2-60. Dallas County: Lancaste 3-40. Hays County: 


54 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


San Marcos, VI-12-40. Hidalgo County: Mission, VII-15-63, VII-31-45; Pharr, 
constant. Nueces County: Corpus Christi, VI. Val Verde County: Del Rio, VI-5-49, 
VII-12-49. mexico. Nuevo Leon: Monterrey, VI-19-35. San Luis Potosi: 6 miles 
south Ciudad Valles (Hotel Covadonga), VI-15-71. Tamaulipas: Ciudad Mante, 
VI-23-64; Ciudad Victoria, VI-23-64, VII-26-66, VIII-16-62; San Francisco, VIII-64. 

This species is usually found in wooded areas, very often in the shade. 
Celia has a rather restricted range as it occurs from the Lancaster, Dallas 
County, Texas area into northcentral Mexico. I have found celia rather 
abundantly in the vicinity of Monterrey, N. L., Mexico. 


26. Amblyscirtes belli Freeman 1941 


Type locality: Vickery, Dallas County, Texas. 

Distribution: UNITED states. Arkansas. Faulkner County: Enders, VII-12-43; 
Hope Hill Farm, VI-24-44, VII-24-44, VIII-1-44; Pinnacle Springs, VII-27-43. Pulaski 
County: Little Rock, VII-21-41, VII-21-43; North Little Rock, VII-5-41. Georgia. 
Madras, VII-23-34. Fulton County: Riverside Drive, Atlanta, V-21-57. IIlinois. 
Southern Illinois, VII. Missouri. Greene County: Willard, VII-25-29. North Carolina. 
Meck County Road, VII-26-70. Oklahoma. McIntosh County: Checotah, VII-25-41. 
Texas. Dallas County: Dallas, IV-12-38, IV-22-40, V-4-40, VII-20-42; Garland, 
IV-28-68; Lancaster, VIII-16-40; Vickery, VIII-5-40, VIII-16-40. 

Belli was recorded as a subspecies of celia by Evans, however there 
are a number of reasons why the two are separate. Biologically their 
habits are different as belli prefers open fields and the edges of woods, 
whereas celia prefers the woods. I have collected both species in the 
same general area at Lancaster, Texas, where their range overlaps. 
Morphologically they are easily separated as the males of belli have the 
spot in space 2 on the primaries V-shaped, while celia has this spot oval. 
On the lower surface of the secondaries celia usually has the discal spots 
lighter and more distinct than in belli where this area is usually rather 
dark and hoary. Celia often has a cell spot on the primaries which is 
completely lacking in belli. There are slight differences in the genitalia, 
however genitalic determinations in the genus Amblyscirtes are practically 
impossible with most species due to the fact that the basic pattern is 
very similar. 


27. Amblyscirtes alternata (Grote & Robinson) 1867 


Synonym: meridionalis Dyar 1905: Georgia. 

Type locality: Atlantic District, Georgia. 

Distribution: unirED states. Alabama. Whistler, IX. Houston County: Cowarts. 
Florida. Orange County: Orlando, III-17-42. Georgia. Scriven County: IV-9-46, 


V-18-46. North Carolina. Leland, VI-17-45. Texas. Smith County: Tyler, IX-5-49; 
Tyler State Park, III-24-59, IV-5-59. 


This small species is readily recognized by the general maculation 
which is characterized by the three apical spots being needle-like points, 


VOLUME 27, NUMBER |] 


Ol 
Ol 


and the indistinct discal spots. On the lower surface of the secondaries 
the wings are uniform dark brownish-black evenly overscaled with small 
grey scales. Specimens that I have collected were found in the general 
vicinity of piny woods. 


28. Amblyscirtes florus (Godman) 1900 


Synonym: mate Dyar 1923: Guerrero, Mexico. 

Type locality: Sierra Madre de Tepic, Nayarit, Mexico. 

Distribution: MeExico. Colima: Comala, VIII-4-67. Guerrero. Jalisco: Ajijic, 
IX-3-66; La Cumbre de Autlan, VII, VIII-67. Morelos: Jantepec, VI-49. Nayarit: 
Sierra Madre de Tepic. San Luis Potosi: 6 miles south of Ciudad Valles (Hotel 
Covadonga), VI-10-66, VIII-6-67. Tamaulipas: Ciudad Victoria, VIII-16-62; 15 miles 
south of Llera, VII-27-66. Veracruz: Catemaco, VIII-10-67. . 


This jungle species is characterized by its uniform brownish-black 
coloration, devoid of any maculation on the upper side. On the lower 
surface of the wings the primaries have very indistinct discal and apical 
spots and the secondaries have indistinct discal spots present. I have 
found florus to be rather abundant at times at Hotel Covadonga just south 
of Ciudad Valles, particularly in the jungle along the Rio Valles. 


Phylace Group 
29. Amblyscirtes anubis (Godman) 1900 


Type locality: Mexico. 
Distribution: MExIcO. Guerrero: Omilteme; Sierra Madre del Sur. Hidalgo: 
Apulco, IV-52. Veracruz: Jalapa; Orizaba. 


This rather rare species can be recognized by the orange-yellow fringe 
of the primaries and the concolorous fringe of the secondaries. The palpi 
are grey in both sexes. There is a narrow broken stigma on the primaries 
of the males. 


30. Amblyscirtes phylace (Edwards) 1878 


Type locality: Southern Colorado. ; 

Distribution: uNrrTED states. Arizona. Cochise County: VIII-99. Colorado. El 
Paso County: Rocky Creek, VII-7-37. Jefferson County: Chimney Gulch; Clear 
Creek Canyon, V-26-21; Lookout Mountain, VI-25-39. Park County: Mill Gulch, 
VI-10-21. Teller County: Rosemont, VI-29-32. New Mexico. Dona Ana County: 
Rincon, VI. Sandoval County: Jemez Springs, VI-9-14, VI-26-14. Texas. Jett Davis 
County: McDonald Observatory, VI-9-49, VII-11-49, VIII-5-62, VIII-10-60. MExico 
Morelos. Puebla: La Malinche. 


This species can be identified by the sordid white fringe of both wings. 

7 . . ir > | »% Are Tir , 
In the males the palpi are orange, while in the female’ are yellowish 
white. The males have a broad, short brand cov - vein 2 near its 


oorits ° ° : (ae Ie (ami-ariad 1) < 
origin on the primaries. Phylace is usually tounc » M-arid mountains, 


56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


31. Amblyscirtes fimbriata (Plotz) 1882 


Synonym: bellus Edwards 1884: Southern Arizona. 

Type locality: Mexico. 

Distribution: UNITED srATEs. Arizona. Cochise County: Chiricahua Mountains, 
VI-28-36, VI-26-36; Onion Saddle Pass, VII-12-60; Pinery Canyon, VII-11-60; Portal, 
VII-10-60; Ramsey Canyon, VI-28-36; Rustlers Park, VI-20-63. New Mexico. Sandoval 
County: Jemez Springs. mexico. Las Vigas. Durango: Milpas. Nuevo Leon: 
Chipinque Mesa, Monterrey, VIII-13-67. Sonora. Valle de Mexico. Veracruz: Jalapa. 


This species can be recognized easily by the presence of an orange 
fringe on both wings in both sexes, also by the palpi being orange in 
both sexes. The males have a narrow, broken, grey stigma from the 
origin of vein 3 to vein 1. Fimbriata usually occurs in wooded moun- 
tainous terrain. 


BIBLIOGRAPHY 


BARNES, W. & F. H. Benjamin. 1926. Check list of the diurnal Lepidoptera of 
boreal America. Bull. S. Calif. Acad. Sci. 25: 3-27. 

Bett, k. L. 1927. Description of a new Amblyscirtes from Texas (Lepidoptera, 
Rhopalocera, Hesperiidae). Bull. Brooklyn Entomol. Soc. 22: 203-204. 

1938. The Hesperioidea. Bull. Cheyenne Mountain Mus. 1: 1-35. 

Brown, F’. Martin, D. Err & B. Rorcer. 1957. Colorado Butterflies. Proc. Denver 
Mus. of Nat. Hist. 368 p. 

Ciark, A. H. 1932. The Butterflies of the District of Columbia and Vicinity. U. S. 
Nat. Mus. Bull. 157. 337 p., 64 pl. 

Evans, W. H. 1955. A catalogue of the American Hesperiidae indicating the 
classification and nomenclature adopted in the British Museum. Part IV. 
Hesperiinae and Megathyminae. London: British Museum. 449 p., pls. 54-88. 

Frecp, W. D. 1938. A Manual of the Butterflies and Skippers of Kansas. Bull. 
Univ. of Kans., Dept. of Ent. 39, 328 p. 

FREEMAN, H. A. 1939. The Hesperiidae of Dallas County, Texas. Field & Lab. 
18 PAI s). 

1941. A new species of Amblyscirtes from Texas. Entomol. News 52: 50-51. 

1942. Notes on some North American Hesperiidae with the description of 

a new race of Polites verna (Edwards). Entomol. News 53: 103-106. 

= aes Two new species of Amblyscirtes from Texas. Entomol. News 54: 

1943b. New Hesperioidea, with notes on some others from the United 

States. Entomol. News 54: 72-77. 

1945. The Hesperiidae of Arkansas. Field & Lab. 13: 60-63. 

——. 195la. New skipper records for Mexico. Field & Lab. 19: 45-48. 

1951b. Ecological and systematic study of the Hesperioidea of Texas. 

So. Meth. Univ. Studies. 6: 1-64. 


1970. A new genus and eight new species of Mexican Hesperiidae. J. N. Y. 
Entomol. Soc. 78: 88-99. 

( /ODMAN, F. D. & O. Satvin. 1887-1907. Biologia Centrali-Americana. Insecta, 
Lepidoptera-Rhopalocera. II: 244-637; LII: plsy 2) 

HOFFMANN, C.C. 1941. Catalogo sistematico y zoogeografico de los Lepidopteros 
Mexicanos, Segunde parte- Hesperioidea. An. Inst. Biol. Mexico. 12: 237-294. 


HoLLAnp, W. J. 1931. The Butterfly Book. New and thoroughly revised edition. 
New York. 424 p., 77 pls. 


VOLUME 27, NUMBER 1 57 


LinpsEy, A. W. 1921. MHesperioidea of America, north of Mexico. Univ. Iowa 

Studies Nat. Hist.9: 1-114. 
1928. Hesperia eos Edwards. Entomol. News 39: 91-93. 

, EK. L. Bett & R. C. WiitiaMs, Jr. 1931. The Hesperioidea of North 
America. Denison Univ. Bull. 31(2); (subtitle). J. Sci. Labs. 26(1): 1-142. 
33 pls. 

McDunnoucu, J. 1938. Check List of the Lepidoptera of Canada and the United 
States of America. Part 1, Macrolepidoptera. Memoirs S. Calif. Acad. Sci. 
Ik, PAG oy 

Seitz, A., ed. 1907-1924. The Macrolepidoptera of the World. Vol. V., The 
American Rhopalocera. Stuttgart. 1139 p., 203 pls. Various authors. 

SKINNER, H. & R. C. WituiAMs, Jr. 1923. On the Male Genitalia of the Hesperiidae 
of North America. Paper III. Trans. Amer. Entomol. Soc. 49: 129-153. 


A REVISION OF THE COLIAS ALEXANDRA COMPLEX 
(PIERIDAE) AIDED BY ULTRAVIOLET REFLECTANCE 
PHOTOGRAPHY WITH DESIGNATION OF A NEW SUBSPECIES! 


CuirFrorD D, FERRIS? 
College of Engineering, University of Wyoming, Laramie, Wyoming 82070 


This paper presents a study of the distribution and taxonomy of the 
Colias alexandra complex. The role of ultraviolet photography as an aid 
to taxonomic studies is discussed and is employed in assigning C. 
alexandra populations to various color groups. Visible light characters 
(pigmentation and facies) are combined with uv reflectance patterns to 
arrive at the taxonomic conclusions presented. One concludes from this 
study that some populations of alexandra can be assigned to specific sub- 
species, while others are best listed as clinal or intergrade forms. Based 
upon uv photography, C. harfordii and C. barbara are assigned to the 
alexandra complex. As a consequence of recent work by Brown (1973), 
a new subspecies of alexandra is proposed. 

Butterfly color patterns are produced by both pigmentation and optical 
effects. The brilliant prismatic colors associated with many tropical 
species are produced by visible light interference with the structures of 
certain wing scales. As shown by po a Torshnya boy (1954) and 
Nekrutenko (1964), certain Coliadinae reflect ultraviolet light from 


particular wing areas such that interference patterns are produced. 
1 Published with the approval of the Director, Wyoming Agricu! Experiment Station, as 
Journal Article no. JA 506. 


2 Research Associate, aie Museum of Entomology, Sarasota, I 


58 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Ghiradella, Eisner, Hinton, and Silberglied (pers. comm. and in review ) 
have determined that these uv reflection patterns are structural and are 
analogous to the white-light-produced brilliant irridescent blues in the 
genus Morpho. Silberglied (pers. comm.) has shown that interference of 
uv light rays in the layered lamellae which comprise the ribs of special 
wing scales is responsible for the “luminous” patches shown in the ac- 
companying figures. Close spacing of the ribs is indicative of strong uv 
reflection. 

Kolyer & Reimschuessel (1969) reported some scanning electron micro- 
scope studies of Colias eurytheme Boisduval but did not interpret the 
structure of the scales. The lamellae are shown in Figs. 2c & d of their 
paper. A simple method for making uv photographs has been described 
by Ferris (1972b ). 

Ultraviolet reflectance photography can be used as a taxonomic aid 
as suggested by Nekrutenko (1964). Some species of Colias are reflective; 
others are not. Reflection is used here in a relative sense and is applied 
to fresh undamaged specimens. There is always some reflection of uv 
light, but only certain species reflect sufficient energy to produce bright 
patterns. Reflection in Colias generally occurs from the discal areas 
(dorsal) of the secondaries and varies considerably for the primaries. 
Males of certain species are reflective, while the females of the North 
American species are non-reflective. Ultraviolet photography of the non- 
reflective species is of no taxonomic value except to separate reflective 
and non-reflective species in questionable cases. 

Colias alexandra males exhibit a uv reflectance pattern which appears 
as a luminous patch on the secondaries and is constant in all of the color 
forms. The term “luminous” is used here to describe the appearance of 
the reflection pattern in a black-and-white photograph. The amount of 
reflectance from the primaries varies from insignificant in the pure yellow 
races to considerable in the orange races. Fig. 3 illustrates the features 
which separate alexandra from other North American Colias. Figs. 4-5 
illustrate examples of C. alexandra as they appear under white light 
photography and uv photography. A dull background has been used 
purposely to eliminate spurious uv fluorescence. 

Ultraviolet photography is used here to assign alexandra populations 
to various color groups. It shows that several populations which appear 
yellow to the human eye, exhibit uv interference patterns characteristic 
of the yellow-orange group. These populations are therefore placed with 
the yellow-orange group rather than with the “pure” yellow group. Uv 
photography cannot be used to make assignments at the subspecies level 
generally, although it does show that C. barbara and C. harfordii belong 


the alexandra complex. 


VOLUME 27, NUMBER 1 59 


Fig. 1. Distribution of Colias alexandra in North America. The outer solid line 
encloses the known areas in which alexandra has been found. The northern boundary 
is still in doubt as indicated by (?). The shaded areas represent distinct subspecies 


as follows: 1, unnamed Alaska-Yukon segregate; 2, christina; 3, columbiensis; 4, 
astraea; 5, krauthii; 6, alexandra; 7, edwardsii; 8, barbara and harfordii; 9, Arizona- 
New Mexico segregate. The remaining areas within the boundary represent intergrade 


forms which cannot be clearly identified as any one given taxon. 


Biology 
The life histories of several members of the alexandra group have been 
published and are cited in Davenport & Dethier (1937). Larval food- 


plants are members of the Leguminosae. There is a paucity of specific 


60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


hostplant records in the literature, but these records include: C. alexandra: 
Astragalus, Thermopsis, Trifolium repens (Davenport & Dethier, 1937); 
Astragalus serotinus (Opler, unpublished ); A. miser (Shields, et al., 1969). 
C. christina: Trifolium (Davenport & Dethier, 1937). C. harfordii: 
Astragalus (Davenport & Dethier, 1937); A. antisellii (Locoweed) (R. C. 
Priestaf, 1972, pers. comm.). Davenport & Dethier list additional authors 
who have reported hostplant preferences for alexandra. 

Colias alexandra is found in a wide variety of habitats. Generally it 
frequents open areas, and in forested land is found in clearings and along 
roads or cuts. Males may be found at puddles along dirt roads where 
they sometimes congregate in large numbers. Some of the subspecies are 
common in open sagebrush regions (Upper Sonoran Desert), while 
others frequent the Transition Zone (aspen-conifer association ), and still 
others are found in meadows or clearings in the Canadian Zone. In the 
Far North, alexandra appears to prefer open clearings in the taiga (spruce- 
scrub biome). Pigmentation in the adults does not appear to be cor- 
related with habitat. To some extent, correlation with latitude exists, 
with more orange color appearing in the north. 


Distribution and Taxa 


The C. alexandra complex is widely distributed in western North 
America (Fig. 1). Three distinct color forms are recognized: yellow, 
yellow and orange, and orange, in addition to clinal forms in which 
specimens from a given geographic location vary from yellow into almost 
pure orange (Ferris, 1972a). Currently recognized taxa belonging to the 
alexandra complex are indicated below according to visible-light color 
(pigmentation ). 


YELLOW POPULATIONS—TAXA 


Colias alexandra alexandra Edwards, 1863 [T. L. Front Range, west of Denver, 
Colorado]. 


Colias alexandra edwardsii Edwards, 1870 [T. L. Virginia City, Storey Co., Nevada]. 
Colias alexandra emilia Edwards, 1870 [T. L. Oregon]. See discussion below. 


YELLOW-ORANGE POPULATIONS—TAXA 


Colias alexandra astraea Edwards, 1872 [T. L. Yellowstone Lake, Wyoming]. 
Colias alexandra christina Edwards, 1863 [T. L. Slave River Crossing, N.W.T., 


Canada]. 
ORANGE POPULATIONS—TAXA 
Colias alexandra krauthii Klots, 1935 [T. L. Black Hills, 12 miles west of Custer, 
Custer Co., South Dakota]. 
he taxon alberta Bowman has been omitted as it appears to describe 
tybric’ situation and suppression of this name has been recommended 


VOLUME 27, NUMBER | 


A 
A 
\ 
— 1 
c (\ 
s° i a | 
@ \ 
. @ ‘ ~ 
ean % g ad 
‘ ~ 1 
o,! Ts 
é ' ie 
4 i SSIS — 
a aie 
| : 'o y 
’ 2¢e 7 
r) ! ' 
‘ UJ | Fo 
\ e ! ! { 
a 1 i} t 
t 1 ! 
ue 1@ , 3 
° tak 
Bee a ees } ' es 
Oo fa f D 
oo! vn --- porn 
ora. ! ! : 
t ~S = beeen, 
aan oNo, | i sis 
( om! i aed a | Q, 
\ ! U Fe iciciesiowien SS 
Sn ! ' ' 
ue "ee Sy isan qt Sop od 
‘ 1 | 
iv ! ! 
] ] 
' 1 
eal 


Fig. 2. Distribution of Colias alexandra isolates. Open circles—yellow populations: 
half-open circles—yellow-orange populations; solid circles—orange populations. 


(Ferris, 1972a). This and other possible alexandra crosses are discussed 
in the paper cited. 
When long series of alexandra of a given subspecies or from a given 


locality are examined, one notices substantial variation in pigmentation 
and, in the females, maculation. Some individual males from yellow- 
orange races appear yellow under white light. When photographed under 


uv illumination, they exhibit luminous patches o primaries which 


61 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


36% 


62 


(c) 


(b) 


(a) 


1 


a 
x) 


(d) 


(g/ 


VoLUME 27, NUMBER l 63 


are characteristic of the yellow-orange group. In the current study, uv 
photography is used to assign various races or populations to one of the 
three designated color groups. Race or population should not be inferred 
as synonymous with subspecies. The uv patterns for each group are 
discussed below. 

There is considerable variation in uv reflectance pattern as well as in 
pigmentation. The former is illustrated by the accompanying figures. 
Only C. a. alexandra and C. a. krauthii, the poles so-to-speak, exhibit 
minimal variation. The angle at which uv radiation strikes the wing 
surfaces affects the reflection pattern (Nekrutenko, 1965). If specimens 
are flat-mounted and illuminated as suggested by Ferris (1972b), this 
problem is minimized. 

In the following paragraphs, reference is made to various isolated 
populations. These represent clinal and intergrade forms which do not 
merit subspecific recognition. Localities are shown in Fig. 2. 


YELLOW POPULATIONS—DISTRIBUTION 

Arizona, California, Colorado, Nebraska, Nevada, New Mexico, Oregon (isolate), 
Utah, Wyoming, Montana (?). 

The yellow races are distinguished by lacking forewing luminosity (or 
exhibiting only a trace at most) under uv illumination and by having 
yellow (concolorous with the ground color) discal spots on the dorsal sur- 
face of the secondaries in the males. This definition differs from previous 
ones which included populations that have orange discal spots. Generally 
the orange-spotted specimens exhibit forewing luminosity. 


YELLOW-ORANGE POPULATIONS—DISTRIBUTION 


California, Idaho, Montana, Nevada (Elko, Nye, Washoe Cos.), Oregon, Utah, 
Washington, Wyoming, Alberta, British Columbia, Manitoba, Northwest Territories, 
Saskatchewan. Recently a single orange-discal-spotted male alexandra was collected 
by M. S. Fisher (Parker, Colorado) in Elbert Co., Colorado, an eastern plains region 
of the state. Further collecting is necessary to ascertain if this specimen is from a 
yellow-orange isolate with possible affinity to the Black Hills krauthii, or a hybrid 
with philodice or eurytheme. Undoubtedly other yellow-orange isolates, not shown 
in Fig. 2, will be found as collectors penetrate into little-collected areas. 


< 


Fig. 3. General extent of luminous patches as they appear on the wings of the 
males in the Colias alexandra complex. Hindwing patches in (a) yellow group; 
(b) orange group; (c) yellow-orange group. The discal spots, shown as open circles, 
are generally black in uv photographs. Forewing patches in (d) transition vellow to 


yellow-orange populations (submarginal band); (e) some yellow-orange populations 
(submarginal band and portions of some cells near veins); (f) other yellow-orange 

1 = c Yi RASS eee we ack awmagcc \. \ 
populations (central portion of wing generally reflects \ some dark areas); (g 


orange populations (wing reflects almost uniformly excep! £0 marginal areas). 


SSK 
\. 
\ 


SOCIETY 


> 


TAL OF THE I[LEPIDOPTERISTS 


st 
oe) 


A 


VOLUME 27, NUMBER l 65 


Some of the yellow-orange races appear yellow under visible light, 
except that the discal spot on the dorsal surface of the hindwings is 
orange. Individual male specimens may show a dark yeliow or a pale 
yellow-orange flush discally and limbally on the upper side of the 
secondaries and females may exhibit considerable orange. Under uv 
illumination, luminous patches appear on the forewings. Individuals of 
C. a. astraea may appear to be pure yellow except for the orange discal 
spot, but this subspecies as a whole ranges from yellow to orange. For 
this reason, populations with orange discal spots in the males are classified 
in the yellow-orange group based upon uy patterns and not visible light 
(pigmentation ) appearance. ? 

Southern Alberta appears to represent a complex blend-zone region. 
In the area from Calgary to Banff and east of the Rocky Mountains, speci- 
mens can be taken which represent alexandra, astraea, christina, and 
krauthii. It is frequently possible to collect two or three good “subspecies” 
at the same locality. This situation is typical of the intergrading which 
occurs in the alexandra complex and is the reason for the restricted ranges 
shown in Fig. 1. 

Northern Utah specimens, especially from Tooele and Wasatch Coun- 
ties, tend toward both astraea (in the males) and christina (in the fe- 
males ). The latter frequently show an overwashed orange coloration. 

Specimens of alexandra from Nevada have generally been determined 
to be subspecies edwardsii. A small series in the collection of the Los 
Angeles County Museum of Natural History taken by A. O. Shields in 
Jett Canyon, Toiyabe Range, Nye Co., Nevada is clearly from a yellow- 
orange population. The discal spots (dorsal secondaries) are orange in 
both sexes and distinct luminous patches show on the forewings of the 
males under uv light. Some specimens from the same locality are pheno- 
typically edwardsii. Peter Herlan (Carson City, Nevada) has found yel- 
low-orange populations in Elko and Washoe Counties as well. The oc- 
currence of these isolates is as yet unexplained. 

Califomia specimens from Lassen (Blue Lake area, Warner Moun- 


< 


Fig. 4. Specimens of Colias alexandra photographed under white (left) and 
ultraviolet (right) light. a & b, C. a. alexandra, Albany Co., Wyoming; ¢, normal &, 
white 9. ec & d, 2 2 of C. alexandra; top, Tooele Co., Utah; bottom, Catron Co., 
New Mexico. e & f, C. alexandra; top ¢, Boundary Co., Idaho; bottom pair, S of 
Golden, British Columbia. g & h, C. alexandra, pair from Utah: Tooele Co., 9 
Wasatch Co. i & j, ¢ 6; top, C. a. “emilia,’ Okanogan Co., Washington (see text); 
middle, C. a. edwardsii, Lander Co., Nevada; bottom, segregate, Apache Co., Arizona. 
k &1, 6 ¢ of C. a. astraea; top, Sublette Co., Wyoming; » f Seebee, Alberta 
in blend-zone region; bottom, Sheridan Co., Wyoming (or rm ) 


66 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


VOLUME 27, NUMBER 1 67 


tains ) and Modoc Counties show the orange discal spot in the males and 
considerable orange scaling in the females. Under uv light, luminous 
patches show on the forewings of the males. 

Oregon specimens are similar, with the exception of a population 
from the Canyon Creek area, Ochocho Mountains, in Crook Co. The 
males from this region are similar to material from British Columbia, but 
the females show considerable dark bordering as in occidentalis Scudder. 
Some are quite similar in pattern and color to this species. Perhaps some 
hybridizing has occurred, but this is speculation. C. occidentalis is a 
non-reflective species. 

The northern Idaho—southern British Columbia segregate is a large 
insect, generally larger than nomenotypical alexandra. This is described 
as a new taxon below. At first, it can be mistaken for gigantea Strecker 
because of the forewing apical rounding, but its habitat is forest clearings 
and roads, not bogs. McDunnough (1928) called this butterfly Eurymus 
emilia. Initially, one would place the population with the yellow races, 
but the discal spot is orange and the forewings exhibit luminous patches 
under uv illumination (Fig. 4f; Fig. 6e, £). This butterfly does not fit 
Edwards's description of emilia, and F. M. Brown (1973) has shown that 
emilia is synonymous with edwardsii which has page priority. 


ORANGE POPULATIONS—DISTRIBUTION 

South Dakota, Wyoming, Montana (?), North Dakota (? reported by Opler, un- 
published), Alberta, Manitoba, Yukon Territory, Alaska. 

A butterfly has been collected in the Yukon Territory (along the Alaska 
Highway ) and in Alaska which appears to be a member of the alexandra 
complex (Fig. 5f-h). In many respects, it is similar to Colias hecla 
Lefébre, but the underside and the uv reflectance pattern from the 
upperside place it tentatively as alexandra. Private correspondence with 
other collectors indicates that F. H. Chermock may have intended to name 
this population. 


< 


Fig. 5. Specimens of Colias alexandra photographed under white (a, c, g, i & k) 
and ultraviolet (b, d, e, f, h, j & 1) light. a & b, 92 of C. a. astraea; top, Johnson 
Co., Wyoming; middle and bottom, S of Seebee, Alberta in blend-zone region. ¢ & d, 
C. a. krauthii, Lawrence Co., South Dakota; top, orange 4; middie, ¢ showing some 
yellow; bottom, @ (note the luminous patches on the female). e, C. a. christina: 
top, 6, Riding Mtns., Manitoba; middle and bottom, pair, S of Seebee, Alberta in 
blend-zone region (note slight luminous patches on forewings of © ). £, C. alexandra 
(?):;.3 64: and 1 @, Yukon Territory. g & h, C. a. christina; top, ¢; middle, Q; 
both S of Seebee, Alberta in blend-zone region; C. alexandra (?); bottom, ¢, Steese 
Highway mile 111, Alaska. i & j, C. a. harfordii; pair, ! »., California. k & I, 
C. a. barbara; Santa Barbara Co., California. 


68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Pure orange races of alexandra have been reported from the area near 
Beulah, Manitoba, and Pocohontas, Alberta. These may be referable to 
the taxon krauthii. The Alberta specimens superficially resemble krauthii, 
but are considerably smaller. 


Taxonomic Studies 


Colias alexandra barbara H. Edwards, new combination 
Colias alexandra harfordii H. Edwards, new combination 


In 1877, Henry Edwards published a paper in which he proposed 
names for two Colias from California. These are the taxa barbara [T. L. 
Santa Barbara, California] and harfordii [T. L. Havilah, Kern Co., and 
Contra Costa Co., California]. They have stood as distinct species until 
P. A. Opler (unpublished) placed barbara as a subspecies of harfordii, 
although Talbot (1935) listed barbara as a form of harfordii. Edwards 
did indicate that both insects were related to alexandra. 

Based upon uv photographs (Fig. 5i-]), it appears that the affinity of 
both butterflies is with alexandra. These subspecies appear to represent 
an intermediate situation. The uv reflectance from the forewings is 
reduced to a trace, as in the yellow populations, but the secondary discal 
spot is orange, as in the yellow-orange populations. 

Additional justification for this assignment lies in range and foodplant. 
C. a. edwardsii is considered rare in California, although it is locally 
common in nearby Nevada (Lander Co.). Another population ( previously 
discussed ) is found locally in Lassen and Modoc Counties. Since alexandra 
is known to the north of central California, it seems odd, based upon 
geology, ecology and geography, that it should not occur centrally and 
along the coast of southern California. The southern and middle coastal 
areas and part of the central portion of the state are the areas in which 
barbara and harfordii occur. As noted earlier, harfordii uses Astragalus 
as a larval hostplant, which is also true of alexandra. Thus from the uv 
reflectance pattern, range, and hostplant affinity, it appears reasonable 
to assign barbara and harfordii to alexandra. 


Colias alexandra columbiensis Ferris, new subspecies 


Browns treatment of emilia (1973) leaves the British Columbia race of 
alexandra without a name. The name columbiensis, derived from the type 
province, is proposed for this butterfly. This subspecies differs from 
other alexandra subspecies in that the apices of the forewings are defi- 
nitely rounded suggesting gigantea. The uv reflection pattern in the males 
places this insect in the yellow-orange group. Comparison with other 


VOLUME 27, NUMBER 1 69 


Fig. 6. Colias alexandra columbiensis Ferris: a, male holotype (upperside); b, 
same (underside); c, yellow female paratype (upperside); d, white female paratype 
(upperside); e, uv photograph of male holotype; f, same, but with specimen tilted 
to show full extent of forewing pattern on upperside. 


members of this group shows that columbiensis differs from christina in 
that the forewings of the males show no orange color. It differs from 
astraea by being much larger, paler yellow in overall color, and is totally 
different in the females. Columbiensis females are pale yellow or white 
with nearly immaculate borders and generally show a large orange discal 
spot on the upperside of the hindwings. 


70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


In the males, the dorsal ground color is bright lemon yellow. Some 
specimens exhibit a dark yellow to orange flush in the discal and limbal 
areas of the hindwings, but do not show the pronounced orange markings 
which frequently occur in astraea and always in christina. The secondary 
discal spot is orange. The black borders are generally narrower than in 
nomenotypical alexandra. The primary cell-end spot is quite narrow. 
Ventrally the ground color is yellow with a slight orange flush. There 
is a dusting of black scales (sometimes heavy) on the secondaries. The 
secondary discal spot is bordered with dark pink scales and has a pearly 
center. Occasionally there is a satellite spot. The wing fringes are pink 
with some yellow as in astraea. 

The females are dimorphic as is the case with other races of alexandra. 
Both yellow and white forms occur, as well as intermediates. The yellow 
females have a lemon yellow ground color frequently overwashed with 
pale orange, less pronounced dorsally than ventrally. In the white forms, 
there may be pronounced yellow-orange overwashing. Dorsally in both 
forms, the dark bordering varies from absent to slight. The primary 
cell-end spot is distinct (more so than in the males). The secondary discal 
spot is bright orange in the yellow forms and varies from orange to white 
in the white forms. Ventrally, the females are generally similar to the 
males, although there is a heavier overscaling of dark scales and the 
ground color is lighter in the white forms. 

This subspecies is generally larger in size than the nominate species. 
The forewing costal margin length of the holotype male is 26 mm, 29 mm 
for the yellow female, and 28 mm for the white female shown in Fig. 6. 
In some males from northern Idaho, the costal margin length is 32 mm. 
Male specimens of C. a. alexandra examined from the Front Range 
(Rocky Mtns.) area measured 23 to 25 mm. 

The holotype and two female paratypes are shown in Fig. 6. In addi- 
tion, the uv reflection pattern of the holotype is presented. It is typical 
of the yellow-orange group. 


Type Series. The type series consists of 6 males and 13 females. Because of the 
female dimorphism, no allotype is designated. 

Holotype 6. The specimen bears two labels. The locality label is machine 
printed black on white, with the exception of part of the date which is handlettered 
in black ink, and carries the following data: Anderson Lake/D’Arcy, B.C./17 June 
1926/J. McDunnough. A second red label, machine printed in black is inscribed: 
Colias alexandra/columbiensis Ferris/Holotype Male. 

Paratypes. 5 6 6, same data as holotype. 1 @ (white), same data as holotype. 
9 29, 100 Mile House, B.C.: 28 June 1938, 4 @@ (white); 29 June 1938, 1 9 
(yellow); 30 June 1938, 3 992 (1 yellow); 4 July 1938, 1 @ (white), leg. J. K. 
Jacob and G. S. Walley. 2 9 2 (white) Lac la Hache, B.C., 5 July 1938, leg. G. S. 
Walley. 1 2 (white) Canin Lake, B.C., 24 June 1938, leg. G. S. Walley. 

Distribution. This subspecies is found in British Columbia south to Washington 


VOLUME 27, NUMBER 1 iat 


(Okanogan Co.) and northern Idaho (Boundary Co.). To the southeast of this 
region, it intergrades with astraea and to the south (southern Washington, Oregon, 
and northern California), it intergrades with edwardsii and possibly harfordii. The 
Canadian Rocky Mountains appear to form an effective barrier against any significant 
intergrading between columbiensis and christina. Specimens collected in the Bitter- 
root Mtns., Ravalli Co., Montana exhibit characters associated with both astraea and 
columbiensis. : 

Colias alexandra columbiensis is figured in Holland (1931), Plate LXVIII, figs. 
22, 23, as C. emilia. The orange discal spot in the male is poorly reproduced. 
The specimens shown were collected by Greene in 1894 at Osyoos, British Columbia 
and are in the Carnegie Museum collection. They came to Holland from W. H. 
Edwards who labeled them as emilia, even though they do not fit his description of 
the taxon. Wright (1907) also figures emilia, Plate XI, fig. 92, but at least two of 
the examples shown are probably philodice enpnale ewandls. 

The type series for columbiensis is placed in the Canadian Newel Collection, 
Ottawa, Ontario. 


CONCLUSIONS 


It is suggested that the taxa associated with the Colias alexandra com- 
plex be arranged as follows: 


Colias alexandra alexandra Edwards 
Colias alexandra edwardsii Edwards 
Colias alexandra harfordii H. Edwards 
Colias alexandra barbara H. Edwards 
Colias alexandra columbiensis Ferris 
Colias alexandra astraea Edwards 
Colias alexandra christina Edwards 
Colias alexandra krauthii Klots 


Unnamed races which possibly merit nomenclatural recognition: 


Colias alexandra Arizona-New Mexico Segregate 
(yellow population ). 

Colias alexandra Yukon Territory-Alaska Segregate 
(orange population ). 

The arrangement is roughly according to pigmentation. The taxa 
alberta and emilia are omitted for the reasons set forth above. Other 
aspects of the alexandra complex have been treated by Ferris (1972a). 

Ultraviolet reflectance photography has been used in this study to 
assign the various alexandra populations to specific color groups. It has 
also been used to identify barbara and harfordii as members of the 
alexandra complex. 


ACKNOWLEDGMENTS 


. ° . ASST BEE Os Last ore —— 
The author expresses his appreciation to the following collectors fo1 
providing material for study: P. J. Conway, J. D. Eff, J. R. Heitzman, 


72 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


J. A. Legge, Jr., J. R. Mori, R. C. Priestaf, J. T. Sorensen, K. B. Tidwell, 
M. Toliver, and R. E. Woodley. Loan specimens from museums were 
supplied by J. P. Donahue, Los Angeles County Museum of Natural 
History and G. E. Ball and D. R. Whitehead, University of Alberta. 

Mr. George Lewis of the Canadian National Collection, Ottawa, kindly 
provided specimens for study from British Columbia, some of which were 
designated in the type series for columbiensis. F. M. Brown and R. E. 
Silberglied kindly read and commented upon preliminary drafts of the 
paper and both generously made results of their studies on Colias avail- 
able. A. B. Klots of the American Museum of Natural History provided 
collection data for Manitoba populations. Special thanks are due H. K. 
Clench, Carnegie Museum, for providing a copy of a scarce paper by 
Henry Edwards. 


LITERATURE CITED 


Brown, F. M. 1973. The types of the pierid butterflies described by William 
Henry Edwards. Trans. Amer. Entomol. Soc. 99 (in press). 

Davenport, D. & V. G. Deruier. 1937. Bibliography of the described life- 
histories of the Rhopalocera of America north of Mexico 1889-1937. Entomol. 
Amer. 17(4): 155-194. 

Epwarps, H. 1877. Notes on the genus Colias, with descriptions of some ap- 
parently new forms. No. 24. Pacific Coast Lepidoptera. Privately published, 
although frequently cited as Proc. Calif. Acad. Sci., Feb. 5, pp. 1-ll. See 
Beutenmuller, W. 1891. List of writings of the late Henry Edwards. Canad. 
Entomol. 23( 12), p. 259, 262. 

Epwarps, W. H. 1863. Description of certain species of diurnal Lepidoptera found 
within the limits of the United States and British America. No. 1. Proc. Entomol. 
Soc. Phila. 2: 78-82. 

1870. Descriptions of new species of diurnal Lepidoptera found within 

the United States. Trans. Amer. Entomol. Soc. 3: 10-22. 

. 1871. Descriptions of new species of North American butterflies. Trans. 

Amer. Entomol. Soc. 3: 266-277. 

1872. Descriptions of new species of diurnal Lepidoptera found within 
the United States. Trans. Amer. Entomol. Soc. 4: 61-70. 

Ferris, C. D. 1972a. Notes on certain species of Colias (Lepidoptera: Pieridae) 
found in Wyoming and associated regions. Bull. Allyn Mus. Entomol. No. 5. 

1972b. Ultraviolet photography as an adjunct to taxonomy. J. Lepid. Soc. 
267 20 = 2115: 

Hoitiann, W. J. 1931. The Butterfly Book. Doubleday, New York. 

Kiors, A. B. 1935. A new Colias from South Dakota (Lepidoptera, Pieridae). 
Amer. Mus. Novitates 767. 

KoLYER, J. M. & A. M. REerMscHuEssEL. 1969. Scanning electron microscopy on 
wing scales of Colias eurytheme. J. Res. Lepid. 8(1): 1-15. 

MAZOKHIN-PORSHNYAKOy, G, A. 1954. Ultraviolet radiation from the sun as an 
oa factor on the habits of insects [In Russian]. J. Obshchey Biologii 
Loe oO2—o0n. 


McDunnoueu, J. 
if ‘ ‘~ | 


1928. Notes on Canadian diurnal Lepidoptera. Canad. Entomol. 


2a 0° 


7 I 


Alo. 


N EKRUTI ©. P. 1964. The hidden wing-pattern of some Palearctic species of 
Gonepteryx and its taxonomic value. J. Res. Lepid. 3(2): 65-68. 


VOLUME 27, NUMBER 1] 


—l 


ies) 


1965. “Gynandromorphic effect” and the optical nature of hidden wing- 
pattern in Gonepteryx rhamni L. (Lepidoptera, Pieridae). Nature 205: 417-418. 

SHIELDS, O., J. F. EMMeEL & D. E. BreEpLove. 1969. Butterfly larval foodplant 
records and a procedure for reporting foodplants. J. Res. Lepid. 8(1): 21-36. 

Taxtpot, G. 1935. Pieridae III. Lepidopt. Cat., ’s-Gravenhage. 66: 385-697 (418). 

Wricut, W. G. 1907. Colored Plates of the Butterflies of the West Coast. San 
Bernardino, Calif. Privately printed. 


THE GENETICS OF FORE AND HINDWING COLOUR IN CROSSES 
BETWEEN DANAUS CHRYSIPPUS FROM AUSTRALIA 
AND FROM SIERRA LEONE (DANAIDAE) 


C. A. CrLarKe, P. M. SHEPPARD AND A. G. SMITH 
Nuffield Unit of Medical Genetics, University of Liverpool, England 


Unlike most warningly colored species, the butterfly Danaus chrysippus 
(L.) is known to be polymorphic in large parts of its range. Before one 
can understand the reason for this it is necessary to determine the genetic 
control of the forms. Recently we obtained a stock of D. chrysippus from 
Sydney, NSW and another from Sierra Leone. This paper gives pre- 
liminary results obtained by crossing the two races. 


MATERIALS AND METHODS 


The material from Australia, which was sent to us as living butterflies 
by post, was monomorphic and typical f. chrysippus of the race D. c. 
petilea (Fig. la). The ground colour of these butterflies is tawny orange 
tending to nutbrown towards the costal margin of the forewing. The 
hindwing upperside is bordered by black, sometimes with a vestige of 
white spotting close to the hindwing border. The apical third of the 
forewing upperside is black, with a variable subapical bar of white spots. 

The specimens from Sierra Leone, f. alcippus, differed from the Aus- 
tralian ones in that the ground colour was more orange and most of the 
hindwing was covered by a patch of white scaling (Fig. lb, c). The 
pale areas of the forewings were of two types—those with a narrow costal 
border of nutbrown pigmentation similar to the Australian butterflies 
(Fig. 1b) and others in which the nutbrown extended over most of the 
forewing (Fig. lc). There was not enough orange on the hindwing to 
determine whether its hue differed in the two Sierra Leone forms. 

Hybrids between the two races were obtained by allowing the males 


74 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


e€ 


i 


Fig. 1. Parental and F; phenotypes: (a) f. chrysippus, Australia; (b) f£. alcippus, 
tawny, Sierra Leone; (c) f. alcippus, nutbrown, Sierra Leone; (d) F:, tawny; 
(e) F,, nutbrown. 


of one subspecies and the females of the other to fly freely in two heated 
greenhouses. Mating took place spontaneously and the females laid on a 
foodplant of the genus Asclepias. Unfortunately in these circumstances 
we were unable to keep the broods separate. The F,; when they emerged 
were allowed to mate and F» generations were readily produced, but here 
igain we did not keep the broods separate. 


VoLuME 27, NUMBER | 75 


Fig. 2. F» phenotypes: (a) f. chrysippus-like, tawny; (b) f. chrysippus-like, nut- 
brown; (c) Fy-like, tawny; (d) Frlike, nutbrown; (e) f. alcippus-like, tawny; 
(£) £. alcippus-like, nutbrown. 


76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TaBLE 1. Phenotypes in the F>». 


Hindwings 
A little white 
Ground colour No white ( F,-like ) White 
Nutbrown 11) Yl 14 
Tawny 58 1) 20 
RESULTS 


In the F, we obtained two distinct types of progeny with respect to the 
ground colour of the fore and hind wings. In one the ground colour of 
the forewings was nutbrown and the hindwings, although slightly paler 
in hue, had scales which also tended towards being nutbrown (Fig. le). 
In the other the ground colour was tawny to tawny orange, the nutbrown 
pigmentation being confined to an area close to the costal border of the 
forewings (Fig. ld). Over the hindwings of both forms there was also 
a scattering of white scales which was variable in extent, being un- 
detectable in some specimens but quite obvious in others. Of 12 butter- 
flies with a tawny ground colour 6 had no detectable white scaling 
whereas of 11 nutbrown individuals only 2 lacked it. The reciprocal Fs 
gave comparable results. 

Both types of F; gave rise to F»s. Among these were nutbrown individ- 
uals (Fig. 2b, d, f) and those of the tawny phenotype (Fig. 2a, ¢, e). 
There was a range of hindwing phenotypes from the Australian form 
without white (Fig. la) through intermediate forms up to a degree of 
white scaling found in the more extreme F\s. With difficulty these could 
be divided into the parental and F, types, but the distinction was not clear 
(Fig. 2a, b, c,d). In addition, there was a phenotype with a white patch 
on the hindwing similar to that found in the Sierra Leone grandparents 
(Fig. 2e, f) (f. alcippus). The extent of the white was variable but 
quite distinct from that in the other insects. Both the nutbrown and the 


tawny phenotypes were to be found among the offspring with the white 
hindwings and those lacking it (Table 1). 


Discussion 


Since both in the F; and Fy, generations the broods were not kept 
separate, we do not know the phenotype (tawny or nutbrown) of the 
Sierra Leone parents. However we do know that the Australian stock 
was monomorphic for the tawny phenotype, consequently the nutbrown 
form must be dominant since it segregated in a clear cut manner in the F. 


VOLUME 27, NUMBER 1 17 


Because the two stocks were both monomorphic with respect to the hind- 
wing pattern we can say that white patch on the hindwing is partially 
recessive and completely so in some individuals. Thus in both the F, and 
F, butterflies white scaling was detectable in a higher proportion of the 
nutbrown phenotype than in the tawny one (p < 0.001). In fact, the 
ratios in the F, and Fs, suggest that the dominance is absent in most 
individuals with the nutbrown phenotype but not in those with the tawny 
ground colour. This difference may be due to the easier detection of a 
few white scales on the darker background. 

The presence or absence of the white patch (as distinct from white 
scaling ) on the hindwing segregated in a clear cut manner and in a good 
approximation to a 3:1 ratio in the F, (Table 1). Thus we can conclude 
that the presence or absence of the white patch on the hindwing is con- 
trolled by a single gene, the heterozygote being variable in expression. 

Since the broods were not reared separately one cannot usefully examine 
the ratio of the nutbrown phenotype to the tawny one in the Fs. However, 
one can examine the data for evidence of linkage. We know that the 
nutbrown form and the white hindwing patch must both have been 
derived from Sierra Leone, and therefore in the presence of linkage the 
two loci would be in repulsion. 

An examination of the ratio of white hindwing patch to its absence 
amongst the F. nutbrown individuals provides no good evidence for a 
departure from the expected 3:1 ratio (non-white to white hindwings ) on 
the assumption of no linkage. Furthermore, if there had been close 
linkage there should have been an excess of the nutbrown phenotype 
amongst the white hindwinged individuals. 


SUMMARY 


In the cross between D. chrysippus material from Australia and that 
from Sierra Leone, it was shown that the white hindwing area of f. 
alcippus is recessive, or nearly so, to the tawny hindwing of f. chrysippus, 
and the character is controlled in the main by a single locus. 

In our Sierra Leone material there were two shades of brown on the 
forewings, tawny and nutbrown. The matings showed that the nutbrown 
coloration is dominant and extends on to the hindwing in the hybrids. 
It also appears to be controlled by a single locus. Segregation of the 
characters in the F, does not suggest close linkage between the two loci 
concerned. 


78 JouURNAL OF THE LEPIDOPTERISTS SOCIETY 


A NEW CALLITHOMIA (LEITHOMIA) FROM AMAZONAS, 
VENEZUELA (ITHOMIIDAE) 


Joun H. Masters 
Lemon Street North, North Hudson, Wisconsin 54016 


During February 1967, Harold Skinner of La Victoria and Albert 
Gadou of Caracas, Venezuela made a collecting expedition to the Ocama 
Mission to the Waika Indians in Amazonas, Venezuela. After their return, 
Senor Skinner sent me a fair number of duplicate specimens. Of particular 
interest among these was a previously undescribed Callithomia, reminis- 
cent of Callithomia lenea (Cramer), but with an unique forewing, unlike 
any other ithomine. Description as follows: 


Callithomia (Leithomia) skinneri Masters, new species 


Female: Has an appearance completely distinct from that of any other Callithomia 
(Fig. 1). This is especially true of the forewings; the hindwings are very much like 
that of a female Callithomia lenea. 

Upperside of forewing: Outlined in black. A large oval-shaped subapical patch 
is lemon yellow and semi-translucent. The balance of the wing is a rather uniform 
soft ochreous brown, slightly transluscent. 

Upperside of hindwing: Broadly outlined in black. A nearly transparent discal 
area has the unique shape, with the hook in cell M:, that is characteristic of the 
lenea group of species. The discal area is surrounded by a black band that is broad 
caudally but quite narrow distally; it is separated from the outer margin by an 
ochreous orange submarginal band. 

Underside of wings: Almost identical to uppersides except for having a number 
of white crescents present in the wing border. There are three well pronounced 
crescents on the forewing, at the termination of cells Ri, Rs and M:. On the hind- 
wing there are seven of them, one at the termination of each primary cell, with the ones 
in cell Mi and Mz being most pronounced. 

Male: Has the general appearance of the female. The forewings have the large 
oval-shaped yellow semi-transluscent patch, but are otherwise more transparent than 
the females. The males forewings are somewhat narrower and more pointed than 
the females and its hindwings have a pronounced hump on the costal margin, similar 
to that of other males in the subgenus. 

Holotype female: Ocama Mission, Amazonas, Venezuela (at the junction of the 
Ocama and Orinoco Rivers, approximately 2°30’N., 65°15’W.), February 1967; 
Harold W. Skinner, collector. Length of forewing: base to tip, 28.5 mm; apex to 
tornus, 16 mm; center of costal margin to tornus, 15 mm. To be deposited at 
Carnegie Museum, Pittsburgh, Pennsylvania. 

Allotype male: Same collecting data as holotype. Length of forewing: base 
to tip, 28.0 mm; apex to tornus 17 mm; center of costal margin to tornus, 13.5 mm. 
in the collection of Harold W. Skinner, La Victoria, Venezuela. 

_ Paratype: One female, same data as holotype. In the collection of Harold Skinner. 
There are additional specimens in the collection of Albert Gadou, Caracas, but I 
e not had the opportunity to examine them. 


VoLUME 27, NUMBER L 79 


Fig. 1. Callithomia skinneri new species, holotype female, Ocama Mission, 
Amazonas, Venezuela, February 1967. Natural scale. 


The general appearance and the relatively narrow male forewings place 
Callithomia skinneri in the subgenus Leithomia which includes xantho 
(Felder), methonella (Weymer), inturna (Fox), epidero (Bates), foxi 
Masters, zingiber Fox, lenea (Cramer) and drogheda (Weeks). Of these, 
only lenea and drogheda in any way resemble skinneri; skinneri can be 
separated from both of them by its distinct more opaque forewings and 
the large oval yellow spot in them. In several respects, drogheda bridges 
the distinctions between skinneri and lenea—these three may be found 
ultimately to be geographic subspecies of a single species. None of the 
species in the nominate subgenus have a similar appearance. 


AN “ALBINO” LYCAENA HELLOIDES (LYCAENIDAE) 


White or whitish specimens of Lycaena phlaeas Linnaeus have been recorded on 
both sides of the Atlantic (e.g. Fuller 1962, J. Lepid. Soc. 16: 59; Martin 1962, J. 
Lepid. Soc. 16: 59-60) but do not seem to be on record for L. helloides Boisduval, 
the most widespread purely Nearctic member of the genus. A fresh female L. 
helloides with the orange coloration completely replaced by creamy white on both 
surfaces was taken 17 April 1972 in the Putah Creek Recreation Area near Davis, 
Yolo Co., California. The blanching includes the normally deep reddish-orange 
ventral subterminal line of the hind wing. It is associated with curled or “frizzled 
scales as previously reported for “albino” L. phlacas. L. helloides is yeaa oes 


near Davis and shows considerable minor variation, especiaily in the maculahon 01 
the females. 


ArTHur M. SHapiro, Department of Zoology, University of California, Davis, 
California 95616. 


80 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


A NEW SUBSPECIES OF CALLITHOMIA HEZIA FROM ZULIA, 
VENEZUELA (ITHOMIIDAE) 


Joun H. MAsTERs 
Lemon Street North, North Hudson, Wisconsin 54016 


Venezuelan lepidopterists have just recently begun penetrating into the 
Sierra de Perija, a northern offshoot of the Venezuelan Andes which forms 
the boundary between Colombia and Venezuela in the northwestern 
Venezuelan state of Zulia. Materials that have been brought back from 
this region have included a number of endemic subspecies and it is not 
suprising to find that the Perija population of Callithomia hezia Hewitson, 
a species known to display marked geographic variation, represents a 
distinctly marked and previously undescribed subspecies. 


Callithomia (Callithomia) hezia perija Masters, new subspecies 


This subspecies differs from the other six subspecies of Callithomia 
hezia by having a row of three separated and relatively small yellow 
patches in the middle of cells M;, Mz and Mz on the hindwing. Subspecies 
baccata Fox and tridactyla Hewitson have yellow patches on the hindwing, 
but they are much larger and touch each other basally. 


Male Fig. 1 A & B: Appearance readily associated with Callithomia hezia. Fore- 
wing length: base to apex, 35 to 37 mm. Forewing breadth: apex to tornus, 20 
to 21 mm; center of costal margin to tornus, 17 to 17.5 mm. 

Upperside of forewing a deep black background color with a reddish brown streak 
in the basal area. Upperside of hindwing light brown in background color except a 
black margin which greatly widens in the apical area. Pattern elements composed 
of opaque yellow spots which include: one or two spots in the forewing cell; a 
double row of forewing spots including a single spot in one of the radial cells and 
in cell Mi, and then two spots each in cells Ms, Mz, Cu: and Cuz; and three relatively 
small, separated spots in the middle of cells M:, Ms and M: on the hindwing. 

Underside of both wings mirrors the appearance of the uppersides, except for the 
presence of five small white crescents present in the wing margins—two on the 
forewing and three on the hindwing. 

Male genitalia Fig. 2 A, B & C: Shows no characteristics which might distinguish 
it from other subspecies of Callithomia hezia. The penis (Fig. 2C) is not forked; 
this is one of the characteristics that separates the nominate subgenus Callithomia 
Bates from subgenus Leithomia Masters. 

Bemale Fig. 1 C, D & E. Appearance almost identical to male except for blunter 
and mre rounded wings. Forewing length: base to apex, 35 to 38 mm. Forewing 
breadtl n: apex to tornus, 20 to 21 mm; center of costal margin to tornus, 19 to 20 
mm. Yellow patches on hindwings, in two specimens, a bit larger than those of 
alee There is a series of six small white crescents along the ventral hindwing 
mar instead of three. 

‘olotype male: El Tucuco, Estado Zulia, Venezuela, September 1964, Albert 
tor. Length of forewing, base to tip, 35.5 mm. 
male: El Tucuco, Estado Zulia, Venezuela, July 1968, Yuri Budokari 


Allo LV 


VOLUME 27, NUMBER | 8] 


LAS. ST 


Fig. 1. Callithomia hezia perija new subspecies. Holotype male, El Tucuco, 
Zulia, Venezuela: (A) upperside, (B) underside. Allotype female, E] Tucuco, Zulia, 
Venezuela: (C) upperside, (D) underside. Paratype female, Como del Tigre, 
Anduze, Colombia: (E) upperside. All specimens natural si 


82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 2. Male genitalia of Callithomia hezia perija: (A) side view with left valve 
removed, (B) left valve, (C) penis. 


collector. Length of forewing, base to tip, 36 mm. Holotype and allotype have been 
placed in the Carnegie Museum, Pittsburgh, Pennsylvania (type series number 674). 

Paratopotypes: 3 males, 2 females in author’s collection; 2 males and 2 females 
in collection of Harold Skinner, La Victoria, Venezuela. There are probably another 
two dozen specimens in other Venezuelan collections, but I have not personally 
examined them and am not including them in the type series. All of these are from 
the same locality as the holotype, various dates, 1964 to 1969, and various collectors. 

Paratype: One female, Como del Tigre, Anduze, Colombia, Sept. 1943 (Fig. 1E) 
in the American Museum of Natural History, New York. 


Callithomia hezia is a very distinct butterfly that can be confused with 
only one other species, its Mullerian mimic Napeogenes peridia (Hewit- 


can ) 


n) which is somewhat smaller. A key to the seven described subspecies 
' Callithomia hezia follows: 


low spots on hindwing 2.2. 3) ee eee 
ow spots not present on hindwing —2 
/ spots of forewing, including the antemarginal series, beyond the end 
cell fused into a large patch extending from the costa to Cur —------ 

__.... C. hezia wellingi Fox, southern Mexico and Guatemala. 


VOLUME 27, NUMBER 1 83 


Antemarginal spots of forewing separated and never included in a patch formed 
by fusion of the spots 
3. Yellow antemarginal and postmedial series of forewing rather large and only 
narrowly separated by black which, especially in the apical part in some 
MlalesmaLcnosntonbertranslucent: 205 ne 
coc 1 C. hezia hedila Godman & Salvin, Guatemala and Honduras. 
Yellow antemarginal spots of forewing smaller than postmedian spots, always 
well separated by opaque black, but the postmedial series sometimes tending 
to fuse with the yellow spots in the end of the cell _... 
nnn r TS C. hezia smalli Fox, Cocle Province, Panama. 
All yellow spots on the forewing tiny and well separated from each other 
6a C. hezia hezia Hewitson, Nicaragua, Costa Rica and Panama. 
4. Yellow spots of the hindwings only slightly larger than those of the forewings 
auamaiscretely separated trom each other 222.2200 
a C. hezia perija Masters, Sierra de Perija, Venezuela and Colombia. 
Yellow spots of the hindwings large, broad and fused together basally, but 
present im a single series _...____. C. hezia tridactyla Hewitson, Colombia. 
Yellow spots of the hindwings large, broad and fused together basally, but 
present in a double series with an outer marginal row of spots present 
enc gE C. hezia baccata Fox, Peru. 


SOME FIELD NOTES ON ISOPARCE CUPRESSI BDV. (SPHINGIDAE) 


During a collecting trip to the Wedge Plantation near McClellanville, Charleston 
County, South Carolina in mid-September of 1972, I had the opportunity to in- 
vestigate the life histories of some of the local Lepidoptera. The larval habits of the 
sphingid, Isoparce cupressi Bdv., were of particular interest to me since my _ host, 
Dr. R. B. Dominick, had successfully reared a brood of this species ex-ovo on 
Taxodium distichum L. (Bald Cypress) in the late summer of 1971. A suspected 
habitat of the species, a small swamp surrounded by T. distichum and located near 
the Wedge Laboratory, was surveyed, and several well exposed and isolated trees 
of the foodplant were chosen for examination. 

Concentrated searching of the lower, accessible branches of eight large cypress 
trees yielded 15 larvae—one 2nd instar, five 3rd instar, seven 4th instar, and two 
5th instar larvae. Six of these were in the process of molting when found. All but 
one of the larvae were on the undersides of the branches of the foodplant, from 
approximately four to seven feet off the ground. The 2nd and 3rd instar larvae were 
always situated near the ends of the branches, on the midribs of the terminal pinnae. 
The location of the 4th instar larvae was usually in the middle of the branches, also 
on the midribs of the pinnae. A large and nearly mature 5th instart larva, was found 
resting in the woody crotch of two branches. Finally, the other 5th instar larva, 
covered with parasitic cocoons, was discovered sitting on the top of a dead branch, 
moribund and obviously near death. 

Two viable ova and several hatched eggshells were collected on the undersides of 
the terminal needles of the Taxodium branches. Since these were always found 
singly and randomly deposited, it appears likely that this is the natural mode of 
oviposition. 


5 5 A ee a be a ca 26 mna- 

Numerous jumping spiders and harvestmen were seen during the daily a Ae 
tions of the cypress branches. Their suspected predatory behavior was confirmed 
when a small grey salticid was observed feeding on a wriggling 3rd instar cupresst 


larva that it had obviously just captured. Judging by the large numbers seen, it 


84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


seems logical to assume that these arachnids contribute considerably to the attrition 
of this sphingid’s early stages. 

Six of the fourteen larvae later produced parasites—a dipterous maggot emerged 
from a 3rd instar larva, while five late 5th instar larvae produced numerous braconid- 
like parasites. Also, several dried corpses of cupressi larvae were seen on the branches, 
with emerged parasitic cocoons found spun in a silken mass at the base of the dead 
larvae. 

To assess the feeding habits and density of I. cupressi larvae, I placed an old 
bedsheet underneath a number of cypress trees. Each morning I examined this sheet, 
and found it covered with large amounts of lepidopterous larval frass. Since there 
are no other known Taxodium-feeding species whose larvae would be as large as 
cupressi, it is assumed that most of the droppings were from this species. Almost 
all of it was identical in color, form, size, and texture to that deposited by the cupressi 
larvae feeding in captivity. Often there were concentrations of large frass pellets on the 
sheet, but searching the lower branches just above these areas revealed no larvae. It 
appears that most of this excreta had filtered down from cupressi larvae on the higher 
branches. Further, the ground beneath every landlocked T. distichum tree on the 
Wedge grounds indicated the presence of cupressi in the form of these large pellets 
of frass. The sheet was shaken clean each morning, and then moved to another 
nearby cypress tree. While the sheet was checked at frequent intervals, almost no 
new droppings were deposited during the day. However, the following morning 
invariably revealed the sheet once again covered with quantities of cupressi-like 
frass. This evidence supports the hypothesis of Bates (1928, Florida Entomol. 12: 
20-21), that I. cupressi larvae feed at night. It seems likely that, at least around 
the Wedge Plantation, the higher cypress branches support healthy populations of 
cupressi larvae. Searching these higher branches at night with a flashlight, and with 
a ladder during the day, was unsuccessful. 


MicHaEL D. VAN Buskirk, 4512 47th S.W., Seattle, Washington 98116. 


INTERESTING FLORIDA BUTTERFLY RECORDS 


The records below represent either range extensions or reinforcements of older 
records as published by Kimball (1965, The Lepidoptera of Florida. Gainesville. 
363p.) and Clench (1970, J. Lepid. Soc. 24: 240-244). Checklist numbers are 
those of dos Passos (1964, Mem. Lepid. Soc. 1) and the nomenclature also follows 
his work. In several cases, primarily those records from the early 1960’s, no specimens 
were taken or the few secured have been destroyed. Therefore, the data in those in- 
stances are of a more general nature. I would like to offer a special thanks to Mr. D. 
W. Bryne who provided support and encouragement during the period that these 
records were being established. 

83. Problema byssus (Edwards). Range extension. In the early 1960’s this Skipper 
could be found in both north Tampa, Hillsborough Co., and at Bishop’s Harbor, 
Manatee Co. 

188b. Staphylus mazans hayhurstii (Edwards). Range extension. Ozello, Citrus 
Co.: 5 September 1970 (5 46 6). This represents a substantial northward extension 
2 te west ae of Florida. The previous records (Kimball) stopped at Sanibel 

sland, Lee Co. 
218. Urbanus dorantes dorantes (Stoll). Range extension. Caxambas, Marco 
nd, Collier Co.: 17 October 1970 (1 6). Branchton, Hillsborough Co.: 29 
1972 (1 9) and 5 April 1972 (1 9). The two specimens from Branchton 
en in open areas within a forested area and represent a one hundred and 
northward range extension over previous records (Clench). This could 


VOLUME 27, NUMBER 1 85 


be only a temporary extension however. The winter of 1971-72 was the warmest in 
many years and could be the reason dorantes suddenly appeared this far north in 
Florida. A normal winter season with numerous days around the freezing mark 
could well result in elimination of the species from this region. 

233. Polygonus leo (Gmelin). New locale. Caxambas, Marco Island, Collier Co.: 
17 October 1970 (1 ¢). This makes more continuous geographically the records 
on the Florida west coast. 

274b. Appias drusilla neumoegenii (Skinner). New locale. Casey Key, Sarasota 
Co.: 6 July 1963 (1 ¢). This capture reinforces an older, uncertain record from 
the same area (Kimball). Also, although I found this species uncommon at Key 
Largo, Monroe Co., in May of 1971, when I returned later in August of that year 
I found it to be not only abundant but also outside its normal hammock environment 
visiting flowers freely in the bright, open, roadside areas. 

307a. Phoebis statira floridensis (Neumoegen). New locale. Bishop’s Harbor, 
Manatee Co.: 27 January 1972 (1 @). 

308. Kricogonia lyside (Godart). Range extension. Casey Key, Sarasota Co.: 
6 July 1963 (1 ¢). This individual, taken at the blossoms of Bidens pilosa Linnaeus, 
represents a considerable northward range extension. 

525a. Anartia jatrophae guantanamo Munroe. Range extension. In the early 
1960’s numerous individuals of this species were taken at Chassahowitzka, Citrus Co. 
The most northerly locale previously recorded on the Florida west coast was Tampa 
(Kimball). 

554a. Phyciodes frisia frisia (Poey). South Bay, Palm Beach Co.: 21 August 1971 
(1 ¢). This is one of the few records for the species north of Dade and Monroe 
Counties. 

639a. Euptychia gemma gemma (Hubner). Range extension. In the early 1960's 
this satyr could be found sparingly but consistently at Chassahowitzka, Citrus Co. 
Also, a colony was discovered at Branchton, Hillsborough Co., in 1972 with the 
species being not uncommon from 20 March through at least 6 April. Moist, grassy 
areas within woods are preferred. 

646b. Euptychia cymela viola (Maynard). Range extension. A colony of this 
species was discovered near Branchton, Hillsborough Co., with individuals on the 
wing from at least 12 March through 17 April in 1971. The habitat consists of an 
oak woods surrounded by swampy cypress stands. 


D. L. Burris, 704 Country Club Drive, Tampa, Florida 33612. 


A NEW SPECIMEN OF CYNTHIA ANNABELLA “AB. MUELLERI” 
FROM CALIFORNIA (NYMPHALIDAE) 


A male Cynthia annabella Field (= Vanessa carye auct.) virtually identical with 
one of “ab. muelleri” figured by Gunder (1929, Pan-Pac. Entomol. 6: 9, pl. 17) 
was taken on 10 May 1972, 214 mi. NE Davis, Yolo Co., California. This phenotype, 
with various minor modifications (cf. Gunder 1927, Entomol. News 38: 263-271, 
pls. 5-10), has now been found between 20 and 30 times in C. annabella throughout 
California. If it has a genetic basis—as suggested by several reported clusters of 


captures—its incidence is probably too high to be ascribed to recurrent mutation 
alone. Extremely similar phenotypes are recorded in Cynthia cardui Linnaeus 
(Gunder, 1927) and C. virginiensis (Drury) (cf. Clark & Clark, 1951 Butterflies of 


Virginia, pl. 30). 


Artour M. SHariro, Department of Zoology, University of California, Davis, 
California 95616. 


86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


BOOK REVIEW 


TropicAL BurreRFLIES, by D. F. Owen. 1971. 214 p., 40 plates. Clarendon Press, 
Oxford, England. $11.50. 


Perhaps because “Tropical Butterflies” is ubiquitous, the book is subtitled, “The 
ecology and behavior of butterflies in the tropics with special reference to African 
species.” The emphasis of the author is summed up even better by two statements 
in the preface: “. . . this book . . . is mainly about African butterflies, because I 
know more about them than about the butterflies of other tropical regions,” and, 
“I advocate throughout the study of living butterflies in their natural environments, 
and although I am not against building up a collection, I would rather see collectors 
devote some of their energy to finding out more about living butterflies.” 

Thus, we have a book consisting of extensive discourse on ecology and behavior 
with only scant references to taxonomy, identification, collecting and mounting tech- 
niques, etc. This is very good, for the author is concentrating on those subjects usually 
neglected in “butterfly books.” Some representative chapter titles include: classifica- 
tion and zoogeography, life histories, population ecology, population genetics, mimicry, 
evolution and conservation. The text is concise, well written, and amply illustrated 
with excellent text figures and photographic plates. There is an index and an exten- 
sive bibliography. 

The author has spent eight years studying butterflies in Africa and much of the 
text is from his personal experiences, including an extensive treatment of the popula- 
tion biology of Acraea encedon which has been his major research interest. Many 
of his observations were made in a small garden in Freetown, Sierra Leone where 
he took about 300 species. This should suggest that elaborate trips into the bush 
are not essential for tropical collecting. Some chapters, such as the one dealing with 
mimicry, take the form of reviews of contemporary research and publication. Although 
most of the exemplification is taken from the African fauna, it is pertinent to pan- 
tropical situations and, in fact, much of it to temperate regions as well. The species 
diversity of the tropical butterfly fauna is treated in detail and numerous com- 
parisons are given with temperate faunas. Seasonal forms, which are most pro- 
nounced in Africa, is the only other purely tropical phenomenon which is discussed 
in detail. 

Without reservation, I regard the book as excellent. For anybody intending to 
collect in the tropics, the book would be a tremendous asset. It is not necessary to 
have any interest in tropical butterflies, however, in order to benefit from the book 
because the emphasis is on the ecology and behavior of butterflies rather than on 
the tropics. I hope that everyone who reads the book will retain some of the author's 
insights regarding the study of living butterflies. I am sure that this will result 
in more meaningful field work whether one lives in Africa, elsewhere in the tropics, 
in temperate regions, or even in the arctic. 


Joun H. Masters, Lemon Street North, North Hudson, Wisconsin 54016. 


NOTES AND NEWS 


As a result of the recent election, it is a pleasure to announce that F. Martin Brown 
and Cyril F. dos Passos were overwhelmingly approved by the membership as honorary 

-mnembers of the Lepidopterists’ Society. The newly elected officers of the 
y are listed inside the front cover. In addition, Dr. Frederick H. Rindge was 
clected as the Jordan Medal Representative (see Vol. 26, p. 208), and the proposed 
constitutional amendments (see Vol. 26, p. 203) were passed. 


VoLUME 27, NuMBER | 87 


OBITUARY 


ALVAH PETERSON (1888-1972) 


Dr. Alvah Peterson, well known entomologist and lepidopterist, died on 
Monday, September 11, 1972, at the Ohio State University Hospital. 

Dr. Peterson was responsible for many publications during his long 
career. His first major work was, “The Head-Capsule and Mouthparts 
of Diptera,” published as an Illinois Biological Monograph in 1916. He 
then launched into research and the publication of many papers on 
economically important insects while he was with the Department of 
Entomology, Rutgers University (1916-1925) and the United States De- 
partment of Agriculture (1925-1928). Dr. Peterson left the Department 
of Agriculture in 1928 to become Professor of Entomology at Ohio State 
University, a position he held until retirement. It was while at Ohio 
State University that he completed the works that most lepidopterists 
will remember him for. These include Larvae of Insects, Entomological 
Techniques, and numerous papers on lepidopteran eggs. His two-volume 
work Larvae of Insects is used throughout the world as the basic work for 
the identification of immature insects. Interestingly, his pioneer work on 
insect eggs was done after his retirement from Ohio State University. 

Dr. Peterson was a member of the Entomological Society of America 
where he served as national president. He also had the distinction of 
being the Society’s first member to be elected to an honorary member- 
ship. He served as consultant or visiting professor of Entomology at 


88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Michigan State, the University of Arizona, Oregon State, the University 
of Florida, and the University of Minnesota. 

Besides his scientific accomplishments, Dr. Peterson was very well 
liked by all who knew him, especially his students to whom he always 
gave generously of his time and help. His passing is mourned by all 
who knew him. 


Cuar.es L. SELMAN, Department of Entomology, Ohio State University, Columbus, 
Ohio 43210. 


: 
- e < 
a Ne 
~~. 3 
- = 
{ 
4 
a { 
4 
a - 
A : 


i i 
| LP ae 


2 | 
ie 
‘ 


i a i 


OT es eS a, ee ee ee ae Re ee 


EDITORIAL COMMITTEE OF THE JOURNAL 


Editor: THEroporE D. SarcENT, Department of Zoology, 
University of Massachusetts, Amherst, Massachusetts 01002 


K. S. Brown, J. M. Burns, R. H. Carcasson, J. P. Donanve, 
J. F. Gates Ciarke, C. D. Ferris, R. O. KENDALL, J. H. MAsTERs, 
L. D. Miter, A. P. Piatt, J. R. G. Turner 


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 LireERATURE CITED, 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 414 inches). Illustrations larger than 8% 
< 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 article’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 approx- 
imate 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. 

Page Charges: Authors with grant or institutional funds are requested to pay a 
charge of $24.00 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 
case of authors who have no grant or institutional funding, as it is not intended that 
any author should pay this charge from personal funds. However, all authors will 
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: Dr. C. V. Covell, Department of Biology, University of 
Louisville, Louisville, Kentucky 40208. 


ALLEN PRESS, INC. SRCTES LAWRENCE, KANSAS 
usr 


CONTENTS 


Lire History oF ISOPARCE CUPRESSI (SPHINGIDAE). Richard B. 
Dominick) 4 
BEHAVIORAL ADAPTATIONS OF Cryptic Motus. VI. FURTHER E:XPERTI- 
MENTAL STUDIES ON BARK-LIKE SPECIES. Theodore D. Sargent — 
Nores oN Lire Histories AND HABITS OF SOME WESTERN THECLINAE. 
E. J. Newcomer 20 a ee 
HABITAT SELECTION AND POPULATION STRUCTURE IN PLEBEJUS 
SAEPIOLUS BoispuvAL (LyYCAENIDAE). Margaret A. Sharp 
and David BR. Parks i000. 
FoopPLANT ECOLOGY OF THE BUTTERFLY CHLOSYNE LACINIA (GEYER) 
(NYMPHALIDAE). I. LarvaAL FooppLants. Raymond W. 
Neok/ sce NS OS 
THE COLLECTION OF BUTTERFLIES MADE BY JACK DENNIS AT BEULAH, 
MAniToBA. John H. Masters 
A REVIEW OF THE AMBLYSCIRTES WITH THE DESCRIPTION OF A NEW 
SPECIES FROM Mexico ( HEspermpAE). Hugh Avery Freeman _ 
A REVISION OF THE COLIAS ALEXANDRA COMPLEX (PIERIDAE) AIDED 


BY ULTRAVIOLET REFLECTANCE PHOTOGRAPHY WITH DESIGNA- 


TION OF A New Susspecies. Clifford D. Ferris —. 


THE GENETICS OF FORE AND HINDWING COLOUR IN CROSSES BETWEEN 
DANAUS CHRYSIPPUS FROM AUSTRALIA AND FROM SIERRA LEONE 
(DanawaE). C. A. Clarke, P. M. Sheppard and A. G. Smith — 


A New CALLITHOMIA (LEITHOMIA) FROM AMAZONAS, VENEZUELA 
(ItHoMupAE). John H. Masters 7 


A New SUBSPECIES OF CALLITHOMIA HEZIA FROM ZULIA, VENEZUELA 
(IrHomMuDAE). John H. Masters 


GENERAL NOTES 


Some field notes on Isoparce cupressi Bdv. (Sphingidae). Michael D. 
Van’ Buskigky i.) te i 


Preliminary report on communal resting of Smyrna karwinskii adults 
(Nymphalidae), Alberto Muyshondit ... ee 


A new specimen of Cynthia annabella “ab. muelleri” from California 
(Nymphalidae), Arthur M: Shapiro 


Bird predation on Papilio polyxenes F,. (Papilionidae). James M. 
Erickson, (10000 Uo a ae 


An “albino” Lycaena helloides (Lycaenidae). Arthur M. Shapiro __ 
Interesting Florida butterfly records. D. L. Burris 


Book Review 


13 


17 


22 


33 


40 


57 


73 


78 


80 


83. 
15 
85 


16 
79 
84 


86 
86 
87 


i 
© 


| 
Na 


Volume 27 1973 Number 2 


- JOURNAL 


of the 


EPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 


16 May 1973 


THE LEPIDOPTERISTS’ SOCIETY 
EXECUTIVE COUNCIL 


J. F. Gates Cxarxe (Washington, D.C.) President 

Harry K. Ciencu (Pittsburgh, Penn.) President-elect 
ALEXANDER B. Kuors (New York, N.Y.) 1st Vice President 
C. F. Cowan (Berkhamsted, England) Vice President 

E. G. Munroe (Ottawa, Ontario) Vice President 

S. S. Nico.ay (Virginia Beach, Va.) Treasurer 

Lee D. Mixer (Sarasota, Fla.) Secretary 


Members at large (three year term): M. C. Nietsen (Lansing, Mich.) 1974 
A. BLANCHARD (Houston, Texas) 1973 D. C. Ferrcuson (Washington, D.C.) 
R. B. Dominick (McClellanville, S.C.) 1975 

1973 R. O. Kenpatt (San Antonio, Texas) 
J. P. DonaunveE (Los Angeles, Calif.) 1973 1975 
J. M. Burns (Cambridge, Mass.) 1974 J. A. Powext (Berkeley, Calif.) 1975 
R. H. Carcasson (Vancouver, B.C.) 1974 


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 $10.00 
Student members—annual dues $5.00 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $15.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia 23455. 


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


by Cyr F. pos Passos 


Price, postpaid: Society members—$5.00, others—$7.50; uncut, unbound signatures 
available for interleaving and private binding, same prices; hard cover bound, mem- 
bers—$8.00, others—$10.00. Revised lists of the Melitaeinae and Lycaenidae will be 
distributed to purchasers free (separately with paper covered copies and unbound 
signatures, bound in with hard covered copies). 


The Lepidopterists’ Society is a non-profit, scientific organization. The office of 
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520. 
Second class postage paid at Lawrence, Kansas, U.S.A. 66044. 


JOURNAL OF 


Tue LEPIDOPTERISTS’ SOCIETY 


Volume 27 1973 Number 2 


LEPIDOPTERA FEEDING AT PUDDLE-MARGINS, 
DUNG, AND CARRION 


J. A. Downes 
Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario 


Butterflies are often seen feeding at the margins of small puddles, 
sometimes in considerable numbers, and it is known also that certain 
species are found at dung and at carrion. In the summers of 1970 and 
1971, in the course of other field work, I was able to make some observa- 
tions on these habits, and came to the conclusion that they are closely 
related expressions of the same phenomenon. I also found that these 
feeding sites are well frequented not only by day but also at night, by 
moths of at least five families. The observations were made at Black 
Lake in North Burgess Township, some fifteen miles south of Perth, 
Ontario, along field tracks through former pasture land and the adjacent 
mixed hardwood and swampy areas. 


Observations by Day 


A list of species observed feeding at the margins of puddles by day 
is given below. For every specimen involved, it was clearly established 
that the proboscis was extended in the feeding position Perhaps, in all, 
twice as many species were seen beside the puddles, but man) individuals 
could not be approached for critical examination. 


NympHatmar. Speyeria cybele Fab.; Boloria bellona Fab 
Drury; Polygonia comma Harris; Vanessa virginiensis Drur 
Cramer. Ga - 

LycaENDAE. Harkenclenus titus Fab.; Feniseca tarquinius Fab.; Lycaend pil 
L.; Celastrina argiolus L. 

Prrmae. Colias philodice Godt.; Pieris rapae L. 

Hesperupar. Erynnis lucilius Scudder and Burge 
P. peckius Kirby; P. mystic Scudder; Wallengrenia ¢ 

Pyratipae. Pyrausta orphrisalis W\k.; Desmia *: 


90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fifty-two individuals of these species were examined for sex; all were 
males with the exception of 2 (out of 5) specimens of Lycaena phlaeas. 

The probing butterflies were found, with few exceptions, around four 
small groups of puddles situated at intervals of a quarter mile or so along 
the field road. At three of these stations a swampy ditch with open water 
ran along one or both sides of the track for a considerable distance, and 
all along the roadway there were patches of thick grassy vegetation 
where dew might remain throughout the forenoon. Often many butter- 
flies were congregated at the puddles for several hours before the dew 
had dissipated. Thus it seemed unlikely, from the outset, that the basic 
function of puddle-visiting was to obtain water, which was usually in 
ample daily supply on a wide scale. 

The butterflies probed, typically, at the moist soil around the margins 
of the puddles, not at the free water itself. They avoided standing on 
the moister regions, and the skippers in particular often perched on a 
dry pebble or twig and, working their way towards the edge, probed 
with the tongue at the damp mud below. The mud remained attractive, 
through not so strongly, even as it dried and ceased to be visibly moist; 
and on occasion Lycaena phlaeas and several of the nymphalids and skip- 
pers were seen probing industriously at normally dry soil more than a 
week after any visible puddle had disappeared. (It is always important, 
in these situations, to establish that the butterfly is engaged in feeding, 
and not merely in sunning—the more so as nymphalids, when they are 
feeding on the ground, usually hold the wings open.) On one occasion 
Vanessa virginiensis was encountered probing steadily among dry duff 
and topsoil beneath pine trees; and Feniseca tarquinius often probed for 
long periods over slightly moist rotten wood lying on the ground. 

Nevertheless, the presence of the puddle clearly increased the attractive- 
ness of the site, and a depression visited only by an occasional individual 
in dry weather became a centre of interest for six or eight species when 
water was again available. The puddles, of course, all lay in depressions, 
and were filled from time to time by water seeping or draining down 
the pathway on either side. Probably they represented areas of con- 
centration of organic debris and solutes, and it seems likely that this was 
the real source of their attractiveness. The decaying organic material 
probably gave off odorous substances, able to attract or arrest passing 
insects, especially when moist; and its attractiveness would decrease as 
it became dry. 

As already mentioned, butterflies are sometimes observed also on dung 
and on carrion. When horse dung, and still more when recently killed 
frogs or turtles or small mammals were added to a puddle, the numbers 


VoLUME 27, NuMBER 2 9] 


of insects visiting it and probing increased very considerably. The in- 
dividuals probing directly at the dung or carrion were remarkably stable 
—especially those on carrion, which were so intent on probing that they 
could usually be approached and picked up with the fingers. Specimens of 
Harkenclenus titus, Celastrina argiolus, Feniseca tarquinius, Phyciodes 
tharos, Vanessa virginiensis and Limenitis arthemis were collected in this 
way from a dead frog. At an ordinary puddle, the butterflies move around 
fairly readily, probing repeatedly with the proboscis as they gradually 
shift their position near the margin. On encountering a piece of dung 
they are at once stabilized, and continue feeding there for a considerable 
period; and still more so on encountering carrion. It seems that the 
ordinary but inevitably somewhat enriched puddle, the horse dung, and 
the dead animals represent successively higher levels of the same stimuli. 
The order of preference was the same for all species present in significant 
numbers, but no additional species, not found at the ordinary puddles, 
visited the dung or carrion. 

Just as some butterflies probed at dry mud that could scarcely have 
yielded any significant amount of liquid, so also, occasionally, they 
visited and probed at desiccated pieces of dung. A specimen of Polygonia 
(? interrogationis Fab.) explored a dry gunny sack that had contained 
horse manure for upwards of 20 minutes, all the time probing between 
the fibres with the proboscis. Lycaena phlaeas also was seen at dry 
horse dung; and Erynnis sp. landed and probed repeatedly on a dry bird 
dropping on a bare rock exposed to the sun. The lepidopteran proboscis 
is adapted for taking up liquids only, and it seemed therefore of interest 
to make some closer observation of these attempts to feed on dry ma- 
terials. 

Specimens of Lycaena phlaeas (3 ¢¢; 1 2), Phyciodes tharos, and 
Polygonia comma, held in glass vials were offered dry clippings of skin 
and muscle from a dead frog, and in direct sunlight they probed readily 
with the extended tongue. The manner in which Lepidoptera feed on 
dry sugar-containing materials is being described elsewhere (Downes, in 
prep.), and the feeding process on other dry materials is very similar. 


Briefly, the tongue is extended and the anterior (dorsal) face of the apical 
region—the only part used for food intake; the main length of the 
proboscis merely enabling the insect to probe at a cistance—is applied 


n the two 


closely to the surface of the skin clipping. The 
galeae that make up the tongue opens slightly 

released along the whole length of the apical r is applied 
to the food. The saliva penetrates among the | kin and muscle 
for a distance of perhaps five tongues’ width 2° hen quickly 


oO} saliva is 


92 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


withdrawn into the tongue, by suction exerted by the muscles of the 
cibarium. At a satisfactory feeding point the cycle of outflow and with- 
drawal of saliva may be repeated 10-20 times in as many seconds, and 
the tongue then probes again nearby. Thus the surface of the food is 
repeatedly washed with saliva, and soluble materials liquefied and 
imbibed. There can be little doubt that this is what happens in nature 
when butterflies probe at dry mud or horse dung. Somewhat similarly, 
individuals of Feniseca tarquinius and Danaus plexippus (L.) have been 
observed salivating and re-imbibing on the skin of the hand, at dry sweat 
and on areas that had earlier been lightly painted with sodium chloride. 
It thus appears that Lepidoptera may obtain not only sugars but also 
other nutrients from dry deposits, by washing them out with saliva. 
[Unfortunately none of the skipper butterflies were observed in detail 
when feeding on dry materials. There are several records in the literature 
of hesperids moistening the food by a droplet of liquid from the anus, 
and then imbibing (see Norris, 1936). I think, however, that this repre- 
sents a special case. | 

The requirement that is satisfied by feeding at puddle-margins and 
the related sources remains unknown. The obvious first suggestion is a 
need for protein or amino acids. There seems, however, to be no indica- 
tion in the literature of the occurrence of proteinases in the digestive 
system of Lepidoptera; and it would, moreover, be remarkable that a 
requirement for nitrogen should be almost restricted to the male sex. 
Other possibilities are a requirement for salt, or some trace substance 
for which the male, with his probably greater flight activity, has more 
need. The same puddles were visited also by many kinds of small bees 
and bombyliid flies; perhaps, therefore, a number of specialized nectar 
feeding insects have certain requirements that floral nectar does not 
supply. 

A few observations were made on the discovery of the puddle and the 
choice of the feeding point. The puddle area seemed to be encountered 
by accident during general flight and then, probably at rather close 
range, the increase in concentration of odorous substances led to settling 
on the ground in the general vicinity of a feeding area. A satisfactory 
feeding point was discovered by random walking, probing meantime at 
the surface with the proboscis; and similar random movements, as al- 
ready noted, led eventually to the feeding points on dung or carrion 
where the insect became more fully stabilized. The discovery of the 
feeding point, of whatever grade of stability, seemed to be mainly ‘acci- 
dental, by unoriented questing. 


More specific stimuli may sometimes be involved. The skippers, both 


VoLUME 27, NuMBER 2 93 


Erynnis and Polites, often landed directly onto small whitish patches of 
bird droppings (which are evidently a favourite feeding site in this family), 
and sometimes also onto horse dung. The precision of landing suggested 
an element of visual control as the approach was completed. Occasionally 
Celastrina argiolus also was seen to land directly on bird droppings. The 
sulphur butterflies often landed within inches of other individuals, evi- 
dently recognising them by sight; and in this way, probably, the familiar 
aggregations of puddle-teeding butterflies are built up. 


Observations at Night 


As noted above, two species of Pyralidae were occasionally seen feeding 
with the butterflies at the puddles. The puddle-visiting habit has seldom 
been recorded except for the Rhopalocera, and this observation suggested 
that it would be interesting to visit the sites by night. On the first night, 
§ July 1971, about an hour after dark, upward of fifty moths, mostly 
Geometridae and Pyralidae but also a few Noctuidae, Pterophoridae and 
Tortricidae, were seen at the puddles. It was curious and striking to 
encounter six or eight conspicuous white or yellow pyralids and geometrids 
perched on the shell of a crushed turtle, busily probing with the proboscis 
between the fractures in the carapace. Good numbers were seen also on 
two occasions later in the month, but few or none on four other nights. 

The moths also were stabilized more strongly by carrion than by the 
normal damp margin of the puddle. Except for a few of the pyralids, 
the moths were generally much more quiescent while feeding than the 
daytime butterflies and usually could be collected directly into individual 
vials. All the geometrids, except for Hydria prunivorata, sat with the 
wings closed, butterfly fashion, above the back. Most of the species 
encountered were long-tongued forms that are probably also nectar 
feeders, except for the tortricids, which perhaps feed on honeydew, and 
the large noctuid Euthisanotia grata. This latter species has, for a 
noctuid, a quite short tongue, and to feed it crouched close to the mud 
with the wings widely spread out to the sides; in this position it evidently 
mimics a large variegated bird dropping. 

As with the daytime visitors, the moths were very predominantly males; 
the single specimen of Tarachidia erastrioides and one of the two Ble- 
pharomastix ranalis were the only females among 63 specimens collected 
and sexed. All these specimens had been clearly observed, before capture, 
with the proboscis extended and probing at the mud or carrion. The 
list of species represented is given below; but again other species were 
seen in apparent feeding situations but were not collected because the 
observation could not be satisfactorily completed. 


94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


NocruiwAr. Leuconycta diphtheroides Gn.; Euthisanotia grata Fab.; Tarachidia 
erastrioides Gn.; 

GEOMETRIDAE. Xystrota rubromarginaria Pack.; Scopula ancellata Hlst.; S. enu- 
cleata Gn.; Euphyia centrostrigaria Woll.; Eupithecia misturata frostiata Swett.; 
Lygris diversilineata Hb.; Hydria prunivorata Ferg.; Euchlaena johnsonaria Fitch; 
Itame exauspicata Wlk.; Xanthotype urticaria Swett.; V. sospeta Drury; Sicya macularia 
Harr.; Nematocampa limbata Haw.; Anacamptodes vellivolata Hlst. 

PyrauwwaE. Argyria nivalis Drury; Crambus sp.; Blepharomastix ranalis Gn.; 
Acrobasis sp.; Tetralopha sp.; Mecyna submedialis Grt.; Herpetogramma pertextalis 
Led.; Pantographa limata G. and R.; Pyrausta orphrisalis Wlk.; Desmia funeralis Hb. 

PrerorpHoRIDAE. Platyptilia pallidactyla Haw. 

TORTRICIDAE. Sparganothis sp. nr. putnamana Free.; Argyrotaenia quadrifasciana 
Fern.; Argyrotoxa semipurpurana Kft.; Choristoneura rosaceana Harris. 


Conclusions from Observations 


The chief results that emerge from these observations are as follows. 

1. The habit of feeding at puddle-margins is not, basically, related to 
water requirements, though it may of course often be the way the need 
for water is supplied. Puddle-visiting occurs even when ordinary water 
is plentiful nearby, and the same species may (a) sometimes probe also 
at the dry sites of former puddles, and (b) prefer dung and carrion to 
the ordinary puddle-margin. The tested species fed from dry materials 
by washing them with repeated pulses of saliva discharged from the 
apical region of the proboscis and then re-imbibing. 

2. Representatives of most of the main groups of butterflies found in 
the study area were observed at the puddles/dung/carrion. At night the 
same feeding sites were visited by moths, principally Geometridae and 
Pyralidae but also a few Noctuidae, Tortricidae and Pterophoridae. Both 
the day- and night-time visitors were very predominantly (approx. 967% ) 
males. 

3. The nutritional requirement satisfied by probing at puddles/dung/ 
carrion is not known, but is unlikely to be a requirement for protein. 
The habit is presumably quite distinct from the sugar-feeding habit 
general in Lepidoptera, and probably represents a secondary development 
related to a specialized need in certain families or under certain circum- 
stances. 

4. The feeding areas are discovered probably by a response to odour 
while in flight. The precise point of feeding is apparently reached, after 
landing, by random questing. There may also be visual responses to 
already established insects, this tending to build up aggregations, and 
sometimes perhaps to special feeding sites, such as bird droppings. 


Discussion 


Many notes have been written on these aspects of the feeding habits 
of adult Lepidoptera, but they are widely scattered and mostly brief. 


I Te 


VoLUME 27, NUMBER 2 95 


An extensive review was published by Norris (1936), and that paper 
remains the basic reference. Initially Norris treats the puddle-visiting 
habit as water drinking; but numerous instances of feeding at dung, 
sweat, salt, the moisture exuding from the eyes of cattle, and such like, 
are then assembled, and it is pointed out that many puddle and riverside 
drinking places are contaminated with animal excreta. It is noted that 
the excreta or soil that the insects visit is sometimes quite dry; and that 
drinking areas are often restricted to contaminated sites even when 
cleaner water is abundantly available. Norris concludes, “The problem 
of water-drinking in the Lepidoptera is [thus] inextricably confused with 
that of their attraction to dung and sweat... There is reason to believe 
that practically all water-drinking may be primarily due to such attrac- 
tion.” My own observations emphasise even further the significance of 
dung and carrion, which are more powerful attractants (or arrestants ) 
than lightly contaminated moist soil. The insects probed at such materials 
even in the dry state and the significant substances were presumably 
extracted by the saliva that they were seen to discharge and re-imbibe. 
It seems clear that decaying animal materials must have a significance, 
and presumably supply a nutritional need, over and above any require- 
ment for water as such. 

Norris’ records nearly all show a great preponderance of males, similar 
to that noted above. Females are well represented (about one third of 
the total) only among the Noctuidae feeding at sweat-impregnated 
clothing in Collenette’s study (1934) in Brazil. The usual great inequality 
itself suggests that some additional, though perhaps minor, nutritional 
requirement is obtained, since it is unlikely that water would be required 
in very different quantity in the two sexes. 

The habit of probing at contaminated soil and decaying animal refuse 
is known throughout the Rhopalocera. The families Papilionidae and 
Satyridae, lacking, presumably by chance, from my own observations, 
are mentioned on many occasions by Norris, and latterly by Payne & King 
(1969). The Riodinidae were mentioned by Bates (1863). Several 
authors have noted a few day-flying Pyralidae, Geometridae or Thyridi- 
dae along with the butterflies, but only Collenette (1934, Brazil) appears 
to have observed a substantial night-time fauna comparable to that 
recorded above. A few records are given also by Payne & King (1969). 
Outside the Rhopalocera the habit seems to be limited mainly to certain 
systematic groupings. Pyralidae, Geometridae and Noctuidae are the 
most frequent, with several observations also of Sphingidae; most of 
the remaining records refer to families related to one or another of the 
above (Thyrididae; Uraniidae; Arctiidae, Agaristidae, Notodotidae). The 


96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


present records of Tortricidae and Pterophoridae, and the old record 
of Psychidae quoted by Norris (p. 81), stand rather apart. 

The strikingly gregarious habits of many puddle-frequenting butter- 
flies receives considerable attention from Norris; and some fine photo- 
graphs of dense groups, chiefly in the tropics, are given by Klots (1958). 
Collenette & Talbot (1928), in experiments with Catopsilia involving 
paper models, have shown that the group is built up by visual recognition 
of the already established individuals. The gregarious habit is especially 
conspicuous in Pieridae. In Nymphalidae and Hesperidae, however, a 
response to odour is highly developed, and the insects tend to respond 
individually to the feeding sites and often do not form notable aggrega- 
tions. Many Nymphalidae likewise find their sources of sugar mainly 
by smell, and tend to feed at ripe fruit rather than at conspicuously 
coloured flowers. 

In discussing the various forms of the habit of feeding at animal excreta 
and refuse, Norris refers to Shannon’s observations (1928) in Argentina 
that certain moths probe at the secretions and pus around the eyes of 
mammals. More recently considerable attention has been given to the 
eye-frequenting moths, in studies in tropical Africa and Asia (Reid, 1954; 
Bittiker, 1967; Banziger & Biittiker, 1969). Irritation and pain are 
caused as the moth probes under the eyelid with its proboscis, and the 
possibility of transmission of infections has been raised. The ingested | 
material includes lachrymal secretions and pus, and sometimes also 
blood from accidental sores or wounds. About 30 species of eye-frequent- 
ing moths have been recorded. The great majority belong to the Noctui- | 
dae, Pyralidae and Geometridae, in somewhat similar numbers; and in | 
addition there are two records of Notodontidae and one of Sphingidae | 
(and one record also of a lycaenid butterfly). Thus the eye-frequenting | 
moths belong to exactly the same array of families as the other excreta | 
feeding forms. The relationship between the two groups is sometimes | 
much closer still. The genus Arcyophora (Noctuidae) which includes | 
several of the most important eye moths, includes also a species taken | 
on the carcass of a bullock (Reid, 1954). Pingasa chlora crenaria Guen. | 
(Geometridae) is recorded both from the eyes of ungulates (Biittiker, 
1967) and from dung and contaminated soil (Banziger, 1971); the genus 
Scopula contains an eye-frequenting species in Thailand (Biittiker, 1967) 
and puddle-margin and carrion species in Ontario (above); and Semio- } 
thisa, again with eye-frequenting species in Thailand, is represented by 
the very closely related genus Itame in the present work. The genus 
Pyrausta figures in Shannon’s (1928) list of eye moths in Argentina and | 


VoLUME 27, NUMBER 2 97 


in the present work. Thus Norris is almost certainly correct in making 
the association. | 

The sex ratio of the eye-frequenting moths is adequately recorded in 
only two cases, the noctuids Arcyophora sylvatica Bittiker and Lobo- 
craspis griseifusa Hampson. In the former, males and females occur in 
similar numbers, and in the latter females exceed males by about three 
to one. This is very different from the usual sex-ratio of puddle-margin 
and dung feeding Lepidoptera, though roughly similar to the particular 
case of the Noctuidae recorded on sweat-impregnated clothing by Col- 
lenette (1934). Collenette’s itemized list of the sex-ratios of the various 
species shows that this habit, in the Noctuidae, requires a much more 
detailed study. It is clear, however, that the usual preponderance of 
males at animal excreta is not an absolute phenomenon; it does not indicate 
a radical difference in the physiology of the two sexes such as is found 
in the mosquitoes and other bloodsucking Diptera, but a more quantita- 
tive difference that might vary with circumstances. Thus the sex-ratios 
of the eye-frequenting noctuids do not tell seriously against the derivation 
of the habit from excreta-feeding. 

Banziger (1968, 1971) has found recently that the noctuid, Calpe 
eustrigata Hamps., is a true bloodsucker, able to pierce the skin of mam- 
mals with its apicaily armoured proboscis. Another species of Calpe, C. 
minuticornis Guen., is recorded as a eye-moth, and yet another eye-moth, 
Mocis undata Fabr., falls into the same sub-family (Catocalinae ) ( Buitti- 
ker, 1962, 1967). It is very likely therefore that this bloodsucking repre- 
sents an extreme case of the group of feeding habits under discussion. 
No information as to sex is given in the work on these insects. 

Banziger, however, suggests that blood-sucking is a derivative of the 
fruit-piercing habit. Fruit-piercing is well known among this section of 
the Noctuidae, and many of the moths have an armoured proboscis 
approaching that of C. eustrigata (Hargreaves, 1936; Biittiker, 1962; 
Banziger, 1971). The suggested relationship of the two habits appears, 
nevertheless, to be very improbable. Fruit-piercing, like flower-visiting, 
provides the insect with sugar; the mud-, dung-, and carrion-visiting 
habits, from which eye-frequenting and blood-sucking have apparently 
evolved, provides the insect with nutrients of an entirely different kind. 
The latter requirement, whatever its nature may be, is typically char- 
acteristic of the male sex; and occurs only in certain families, for the 
most part highly specialized ones and frequently ones in which the sugar- 
feeding habit is also highly developed (e.g. Rhopalocera, Noctuidae). 
Indeed, fruit-piercing has been observed in the blood-sucking Calpe 
eustrigata itself (Banziger, 1968, 1971), in the eye-visiting C. minuticornis, 


98 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in other species of Calpe, and in many other genera of the sub-family. 
Thus the two types of habit are by no means alternatives, one of which 
has evolved from the other; rather sugar-feeding is a basic (plesiotypic) 
habit of the Lepidoptera that has been maintained in the majority of 
the component groups, while the ‘animal excreta’ group of habits appears 
to be a development sui generis that exists alongside sugar-feeding in a 
limited range of forms. (Banziger is further mistaken in drawing a com- 
parison with the mosquitoes. Here again there are two kinds of feeding 
habit, sugar feeding and blood sucking; but they exist side by side in 
many species and are indeed both plesiotypic features of the order 
Diptera. In no sense are they alternatives one of which has evolved 
from the other (see Downes, 1958).) There is however only one feeding 
organ and one mode of food uptake in Lepidoptera; and it would not 
be unexpected that a modification of the proboscis associated with a 
change in the manner of taking sugar (from probing at floral nectaries 
or rotting fruit to piercing the rind of sound fruit) might elicit a change 
in the other feeding habit also (from absorbing liquid in contaminated 
mud or dung to probing under the eyelid and finally to piercing the 
skin). This is what appears to have taken place in Calpe eustrigata. 


General Summary 


An accessory feeding habit, that of feeding at contaminated liquids 
and decaying animal material, is found in several groups of Lepidoptera, 
especially Rhopalocera, Noctuidae, Geometridae and Pyralidae. The habit | 
is usually almost restricted to the male sex. Its significance is not known | 
but it is probably not related to a need for protein and almost certainly 
not, in essence, a requirement for water; well dried materials can be 
utilized by repeated salivation. Its most common expression is the habit | 
of feeding at the margins of contaminated puddles, but dung and carrion | 
are preferred if available. In the tropics various noctuids, geometrids _ 
and pyralids probe at the exudates of the eyes of mammals, and it seems _ | 
likely that this is a specialized form of the same habit; it is proposed | 
also that the blood sucking of the noctuid Calpe eustrigata is a further | 
specialization of the same nature. | 


ACKNOWLEDGMENTS 


I am much indebted to my colleagues W. C. McGuffin, Akira Mutuura, | 
G. G. Lewis and E. W. Rockburne for assistance with the identifications, | 
and to C. F. Hinks, E. H. Salkeld and W. C. McGuffin for commenting 
on a draft manuscript and for many interesting discussions. : 


VoLUME 27, NUMBER 2 99 


LITERATURE CITED 


BAnziceR, H. 1968. Preliminary observations on a skin-piercing blood-sucking 
moth [Calyptra eustrigata (Hmps.) (Lep., Noctuidae )] in Malaya. Bull. Entomol. 
Res. 58: 159-163. 

1971. Bloodsucking moths of Malaya. Fauna 1: 5-16. 

. & W. Burtixer. 1969. Records of eye-frequenting Lepidoptera from man. 
J. Med. Entomol. 6: 53-58. 

Bates, H. W. 1863. The Naturalist on the River Amazons. 2 vols. John Murray, 
London. 

Burtiker, W. 1962. Biological and morphological notes on the fruit-piercing and 
eye-frequenting moths. Verhandl. XI Intern. Kongr. Entomol., Wein (1960) 2: 
10-15. 

1967. Biological notes on eye-frequenting moths from N. Thailand. Mitt. 
Schweiz. Entomol. Ges. 39: 151-179. 

CoLLENETTE, C. L. 1934. On the sexes of some South American moths attracted 
to light, human perspiration and damp sand. Entomologist 67: 81-84. 

.& G. Tatsor. 1928. Observations on the bionomics of the Lepidoptera of 
Matto Grosso, Brazil. Trans. Entomol. Soc. London 76: 391-414. 

Downes, J. A. 1958. The feeding habits of biting flies and their significance in 
classification. Ann. Rev. Entomol. 3: 249-266. 

Harcreaves, E. 1936. Fruit-piercing Lepidoptera in Sierra Leone. Bull. Entomol. 
Res. 27: 589-605. 

Kuots, A. B. 1958. The World of Butterflies and Moths. Harrap, London, 207 p. 

Norris, M. J. 1936. The feeding habits of the adult Lepidoptera Heteroneura. 
Trans. Roy. Entomol. Soc. London 85: 61—90. 

Payne, J. A. & E. W. Kine. 1969. Lepidoptera associated with pig carrion. J. 
Lepid. Soc. 23: 191-195. 

Rew, E. T. M. 1954. Observations on feeding habits of adult Arcyophora. Proc. 
Roy. Entomol. Soc. London, B, 23: 200-204. 

SHANNON, R. C. 1928. Zoophilous moths. Science 68: 461-462. 


A NEW GENUS AND SPECIES OF OECOPHORIDAE FROM 
TROPICAL AMERICA 


J. F. Gates CLARKE 


National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560 


The species described below is so striking, and such an unusual 
oecophorid, that its presence in the Neotropical fauna should be recorded. 
It is nearly related to the genus Filinota Busck and adds another link in 
the growing classification of the South American fauna. 

The drawings for this paper were made by Mrs. Elsie H. Froeschner, 
staff artist, and the photograph was produced by Victor Krantz, both 
of the Smithsonian Institution. 


100 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Profilinota phillita, new species: adult male paratype. 


Profilinota, new genus 


Type species: Profilinota phillita, new species (by monotypy and present 
designation ). The generic name is of feminine gender. 


Labial palpus very long, slender, recurved, exceeding vertex; third segment slightly 
longer than second, acute. Maxillary palpus minute, appressed to base of tongue. 
Head roughened; ocellus absent. Antenna as long as forewing, long ciliated for more 
than three-fifths its length; scape with pecten. Forewing with 12 veins; 1b furcate; 
2 from near outer three-fourths; 3 from angle; 3, 4 and 5 nearly equidistant; 7 and 8 
stalked, 7 to termen slightly below apex; 10 nearer to 9 than to 11; 11 from middle; 
termen very slightly convex. Hind wing with 7 veins; 2 from three-fifths; 3 and 4 
coincident, from slightly before angle; 5, 6 and 7 about equidistant. Posterior tibia 
roughened above with long hairlike scales. Male genitalia with uncus absent; gnathos 
well developed. 

This genus is very closely related to Filinota Busck (1911: 206), but 
differs from it by the separate veins 3 and 4 of the forewing, by having a 
well developed antennal pecten, the absence of an uncus; and the length 
of third segment of palpus is longer than second. 

Busck says of the antenna of his type “basal joint without pecten.” 
The type, and three additional specimens before me have no pecten, al- 
though two examples have a single scale, somewhat out of place, on the 
scape of one side, that might be considered a weak development of that 
character. Busck further states “7 to costa” [of forewing]. I would deter- 


101 


e 


1/ 


OA. Frceschnar 


c 
c 


aspect of head showing palpus; 


lateral 


? 


a 


VQ 
\ 


Zw 
LL a \ 


a g 
Sf 
ee N Yj 


Ss 
SSS 


YY LE 


Sr ae 
Se oe ee Sify 
ES EW Go 
RtELLZE 
LEELA, 


Fig. 2. Profilinota phillita, new species: 
b, section of antenna showing pecten and cili 
ventral view of male genitalia with left harpe 


VoLUME 27, NuMBER 2 
equals 1 mm.); e, aedeagus. 


d, 


s: c, venation of right wings; 
and aedeagus omitted (scale shown 


ation 


-_—— —— - 


102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


mine the position of vein 7 as at apex. Vein 7 of Profilinota extends to 
termen just below apex. 

Meyrick (1922: 158) redescribed Filinota, apparently from a combina- 
tion of characters of several species. He stated “ocelli posterior” but they 
are absent from the type species. He also gave a rather startling array 
of venational characters, but these various combinations are suspect as 
far as the generic limits of Filinota are concerned. 


Profilinota phillita Clarke, new species 
Figs. 1-2 


Alar expanse 32-34 mm. Labial palpus buffy brown; second segment olive brown 
on outer side at apex; third segment olive brown beyond basal third; apex light buff. 
Antenna olive brown shading to buffy brown apically. Head sordid white on crown 
and vertex; face buffy brown; lateral tufts olive brown. Thorax light buff; collar 
and base of tegula and spot posteriorly olive brown; apex of tegula light buffy brown. 
Forewing ground color buffy brown shading to olive brown dorsally; on dorsal 
edge two elongate white marks, the first reaching to near middle, the second to 
tornus; at end of cell a faint olive brown spot; cilia white at tornus, then concolorous 
with forewing. Hingwing basal two-thirds white; outer third buffy brown; cilia 
white adjacent to white portion of wing, buffy brown beyond; a few white cilia at 
apex. Foreleg pale buffy brown; tibia shaded olive brown on outer side; tarsal 
segments olive brown on outer side; midleg similar; second to fourth tarsal segments 
buff basally; hindleg pale buffy brown; tarsal spurs buff; tarsal segments marked 
buff on outer side. Abdomen buffy brown dorsally, ocherous white ventrally. 

Male genitalia (slides JFGC 12252, 12253): Harpe elongate, triangular, broad 
basally, tapering to a bluntly pointed cucullus; on outer side of harpe, about middle, 
a cluster of long setae. Gnathos large, consisting of two spined lobes. Uncus absent. 
Vinculum rounded, thickened medially. Tegumen about as long as harpe, truncated 
posteriorly. Anellus an oval plate with a sclerotized circular band posteriorly. 
Aedeagus stout, slightly curved, pointed and armed with two sets of teeth, one 
ventrally, one apically; vesica unarmed. 

Holotype: U.S. National Museum No. 72172. 

Type locality: Venezuela, Aragua, Rancho Grande, 1100 m. 

Distribution: Venezuela, Peru. 

Food plant: Unknown. 

Described from the holotype male (31 October 66, S. S. and W. D. Duckworth) 
and one male paratype (Peru, R. Huacamayo, Carabaya, dry s., 3100 ft., June 04, 
G. Ockenden). Female unknown. Paratype in the British Museum (Natural History). 


There is no described species, except the type species of Filinota, F. 
hermosella Busck, with which this striking moth can be compared struc- 
turally. This brown and white species presents a great contrast to the red, 
yellow and white hermosella, but reminds one of an oversized Hastamea 
argentidorsella Busck, from which it is abundantly distinct structurally. 


LITERATURE CITED 


Buscx, A. 1911. Descriptions of Tineoid Moths (Microlepidoptera) from South 
America. Proc. U. S. Nat. Mus. 40(1815): 205-230, pl. 8, 9. 

Meyrick, E. 1922. In Wytsman, Genera Insectorum. Lepidoptera, Heterocera. 
Fam. Oecophoridae. Fasc. 180: 1-224, pl. 1-6. 


VOLUME 27, NuMBER 2 103 


RECORD AND ILLUSTRATION OF SOME INTERESTING 
MOTHS FLYING IN TEXAS (SPHINGIDAE, CTENUCHIDAE, 
NOCTUIDAE, NOTODONTIDAE, GEOMETRIDAE, 
PYRALIDAE, COSSIDAE) 


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


This paper illustrates and records the presence in Texas of some moths 
which are considered interesting because they are either rare, or not 
included in the McDunnough check list. Some of them are probably 
new records for the United States. 


SPHINGIDAE 


Amplypterus donysa (Druce). Fig. 1. 
Ann. Mag. Nat. Hist., Ser. 6, 4: 78, 1889. 

Big Bend Nat. Park, Green Gulch, 3 May 1972, 1 6; 6 May 1972, 1 9; 12 May 
1972, 1 2. The male is rubbed but the females are very nice, and it seems probable 
that Green Gulch or some other place nearby was the breeding area. Dr. J. G. 
Franclemont who was with us when we discovered these insects in our traps says 
(in litt.) “. . . from what little is known about the host plants of the group, I judge 
that larva feeds on some member of the Anacardiaceae, possibly Rhus microphylla 
or R. triloba.” Both of these were growing where the moth was taken. The forewings 
are different shades of ashy green, the hindwings mostly of the color of crushed 
strawberries. 


CTENUCHIDAE 


Syntomeida melanthus (Cramer). Fig. 2. 
Paparxouo O45 b> (index); pl. 248, fig.c, 1779. 
Big Bend Nat. Park, 21 specimens, all taken in the desert around the Chisos 
Mountains, from early April to late June, and from late August to early October. 
The wings are bluish black with yellowish-white spots. 


Episcepsis inornata Walker. Fig. 3. 
List Lep. Ins. Coil. Brit. Mus. 7: 1636, 1856. 
Santa Ana National Wildlife Refuge, 14 November 1971, 2 22. The specimen 
pictured is badly rubbed. The wings are powdery black. The abdomen is sprinkled 
with shinning steel-blue scales. 


NOCTUIDAE 


Euxoa xasta Barnes & McDunnough. Fig. 4. 
Can. Entomol. 42: 429, 1910. 

The rarity of this Agrotinae is my reason for giving my record of captures, al- 
though it is included in McDunnough’s check list and Kerrville, Texas is the type 
locality. Fort Davis, 17 May 1966, 1 ¢; 18 May 1971, 1 ¢. Big Bend National 
Park, Government Spring, 27 March 1971, 1 ¢; 6 May 1972, 1 6, 3 QQ. Sierra 
Diablo Wildlife Management Area, 14 July 1971, 1 @. 

Eriopyga mulina Schaus. Fig. 


Trans. Amer. Entomol. Soc. 21: 237, 1894. ; 
Big Bend National Park, where trees and brush are growing: Green Gulch, Oak 


oy | 


104 JouURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-7. 1, Amplypterus donysa (Druce) @, Big Bend Nat. Pk., Green Gulch, 
6 May 1972; 107 mm. 2, Syntomeida melanthus (Cram.) ¢, Big Bend Nat. Pk., 
Chihuahua Desert near Nugent Mt., 17 Sept. 1971; 45 mm. 3, Episcepsis inornata 
Wlk. 2, Santa Ana Nat. Wildl. Ref., 14 Nov. 1971; 32 mm. 4, Euxoa xasta B. and 
Mc.D. 6, Big Bend Nat. Pk. Government Spring, 27 March 1971; 36 mm. 5, 
Eriopyga mulina Schaus 9, Big Bend Nat. Pk., Green Gulch, 4 October 1967; 34 mm. 
6, Emariannia cucullidea Benj. 6, Sierra Diablo Wildl. Mgt. Area, 1 Sept. 1970; 
36 mm. 7, Neophanis respondens Wik. 9, Brownsville, 7 Nov. 1969; 42 mm. 


VOLUME 27, NuMBER 2 105 


Spring, Government Spring; 13 specimens, one in March, one in May, the others 
from late September to late October. Forewings different shades of lustrous brown, 
hindwings more dusky. 


Emariannia cucullidea Benjamin. Fig. 6. 
Pan Pac. Entomol. 9: 3, 1933. 

Alpine, 17 Sept. 1962, 1 9. Fort Davis, 27-29 Aug. 1964, 2 6 4,2 9 2; 6 Sept. 
1964, 1 6; 28-29 Aug. 1970, 2 6 6. Sierra Diablo Wildlife Management Area, 15 
July 1969, 1 ¢; 1 Sept. 1970, 1 ¢. The forewings are patterned in dusky brown 
over a lighter brown background. The hindwings are white. 


Neophanis respondens Walker. Fig. 7. 
List Lep. Ins. Coll. Brit. Mus. 15: 1720, 1858. 
Brownsville, 7 Nov. 1969, 1 2. Forewings patterned in black over mossy green. 
Hindwings shiny orange-yellow with black margin. 


Iscadia daemonalis Dyar. Fig. 8. 
Proc. U.S. Nat. Mus. 38: 258, 1910. 
iaae Bend National Park, Green Gulch, 6 Oct. 1966, 1 ¢. Artesia Wells, LaSalle 
, Chaparral Wildlife Management ARR. JS=aSeptalOilevors6 os lO Nove 19715 
: * dé; 12 June 1972, 1 6, 1 @. Santa Ana National Wildlife Benne! 14 Nov. 
1971, 1 @. Forewings dark blackish brown, tipped with gray, crossed with fine, 
wavy, transverse black lines. Hindwings white, with black margin in the female. 


Meropleon titan Todd. Fig. 16. 
J. Wash. Acad. Sci. 48(1): 27, 1958. 
Canadian, 27 September 1968, 3 ¢ ¢. This collection extends the known distribu- 
tion of this species far to the west. Forewings dark brown; hindwings lghter. 


Sigela basipunctaria Walker. Fig. 9. 
List Lep. Ins. Coll. Brit. Mus. 23: 785, 1861. 
San Antonio, Mountain View Acres (Roy Kendail’s Ebony Hill Laboratory), 9 
March 1972, 1 ¢. Fishing camp on Guadalupe River, north of New Braunfels, 12 
March 1972, 1 ¢. Black dots on whitish background. 


Matigramma psegmapteryx Dyar. Fig. 17. 
Proc. U.S. Nat. Mus. 44: 301, 1913. 
Garner State Park, 24 March 1965, 1 ¢, (identified by J. G. Franclemont who 
prepared slide A.B. 4). The wings are different shades of neutral gray. 


Herminodes stigmaphiles (Dyar). Fig. 18. 
Proc. U.S. Nat. Mus. 47: 386, 1914. 
Big Bend National Park, ren Gulch, Basin, and desert near Nugent Mountain, 
end of September 1971, 3 44, 3 2 2; Green Gulch, early May 1972, 2 $4. The 
insect is different dhadles of duis: lncanrein 


Orthogramma prona Moeschler. Fig. 10. 
Verhandl. d.k.k. zool bot. Ges. Wien 30: 443, 1880. 
Welder Wildlife Refuge near Sinton, 14 November 1968, 1 ¢. The dark areas 
are dusky brown, the lighter areas yellowish. 


Gonodonta sinaldus Guénée. Fig. 11. 
Spec. Gen. Lepid. 6: 372, 1852. 

Santa Ana National Wildlife Refuge and Brownsville, late October to early 
November, twelve specimens. The dark areas of the forewings are deep chocolate 
brown, the light areas purplish powdery gray. The hindwings are orange-yellow 
with a gray border. } 

I have also a battered specimen of Gonodonta sicheas Cramer, taken at Garner 
State Park, 17 September 1961. It might have been a gooc specimen when it came, 


106 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


YH 


Figs. 8-15. 8, Iscadia daemonalis Dyar 6, Artesia Wells, 28 Sept. 1971; 48 mm. 
9, Sigela basipunctaria Wlk. 6, New Braunfels, 12 March 1972; 11.5 mm. 10, 
Orthogramma prona Moesch. 6, Welder Wildl. Ref., 14 Nov. 1968; 42 mm. 11, 
Gonodonta sinaldus Gn. 6, Santa Ana Nat. Wildl. Ref., 13 Oct. 1971; 35 mm. 
12, Rhescipha servia Cram. 6, Welder Wildl. Ref., 16 Nov. 1966; 38 mm. 13, 
Glympis concors Hbn. 6, Brownsville, 5 Nov. 1969; 23 mm. 14, Radara anartoides 
Wilk. 2, Welder Wildl. Ref., 20 March 1969; 27 mm. 15, Pentobesa valta Schaus 
6, Santa Ana Nat. Wildl. Ref., 7 April 1972; 36 mm. 


VoLUME 27, NUMBER 2 107 


but I overlooked it when I folded the sheet on which I had been collecting, and 
discovered it only the next morning. 


Rhescipha servia Cramer. Fig. 12. 
Pap. Exot. 4: 66; pl. 321, fig. e, 1782. 

This insect with odd looking palps and oddly shaped wings, iicluded in Mc- 
Dunnough’s check list as R. obtusa Walker, is relatively rarer in collections than in 
nature. Of 34 specimens in my collection, 23 were taken in the Welder Wildlife 
Refuge and 8 in the Santa Ana National Refuge; two were taken in February, one 
in March, two in April, one in May, all the rest in October and November. The 
record of the other three is as follows: San Antonio, Ebony Hill Laboratory, 9 Sept. 
1971, 1 2; Guadalupe River near New Braunfels, 12 March 1972, 1 64; Artesia 
Wells, LaSalle Co., Chaparral Wildlife Management Area, 11 June 1972, 1 @. The 
color is brown, more or less dusky, an extremely variable species. 


Radara anartoides Walker. Fig. 14. 
List Lep. Ins. Coll. Brit. Mus. 33: 843, 1865. 

Welder Wildlife Refuge, near Sinton, 13 and 14 Nov. 1968, 7 ¢ 8: 20 March 
1969, 1 ¢. The forewings are iiltenent shades of reddish bro, the hindwings 
senegal 

_ Glympis concors Hubner. Fig. 13. 
Zutrag. Samml. exot. Schmett. Zweites Hundert, p. 22, figs. 315-316, 1823. 

Of 18 specimens, 15 were taken in the two southernmost counties of Texas, 
Cameron Co. and Hidalgo Co., all in October and November; the record of the 
other three is as follows: Houston, 27 Nov. 1964, 1 9; Fort Davis, 5 Oct. 1969, 
1 @; San Antonio, Ebony Hill Laboratory, 9 Sept. 1971, 1 ¢. The wings are 
brownish gray, patterned with black. 


NOTODONTIDAE 


Pentobesa valta Schaus. Fig. 15. 
Trans. Entomol. Soc. London 49, p. 269, 1901. 
Santa Ana National Wildlife Refuge, 7 April 1972, 3 ¢¢. The forewings are 
pale brownish gray, the hindwings whitish. 


GEOMETRIDAE 


Scordylia atalanta Guénée. Fig. 19. 
Spec. Gen. 10: 383, 1857. 
Santa Ana National Wildlife Refuge, 9 April 1966, 1 ¢; 18 November 1966, 2 2 @. 
The wings are egg-yolk yellow and velvety black. 


Aeschropteryx olivata Warren. Fig. 20. 
Nov. Zool. 11: 128, 1904. 

Brownsville, 17 specimens taken in April, May, October and November. The 
insect resembles Prochaerodes transversata, but is smaller, more olivaceous, and the 
wings are shaped differently. 

PYRALIDAE 


Evergestis consimilis Warren. Fig. 21. 
Ann. Mag. Nat. Hist., Ser. 6, 9: 433, 1892. 
Davis Mountains, Mount Locke, 27 Aug. 1970, 2 é 6; Sierra Diablo Wildlife 
Management Area, 30 and 31 Aug. 1970, 3 646, 3 22. The wings are straw- 
yellow, blotched with purplish brown. 


Polygrammodes sanguinalis Druce. Fig. 22. 
Biol. Cent. Amer. 2: 218, 1895. 
Brownsville, 27 Oct. 1970, 1 ¢. The wings are pale yellow and brownish red. 


108 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 16-23. 16, Meropleon titan Todd ¢, Canadian, 27 Sept. 1968; 40 mm. 
17, Matigramma psegmapteryx Dyar 6, Garner State Pk., 24 March 1965; 31 mm. 
18, Herminodes stigmaphiles Dyar 9, Big Bend Nat. Pk., Chihuahua Desert near 
Nugent Mt., 21 Sept. 1971; 38 mm. 19, Scordylia atalanta Gn. 9, Santa Ana Nat. 
Wildl. Ref., 18 Nov. 1966; 29 mm. 20, Aeschropteryx olivata Warren ¢, Browns- 
ville, 8 May 1967; 40 mm. 21, Evergestis consimilis Warren ¢, Sierra Diablo Wildl. 
Met. Area, 30 Aug. 1970; 25.5 mm. 22, Polygrammodes sanguinalis Druce 6, | 
Brownsville, 27 Oct. 1970; 29 mm. 23, Givira redtenbacheri Hammerschmidt 6, | 
Big Bend Nat. Pk., Green Gulch, 28 March 1971; 31 mm. | 


VoLUME 27, NUMBER 2 109 


COSSIDAE 


Givira redtenbacheri (Hammerschmidt). Fig. 23. 
Naturwissenschaftliche Abhandlungen Haidinger 2: 151, 1847. 

Common in Big Bend National Park from late March until early May. It flies 
sympatrically with Heterocoma albistriga B. and McD., which it resembles, although 
it is a much darker gray. G. redtenbacheri is more common in Green Gulch, H. 
albistriga in the Chihuahua Desert. 


This random selection of 23 species, chosen among those which have 
already been described, does not exhaust my reserve of interesting moths 
flying in Texas. As time permits I intend to carry on with another 
installment. 

ACKNOWLEDGMENTS 


I am deeply indebted to many individuals who generously contributed 
much time and information, and without whose careful work of identifi- 
cation this paper would not have been possible. To all of the following 
I express my sincere thanks: Drs. D. C. Ferguson, J. G. Franclemont, 
D. F. Hardwick, R. W. Hodges, E. G. Munroe, F. H. Rindge and E. L. 
Todd. 

It is a pleasure to acknowledge with thanks the always gratifying 
cooperation of the administration and managers of the National Parks 
and Refuges and State owned Wildlife Management Areas. 


CHROMOSOME NUMBERS FOR PLEBEJUS (ICARICIA) ACMON, 
P. LUPINI, AND P. NEURONA (LYCAENIDAE) 


CaARLL GOODPASTURE 
Department of Entomology, University of California, Davis, California 95616 


The Plebejus acmon (Westwood & Hewitson) group is comprised of 
one easily recognized species, P. neurona (Skinner), and two closely 
related, less distinct entities, P. acmon and P. lupini (Boisduval). The 
present paper is a report of the results of cytological examinations of 
certain populations of these species. 3 

Chromosome complements of one population of P. neurona, two of 
P. acmon, and three of P. lupini from southern and central California 
have been examined. Field collected adults and laboratory reared pupae 
were used in this study. Specimens fixed in the field were treated and 


subsequently examined following the procedure of Emme! (1965). 


110 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


sae: 3 pe 


Figs. 1-4. Chromosomes of the Plebejus acmon group: 1, P. lupini lupini, N = 
24, metaphase; 2, P. neurona, N = 24, metaphase; 3, P. acmon acmon, N = 24, 
late diakinesis; 4, P. acmon acmon, bivalents from a cell in diakinesis stage, upper 
bivalent showing probably one nonterminalized chiasma, lower showing terminalized 
chiasma (note four strands at arrow). All cells primary spermatocytes from laboratory 
reared pupae, 0.5% lacto-acetoorcein, phase contrast. 


Laboratory reared material was treated as follows: tests were removed 
from pupae one day prior to time of adult eclosion; dissections were 
carried out in Hayes saline (Hayes, 1953), transferred to 1% sodium 
citrate for 20 minutes, fixed 5 minutes in 3:1 absolute ethanol: glacial 


VOLUME 27, NuMBER 2. ee 


acetic acid, and stained in 0.5% lacto-acetoorcein (Breland, 1961). 
Temporary squash preparations were made using thumb pressure. 

Slides were examined under oil using phase contrast illumination at 
a magnification of 960X. Photographs were taken on Kodak High 
Contrast Copy 35 mm film at a film plane magnification of 400x. 

Plebejus neurona. N = 24. Counts were made in 40 well spread nuclei 
in meiotic metaphase and two in dikinesis from four individuals from 
Chuchupate Ranger Station, Ventura Co., California. 

Plebejus acmon acmon. N = 24. Counts were made in 75 nuclei in 
meiotic metaphase, nine in diakinesis, and 10 in mitotic metaphase from 
eight individuals from Putah Creek, University of California at Davis, 
Yolo Co., and Monticello Dam, Napa Co., California. 

Plebejus lupini lupini. N = 24. Counts were made in 12 meiotic meta- 
phase nuclei from two pupae reared from stock collected at Echo Lake, 
8,000 feet, El Dorado Co., California. 

Plebejus lupini monticola. N = 24. Counts were made in 31 meiotic 
metaphase nuclei from six pupae reared from stock collected at La Posta 
Creek, 3,100 feet, San Diego Co., and Sierra Pelona Road, Mint Canyon, 
Los Angeles Co., California. 

Twenty-four chromosomes were usually counted in primary and 
secondary spermatocyte metaphase figures from all populations sampled. 
An occasional cell encountered with more or less than this number was 
probably due to observation of superimposed or displaced chromosomes 
within a single cell. Twenty-four bivalents were easily recognized in all 
cells found in diakinesis (Fig. 3). 

Exceptionally favorable diakinesis-diplotene figures were obtained 
from P. acmon and P. neurona. At least one chiasma was usually dis- 
tinguishable in all bivalents at diakinesis and individual chromatids were 
sometimes resolved in bivalents where chiasmata had terminalized (Fig. 
4, arrow). | 

A haploid chromosome number of N = 24 conforms to that previously 
reported for one other member of the subgenus Icarica; Plebejus icarioides 
(Boisduval) (Maeki & Remington, 1960). Although cytologically de- 
tectable chromosomal differentiation has not been found in this group, 
examinations of populations of the P.acmon-P. lupini species complex from 
areas outside of California where intergradation and possible hybridiza- 
tion occurs may be worthwhile. 


LITERATURE CITED 


BRELAND, O. P. 1961. Studies on the chromosomes of mosquitoes. Ann. Entomol. 
Soc. Amer. 54: 360-375. . 

Emmez, T. C. 1968. Methods for studying the chromosomes of Lepidoptera. J. 
Res. Lepid. 7: 23-28. 


112 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Hayes, R. O. 1953. Determination of a physiological saline solution for Aedes 
aegypti (L.). J. Econ. Entomol. 46: 624-627. 

Maext, K. & C. L. Remincron. 1960. Studies on the chromosomes of North Ameri- 
can Rhopalocera. 3. Lycaenidae, Danaidae, Satyrinae and Morphinae. J. Lepid. 
Soc. 14: 127-142. 


CALLOPHRYS (INCISALIA) POLIOS (LYCAENIDAE): 
DISTRIBUTION IN NORTH AMERICA AND 
DESCRIPTION OF A NEW SUBSPECIES! 


CiirForD D. FERRIS? 
College of Engineering, University of Wyoming, Laramie, Wyoming 82070 


AND 


MiIcHAEL S. FISHER 
1200 Summitt Road Rt. 2, Parker, Colorado 80134 


Callophrys (Incisalia) polios was described by Cook and Watson in 
1907. Subsequently Cook published two papers (1907, 1908) in which 
he identified the larval foodplant and discussed part of the life history 
of this insect. The egg is shown in a plate and described in the text of 
the 1908 paper, but the paper ends at this point with the statement “To 
be continued.” Apparently the proposed continuation was not published. 
In his 1907 paper, Cook mentioned having reared polios to the pupal 
stage, but did not describe the larva or pupa. A footnote in the same 
paper mentions that William P. Comstock had reared polios from ova to 
maturity. 

Cook and Watson described polios from a series of 84 specimens taken 
at Lakewood, New Jersey. Mention was also made of specimens from 
Calgary, Alberta, “Graham’s Park on Rio de los Pinos, Cal.,” and Colorado. 
These were not included in the type series. Later Cook (1908) corrected 
the Graham’s Park locality to Colorado and indicated that “Cal.” was a 
misprint in the earlier paper. 

In the 1908 paper, Cook noted the known distribution of polios as New 
Jersey, Massachusetts, New Hampshire, Maine, Nova Scotia, Indiana, 


' Published with the pptovall of the Director, Wyoming Agricultural Experiment Station, as 
Journ il Article no. JA 5 
* Research Associate, aan Museum of Entomology, Sarasota, Florida. 


VoLUME 27, NuMBER 2 lee 


Alberta, and Colorado. He also listed Puget Sound (Washington), based 
upon Wright (1905). 

Since Cook’s papers, little has been published on polios. The butterfly 
is listed in a variety of checklists. Brief descriptions are included in 
regional works (Clark & Clark, 1951; Brown et al., 1957; Shapiro, 1966). 
Klots (1951) provided additional distribution data for the eastern United 
States as did Clench for North America (1961). Holland (1931) said 
little about the insect. 

The larval foodplant was identified by Cook as Arctostaphylos uva-ursi 
(L.) Spreng. (Ericaceae). Clench (1961) also suggested Epigaea repens 
in Pennsylvania, as did Shapiro (1966). 

In 1968, we undertook a study of C. polios in the Rocky Mountain 
region, and later extended our work to include the distribution of this 
species in North America. Ferris has attempted several times to rear 
polios, but with little success. The foodplant in Wyoming has been 
identified as A. wva-ursi, but the females appear quite reluctant to oviposit 
in captivity. Oviposition has been observed at the bases of the flower 
buds and on the leaves of the hostplant. The ova have been adequately 
described by Cook (1908). In the west, the first instar larvae are pale 
green and covered with long fine hairs. To date, we have not succeeded 
in rearing beyond the first instar. 


Distribution 


During the course of this study, hundreds of specimens of polios from 
both museum and private collections were examined. Additional locality 
data were provided by a number of individuals. It soon became apparent 
that polios could be divided into two subspecies. The nominate sub- 
species occurs east of the 95th meridian. In the continental United States, 
the Great Plains appear to form a barrier between the eastern and western 
populations. To the north, a blend-zone occurs in Manitoba. We have 
reliable records of nominate polios from Nova Scotia, Quebec, Ontario, 
Maine, New Hampshire, Massachusetts, Connecticut, Rhode Island, New 
York, New Jersey, Pennsylvania, Virginia, West Virginia, Indiana, Illinois, 
Michigan, Wisconsin, and eastern Minnesota. As yet, we have no records 
from New Brunswick, Vermont, Delaware, Maryland, and Ohio, although 
one might expect the butterfly to occur in the mountainous areas of 
western Maryland, and it undoubtedly occurs in Vermont. The foodplant 
is rather scarce in Ohio. 

The western subspecies, which is described below as a new subspecies, 
occurs in Arizona, New Mexico, Utah, Colorado, Wyoming, western South 
Dakota, Idaho, Montana, Oregon, Washington, Manitoba, Saskatchewan, 


114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Distribution of Callophrys polios in North America. Cross-hatched area in 
Manitoba indicates a blend-zone between the eastern and western subspecies. 
Solid dots indicate states, provinces, and areas from which collection records exist. 


Alberta, British Columbia, Northwest Territory (District of Mackenzie), 
Yukon Territory, and Alaska. Until recently, there were no records from 
Montana, although the specimens listed by Elrod (1906) as irus (Godart) 
are probably polios, or possibly fotis schryveri Cross. Cook (1908) men- 
tioned confusion among various authorities over polios, irus, and henrici 
(Grote & Robinson). In 1971, S. Kohler found both polios and fotis 
schryveri in Missoula Co., Montana. The occurrence of polios in North 
Dakota appears doubtful. Occasional references to the occurrence of 
polios in California probably relate to the misprint in the Cook and 
Watson paper, or to confusion of polios with fotis (Strecker). 


VoLUME 27, NUMBER 2 115 


Fig. 1 illustrates the presently known distribution of C. polios in North 
America. Exact locality citations are too numerous to include. The insect 
does not occur in the total area delineated, but is restricted to those 
areas within the boundary where the foodplant grows. In the west, this 
is generally in foothills regions; in the east, at the edges otf woods and 
in boggy areas near the hostplant. There are certain areas where the 
foodplant occurs, but from which polios has not yet been collected. These 
are noted in the figure by the hatched areas. Within the overall area 
denoted for polios, various isolates occur and certain of these are noted 
by the inner closed lines in Fig. 1. Some of the areas between isolates 
appear to be unsuited to polios. The principal isolates encompass the 
following regions: northeast coastal area; Virginia-West Virginia; Illinois- 
Indiana; Black Hills, South Dakota; Utah; Arizona; and southern Michi- 
gan. It would appear, from Kohler’s recent discovery, that polios has 
probably been overlooked in many areas in the Rocky Mountains and 
may be much more widely distributed in this region than present records 
indicate. 


Callophrys (Incisalia) polios obscurus, Ferris and Fisher, 
new subspecies 


The new subspecies differs from C. p. polios in two consistent char- 
acters. The dorsal color in both sexes is uniformly a distinct gray brown 
rather than the warm rufous brown of the eastern subspecies. The scent 
pad on the forewing of the males of obscurus is black, or darker than 
the ground color. In polios, the scent pad is generally pale, or lighter 
than the ground color, although there are individual exceptions in both 
subspecies. The maculation of the undersides in both subspecies is highly 
variable, and we could find no consistent characters which allow separa- 
tion of obscurus from polios. Generally speaking, the hoary patches on 
the secondaries are brighter and more clearly defined in obscurus than 
in polios, and the other markings are more crisply defined. Fresh speci- 
mens should be used for comparison as fading occurs rather rapidly under 
exposure to strong light or elevated temperature. 


Variation: The occurrence of fulvous markings in the anal area of the DHW 
is infrequent in males from Colorado and Wyoming. This coloring is occasionally 
observed in females from Colorado and Wyoming, and fulvous markings are common 
in both sexes of polios from New Jersey. Some specimens from northern Idaho exhibit 
about as much ruddy color as eastern polios, but the males have the dark scent pads 
of obscurus. Specimens from Michigan have well pronounced fulvous areas over the 
outer two-thirds of the DHW in a majority of the specimens examined. Some have 
this character on the FW as well, giving these specimens a somewhat bright appear- 
ance. Fewer Michigan specimens are dark as in New Jerse) material. Ventrally, 
Michigan specimens tend to be lighter than either eastern or western populations, 


116 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 2. Callophrys polios obscurus Ferris and Fisher: a. holotype ¢ (right) and 


allotype 2 (left); b. same, undersides; ¢. specimens from Goldstream Valley, Alaska 
(male at left); d. same, undersides. 


VoLUME 27, NUMBER 2 LUT 


and the maculation appears to be somewhat frosted in aspect, producing a much less 
clearcut maculation than is found in specimens from other areas. This is also true, 
to some extent, of specimens from central Maine. Ontario specimens also have 
characters as described for the Michigan area, but somewhat less pronounced, which 
gives Ontario material a darker aspect, but not so dark as New Jersey polios, which 
is still lighter than western obscurus. 

Specimens from Maine are generally lighter in color than New Jersey material. 
Manitoba specimens, especially from Sandilands Provincial Forest, are rather dark. 
The fulvous areas are not pronounced, but the scent pads are pale as in eastern 
polios. Some populations from eastern Manitoba may be referred to polios and some 
from western Manitoba to obscurus, but in the broad sense, Manitoba represents a 
blend-zone region. Alaskan specimens tend to be a warmer brown than obscurus 
from Colorado and Wyoming. Pale scent pads occur more frequently than in other 
western populations. They are closer to Maine specimens than any other of the 
western material. A similar situation occurs with C. augustinus (Westwood) from 
Alaska. : 

Holotype male. The holotype was collected on Lookout Mountain, Jefferson Co.., 
Colorado. The male holotype bears a red label, machine printed in black ink: C. 
polios obscurus/ Ferris & Fisher/HOLOTYPE ¢; and a white data label, machine 
printed in black ink with handwritten date: Lookout Mt., Jefferson Co., COLO./ 
above Golden 6500-7200 ft./leg. Mike Fisher/14 May 68. 

Allotype female. The female allotype bears a green label, machine printed in 
black ink: C. polios obscurus/Ferris & Fisher/ALLOTYPE @, and a data label similar 
to that of the holotype with the date 12 May 68. 

The forewing length of both the holotype and allotype is 12.5 mm. There is 
remarkably little size variation in the type series. 

Type series. The new subspecies is being described from a series of 168 specimens, 
including the holotype and allotype, from the following areas: Colorado, 86 speci- 
mens from Boulder, Clear Creek, El Paso, Jefferson, and Park Counties, 14 
May-—2 June 1957, 1960-61, 1967-71, leg. J. D. Eff, M. S. Fisher, R. J. Jae, 
J. Scott; Wyoming, 82 specimens from Pole Mountain, Medicine Bow National Forest, 
Albany Co., 14 May—6 June 1969-71, leg. C. D. Ferris and M. S. Fisher. 

The holotype and allotype are deposited in the collection ot the Los Angeles 
County Museum of Natural History. Paratypes are being distributed to the following 
museum collections: Allyn Museum of Entomology, American Museum of Natural 
History, Canadian National Collection, Carnegie Museum, Los Angeles County 
Museum of Natural History, United States National Museum, Michigan State Uni- 
versity, University of Wyoming. 

Included under the taxon obscurus are specimens from Washington. There is 
some indication (D. Frechin, Seattle, Washington, pers. comm.) that F. H. Chermock 
had intended to name the polios population from the vicinity of Tenino, Thurston 
Co., but we can find no record in the literature. 


ACKNOWLEDGMENTS 


The authors wish to express their appreciation to the following in- 
dividuals for supplying collection data and the loan of specimens for 
study: Mr. R. Bailowitz, Portland, Ore.; Mr. W. A. Bergman, Chicago, 
Ill; Mr. H. K. Clench, Carnegie Museum, Pittsburgh, Penn.; Mr. P. J. 
Conway, Chicago, Ill.; Dr. J. P. Donahue, Los Angeles County Museum 
of Natural History, Los Angeles, Calif.; Mr. J. D. Eff, Boulder, Colo.; 
Mr. J. H. Fales, USDA, Beltsville, Md.; Mr. W. D. Field, United States 
National Museum, Washington, D.C.; Mr. R. W. Fischer, Michigan 


118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


State University, East Lansing, Mich.; Mr. D. Frechin, Seattle, Wash.; 
Mr. L. P. Grey, Enfield, Maine; Mr. H. Hensel, Edmundston, New 
Brunswick; Mr. J. R. Heitzman, Independence, Mo.; Mr. R. L. Huber, 
St. Paul, Minn.; Mr. R. J. Jae, Denver, Colo.; Mr. S. Kohler, Missoula, 
Mont.; Mr. I. Leeuw, Cary, IIl.; Mr. G. Lewis, Canadian National Col- 
lection, Ottawa; Mr. L. L. Martin, Prescott, Arizona; Mr. B. Mather, 
Clinton, Miss.; Mr. M. C. Nielsen, Lansing, Mich.; Dr. K. W. Philip, 
Fairbanks, Alaska; Dr. F. H. Rindge, American Museum of Natural 
History, New York, N.Y.; Mr. A. C. Sheppard, Lyman Entomological 
Museum, Ste. Anne de Bellevue, Quebec; Mr. J. T. Sorensen, Cedar Falis, 
Iowa; Mr. G. D. Willis, Wilmington, Del.; and Mr. B. Wright, Nova 
Scotia Museum, Halifax, Nova Scotia. 

Mr. Harry K. Clench most kindly read and commented upon a pre- 
liminary draft of the manuscript and provided considerable distribution 
data which are incorporated into Fig. 1. 


LITERATURE CITED 


Brown, F. M., J. D. Err & B. Rorcer. 1957. Colorado Butterflies. Denver Mu- 
seum, Denver. 

Ciark, A. H. & L. F. Crarx. 1951. The Butterflies of Virginia. Smithsonian 
Mise. Coll. 116(3). 

Crencu, H. K. 1961. In P. R. & A. H. Ehrlich, How to Know the Butterflies. 
Brown, Dubuque, Iowa. 

Coox, J. H. 1907. Studies in the genus Incisalia. Can. Entomol. 39: 405-409. 

1908. Studies in the genus Incisalia, V—Incisalia polios. Can. Entomol. 
39: 202-204. 

Exrop, M. J. 1906. The Butterflies of Montana. Bull. No. 10, Biological Series. 
Univ. Montana. 

oe W. J. 1931. The Butterfly Book (revised edition). Doubleday, New 
York. 

Kuots, A. B. 1951. A Field Guide to the Butterflies. Houghton Mifflin, Boston. 

SHaprro, A. M. 1966. Butterflies of the Delaware Valley. Amer. Entomol. Soc., 
special publication. 

Wricut, W. G. 1905. The Butterflies of the West Coast of the United States. 
Whitaker and Roy, San Francisco. (see plate XXVII, fig. 331). 


VoLUME 27, NUMBER 2 119 


THE LARVA OF LOXAGROTIS KYUNE (BARNES) (NOCTUIDAE) 


GrorcE L. GopFREY 


Illinois Natural History Survey and Illinois Agricultural Experiment Station, 
Urbana, Illinois 61801 


Loxagrotis kyune (Barnes) was described on the basis of a single 
female collected in the Huachuca Mountains, Arizona (Barnes, 1904). 
Barnes originally placed kyune in the genus Hadena. Except for changes 
in generic reassignments of the species, no additional published informa- 
tion is available. The following notes constitute the first information 
about the larval instars of kyune. 

Dr. John G. Franclemont captured a female moth of kyune on the night 
of 12 July 1967, at Onion Saddle, 7600 feet, Chiricahua Mountains, 
Cochise County, Arizona. Eggs obtained from the specimen the same 
night hatched on 19 July. The larvae subsequently passed through five 
instars before burrowing into the peat moss and sand filled rearing boxes 
during prepupal activity. 

The natural host is unknown. Newly hatched larvae were confined 
with cuttings of commercial lettuce (Lactuca sp.), Astragalus sp., Rubus 
sp., Quercus spp., Brickellia sp., Viguiera multiflora (Nutt.) Blake, and 
Pseudotsuga douglasi Carr. (Douglas fir). Except for the last plant, all 
cuttings showed some feeding signs by the first instars. The greatest 
feeding was on Viguiera multiflora. The second instars fed only on V. 
multiflora, and I eventually discontinued offering the other plants. 

The illustrations that accompany the following description of the last 
larval instar were drawn to scale by a grid system. All scale lines repre- 
sent 0.5 mm. The terminology and abbreviations follow those adopted 
earlier (Godfrey, 1972). 


General. Head 3.0-3.2 mm wide. Total length 38-43 mm. Abdominal prolegs 
present on third through sixth segments. Head and body smooth. Dorsal abdominal 
setae simple; setal insertions minute. Dorsal setae on seventh and eighth abdominal 
segments 0.75-0.88 times height of seventh abdominal spiracle. 

Head (Fig. 1). Epicranial suture 0.51 times height of frons. Distance from frontal 
seta (F-1) to frontoclypeal suture 0.33 times distance between F-1’s. Anterior setae 
(A 1-3) forming slightly less than 90° angle. Adfrontal puncture (AFa) anterior 
and second adfrontal seta (AF-2) posterior to apex of frons. First posterior setae 
(P-1’s) slightly anterior to apex of frons. Lateral setae (L’s) transversely anterior 
to AFa’s. Ocellar interspaces between Oc-1-—Oc-2 0.50 times that of Oc-2—Oc-3; 
Oc-2—Oc-3 4.0 times that of Oc-3-Oc-4. 

Mouthparts. Oral surface of labrum unspined. Hypopharyngeal complex (Fig. 2): 
spinneret with distal lip bearing long fringes, short, not surpassing first segment of 
labial palpus (Lps-1); stipular seta (S) about 0.25 length of Lps-1, twice length of 
seta (Lp-1) borne by Lps-1 and second segment of labial palpus (Lps-2), about 
0.33 length of seta (Lp-2) borne by Lps-2; Lp-1 positioned laterad of Lps-2; distal 


first abd 


120 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


4 


3. = 


2 Wt eee 


Figs. 1-4. Loxagrotis kyune, Chiricahua Mts., Arizona: 
capsule; 2. left aspect of hypopharyngeal complex; 3. 
ominal segment; 4. posterior aspect of tarsal c 


1. frontal aspect of head 
ventral setal arrangement of 
law. 


VoLUME 27, NUMBER 2 123 


SVE2 ees: 
DD 


6 


SV-1 


aS ee 


Figs. 5-6. Loxagrotis kyune, Chiricahua Mts., Arizona: 5. left dorsolateral setal 
arrangement of prothorax; 6. oral aspect of left mandible. 


region of hypopharynx densely covered with fine spines becoming stouter proximad; 
proximomedial region without spines; proximolateral region bearing single row of 
about 15 distinct spines. Mandible (Fig. 6): inner ridges distinct; no inner tooth; 
outer teeth 1-5 forming distinct angles; sixth outer tooth low, divided into smaller 
subteeth. 

Thoracic segments. Segment T-1: seta D-2 posterior of line formed by D-1 and 
XD-2 (Fig. 5); major axis of prothoracic spiracle passing through base of D-2 and 
distinctly posterior of subventral setae (SV 1-2); SD-1 transversely anterior of D-1 
and D-2. Lateral setae (L 1-2) and SV 1-2 enclosed by pinacula. Segments T 2-3: 
seta L-1 located above and slightly posterior of L-2. Tarsal claw (Fig. 4) with 
reduced basal angle; tarsal setae simple, tapering distad. 

Abdominal segments. Ab-1 (Fig. 3): only two subventral setae (SV-1, SV-3) 
present; SV-1 laterad of line formed by setae V and SV-3. Ab-2-6: three subventral 
setae present. Ab-8: only one seta in each subventral group. Ab-9: seta SD-1 
much finer than setae D 1-2. Anal and subanal setae subequal to lateral setae on anal 
proleg. Crochets: uniordinal, 18-22 per third abdominal proleg, 22-26 per fourth, 
24-28 per fifth, 25-32 per sixth. 

Coloration. Head pale brown with darker brown freckles. General body color 
brownish green. Middorsal and subdorsal lines, thin, whitish. Subdorsal area dark 
olive green with thin white line passing length of body. Lateral area diffusley white 
with indistinct brown center stripe, passing to tip of anal proleg. Spiracles black. 

Material examined. 10 specimens, Onion Saddle, 7600 ft., Chiricahua Mountains, 
Cochise County, Arizona, July-August 1967, from ova of female collected and de- 
termined by J. G. Franclemont, reared by G. L. Godfrey. Hypopharyngeal complex 
on slide G-0203. 


ACKNOWLEDGMENTS 


This research was supported by USDA AGR RMA Grant No. 12-14-100- 
8031 (33) awarded to Dr. John G. Franclemont of Cornell University. 
I thank Dr. Franclemont for his assistance. Additional funds and facilities 


122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


were supplied by the Illinois Natural History Survey, the Illinois Agri- 
cultural Experiment Station, and the Office of International Agriculture, 
College of Agriculture, University of Illinois, Urbana. 


LITERATURE CITED 


Barnes, W. 1904. New species of North American Lepidoptera. Can. Entomol. 
36: 165-173. 

Goprrey, G. L. 1972. A review and reclassification of larvae of the subfamily 
Hadeninae (Lepidoptera, Noctuidae) of America north of Mexico. U.S. Dept. 
Agr. Tech. Bull. 1450. 265 p. 


METHODS FOR EXTERNALLY SEXING MATURE LARVAE 
AND PUPAE OF LIMENITIS (NYMPHALIDAE) 


Puitie J. KEAN AND AvusTIN P. PLATT 


Department of Biological Sciences, University of Maryland Baltimore County, 
Catonsville, Maryland 21228 


Little published information exists regarding accurate methods for 
sexing the larvae and pupae of butterflies. It is well-known, however, 
that larger larvae and pupae within a brood generally develop into 
females, whereas, the smaller ones usually turn out to be males. In 
addition, the majority of males often will eclose at the beginning of a 
brood, whereas, the latter portion of the brood will consist almost ex- 
clusively of females. Nevertheless, numerous exceptions occur, and such 
methods cannot be considered to be very accurate. 

Among the larger moths (Saturnidae and Sphingidae), morphological 
differences such as the relative breadth of the pupal antennae and subtle 
differences in the ventral genital plates of pupae have been used for 
predicting the sex of the imago (Villiard, 1969), and methods for sexing 
mature larvae of the tobacco hornworm (Manduca sexta Johanssen) are 
known (W. Bowers, personal communication). Recently, other methods 
have been reported for sexing both the larvae and pupae of the codling 
moth, Laspeyresia pomonella L. (MacLellan, 1972). 

During laboratory hybridization studies on the Nearctic Limenitis, 
which were initiated in 1966 and are still in progress (Platt, 1969; Platt, 
Frearson & Graves, 1970; Platt & Greenfield, 1971), it became apparent 
that in inter-specific hybrid crosses, either excessive or complete hetero- 
gametic (female) inviability often occurs (Haldane, 1922; Remington, 


VoLUME 27, NuMBER 2 123 


1958). Since egg hatching within hybrid broods sometimes exceeds 50%, 
it was at once apparent that some female hybrid larvae were initially 
viable, but then died at certain stages during development. The need 
for positively determining the sexes of dying hybrid larvae and malformed 
pupae prompted this study. 

The purposes of this paper are to report means for positively sexing 
the mature larvae and pupae of North American Limenitis, and to point 
out how these and similar methods can add a new dimension to studies 
of lepidopteran genetics. 


MATERIALS AND METHODS 


Larvae of L. archippus Cramer and L. arthemis Drury were used princi- 
pally in these studies. The former were collected as over-wintering third 
instar larvae found in hibernacula during January and February, 1972, 
and represent stocks collected from Colchester (Chittenden Co.) and 
Starksboro (Addison Co.), Vermont, as well as a Maryland stock collected 
from the University of Maryland Baltimore County (Catonsville) campus. 
The latter species is represented by two broods of over-wintering third 
instar larvae, which represent F, crosses obtained from wild-caught L. 
arthemis—astyanax females collected from the intergrading population at 
Shutesbury (Franklin Co.), Massachusetts (see Platt & Brower, 1968), 
and by a third inbred L. a. astyanax Fabricius stock obtained several 
years ago from the Baltimore, Maryland vicinity. 

All hibernating larvae were kept in total darkness in a refrigerator 
within their hibernacula under moist conditions for a minimum of several 
weeks prior to study. Each hibernaculum then was individually isolated 
in a labeled plastic-covered styrofoam cup containing leaves of foodplant 
(Salix babylonica L. for L. archippus larvae, and Prunus serotina Ehrh. 
for L. arthemis—astyanax larvae). The cups were kept in covered trans- 
parent shoeboxes containing moist paper towels. The larvae were reared 
at room temperature in a closed photoperiod chamber under 20 hrs. of 
fluorescent illumination per 24 hr. day. Mature larvae were permitted 
to pupate on sticks placed across the tops of the cups. 

As each larva emerged from its hibernaculum, it was assigned an 
individual number. The next day each was sexed independently, using 
external markings (Fig. 1), which will be described in detail in the results 
section of this paper. A Wild 3M stereoscope with a Pentax camera back 
was used for examining and photographing the larvae and pupae. The 
day following each subsequent larval moult (to the fourth and fifth in- 
stars), each larva was again sexed independently, without reference to 
the previous sexing information. In each instance, the presence or 


124 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


— 
Imm 


Fig. 1. Morphological sex differences on the ventral surface of the eighth ab- 
dominal segment of female (A) and male (B) Limenitis larvae. The larvae are 
oriented with their posterior regions toward the top of the photographs. gp = genital 
pore; cw = chitin windows of female larva; b = pair of single bristles of male larva. 


absence of the same external markings was used to determine the larval 
sex. Upon metamorphosis, both pupae and adults likewise were sexed 
independently, using different morphological characters. Fig. 2 shows 
the morphological traits used for sexing Limenitis pupae. All larvae and 
pupae which died during development were deleted from the data, 
because the sex of the imago could not be positively determined. 


RESULTS 


External Sex Characters of Larvae. Third through fifth instar female 
larvae always possess two small transparent cuticular patches, which we 
have termed “chitin windows.” They are located on the mid-ventral 
surface of the eighth abdominal segment (Fig. 1A). These transparent 
spots appear to be dark against the grey-white mid-line, because the 
green gut of the larva shows through from beneath them. These spots, 
or chitin windows, lie anterior to a similar bifurcate dark crescentic patch 
(Figs. 1A, B) located just behind the anterior edge, in the middle of 
the ventral region of the ninth abdominal segment. The latter spot is 


| 
! 
| 
’ 


VoLUME 27, NUMBER 2 125 


14 
Imm 


Fig. 2. Morphological sex differences on the ventral genital plates of the eighth 
and ninth abdominal segments of female (A) and male (B) Limenitis pupae. The 
differences described in the text are indicated by the arrows. 


present in larvae of both sexes and apparently marks the primordial 
genital region. The functions and adult structures (if any) arising from 
the chitin windows presently are not known, although the female genital 
structures form in this region of the pupa. Male larvae (Fig. 1B) lack 
the chitin windows, and so appear to possess an unbroken white mid-line 
on the ventrum of the eighth abdominal segment. This difference can 
be determined readily with the naked eye in mature larvae. However, 
it is best to check all larvae with a stereo-microscope, using low to medium 
magnification. When this is done, two very small dark bumps, each 
containing a single short curved bristle, can be seen on the white ventral 
mid-line of the male larva’s eighth abdominal segment. The position of 
these bumps is comparable to that of the chitin windows of female larvae. 

External Sex Characters of Pupae. Limenitis pupae, likewise, can be 
sexed externally by examination of the morphology of the genital plates 
located ventrally, and immediately anterior to the base of the lateral 
processes of the cremaster. These plates cover the mid-ventral region 
of the entire eighth, and the anterior margin of the ninth, abdominal 
segments. Microscopic examination is necessary to observe the male and 


126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Taste 1. Results of externally sexing Limenitis larvae and pupae by independent 
examination at each stage. 


A. Correct sexing observations (percent accuracy given in parentheses): 
Developmental Stages 


Numbers Larval instars 
Species Vee Oe) Teal ed: ER nS Pupae Adults 
L. archippus ——32 26 58 Al BD 58 58 58 
(71%) (95%) (100%) (100%) (100%) 
L. arthemis— 
astyanax' 10 LA Dil 1S} 16 21 21 21 
(62%) (76%) (100%) (100%) (100%) 
Totals 42, Sil 79 54 71 79 79 79 


(68%) (90%) (100%) (100%) (100%) 


B. Inaccurate sexing observations, given by sex of adult individual (based on a 
total of 237 observations: 
Larval Instars 


ord 4th 
sub sub Grand 
Species Cites 22 totals oS Qe totals Totals 
L. archippus 6 Tal Ia 2 1 3 20 
L. arthemis— 
astyanax* 4 4 8 3 2 5 13 
Column Totals 10 15 25 5 3 8 33 


1Includes 3 L. arthemis arthemis Drury, 14 L. arthemis prosperpina Edwards, and 4 L. 
arthemis astyanax Fabricius. 


female differences clearly. Female pupae possess a longitudinally slotted 
mid-ventral pad (Fig. 2A). The slot always extends forward across the 
eighth segment (where the larval chitin windows were located) to the 
posterior margin of the seventh abdominal segment, appearing to arise 
from the segmental fold, at right angles to it. This slotted pad seems to 
be homologous with the distal ends of the female genital ducts in the 
imago. Male pupae (Fig. 2B) lack this vertically slotted pad, but possess, 
instead, two small paired swellings, located posterior to a prominent 
horizontal fold, which forms the base of a medial flattened isoceles 
triangular plate. This plate is circumscribed by two anterolateral oval 
plates. This entire small crescentic structure is restricted to the ninth 
abdominal segment, leaving the mid-ventral region of the eighth ab- 
dominal segment intact. 

Accuracy of the Sexing Methods. The validity of using the external 
morphological features described above to sex the larvae has been veri- 
fied in two ways. First, a small number of larvae of both sexes were 


VoLUME 27, NUMBER 2 127 


dissected, and the developing gonads identified. (In mature larvae of 
Lepidoptera, the gonads lie beneath the body wall in the dorsal region 
of the fifth abdominal segment. In Limenitis, this region is the central 
area of the grey-white larval saddlepatch. The developing ovaries of 
female larvae are paired flattened whitish strands of delicate wavy tissue, 
often imbedded in fat. The paired reddish-brown oval male testes are 
somewhat easier to locate in larvae; the latter fuse into a single medial 
rounded testis in the adult butterfly.) Secondly, 79 larvae of L. archippus 
and L. arthemis—astyanax were independently sexed during the third, 
fourth, and fifth larval instars, and later both as pupae and as adults 
(Table 1A). External sexing of mature larvae and of pupae was ac- 
complished with an accuracy of 100%. Third and fourth instar larvae 
were sexed with 68% and 90% accuracy, respectively. Only a total of 33 
(14%) inaccurate sexing observations were made in 237 independent 
larval observations. The majority of erroneous sexings occurred when 
examining third instar larvae (Table 1B). In most cases, this happened 
because the chitin windows (which are present) were not clearly visible 
in the small female larvae. 


DISCUSSION AND CONCLUSIONS 


These methods for externally sexing the fifth instar larvae and the 
pupae of North American Limenitis are 100% reliable. Fourth instar 
larvae have been sexed with an over-all accuracy of 90%. The methods 
may be somewhat more reliable for L. archippus larvae than for the 
larvae of L. arthemis at this instar (Table 1A). Because of their smaller 
size (approximately 12 mm long by 3 mm wide, or less), third instar 
larvae are much more difficult to sex. Our over-all accuracy is only 687% 
for this stage, with the sexing observations on L. archippus larvae again 
being more reliable than those carried out on larvae of L. arthemis— 
astyanax. We conclude that our methods cannot be considered com- 
pletely reliable for the third instar. Because of the small size and deli- 
cateness of younger larvae, attempts were not made to sex them in the 
first and second instars. 

These larval and pupal sexing methods have been tried on a few larvae 
of the Western species of Nearctic Limenitis (L. lorquini Boisduval and 
L. weidemeyerii Edwards). The immature stages of both of these species 
also can be sexed accurately using these characters. In fact, it is probable 
that species in closely related genera, such as Adelpha, Apatura, Neptis, 
Parthenos, etc., and, perhaps even more remotely related nymphalines, 
can be accurately sexed in their immature stages using either these, or 
similar morphological criteria. 


128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Since sex is a genetically determined trait, the ability to sex larvae 
and pupae of certain Lepidoptera externally, provides a means of estab- 
lishing genetic ratios prior to the adult stage. As has been implied earlier 
in this paper, this becomes a most useful tool in cases involving inter- 
specific hybridization. 

For example, accurate sex ratios now can be established for crosses in 
which heterogametic inviability occurs during larval growth. Among the 
species of Nearctic Limenitis, inter-specific hybridization occurs only 
rarely in nature, but can be accomplished readily in the lab by hand- 
pairing the insects (Remington, 1958; Platt, 1969; Platt & Greenfield, 
1971). Although the number of larvae hatching from eggs of such crosses 
sometimes exceeds 50%, a number of the developing larvae often die 
prior to, or during pupation. Are these, perhaps, mostly females, since 
only males are found to emerge from normally formed pupae (especially 
in hybrid crosses involving the broadly sympatric species, L. archippus)? 

Preliminary evidence suggests that the answer to this question is yes: 
Recently, 21 hybrid larvae, representing three broods of L. arthemis— 
astyanax X L. archippus crosses have produced malformed pupae at the 
end of the fourth instar. Each pupa has been sexed as female externally, 
and no definite males have been found among them, although it is 
possible that some of the most malformed ones may represent inter-sex 
pupae. Several other hybrid larvae have grown slowly to immense size 
in the fifth instar, but then have failed to metamorphose. Each of these, 
likewise, has been externally sexed as a female. 

Therefore, our methods of externally sexing the immature stages of 
Limenitis provide a means of confirming the developmental stages during 
which female inviability occurs. Within hybrid female larvae, species 
incompatibilities in juvenile hormone and ecdysone are implicated, but 
must be substantiated by further work. Using the external sexing methods 
to identify female hybrid larvae prior to death, it may be possible to 
treat them with the appropriate synthetic insect hormones, in order to 
enable them to pass through a normal metamorphosis. Such techniques 
may prove exceedingly useful in furthering our present knowledge of 
evolution among related species of Lepidoptera. 


SUMMARY 


Methods for accurately sexing the fourth and fifth instar larvae and 
the pupae of the North American species of Limenitis by means of ex- 
ternal morphological differences are reported. Female larvae possess a 
pair of prominent dark spots (termed chitin windows) on the ventral 
white mid-line, located centrally on the eighth abdominal segment; male 


VoLUME 27, NUMBER 2 129 


larvae lack these spots. Female pupae possess an antero-posterior slotted 
pad on the ventrum of the eighth abdominal segment, whereas male 
pupae possess a smaller genital pore, consisting of two bulbous swellings 
lying just posterior to a prominently horizontal fold, the entire structure 
being restricted to the ninth abdominal segment. The eighth abdominal 
segment of male pupae possesses no mid-ventral pad with an antero- 
posterior slit, as it does in female pupae. These methods proved 100% 
effective for sexing mature larvae and pupae. Fourth instar larvae were 
sexed with an accuracy of 90%. However, small third instar larvae could 
be sexed only with an accuracy of 68%. The genetic implications of being 
able to externally sex immature stages of Limenitis, with respect to inter- 
specific hybridization studies, are discussed briefly. 


ACKNOWLEDGMENTS 


We are greatly indebted to F. M. Mills, III of the University of Mary- 
land Baltimore County for his initial studies on this topic, and to Pamela 
C. Platt for assistance in collecting the larval stocks. We wish to thank 
Dr. William Bowers, Department of Entomology, of Cornell University 
at Geneva, New York, for providing the information regarding sexing 
procedures of certain sphingid larvae. Dr. L. P. Brower of Amherst 
College and Dr. T. D. Sargent of the University of Massachusetts at 
Amherst provided helpful comments regarding certain portions of this 
manuscript. 

LITERATURE CITED 


Haxpang, J. B. H. 1922. Sex-ratio and unisexual fertility in hybrid animals. J. 
Genet. 12: 101-109. 

MacLetian, C. R. 1972. Sex ratio in three stages of field collected codling moth. 
Can. Entomol. 104: 1661-1664. 

Puatr, A. P. 1969. A simple technique for hand-pairing Limenitis butterflies. 
(Nymphalidae). J. Lepid. Soc. 23: 109-112. 

. & L. P. Brower. 1968. Mimetic versus disruptive coloration in inter- 

grading populations of Limenitis arthemis and astyanax butterflies. Evolution 

22: 699-718. 

. S. D. Frearson & P. N. Graves. 1970. Statistical comparisons of valval 

structure within and between populations of North American Limenitis (Nym- 

phalidae). Can. Entomol. 102: 513-533. 

& J. C. GreenFiELp, Jr. 1971. Inter-specific hybridization between 
Limenitis arthemis astyanax and L. archippus (Nymphalidae). J. Lepid. Soc. 
25: 278-284. 

Remincton, C. L. 1958. Genetics of populations of Lepidoptera. Proc. 10th 
Intern. Congr. Entomol. 2: 787-805. 
Vintiarp, P. 1969. Moths and How to Rear Them. Funk & Wagnells, New York. 

p. 242. 


130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ILLUSTRATIONS OF HELICONIUS (NYMPHALIDAE): 
SOME RARE AND IMPORTANT SPECIMENS 


Joun R. G. TURNER’ 


Photographed by 
RICHARD HUNTER 
Department of Biology, University of York, England 


Heliconius butterflies are now being intensively used as research 
animals for varied projects in genetics, ecology, behavior, and physiology. 
The taxonomy of the genus is extremely difficult, largely because of 
mutual mimicry within the genus and extreme geographical variation 
and polymorphism within some species. Its study has been somewhat 
hampered in the past by the absence of iilustrations, which in addition 
makes it difficult for both the amateur butterfly collector and the pro- 
fessional researcher to identify the material with which he is working. 
Presented here are some pictures of important specimens, most of them 
types, and most of them never previously illustrated. These are in order: 
type specimens designated by Felix Bryk, from the Swedish Amazon 
Expedition, in the Naturhistoriska Riksmuseet, Stockholm; forms of the 
rare and little known species Heliconius demeter; and type specimens of 
another comparatively rare and difficult species, Heliconius elevatus, 
from the Naturhistorisches Museum, Wien. 

Other works illustrating large numbers of Heliconius specimens are the 
two standard monographs (although by no means every species is illus- 
trated) by Stichel & Riffarth (1905) and by Stichel (1906), and the 
profuse color illustrations by Seitz (1913); all these works are well out 
of date in their classification. All the Trinidadian species have been 
illustrated by Beebe, Crane & Fleming (1960), reprinted by Emsley 
(1963), and all of these appear in color in the recent book by Barcant 
(1971). Sixteen of the commoner species are figured by Turner 
(1968 )”, who has also figured some of the subspecies of H. elevatus and 
H. melpomene (Turner, 1967). A great variety of Brasilian species have 
recently been illustrated by Brown & Mielke (1972) and Brown (1972). 
The geographical variation of Heliconius melpomene and H. erato has 
been illustrated by Brown & Mielke (1972) and in color by Turner 
(1971). 

The butterflies are fully described in the figure legends, but a few 

1 Present address: Department of Ecology and Evolution, Division of Biological Sciences, State 


University of New York at Stony Brook, Stony Brook, New "York 11790. 
* The alleged egeria in that paper is in fact burneyi. 


VoLUME 27, NuMBER 2 130 


Figs. 1-7. Type specimens designated from the Swedish Amazon Expedition, 
and preserved in the Naturhistoriska Riksumuseet, Stockholm: 1. H. elevatus tara- 
cuanus Bryk, holotype male (Brasil, Est. do Amazonas, Rio Uaupés, Taracua); 
2. H. erato reductimacula Bryk, holotype male (data same as 1); 3. H. aoede 
aoede from postalbimacula Bryk, holotype male (Brasil, Est. do Amazonas, Manaos); 
4. H. erato estrella form aurivillii Bryk, holotype male (Brasil, “Rio Autaz”); 5, 6, 7. 
H. egeria homogena Bryk, male syntypes, the specimen at figure 6 is hereby 
designated lectoholotype (Brasil, Est. do Amazonas; 5-6, Rio Uaupés, Taracua; 
7, Rio Negro, Sao Gabriel). 


132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


points are worth noting. Among the type specimens from the Swedish 
Museum in the first photograph are three species (Figs. 1, 2, 5, 6, 7) 
from the upper Rio Negro; all these forms show very similar patterns, 
with the yellow marks in the outer area of the forewing more or less 
fused into a large patch, a feature which tends to be rare to the south 
on the Amazonas, where the yellow patch tends to be broken up into 
spots or reduced to a broad yellow bar; there is a strong suggestion here 
of mutual mimicry among these three species, a mimicry which is known 
to occur in other parts of their ranges. 

Illustrated here for the first time are type specimens of H. elevatus 
carrying the designations taracuanus, pseudocupidineus, griseoviridis, 
noeldneri, and aquilina. These represent a slight extension of the work 
on the taxonomy of this genus in a previous paper (Turner, 1967). The 
status of the forms appears to be as follows: 

taracuanus (Fig. 16): This is probably a good subspecies found in the 
north Amazon Basin, replaced to the south by nominotypic elevatus, and 
closely resembling the form perchlora which is found on the southern 
border of the Amazon Basin in Bolivia. There is however some variability 
of patterns in the central Amazon Basin, and the division into these three 
races may be to some extent arbitrary. 

pseudocupidineus (Figs. 17, 18): This is a good subspecies confined 
apparently to the valley of the Rio Huallaga in Peru; it differs from 
nominotypic elevatus in having the yellow band on the forewing ex- 
tremely narrow (in nominotypic elevatus it is of a width intermediate 
between pseudocupidineus and taracuanus ). 

griseoviridis and noeldneri (Figs. 19, 22): These are probably to be 
regarded as mere aberrations, although there is no hint about their 
causation. 

aquilina (Figs. 20, 21): Populations of H. elevatus in the Guianas, 
lower Amazon and Mato Grosso have the yellow marks on the forewing 
broken up into dots. The more northerly of these populations have yellow 
apical spots on the forewing in addition (see illustrations by Turner, 
1967); the populations in the south of the range lack these apical spots, 
as can be seen from the illustration. If one ignores the presence or 
absence of spots, as being unworthy of producing a subspecific separa- 
tion, then aquilina is a junior synonym of bari (Oberthiir), the form 
found in the Guianas; if one wishes to split these populations into two 
subspecies then aquilina is a synonym of schmassmanni Joicey & Talbot, 
which also comes from the Mato Grosso. These names were published 
during the same year, and date priority has not yet been established. 

The full taxonomic references are given elsewhere (Turner, 1967) and 


VoLUME 27, NUMBER 2 133 


Figs. 8-16. Specimens of H. demeter from the British Museum ( Natural History ), 
London (uppersides on left, undersides on right): 8, 9. H. d. bouqueti Noéldner, 
males (Guyane francaise, St. Jean de Maroni); 10. H. d. beebei Turner, holotype 
male (central Guyana—see type description); 11, 12. H. d. beebei, paratype 
females (central Guyana—see type description); 13, 14. H. d. demeter Staudinger, 
males (Peru, Iquitos); 15, 16. H. d. demeter, females (Colombia, upper Rio 
Putumayo, Florida). 


134 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 17-22. Type specimens of H. elevatus designated by Neustetter in the 
Naturhistorisches Museum, Wien, Austria: 17. H. e. pseudocupidineus Neustetter, 
lectoholotype male (Peru, Yurimaguas); 18. H. e. pseudocupidineus, lectoparatype 
female (Peru, Yurimaguas); 19. H. e. elevatus form griseoviridis Neustetter, holo- 
type male (Peru, Yurimaguas); 20. H. e. aquilina Neustetter, lectoholotype male 
(Brasil, Rio Machados, Mato Grosso); 21. H. e. aquilina, lectoparatype female 
(Brasil, Rio Machados, Mato Grosso); 22. H. e. elevatus form noeldneri Neustetter, 
holotype male (Peru, Yurimaguas). 


need not be repeated here; a distribution map of the main forms can be 
found in Turner (1971). 

The second photograph illustrates Heliconius demeter, which is a very 
rare species, except possibly in the upper Amazon Basin. Three of the 
four subspecies are illustrated here: H. d. bouqueti from the lower 
Amazon and Guianas, H. d. beebei from the region of the Guiana Shield 
(these three are the type specimens) and the nominotypic subspecies 
H. d. demeter from the upper Amazon. The fourth subspecies H. d. 
eratosignis is not illustrated, as at the time of taking the photograph only 


VoLUME 27, NUMBER 2 135 


the two type specimens were known. The forewing resembles that of 
bouqueti and beebei, but the rays on the hindwing are separated and 
not fused into a red patch, even in the male. In the other subspecies 
(except beebei) there is more or less extensive fusion of the rays in the 
male, although not in the female. The female of bouqueti is not illus- 
trated as no specimen could be found to photograph; it has previously 
been illustrated by Neustetter (1931), under the name H. eratoformis. 
In collections it is about twenty times as rare as the male; the only speci- 
mens known to me apart from the type (originally in the Larsen collec- 
tion and not traced), are one in the collection of Drs. E. H. Jonkers of 
the Netherlands Government Economic Mission to Suriname, in Para- 
maribo, and one in the British Museum (Natural History ).. An account 
of the complicated synonymy is given elsewhere (Turner, 1966). 

We are very grateful to the following for the loan of the specimens 
photographed: The Naturhistoriska Riksmuseet, Stockholm; the British 
Museum (Natural History), London; and the Naturhistorisches Museum, 
Wien. Where necessary, designations of lectotypes are made in the 
figure legends. The colors are black, red and yellow. 


SUMMARY 


Twenty-two specimens of Heliconius, most of them belonging to forms 
previously not figured, from museums in Austria, England, and Sweden, 
are illustrated with photographs. Sixteen of these are type specimens of 
forms in the species elevatus, demeter, erato, aoede, and egeria. One 
lectoholotype is designated, and a short account is given of the importance 
of these specimens. The photographs show a mimicry ring found on 
the upper Rio Negro, and for the first time, the major subspecies of 
Heliconius demeter. 


LITERATURE CITED 


Barcant, M. 1971. Butterflies of Trinidad and Tobago. London, Collins. 

BEEBE, W., J. CRANE & H. Fieminc. 1960. A comparison of eggs, larvae and 
pupae in fourteen species of Heliconiine butterflies from Trinidad, W.I. Zoo- 
logica, New York 45: 111-154. 

Brown, K.S. 1972. The heliconians of Brazil (Lepidoptera: Nymphalidae). Part 
III. Ecology and biology of Heliconius nattereri, a key primitive species near 
extinction, and comments on the evolutionary development of Heliconius and 
Eueides. Zoologica, New York 57: 41-69. 

_ & O. H. H. Mrerxe. 1972. The heliconians of Brazil (Lepidoptera: 
Nymphalidae). Part II. Introduction and general comments, with a supple- 
mentary revision of the tribe. Zoologica, New York 57: 110. 

Bryk, F. 1954. Lepidoptera aus dem Amazonasgebiete und aus Peru gesammelt 
von Dr. Douglas Melin und Dr. Abraham Roman. Ark. Zool. 5: 1-265. | 

Emstry, M. 1963. A morphological study of imagine Meliconiinae (Lep.: Nym- 


136 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


phalidae) with a consideration of the evolutionary relationships within the group. 
Zoologica, New York 48: 85-130. 

NeustTeTTer, H. 1931. Neue Heliconius. Intern. Entomol. Zeit. Guben 25: 165- 
WAS 

Seirz, A. 1913. MHeliconiinae, in A. Seitz, ed., The Macrolepidoptera of the world. 
Stuttgart, Kernen. 

SticHEL, H. 1906. Lepidoptera, fam. Nymphalidae, subfam. Heliconiinae. Genera 
Insectorum 37: 1-74. Bruxelles, Wytsman. 

. & H. Rirrarru. 1905. Heliconiidae. Das Tierreich 22: 1-290. Berlin, 
Friedlander. 

TurRNER, J. R. G. 1966. A rare mimetic Heliconius (Lepidoptera: Nymphalidae). 
Proc. Roy. Entomol. Soc. London. (B) 35: 128-132. 

1967. A little-recognised species of Heliconius butterfly (Nymphalidae). 

J. Res. Lepid. 5: 97-112. 

1968. Some new Heliconius pupae: their taxonomic and evolutionary 

significance in relation to mimicry (Lepidoptera, Nymphalidae). J. Zool. 

(London) 155: 311-325. 

1971. Studies of Millerian mimicry and its evolution in burnet moths and 

heliconid butterflies. in, E. R. Creed, ed., Ecological Genetics and Evolution. 

Oxford, Blackwell. p. 224-260. 


POPULATIONS OF PAPILIO ANDRAEMON BONHOTEI SHARPE 
AND PAPILIO ARISTODEMUS PONCEANUS SCHAUS 
(PAPILIONIDAE) IN BISCAYNE NATIONAL 
MONUMENT, FLORIDA 


Larry N. BROWN 
Department of Biology, University of South Florida, Tampa, Florida 33620 


A survey of the Lepidoptera found on the islands of Biscayne National 
Monument, Florida, in April and May 1972, revealed sizeable breeding 
populations of two rare papilionid butterflies. These are the Bahaman 
Swallowtail (Papilio andraemon bonhotei Sharpe) and Schaus’ Swallow- 
tail (Papilio aristodemus ponceanus Schaus). The former species has 
been recorded only a few times in Florida (Holland, 1902; Clarke, 1940; 
Kimball, 1965) and until now has been considered only a stray or acci- 
dental visitor to United States shores following hurricanes. The scarceness 
of the latter species in southern Florida has also been previously docu- 
mented by many authorities including Bates (1934), Grimshawe (1940), 
Henderson (1945a, 1945b, 1946), Klots (1951) and Rutkowski (1971). 
The majority of records for Schaus’ swallowtail are from Key Largo and 
Lower Matecumbe Key (Kimball, 1965) which suggests that these islands 
house the remaining remnant population of this species found in the 


VoLUME 27, NUMBER 2 137 


United States. However, this proves not to be the case since the species 
is now found to be prevalent on several remote keys to the north and 
east of Key Largo. 

From 20-24 April 1972, and again from 27-30 May 1972, I surveyed 
the major islands of Biscayne National Monument. Swallowtails were 
quite abundant, and in a single day I often encountered as many as 100 
P. aristodemus ponceanus and a like number of P. andraemon bonhotei 
on these islands. In April large numbers of adult Schaus’ and Bahaman 
swallowtails had recently emerged, as indicated by their perfect condi- 
tion and unworn scales. Adults were flying in substantial numbers in 
both the dense hammocks and along the narrow trails which cut through 
the jungle-like vegetation of the islands. A limited number of reference 
specimens of each species was collected and preserved. 

In late May, nearly all specimens of both species were badly tattered 
or worn, and many flew rather weakly compared to the previous month. 
This suggested that the peak emergence had occurred in April or early 
May, and that most of the adults were nearing senesence. Approximately 
half of the swallowtails observed and collected in April were P. andraemon 
bonhotei, but in May they comprised only about one-third of the total 
number encountered. 

On two occasions female P. andraemon bonhotei were observed in late 
April laying eggs on the leaves on Key lime (Citrus aurantifolia) and sour 
orange (Citrus aurantium) trees. Citrus trees are scattered sparingly 
throughout the jungle-like forests which cover these keys. Also, one 
P. aristodemus ponceanus was seen laying eggs on the leaves of torchwood 
or sea amyris (Amyris elemifera), which is an abundant understory shrub 
or small tree on most of the islands surveyed. This agrees with the 
Grimshawe (1940) and Rutkowski (1971) reports that torchwood is the 
foodplant for P. aristodemus ponceanus. Also, two late instar larvae of 
this species were located on torchwood in late May 1972. 

The relative abundance of these papilionids on the Biscayne Bay Keys 
suggests that they have a high potential for survival on a long term basis. 
The two greatest threats to their populations would seem to be: 1) habitat 
destruction by man or hurricane, and 2) overcollecting by dealers and 
lepidopterists. Since their prime habitat is now fully under the control 
and protection of the National Park Service, there seems little chance of 
total habitat destruction due to man. There is always the possibility of 
overcollecting, but since a federal collecting permit is required to legally 
take anything from a national park of monument, it should be relatively 
easy for the Park Service to police Lepidoptera poaching once they are 
made aware of the presence of these rare species within their jurisdiction. 


138 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Two Papilio andraemon bonhotei Sharpe taken in Biscayne National 
Monument, Dade Co., Florida, April 1972. Underside (above); upperside (below). 


VoLUME 27, NUMBER 2 139 


One obvious factor which operates to protect these populations and 
until now prevented their detection, is the relative isolation of these 
islands which are located between seven and eight miles from the 
adjacent Florida mainland. The fact that they can be reached only by 
boat reduces the probability that collecting pressure will be great. This 
pressure will probably increase however, once the National Park Service 
institutes its announced plan of establishing a regular ferry to some of 
the islands for the benefit of tourists. 

Evidence of the ability of these papilionids to colonize adjacent keys 
or the mainland was noted during the survey. On several occasions while 
traveling by boat between islands in the chain, both Schaus’ and Bahaman 
swallowtails were seen flying across the open water. On two occasions 
we collected the butterflies with hand nets after pursuing them by boat 
for one-fourth mile or more across Biscayne Bay. On one occasion a 
P. aristodemus ponceanus eluded our pursuit and flew safely from the 
south end of one key and entered the jungle on the northeast side of an 
adjacent key in a route that covered over one-half mile. It therefore 
seems certain that movement of these rare species by island-hopping 
along the chain of Florida Keys occurs routinely. However, since both 
the Schaus’ and Bahaman swallowtails have been recorded somewhat 
sparingly from Key Largo, Lower Matecumbe Key, and the mainland of 
the southern Florida, these areas could presently be considered rather 
marginal when compared to the populations located in Biscayne Bay 
National Monument. The foodplants for these species are readily avail- 
able throughout the middle and lower Florida Keys as well as on the 
mainland; therefore, the explanation for their scarcity there is probably 
related to overzealous collecting activities by man in past years. 

Other species of Lepidoptera which were collected on keys of the 
Biscayne National Monument during the April-May 1972 survey included 
the following: Battus polydamas (Linnaeus ), Papilio cresphontes Cramer, 
Phoebis agarithe Boisduval, Appias drusilla Cramer, Phyciodes tharos 
(Drury), Cynthia virginiensis (Drury), Junonia evarete zonalis (Felder 
& Felder), Metamorpha stelenes (Linnaeus), Eunica tatila (Herrich- 
Schaffer), Hemiargus ammon (Lucas), Heliconius charitonius Linnaeus, 
Dryas julia (Fabricius), Agraulis vanillae Linnaeus, Phocides pigmalion 
(Cramer), Polygonus leo (Gmelin), and Urbanus proteus (Linnaeus ). 


ACKNOWLEDGMENTS 
I would like to thank my graduate students, Richard McGuire and 
Ronald Pscion, as well as members of the University of Terrestrial Ecology 
. ° . . i b = . event = > ae ~ 
class, for assistance in surveying the Lepidoptera of the Biscayne Bay 


140 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


islands. I also wish to express gratitude to the personnel of the National 
Park Service and the Dade County Parks Department for cooperation 
and assistance during this survey. 


LITERATURE CITED 


Bates, M. 1934. Notes on some tropical Florida butterflies. Entomol. News 45: 
166-169. 

Cuarke, J. F. 1940. United States records of tropical American Lepidoptera. 
Proc. Entomol. Soc. Wash. 42: 155-156. 

Grimsuawe, F. M. 1940. Place of sorrow: the world’s rarest butterfly and 
Matecumbe Key. Nature Mag. 33: 565-567, 611. 

HeNvEeRson, W. F. 1945a. Papilio aristodemus ponceana. Entomol. News 56: 
29-32. 

1945b. Additional notes on Papilio ponceana. Entomol. News 56: 187- 


188. 

. 1946. Papilio aristodemus ponceana Schaus notes. Entomol. News 57: 
100-101. 

Ho.Luanp, W. J. 1902. Two new species of Bahaman lepidoptera. Ann. Carnegie 
Mus. 1: 486-489. 

Krmpaxz, C. P. 1965. Lepidoptera of Florida. Division of Plant Industry, Gaines- 
ville, Fla. 363 p. 

Kiots, A. B. 1951. A Field Guide to the Butterflies. Houghton Mifflin, Boston. 
349 p. 

Rutkowski, F. 1971. Observations on Papilio aristodemus ponceanus (Papilioni- 
dae). J. Lepid. Soc. 25: 126-136. 


ON GLENOIDES TEXANARIA (GEOMETRIDAE) WITH DESIGNATION 
OF THE LECTOTYPE 


As Mr. André Blanchard is describing new species of Glenoides McDunnough from 
Texas, it became necessary to definitely establish the identity of texanaria (Hulst), the 
heretofore sole included species of this genus. 

Hulst described Tephrosia texanaria from a series of eight males and six females, 
according to the original description. The Hulst collection contained one male; this 
specimen is hereby designated as the lectotype. It is in the collection of the American 
Museum of Natural History, and its genitalia are mounted on slide FHR No. 16712. 

The species is widely distributed across the southeastern United States. Specimens 
are before me from eastern Texas, Louisiana, Mississippi, Arkansas, Missouri, Ken- 
tucky, and South Carolina. It may occur in Florida, but more study is necessary 
to determine the status of specimens from that state. 

There appears to be some variation in the size of the adults of texanaria, being 
apparently correlated with the time of flight. Moths captured in late fall and early 
spring months are larger and appear somewhat darker than those taken from May 
through September. 


Freperick H. Rinpce, Department of Entomology, The American Museum of 
Natural History, New York, New York 10024. 


VoLUuME 27, NUMBER 2 14] 


A NEW SPECIES OF THE 
GENUS GLENOIDES McDUNNOUGH (GEOMETRIDAE) 


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


Hulst (1888) described Tephrosia texanaria, which he later (1896) 
moved to his new genus Glena (orthotype cognataria Hbn.). McDun- 
nough (1920) recognized that texanaria is not congeneric with cognataria 
and created for it the monotypic genus Glenoides. 


Glenoides lenticuligera A. Blanchard, new species 


Head: Smoothly scaled; vertex pale brownish; front broadly black, with upper and 
lower, narrow, white borderlines; palpi short, porrect, projecting only slightly beyond 
front; male antennae bipectinate with short, simple apical section; each pectination 
bearing a double row of cilia and a longer seta at apex; female antennae roughly 
scaled above and laterally, cilate below. 

Thorax: Pale brownish, spotted with dark brown above; legs slender, smooth, 
male without hair pencil, blackish externally, all segments of tarsi distally ringed with 
white. 

Abdomen: Pale brownish, except segments three and four which are dark brown 
above. 

Pattern of maculation (Figs. 1-4): Ground color of forewing pale brownish, 
sprinkled with brown scales, more heavily along costa, and in subterminal and 
terminal spaces: four dark brown blotches about equally spaced on costa, three 
innermost ones mark the origins of a.m. line, median shade, and p.m. line, fourth 
blotch adnate to and basad of s.t. line; a.m. line brown starting on costa one fourth 
distance from base to apex, regularly and outwardly convex, reaching inner margin 
one-fifth distance from base to tornus; p.m. line brown, starting on costa two-third 
distance from base to apex, roughly parallel to outer margin; s.t. line of ground color, 
irregular, slightly retracted between veins, more so in cell Cu, inwardly bordered 
by dark brown blotches, of which the most conspicuous straddles vein M2, terminal 
black dots in all cells between, and generally including, R: to Cuz. Hindwing patterned 
in direct continuation of forewing, with well marked black discal dot. Pattern of 
maculation beneath similar but fainter on paler, less freckled background. 

Length of forewing: Male 7.2 to 8.0 millimeters (average 7.7 mm); female 8.0 
to 8.8 millimeters (average 8.4 mm). 

Male genitalia (Fig. 5): Valves unarmed; juxta replaced by two spinose processes, 
one on each side; aedeagus with a row of four to six teeth on outer margin; vesica 
armed with numerous cornuti. 

Female genitalia (Fig. 6): Ovipositor lobes appear membranous; a_ heavily 
sclerotized, hourglass shaped combination of sterigma and ductus bursae presents a 
short ventral fold; small star-shaped signum on lateroventral right side of nearly 
spherical bursa. 

Holotype: Male, Santa Ana National Wildlife Refuge, Hidalgo Co., Texas, 15 
February 1971, deposited in the National Museum of Natural History (No. 72326— 
genitalia slide A.B. 2633). 

Paratypes: Santa Rosa (Longoria unit of Las Palomas Wildlife Management 
Area) Cameron Co., Texas, 21 Nov. 1966 (one ¢). Brownsville (Voshell unit of 
Las Palomas Wildlife Management Area) Cameron Co., Texas, 10 & 12 Noy. 1965 


142 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-4. G. lenticuligera: 1, male holotype, Santa Ana Refuge, 15 Feb. 1971; 
2, male paratype, Santa Ana Refuge, 15 Feb. 1971; 3, female paratype, Brownsville, 
12 Nov. 1968; 4, female paratype, Santa Ana Refuge, 14 Novy. 1971. 


Figs. 5, 6. G. lenticuligera, genitalia: 5, male holotype (A.B. 2633); 6, female 
paratype, Santa Ana Refuge, 13 Nov. 1971 (A.B. 3033). (Linear segments represent 
one millimeter. ) 


VoLUME 27, NUMBER 2 143 


(two 22); 5 to 9 Nov. 1969 (two ¢ 6, one 2); 26 Oct. 1970 (one ¢); 18 Nov. 
1971 (two 6 6). Santa Ana National Wildlife Refuge, Hidalgo Co., Texas, 23 Oct. 
1970 (one ¢); 15 & 16 Feb. 1971 (three 6 6, one 2), 13 to 16 Nov. 1971 (four 
66, ten 22); 7 April 1972 (five 6 3, eight 92). Paratypes will be deposited 
in the National Museum of Natural History, in the American Museum of Natural 
History and in the British Museum (Natural History). 

The new species is quite close to Glenoides texanaria, the only other 
taxon in the genus; the pattern of maculation is nearly the same, but the 
transverse lines of texanaria are much better defined and its background 
is nearly clean of scattered brown scales and dark blotches; G. texanaria 
is appreciably larger; the unmistakable differences between the two 
species are however in the genitalia: the vesica of the male texanaria 
is unarmed and the postvaginal plate of its female presents an elongated 
sclerotization which does not exist in G. lenticuligera. 


ACKNOWLEDGMENTS 


It is a pleasure to acknowledge with thanks the generous help which 
I received from Drs. D. C. Ferguson of the Entomology Research Di- 
vision, U.S.D.A. and F. H. Rindge of the American Museum of Natural 
History in preparing this paper. I am also grateful to the Bureau of Sport 
Fisheries and Wildlife (Albuquerque, New Mexico) and to the Park 
and Wildlife Department (Austin, Texas) for the authorization that they 
gave me to set traps in the territories under their jurisdiction. 


LITERATURE CITED 


Huust, G. D. 1888. New species of Geometridae (No. 4). Entomol. Amer. 3: 
213-217 (p. 216). 

1896. A classification of the Geometrina of North America, with descrip- 
tions of new genera and species. Trans. Amer. Entomol. Soc. 23: 245-386 (p. 
SS) e 

McDunnovucu, J. 1920. Studies in North American Cleorini (Geometridae). Can. 
Dept. Agric. Bull. 18 (p. 38). 


MIDGES (DIPTERA: CERATOPOGONIDAE) SUCKING BLOOD 
OF CATERPILLARS 


With reference to Willis W. Wirth’s note under this heading (1972, J. Lepid. Soe. 
26: 65), I have a record of a larva of Acherontia atropos L. (Sphingidae) bearing 
seven of these small midges. The larva was found in Kampala in July 1950, and 
was carried by car for over a mile clinging to a twig without disturbing the midges. 
My notes state that the larva appeared to suffer no inconvenience and that there was 
no exudation of fluid from the punctures, which were invisible under a hand lens, 


when the midges were removed. 


144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


NOTES ON VIRGINIA BUTTERFLIES, WITH TWO 
NEW STATE RECORDS? 


CHARLES V. COVELL, JR. 
Department of Biology, University of Louisville, Louisville, Kentucky 40208 


AND 


GERALD B. STRALEY 
Botany Department, Porter Hall, Ohio University, Athens, Ohio 45701 


In 1951 Austin and Leila Clark published The Butterflies of Virginia. 
This work crowned 20 years of avid collecting and observation in the 
Old Dominion by the Clarks and their colleagues, and brought together 
all the records and knowledge about the Virginia butterfly fauna up to 
that time. According to the nomenclature used by the Clarks, a total 
of 154 species and subspecies (144 species) was recorded as having 
been collected in Virginia. Keys, diagnostic features, range, variation, 
occurrence, season, and interesting discussions were given for the butter- 
flies and a complete bibliography of literature on Virginia butterflies 
rounded out the work. 

Since the appearance of Clark & Clark (1951) very little has been 
published to add to our knowledge of the Virginia butterfly fauna. Covell 
(1962) added Satyrium kingi (Klots & Clench) to the State list and 
Straley (1969) recorded Thymelicus lineola (Ochsenheimer) for the 
first time from the State. Nomenclature and arrangement of species 
were brought into line with dos Passos (1964), the present standard for 
North American Rhopalocera classification, by Covell (1967). The latter 
list also included the first record of Megathymus yuccae (Boisduval & 
Le Conte) for Virginia. 

In this paper we include two more first records for the State—Problema 
bulenta (Boisduval & Le Conte) and Satyrium caryaevorus (McDun- 
nough). In accordance with the dos Passos arrangement, plus recent 
changes in the nomenclature of Lethe, 158 species and subspecies (149 
species) of butterflies are now known to have been taken in Virginia. 

Nomenclature and arrangement of species employed in this paper 
follow dos Passos (1964, 1970) and other recent works; but references 
are made to coverage of the species in Clark & Clark (1951). Botanical 
nomenclature follows Massey (1961). 

Our purpose in writing this paper is to add our records and observa- 


1 University of Louisville Contribution in Biology, No. 155 (New Series). 


VoLUME 27, NUMBER 2 145 


tions on certain butterfly species to those of the Clarks, thus bringing 
The Butterflies of Virginia more nearly up to date. In addition to our 
own records we include information from others who have collected in the 
State in the past 20 years and records from the Field Season Summaries 
of the News of the Lepidopterists’ Society. 

We concentrate our attention on the rarer species in the State, giving 
range extensions and new county records for species previously known 
from only a few counties. We hope that this additional information 
will be of value in helping other collectors find new collecting areas, 
especially for the rarer species in Virginia. 

Covell’s records from the Commonwealth span the years 1952 to 1971, 
with most of his collecting having been done between 1958 and 1964. 
He visited 81 counties, taking at least 5 butterfly species in each. Straley 
has actively collected from 1963 to the present. We have both resided 
in several different areas of the State and have concentrated on those 
areas; both our efforts, combined with those of other collectors, have 
encompassed most of the State. We have both done our most intensive 
collecting in the Norfolk-Virginia Beach area, and in Giles and Mont- 
gomery counties. 

Specimens have for the most part been retained in our private col- 
lections. However, a large amount of representative material has been 
placed in the collection of the Entomology Department of Virginia 
Polytechnic Institute and State University, Blacksburg, Virginia. Further 
information on species treated is available from the authors. 

In recent years there have been several changes in political boundaries 
in Tidewater Virginia, which might cause some confusion when one 
compares records in this paper with those of Clark & Clark (1951). The 
counties of Norfolk, Princess Anne, Warwick, and Elizabeth City no 
longer exist. The City of Norfolk is the same; but the remainder of old 
Norfolk County is now the City of Chesapeake, Princess Anne County 
is now the City of Virginia Beach, Warwick County is the City of Newport 
News, and Elizabeth City County is now the City of Hampton. 


Treatment of Selected Species 
MEGATHYMIDAE 


Megathymus yuccae (Boisduval & Le Conte). We know of no other records of 
this skipper since the sighting of larval tents by the late F. H. Chermock near Sand- 
bridge, Virginia Beach [reported in Covell (1967), p. 21], although several colonies 
of Yucca filamentosa L. have been checked in Virginia Beach on several occasions 
by Straley and Anderson. 

HESPERIIDAE 


Panoquina panoquin (Scudder). Panoquin has been observed quite often straying 


: z ° e he aan te 8 » rdoec : . 
for some distances from its salt marsh habitat into open | ind the edges of woods. 


146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


We have taken it commonly on flowers of white clover (Trifolium repens L.), sweet 
pepper bush (Clethra alnifolia L.), and the blue mist-flower (Eupatorium coelestinum 
L.). Covell added a new county record: King and Queen Co., near West Point, 15 
Sept. 1962. 

Panoquina ocola (Edwards). We have three new county records in the State for 
ocola: Giles (Straley), Montgomery and Middlesex (Covell), which support the 
Clarks’ observation that ocola occurs in the Coastal Plain and southwestern mountains, 
but not in the heart of the Piedmont (p. 186). 

Amblyscirtes aesculapius (Fabricius). We have found this skipper, known by the 
Clarks as textor Hiibner, common in many areas of southeastern Virginia. The Clarks 
(p. 180) mention that it is attracted to the flowers of Prunella and Elephantopus. 
In addition to these we have taken it frequently on white clover, dogbane (Apocynum 
cannabinum L.), and sweet pepper bush, mostly at the edges of woods and along 
forest paths. We agree with the Clarks that there is a spring brood, as we have 
found it in late April and early May. 

Amblyscirtes carolina (Skinner). Carolina, another flower visitor, tends to hug 
the ground in flight. We have found it especially attracted to flowers in the rose 
family, ie. blackberry (Rubus), cinquefoil (Potentilla), and strawberry (Fragaria), 
as well as sweet pepper bush, swamp milkweed (Asclepias incarnata L.) and others. 
The form reversa Jones needs a great deal more critical field and laboratory study. 
A number of collectors feel that reversa may be a distinct species rather than a form 
of carolina. 

Atrytonopsis hianna (Scudder). Records of this species from Montgomery (VPI 
Collection) and Giles (Straley) extend the range somewhat westward in Virginia. 
Even more interesting, however, are Anderson’s records from Nansemond, Chesapeake, 
and Virginia Beach (6 to 24 May) which are the first known captures of hianna in 
southeastern Virginia. It probably occurs throughout the State. 

Euphyes palatka (Edwards). Although several collectors have searched diligently 
for palatka in southern Virginia Beach where Otto Buchholz took it, none had been 
found since his records of 1944. Finally, in 1971, Anderson succeeded in collecting 
a male on 7 June on pickerelweed (Pontederia cordata L.) in Currituck County, North 
Carolina, just across the State line from Virginia Beach, and another male on 11 June 
at Blackwater in Virginia Beach. Palatka may venture this far north only sporadically. 

Euphyes dion dion (Edwards). A record of dion in New Kent County (Straley) 
extends the range of this subspecies slightly northward in Virginia. It was previously 
known only from three counties bordering North Carolina. 

Euphyes dion alabamae (Lindsey). Although it is now generally considered a 
subspecies of dion, alabamae was thought to be a distinct species by the Clarks (p. 
174). Dr. Lee D. Miller (pers. comm.) has collected and examined individuals from 
the Dahl Swamp, Accomack County, that the Clarks considered alabamae. He is 
of the opinion that this population may be an undescribed subspecies—not alabamae. 
This is another species-subspecies complex which needs more careful study. 

Euphyes dukesi (Lindsey). Dukesi can be collected in quantity on pickerelweed 
flowers along the North Landing Road and at Blackwater in Virginia Beach. The 
Clarks (p. 176) did not realize that it has a fall brood. We have fall records from 
2 August to 9 September in the localities mentioned by the Clarks. 

Poanes yehl (Skinner). We must disagree with the statement by the Clarks (p. 
172) that yehl is “exceedingly shy and difficult to catch.” We have found it very 
easy to net on numerous occasions, especially when it is visiting its favorite flowers— 
pickerelweed, swamp milkweed, and sweet pepper bush. Yehl is one of the most 
common skippers flying in southeastern Virginia during late July and August. 

Problema bulenta (Boisduval & Le Conte). Figs. 1-4. The discovery of this 
little-known skipper in Virginia is without doubt the most exciting addition to the 
butterfly fauna of the State in recent years. It was first collected in the State by 
John Bauer and Bruce Dixon, 21 August 1967, about two miles south of the town of 


VoLUME 27, NUMBER 2 147 


i. 6 | 7 


Figs. 1-7. Virginia butterfly specimens: 1, 2. Problema bulenta, male, New 
Kent Co., 17 Aug. 1970, upper and lower aspects; 3, 4. P. bulenta, female, same 
data as male, upper and lower aspects; 5, 6. Thymelicus lineola, male, Eggleston, 
Giles Co., 21 June 1968, upper and lower aspects; 7. Satyrium caryaevorus, female, 
Eggleston, Giles Co., 4 July 1963, lower aspect. 


Lanexa, along the Chickahominy River in the extreme southeastern corner of New 
Kent County. One female was placed in the Camegie Museum in Pittsburgh and 
presumably forgotten until July 1970, when Nicolay and Straley were visiting the 
museum and noticed the specimen. 

The Virginia locality was visited several times that August by Anderson, Nicolay, 
Covell, and Straley; and after many hours of collecting, an additional 14 males and 
A females were taken, all on the flowers of swamp milkweed. Hoping to find an 
earlier brood, Anderson searched the area unsuccessfully on 19 July 1971; but he 
and Nicolay did take two more males in August, 1971. 


148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The habitat for bulenta is the broad expanse of marshy flood plain of the Chicka- 
hominy River. The dominant plants along the river are pickerelweed, swamp milk- 
weed, buttonbush (Cephalanthus occidentalis L.), rose mallows (Hibiscus moscheutos 
L.), cattails (Typha sp.), and various grasses and sedges, all partially submerged 
at high tide. Islands and peninsulas of higher ground support shrubby dogwoods 
(Cornus sp.), bald cypress [Taxodium distichum (L.) Richard], and black gum 
(Nyssa sylvatica Marsh). Due to the extensiveness of the marshy area along the 
Chickahominy and the James River into which it flows, it seems likely that bulenta 
may occur widely in the area. Accessibility to much of the area is limited—a problem 
which may be solved by use of a boat for collecting. 

Wallengrenia otho otho (Smith). The Clarks (p. 167) recognized two subspecies: 
otho otho (with dull orange or orange-red ground color on the hindwings beneath), 
and otho egeremet (Scudder) (with dark purplish-brown ground color on the hind- 
wings beneath). According to the records cited by the Clarks, otho otho occurs 
in the Piedmont and Coastal Plain, and otho egeremet flies throughout the State, 
including areas where otho otho has been collected. Our own records from the 
Dismal Swamp, New Kent County, and Virginia Beach indicate that the two forms 
are sympatric in these areas. We feel that the differences in pigmentation may be 
due to diet or otherwise environmental in nature, and do not indicate subspeciation. 
Sympatry could indicate that the two are separate species. Perhaps careful study 
of the differences will clarify the situation. 

Hesperia metea Scudder. The Clarks list six county records for this species and 
state (p. 160) “undoubtedly much more generally distributed in the western part 
of the State than these records would indicate.” We support this statement with 
four additional county records: Stafford (Nicolay), Henry and Giles (Straley), and 
Rappahannock (Ferris). We have found it attracted to low-growing flowers especially 
those in the Rosaceae. It can also be found flying about grassy clearings among cedar 
groves on limestone outcrops, as Covell found in Montgomery Co. 

Hesperia leonardus Harris. Covell has taken leonardus in three additional counties 
—Powhatan, Prince William, and Stafford. The Powhatan County record shows its 
range extends more into central Virginia than indicated by the Clarks (p. 160). 

Thymelicus lineola (Ochsenheimer). Figs. 5-6. Straley (1969) reported the 
first known occurrence of the European Skipper in Virginia in the mountains in 
Giles County. Mitchell took two males in Poverty Hollow in neighboring Mont- 
gomery County on 22 June 1969. Straley found it fairly common but worn on 3 
July 1971, in a meadow near the original locality; and Showalter also took it in 
1971 at Blacksburg in Montgomery County. Lineola is probably well established in 
western Virginia and probably now even farther south and east. 

Staphylus mazans hayhurstii (Edwards). The Clarks (p. 153) mention that 
hayhurstii is confined to the Coastal Plain. We have additional records in the eastern 
part of the State for Stafford (Nicolay), Virginia Beach and Northumberland 
(Straley). A most surprising capture of this species was made by Straley in Henry 
County in the western edge of the Piedmont. A male was first taken in a weedy 
area in Martinsville on 30 June 1969; three males and a female were taken in the 
same locality on 30 July 1969; and a male was caught on 23 May 1970, near the 
town of Figsboro, also in Henry County. These records extend the range of hay- 
hurstii about two hundred miles westward in Virginia. 

Urbanus proteus (Linnaeus). Covell has three additional Coastal Plain county 
records: Middlesex, Nansemond, and Chesapeake, all in late September 1959. 


PIERIDAE 


Colias eurytheme Boisduval. Figs. 12-15. There is always a great deal of variation 
in eurytheme and its hybrid with philodice, both seasonally and among specimens 
caught at one time in any one locality. Straley has taken four aberrations of this 


VoLUME 27, NUMBER 2 149 


11 


Figs. 8-11. Satyrium kingi from Virginia: 8, 9. male, near Suffolk, Nansemond 
Co., 21 June 1970, upper and lower aspects; 10, 11. female, same data as male, 
upper and lower aspects. 


common butterfly while searching for more choice species. All were collected in 
Open grassy meadows just north of Buckeye Mountain about three miles west of 
the town of Eggleston, in Giles County. 

The most unusual was a melanic male taken on red clover (Trifolium pratense L.) 
on 6 September 1965 (Figs. 12, 13). The upper surface is a dark sooty brown, 
lightly dusted with yellow scales basally. The normally black borders are lighter 
brown than the rest of the wings. On the lower surface the hindwings are a uniform 
dull yellow and the forewings are dull brown with yellow tips and costal margins. 

The other three aberrations are all similar on the lower surface. All have black 
streaks from the submarginal row of spots inward on the underside of the forewings 
and a brown patch or broad brown streaks from the submarginal spots inward on 
the hindwings. The first specimen of this aberration was collected on 19 June 1962. 
The upper surface was normal except for a noticeable pink iridescence in the orange 
ground color. The second specimen (Figs. 14, 15) was an orange female taken 
on 22 September 1965. It has wide black borders extending into the spot at the 
end of the cell on the upper surface of the forewings. The submarginal yellow spots 
in the black border are greatly reduced and smudged. A white female was collected 
on 24 April 1966, with the upper surface normal. The under surface, however, has 
the same black-streaked forewings and brown-streaked hindwings. 


RIODINIDAE 


Calephelis virginiensis (Guérin—-Méneville). This small metalmark was recorded 
from four localities in only two counties—Princess Ann (now Virginia Beach) 


150 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 12-15. Colias eurytheme from Virginia: 12, 13. melanic male, Eggleston, 
Giles Co., 6 Sept. 1965, upper and lower aspects; 14, 15. aberration female, Eggles- 
ton, Giles Co., 22 Sept. 1965, upper and lower aspects. 


and Nansemond. We have found it in three other locations in Virginia Beach 
(Anderson & Straley), three in Chesapeake (Anderson & Covell), and one in the 
City of Norfolk (Anderson). The best locale we have found for virginiensis is a 
low, weedy roadside ditch along London Bridge Road at Oceana Naval Air Station 
in Virginia Beach, where eighteen specimens were taken in a short time one day. 
This species is probably more widely distributed than our records indicate, but is 
easily overlooked owing to its small size, restricted occurrence, and probably short 
flight period for each brood. Adults seem to prefer alighting on broad leaves, but 
they often visit the flowers of the blue mist-flower (Eupatorium coelestinum L.). 

Calephelis borealis (Grote & Robinson). The Poverty Hollow (Montgomery 
County) colony of borealis continues to flourish. It has been found there regularly 
in recent years by a number of collectors. Covell found another colony in Mont- 
gomery County along Slusser Chapel Road. Kenneth Frank discovered a colony at 
Nature Camp, near Vesuvius, in Rockbridge County, which is a new county record. 
This brings to five the number of counties in Virginia in which borealis is known 
to occur. 


LYCAENIDAE 


Harkenclenus titus mopsus (Hubner). The Clarks (p. 79) record mopsus from 
seven counties only in the Piedmont and Shenandoah Valley of the State. We have 
additional records from Giles (Straley), Rappahannock (Ferris), Stafford (Nicolay), 
Nansemond (Anderson), Middlesex and Gloucester (Covell). The latter three 
counties extend the range of mopsus into the Coastal Plain. It should be looked for 


VoLUME 27, NuMBER 2 ESI 


throughout the State, especially on the flowers of butterflyweed (Asclepias tuberosa 
L.) and common milkweed (Asclepias syriaca L.). 

Satyrium liparops strigosa (Harris). This subspecies is recorded from five counties 
in the State by the Clarks (p. 81). We have three additional records: Rockbridge 
(Covell), Alleghany (Clench), and Giles (Straley). Although it is probably not 
“rare as the Clarks say, it is certainly not easily encountered. 

Satyrium kingi (Klots & Clench). Figs. 8-11. Covell (1962) recorded the first 
capture of kingi in Virginia on 11 June 1958. Unfortunately the locality near the 
Norfolk Airport is in the heart of the urban expansion of the cities of Norfolk and 
Virginia Beach, and an elementary school now stands where kingi once flew. Several 
other localities have, however, been discovered by other collectors in the State. 
Miller collected one worn female five miles south of Suffolk on the west side of the 
Dismal Swamp. 

Anderson found it near the Suffolk Airport on 16 July 1967. This spot has been 
collected every year since then by several collectors and has yielded a large number 
of specimens. The locality is a second growth mixed deciduous woods. Males have 
been noted visiting the flowers of sourwood [Oxydendrum arboreum (L.) DC.] where 
they are easily taken, especially with a long-handled net. More commonly they sit 
on the tops of broad leaves and sun themselves. The females tend to fly lower among 
the underbrush and they might even be called sluggish for a hairstreak. We have 
rarely seen the females visit flowers. 

Another colony of kingi was found by Straley in July 1971, in Chesapeake on the 
east side of the Dismal Swamp ai the junction of routes 104 and 190, in the same 
type of cut-over deciduous woods. There are undoubtedly other colonies in the 
Coastal Plain of Virginia which should be sought beginning about 10 June. 

Satyrium calanus falacer (Godart). We have found falacer present, although not 
common, in the southeastern counties and cities of Norfolk, Chesapeake (Anderson), 
Nansemond and Virginia Beach (Straley), where it was previously unrecorded. 
Falacer is at times very common in the mountains of the southwestern part of the 
State, especially attracted to New Jersey tea (Ceanothus americanus L.), common 
milkweed, and dogbane. 

Satyrium caryaevorus (McDunnough). Fig. 7. The first known record of this 
northern hairstreak in Virginia is from Giles County in the mountains. One female 
was taken on New Jersey tea by Straley on 4 July 1963. The specimen was thought 
at that time to be a female falacer and was papered and not identified until 1970. 
The determination was confirmed by Nicolay. It has been compared with a number 
of caryaevorus from Connecticut and New York and matches them quite closely. 

The locality is a steep dry hillside just north of Buckeye Mountain about three 
miles west of the town of Eggleston. The foodplant, hickory (Carya sp.), is 
scattered. over the hillside and is common on Buckeye Mountain. The area has been 
collected briefly on two occasions since 1963, but no more caryaevorus have been 
taken. A more thorough search of the area and similar habitats in early July should 
turn up more specimens. 

Satyrium edwardsii (Saunders). The Clarks (p. 80) record edwardsii from only 
four counties. Covell has an additional four: Montgomery, Middlesex, King and 
Queen, and Gloucester. This is another species of hairstreak which has probably 
been overlooked in most of the State, and is sometimes locally common in season. 

Callophrys irus (Godart). We have additional records for irus in Stafford 
(Nicolay) and two locations in Montgomery (Mitchell). Jrus is probably more 
widespread than our records indicate. It should be sought where the foodplant, 
lupine (Lupinus perennis L.), grows. 

Callophrys henrici (Grote & Robinson). We have new county records for henrici 
from nine counties, four of which are in the Coastal Plain, from which there were 
previously no records. The counties are: Giles and Northumberland (Straley), 
Stafford (Nicolay), Rappahannock (Ferris), Goochland (Powell), Virginia Beach 


152 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 


(Anderson), Middlesex, King William, and Nansemond (Covell). It should be 
considered common on the Coastal Plain and generally distributed across the State 
as the dominant Elfin species. 

We have found henrici most common along the wooded trails in Seashore State 
Park, Virginia Beach, on the very first warm days of spring. It rests on sunny spots 
on the trails or on broad leaves where it is easily netted. In other localities it may 
be caught on redbud blossoms. 

Callophrys augustinus croesioides Scudder. Augustinus was also not recorded from 
the Coastal Plain of Virginia until Covell took it in Chesapeake on 23 April 1960. 
It was taken by Straley in Northumberland and in a second locality in Chesapeake 
in 1971. It should be expected throughout the State, often flying with henrici. 

Callophrys gryneus (Hubner). Covell took a striking aberrational female among 
the red cedars on a limestone outcrop near the tunnel at Pepper Station, Montgomery 
Co., 26 April 1962. The underside is generally brown with dark olive suffusion, and 
all the white lines and other markings are very obscure and dark, contrasting very 
little with the ground. 

Atlides halesus (Cramer). We agree with the Clarks (p. 77) that halesus is not 
very common in southeastern Virginia, but is regularly present and easily overlooked 
because of its tendency to stay high up in the trees. We found it most common on 
the flowers of devil’s club (Aralia spinosa L.) where a long handied net is a necessity 
for collecting any number of specimens. Occasionally it comes down to the flowers 
of sweet pepper bush and climbing hempweed [Mikania scandens (1L.) Willd.]. 

Euristrymon ontario (Edwards). Clench (1971) recorded ontario from Alleghany 
County, Virginia. His five specimens represent the largest number of this species 
taken at any one locality in the State. It has also been taken in Middlesex (Covell), 
bringing to five the total number of counties in which it has been found in Virginia. 
It is probably not an “infrequent casual” as the Clarks (p. 80) say, but is present 
in isolated colonies which have been mostly overlooked. 

Panthiades m-album (Boisduval & Le Conte). We have additional records for 
m-album in the following counties: Giles (Straley), Rappahannock (Ferris), Rock- 
bridge (Covell), Chesapeake (Anderson & Straley), Stafford and Virginia Beach 
(Nicolay). Most of these records represent captures of single specimens, supporting 
the Clarks’ statement (p. 78) that m-album is apparently a permanent resident, but 
often of irregular occurrence in any one place. We have taken it most commonly 
in Chesapeake, attracted to the flowers of sourwood and especially sweet pepper bush. 

Erora laeta (Edwards). At the time the Clarks completed their work on the 
Virginia butterflies in 1951, only one specimen of the Early Hairstreak was known 
to have been taken in Virginia (Mountain Lake, Giles County, 23 June 1938). A 
second specimen, a fresh female, was collected by Straley with his fingers at Eggleston, 
also in Giles County, on 7 May 1964, on moist sand on the bank of New River. The 
location was near a wooded area, but not a typical habitat where one might expect 
laeta. Mitchell also took a specimen in neighboring Montgomery County in Poverty 
Hollow, 27 April 1969. 

The only multiple capture of laeta in Virginia was by Wagner in July 1970, again 
in Giles County, on the road between Mountain Lake and West Virginia. In a letter 
dated 19 July 1970, Wagner stated, “I have seen over two dozen Erora laeta! .. . 
There is a spot along the West Virginia Road where you can see two or three at a 
time.” He took nine specimens, but spent a great deal of time just observing Iaeta, 
hoping to learn more about the habits of this elusive hairstreak. Unfortunately, 
because of a sudden change of weather to dark and rainy lasting for more than a 
week, the butterflies disappeared and the investigations ceased. 


NYMPHALIDAE 


Speyeria diana (Cramer). This favorite of collectors can stil! be collected com- 
monly in the vicinity of Brush Mountain, just west of Blacksburg in Montgomery 


VOLUME 27, NUMBER 2 153 


Co., and probably in many other localities in the lower mountainous regions of 
Virginia. Covell found that the males appeared in the first week in July, not finding 
any in late June, during 1960-62. They at first kept to the higher elevations (around 
2,000—2,500 ft.), flying along and alighting in the dirt road atop Brush Mountain. 
There they might be found visiting animal excrement or carrion. Arter a few days 
they would be found visiting flowers of the milkweeds Asclepias tuberosa L. and 
A. syriaca L., growing in bends in Rt. 460 down the eastern slope, and in Poverty 
Hollow at the bottom of the western slope of Brush Mountain. The affinity of diana 
for milkweed blossoms—especially A. tuberosa, the Orange Milkweed or Butterfly- 
weed—seems general for the species, as Covell has collected both sexes on these 
flowers in North Carolina and Kentucky as well. Mitchell has reported good catches 
of diana in this general area in more recent years. The middle of July seems the best 
time to seek both sexes in good condition. 

The Clarks (p. 57) also mentioned Coastal Plain populations of diana, recorded 
from counties south of the York River (Chesterfield and James City south into 
Nansemond counties). The continued presence of the species in this general area was 
indicated by the capture by Covell of males on A. tuberosa beside Rt. 33 a few 
hundred yards from its junction with Rt. 17 at Glenns in Gloucester Co. The species 
was seen there on 19 June 1958, and males were collected there on 25, 26, and 27 
June. As in the mountains, species visiting Asclepias with diana included S. cybele 
and Harkenclenus titus mopsus. These captures represent a new county record as 
well as a slightly northern range extension in the Coastal Plain, apparently the first 
taken north of the York River. 

SATYRIDAE 

Lethe portlandia anthedon (Clark). The Clarks (p. 31) record this butterfly 
from only four mountain counties in the western part of the State. We can add 
two additional counties: Montgomery (Covell & Straley, four locations) and Rock- 
bridge (Kenneth Frank). The Clarks mention only one brood; but we have records 
for every month from 20 May to 4 Aug., indicating at least two broods, maybe three. 
All our records are for captures of single specimens. 

Lethe appalachia R. L. Chermock. This species was referred to by the Clarks and 
by Covell (1967) as a subspecies of L. eurydice (Johansson). In accordance with 
Cardé, Shapiro & Clench (1970), we here refer to appalachia as a valid species. 
Note the Virginia records, including paratypes, cited on p. 87 of Cardé, Shapiro & 
Clench, adding to those of the Clarks (p. 32). Covell found a small colony in grassy 
backwaters of a pond in Poverty Hollow, Montgomery Co., in 1960, and the two 
following years (18-31 July). 


ACKNOWLEDGMENTS 


We wish to express our gratitude to the following for their help in the 
form of records and other valuable information and assistance in prepar- 
ing this manuscript: LCDR Richard A. Anderson, Fayetteville, N.C.; 
Mr. Harry K. Clench, Carnegie Museum, Pittsburgh, Pa.; Mr. William 
D. Field, U.S. National Museum, Washington, D.C.; Mr. Kenneth Frank, 
Boston, Mass.; Dr. Lee D. Miller, Allyn Museum of Entomology, Sarasota, 
Fla.; Mr. Judson T. Mitchell, Blacksburg, Va.; Col. Stanley S. Nicolay, 
Virginia Beach, Va.; Mr. Richard Pine, Bayside, N.Y.; Mr. Llewellyn 
Powell, Richmond, Va.; Mr. Amos Showalter, Waynesboro, Va.; and Dr. 
Warren H. Wagner, Jr., University of Michigan, Ann Arbor, Mich. For 
the photography we are indebted to Warrant Officer G. G. Thomas and 
S/Sgt. Richard E. Banzal, USMC. 


154 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


LITERATURE CITED 


CarpeE, R. T., A. M. SHapmo, & H. K. Crencw. 1970. Sibling species in the 
eurydice group of Lethe (Lepidoptera: Satyridae). Psyche 77(1): 70-103. 
Cuark, A. H. & L. F. Crarx. 1951. The Butterflies of Virginia. Smithsonian Misc. 
Coll AIG CA z2iapeneO mols: 

Ciencu, H. K. 1971. Some records of Euristrymon ontario (Lycaenidae). J. 
Lepid. Soc. 25: 80-82. 

CovELL, C. V., Jr. 1962. The occurrence of Satyrium kingi (Lycaenidae) in 
Virginia. J. Lepid. Soc. 18: 197-198. 

. 1967. A checklist of the butterflies and skippers of Virginia. Va. J. Sci. 

18: 21-24. 


Massey, A. B. 1961. Virginia Flora. Tech. Bull. 155, Va. Agr. Exp. Sta., Blacks- 
burg, 258 p. 


pos Passos, C. F. 1964. A Synonymic List of the Nearctic Rhopalocera. Mem. 
Lepid. Soc. 1. 145 p. 

1970. A revised synonymic catalogue with taxonomic notes of some nearctic 
Lycaenidae. J. Lepid. Soc. 24: 26-38. 

SrrALey, G. B. 1969. Occurrence of Thymelicus lineola (Hesperiidae) in Virginia. 
J. Lepid. Soe: 233 76. 


A MASSIVE MIGRATION OF KRICOGONIA (PIERIDAE) IN 
CAMPECHE, MEXICO 


Byers (1971, J. Lepid. Soc. 25: 124-125) notes that, “. . . this is the first record 
of a migration of a species of Kricogonia,” and, “Recent books on insect migration. . . 
do not mention the genus.” My article on migrations (1959, J. Lepid. Soc. 13: 62-64) 
must have been overlooked, as I recorded the migration of Kricogonia lyside Godart. 
Attention must also be called to articles by Clench (1965, J. Lepid. Soc. 19: 223- 
224), Heitzman (1962, J. Lepid. Soc. 16: 249-250), Howe (1964, J. Lepid. Soc. 
18: 26), and Welling (1964, J. Lepid. Soc. 18: 229-230). 

dos Passos (1964, A Synonymic List of the Nearctic Rhopalocera. Mem. Lepid. 
Soc. 1, p. 46) refers to two species, K. lyside Godart 1819, and K. castalia ( Fab.) 
1793. de la Torre y Callejas (1958, Reconsideracién Taxonémica de las Especies del 
Género Kricogonia Reakirt con Vista al Estudio de sus Organos Genitales. Publ. 
Univ. Oriente, Santiago, Cuba) refers to K. lyside and K. castalia as being a single 
species, and K. cabrerai Ramsden 1920 as the other. As to which species we may 
be referring to, much must be left to speculation. Klots (1951, A Field Guide to 
the Butterflies. Houghton-Mifflin, Boston) notes that a thorough study of the com- 
plex needs to be made, most records being untrustworthy. 

I take this opportunity to record another great migration of Kricogonia. I first 
note that 1971 seemed to be a year of intense rainfall in the Yucatan peninsula, after 
a few years of fair to mediocre precipitation. In my prior article (1959, op. cit.) I 
refer to dry year cycles alternating with wet year cycles, and migrations seeming to be 
associated with the latter. On 9 June 1971, on passing south of the city of Campeche, 
Campeche, and until reaching Escarcega, Campeche, I observed the heaviest migration 
I have ever seen of any kind of insect. Once again it was our commonly-reported 
Kricogonia sp. The heaviest part of the migration was slightly south of Lerma, 
through the villages of Sihochac, Seybaplaya, Haltunchén, Champotdon, to about 
X-bacab, all in the state of Campeche. The slight wind was from the east, with 
the migration going straight against it. Where these butterflies originated, I could 
not guess, as all the way south to X-bacab the migration direction was straight east- 


VoLUME 27, NUMBER 2 155 


wardly. These great swarms were seen as coming from the sea, as the highway follows 
the shore very closely most of the way. Slightly before mid-day a great thunderstorm 
arose from the east, but in spite of the heavy rain, the butterflies did not stop to 
take shelter or rest—they just kept coming. 

It is difficult to calculate the immense number that might have passed 100 meters 
of roadway per minute, as they formed a veritable yellow-white cloud. A conservative 
attempt would say there were 10,000 specimens per 10 meters of roadway, taking 
into account the width of the road, and the fact that the bulk of them were flying 
from ground level to 6-8 meters high. There were so many dead butterflies along 
the road, having been hit by vehicles or killed by other factors, that it looked as if 
it had just snowed. As I drove through the cloud of migrants I must have been 
killing at least 100 or more every split second; the din of their crushed bodies against 
my vehicle stirred remorse within me—I felt like a treacherous assassin. This migra- 
tion began to thin out near X-bacab, and by the time I arrived at Escarcega, it was 
possible to calculate about 1000 specimens crossing 100 meters of road per minute. 

When I returned through the same area about 3 July 1971, the bulk of the migra- 
tion had waned. Now the strongest concentration was south of X-bacab, through 
Escarcega, and southwest to Pital and Rio Candelaria. The numbers were about 500 
per 100 meters of roadway per minute, and the direction had changed towards the 
southeast. 

I will here add that when passing through Tamaulipas in northeast México in early 
June after leaving Campeche, I noticed Kricogonia flying westwardly, but in minimum 
numbers, about 1 or 2 in sight at any given moment, crossing the road at ground 
level. 


Epuarpo C. WELLING M., Apartado Postal 701, Mérida, Yucatan, México. 


AN ALTERNATIVE CAUSE OF DIMORPHISM IN PAPILIO PUPAE 
(PAPILIONIDAE ) 


D. F. Owen’s paper, “Pupal Colour in Papilio demodocus (Papilionidae) in Rela- 
tion to the Seasons of the Year,” (1971, J. Lepid. Soc. 25: 271-274), has prompted 
me to make the following observations. 

The idea of seasonal variation appears to have one fatal flaw. The Rutaceae, on 
which Papilio demodocus and many other Papilio species with dimorphic pupae feed, 
are always in green leaf and leaves fall off while still green. Furthermore, a recent 
note by Vaidya (1971, J. Bombay Nat. Hist. Soc. 68(2): 477-478) points out that 
the ovipositing female of Papilio demoleus L. requires a visual stimulus connected 
with leaf colour as well as the olfactory one connected with the scent of the food- 
plant. In my experience females of the citrus-feeding Papilio species invariably lay 
on new growth. 

Generally speaking, what may be considered as typical pupae, such as Papilio 
polytes L., P. demoleus L., P. demodocus Esp., P. nireus L. and many others, are 
dimorphic, either green or otherwise, the ‘otherwise’ varying from pale to dark 
brown, various shades of grey, etc. Specialised pupae, such as the cylindrical, dark, 
stick-like pupa of Chilasa clytia L., the flattened, green, leaf-like pupa of Papilio 
dardanus Brown, and the brown pupae of Byasa hector L. and B. aristolochiae F., 
with their leaf-like subdorsal projections, are monomorphic. 

If the colour of the pupa is controlled genetically, it is possible that there are three 
genotypes, a green, a brown and an optional green or brown controlled by external 


156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


stimuli. Such a hypothesis might explain the discrepancies in the figures for various 
broods which fail to conform to the usual Mendelian ratios. 

The ‘matching’ theory has never attracted me, there are far too many exceptions 
to be found both in pupae formed naturally in the wild and those formed in the 
somewhat unnatural conditions of captivity; and experiments I carried out in 
Calcutta with Papilio polytes L. and P. demoleus L. (Sevastopulo 1948, Proc. Roy. 
Entomol. Soc. London (A) 23: 93) which, although not conclusive, certainly did not 
support the theory. It was noted in the course of these experiments that when pupating 
larvae slipped through their girdles and fell, the proportion of brown to green pupae 
was far higher than when larvae suspended themselves and pupated successfully. 
This suggested that movement shortly before pupation might be the determining 
factor, resulting in the formation of melanin. 

Recently a paper by Oldroyd (1971, Entomologist 104: 111-123) gave the 
interesting information that the difference in colour between green and brown pupae 
of Papilio memnon L. and others appeared to be due to the presence of melanin in 
the latter. 

I have, therefore, put forward the suggestion that it is the amount of pre-pupational 
travel that determines whether a pupa will be green or brown. Such a theory would 
account satisfactorily for the fact that green pupae are usually found among leaves, 
as the larvae producing these would not have to travel far, as well as explaining the 
occasional brown pupa found in such situations, whilst larvae that travelled some 
distance would be more likely to pupate on a branch or tree trunk and would be 
brown. Owen’s own experiment would appear to support this idea. 

Other supporting examples can be quoted from non-papilionid species. Recently 
at Mackinnon Road in Kenya I found enormous numbers of the pupae of the pierid 
Anapheis aurota F. These were placed nose-to-tail all along bare twigs of the stripped 
foodplant and side-by-side all around them. In many cases earlier pupae had been 
used as the substrate for later ones. All these pupae were a sooty black with varying 
amounts of white patches, and quite unlike the usual form of pupa formed in un- 
crowded situations, which is very like the norma! forms of pupa of the European, 
and probably American, species of the genus Pieris. There can be no doubt that 
these vast numbers of larvae must have interfered with each other immediately prior 
to pupation and the amount of wriggling must have been tremendous. 

Another example is provided by the nymphalid Aglais urticae LL. When solitary 
larvae pupate amongst the leaves of the foodplant, they produce a brilliant golden 
pupa, but when they pupate on a neighbouring fence or similar object the pupae are 
blackish. Poulton, who studied the question extensively (1892, Trans. Entomol. Soc. 
Lond., 293-487), considered that the pupal colour was determined by the light 
falling on the larva immediately prior to hanging up for pupation, a yellow light 
producing golden pupae and a blue light dark ones, and suggested in another 
paper, which I am not able to trace at present, that the effect of the dark colour 
of a number of clustered larvae immediately before pupation was the cause of the 
blackish pupae formed in such situations. Is it not possible that the excessive move- 
ment of such a group of suspended larvae resulted in the production of extra melanin 
through oxidization? 

Another supporting factor is that such a hypothesis would provide a satisfactory 
explanation for the fact that many species of larvae, when reared in overcrowded 
conditions, produce a form darker than the normal, the overcrowding resulting in 
abnormal disturbance and subsequent movement. (Long, 1953, Trans. Roy. Entomol. 
Soc. Lond. 104: 543-585). 

Finally, to go outside the Lepidoptera, apart from the migratory urge, the main 
difference between the solitary and gregarious phases in the hopper stage of locusts 
is the darker colour of the latter, again the result of excessive melanin. 


D. G. SEvAsropu.o, F.R.E.S., P.O. Box 95026, Mombasa, Kenya. 


VoLUME 27, NUMBER 2 157 


ABNORMALITIES AND HEREDITY 


Mr. Manley’s paper, “Iwo Mosaic Gynandromorphs of Automeris io (Satumiidae),” 
(1971, J. Lep. Soc. 25: 234-238), has, to me, a most surprising omission—he does 
not mention whether or not the two specimens he describes came from the same 
brood. Many abnormalities, spiral segmentation for example, would appear to have 
some hereditary basis, and Ford (1955, Moths, p. 39) quotes a case where a brood 
of Hemerophila abruptaria Thnbg. (Geometridae) contained no fewer than four 
gynandromorphs. 

This phenomenon appears to be far more common, or far more often observed, in 
Great Britain than in America, to judge from reading Mr. Manley’s paper. Hardly 
a year goes by without there being some reference to one or more cases in the 
various British entomological journals. Cockayne, in a paper published in the 
Transactions of the Entomological Society of London in 1916, illustrated no fewer 
than twenty-one gynandromorphs of the lycaenid Lysandra coridon Poda. 

Reverting to my remarks on spiral segmentation, the only two cases which have 
come to my personal knowledge have both involved more than one individual in a 
brood. In one case, a brood of the nymphalid Euxanthe wakefieldi Ward bred by a 
friend, the majority of the larvae died of disease before developing the characteristic 
and revealing dorsal markings; out of the five or six that reached maturity, two were 
examples of spiral segmentation, and it is more than probable that there were other 
examples undetected among the larvae that died earlier. The other case was in a 
brood of the noctuid Leucania irregularis Wlk., which contained at least four examples 
of spiral segmentation. Unfortunately the brood, which was divided into three batches 
at an early stage, suffered severe casualties; two of the three batches were wiped out 
by virus disease when still small, and a large number of the third batch was used 
to provide live food for some insectivorous birds before the abnormalities were 
detected, so that, here again, it is more than probable that a considerably larger 
number of spirals were actually present in the brood. 

Might I end on a note of criticism of Mr. Manley’s paper. Surely it is incorrect 
to write (top of p. 235) “Thus cells of the male with ZZ chromosomes are expressed 
as yellow, while those of the female with a ZO chromosome complement are rosy 
brown.” Would it not be more correct to state that the scales and hairs arising from 
cells with ZZ chromosomes are yellow and those from cells with ZO chromosomes are 
rosy brown? It is unfortunate that the figures are too dark to allow the pattern of the 
‘broken eye blotch’ to be seen. 


D. G. SEvAsropuLo, F.R.E.S., P.O. Box 95026, Mombasa, Kenya. 


HOST RECORDS FOR BREPHIDIUM EXILIS (LYCAENIDAE) 


The Western Pygmy Blue, Brephidium exilis Boisduval, is generally recorded as 
feeding on Chenopodiaceae, but specific host identifications are few. Downey (1961, 
in Ehrlich & Ehrlich, How to Know the Butterflies) lists, “Atriplex bracteosa ( lamb’s 
tongue), Chenopodium (pigweed), Petunia parviflora.” Petunia is in the Solanaceae, 
a most unusual group to be fed upon by a Lycaenid; one species, P. violacea Lindl. is 
recorded as toxic to various insect larvae (Shapiro 1968, Ann. Entomol. Soc. Amer. 
61: 1221). This paper presents information on host plants for B, exilis in lowland 
central California; plant names are from Munz & Keck (1970, A California Flora). 


158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Chenopodiaceae 


Atriplex patula L. “ssp. hastata (L.) Hall. & Clem.” (= A. hastata L.).—This 
abundant plant of brackish marshes and moist alkaline situations is the only known 
host for the large population of B. exilis in the marshes along Suisun Slough near 
Fairfield, Solano Co., California. It is probably the principal host in salt marshes 
throughout the range of B. exilis and is also used locally in the interior, e.g. in the 
Sacramento Valley east of Woodland, Yolo Co. The larvae feed on leaves, flowers, 
and fruit. 

Atriplex rosea 1.—An abundant weed of dry, alkaline, usually disturbed soils 
throughout interior California; naturalized from Eurasia. The most frequent host of 
B. exilis throughout the Sacramento Valley. Larvae feed on buds, flowers, and fruit 
and can only be found near the tips of branches. 

Atriplex cordulata Jeps.—Occasional native species of dry, compacted alkaline soils; 
used by B. exilis north of Davis, Yolo Co. 

Atriplex semibaccata R. Br.—Occasional weed of alkaline or subsaline soils and 
roadsides; naturalized from Australia. Fed upon by B. exilis at Fairfield, Solano Co., 
and north and west of Davis, Yolo Co. Larvae on leaves and (especially) the female 
flowers. 

Suaeda fruticosa (L.) Forsk. (= S. moquini Greene ).—Local but often common, 
a perennial weed of dry alkaline and subsaline situations as at the Davis municipal 
landfill and along the levees at Willow Slough, Yolo Co. B. exilis larvae abundant 
on the plant, feeding on inflorescence, fruit, and upper leaves; seemingly preferring 
Suaeda to Atriplex rosea where the two occur together. Populations of larvae are so 
dense on this plant as to constitute a potentially significant threat to its seed produc- 
tion, at least locally. 

Brephidium exilis has not been found feeding or ovipositing on any of the following 
Chenopodiaceous weeds, although all are common within its range and often in close 
proximity to Atriplex or Suaeda bearing many larvae: Salsola kali L. var. tenuifolia 
Tausch. (“Russian thistle”); Cycloloma atriplicifolium (Spreng.) Coult.; Cheno- 
podium ambrosioides L. and its varieties anthelminticum (L.) Gray and vagans 
(Standl.) Howell; C. botrys L.; C. californicum Wats.; C. album L.; C. murale L. 
Larvae taken from Suaeda fruticosa accepted Chenopodium californicum and C. 
murale but refused C. ambrosioides and C. botrys. They also accepted Cycloloma and 
Kochia scoparia (L.) Schrad. One larva taken from Atriplex rosea accepted C. botrys 
flowers. I have not seen B. exilis oviposit on any species of Chenopodium, but have 
taken females hovering around C. murale (banks of the Sacramento River) and C. 
botrys (Cache Creek, Yolo Co.), once each. Records on Chenopodium may well be 
due to misidentification of Atriplex rosea, which resembles that genus strongly. 

Although the Eastern Pygmy Blue, Brephidium pseudofea Morrison, is recorded 
as feeding on Salicornia (Downey 1961, op. cit.), B. exilis showed no interest in 
S. virginica L. in the Suisun marshes where extensive patches of it are near colonies 
of Atriplex patula ssp. hastata, and no larvae were found by sweeping. 

Larvae of the Western Pygmy Blue pupate on the host plant. As many of the 
hosts are tumbleweeds, passive dispersal of pupae on the plants in fall and winter 
may be a significant factor in the colonizing ability of the insect. 


Solanaceae 


Petunia parviflora Juss. was not available for testing, but leaves, buds, and flowers 
of two horticultural varieties of P. violacea were completely unacceptable to 23 larvae 
of mixed sizes taken from Suaeda. Although less viscid than the garden species, 
P. parviflora is glandular-puberulent and should be suspected of the same toxic 
properties as its congener until shown otherwise. 


ArtHuR M. Swaprro, Department of Zoology, University of California, Davis, 
California 95616. 


VOLUME 27, NUMBER 2 159 


AN ATTEMPTED INTERFAMILIAL MATING 
(LYCAENIDAE, NYMPHALIDAE) 


Interspecific courtships and matings are seldom reported in butterflies (Downey 
1962, J. Lepid. Soc. 16: 235-237). The rarity of such reports involving pheno- 
typically similar sympatric species suggests the widespread occurrence of effective 
prezygotic reproductive isolating mechanisms, at least some of which may have 
evolved as a result of selection against deleterious hybridization (Remington 1968, 
Evol. Biol. 2: 321-428). Although species from widely different groups might be 
expected to differ in so many ways as to make courtship and mating very unlikely, 
it is conceivable that such taxonomically wide behavioral “mistakes” may be at least 
as frequent as those between sympatric congeners. The actual records are far too 
fragmentary to allow a conclusion one way or the other. This note reports a mistaken 
courtship involving representatives of two butterfly families which, however, did not 
result in copulation. 

On 26 May 1972 a fresh male Lycaena helloides (Boisduval) (Lycaenidae) was 
seen courting a fresh female Cynthia annabella Field (= Vanessa carye auct.) 
(Nymphalidae) in a vacant lot at Southport, Yolo County, California. The pair was 
first encountered at 1357 hours. The male fluttered behind the female, which sat 
on a leaf just above the ground. Repeated attempts at genital contact elicited only a 
shuffling sideways movement in-the female, which finally flew desultorily some ten 
feet and lit on the ground, only to be overtaken at once by the male. This sequence 
was repeated four times in nine minutes. The courtship was terminated when the 
observers shadow was inadvertently passed over the female and she left the area. 
The behavior of the female C. annabella was in no way dissimilar to that shown by 
the species when being courted by its own males, but not disposed to mating. How- 
ever, C. annabella, like most “Vanessas,” seems to mate principally in the late 
afternoon and at dusk. 

Lycaena helloides was abundant in the area and several unsuccessful courtships 
within that species were seen during the early afternoon; female L. helloides invariably 
fanned their wings intermittently in response to the activity of the male, differing 
in this respect from the response of the C. annabella. The female annabella ap- 
proached was the only individual of its species in the field during most of the after- 
noon. The pheromones, if any, of neither species have been studied. The C. anna- 
bella was fully twice the size of an L. helloides female, and resembled it superficially 
only in general color. 


ArtHur M. SHapimo, Department of Zoology, University of California, Davis, 
California 95616. 


ISSIKI COLLECTION OF MICROLEPIDOPTERA TO THE 
SMITHSONIAN INSTITUTION 


With considerable assistance and cooperation from the United States Department 
of Agriculture and the United States Forest Service, the Smithsonian Institution has 
recently acquired the finest collection of Japanese and Formosan Microlepidoptera 
ever assembled. The collection represents the lifelong effort of Professor Syuti Issiki, 
a renowned Japanese entomologist, who, with his famous teacher S$. Matsumura, 
pioneered the study of Oriental Microlepidoptera. 

The total size of the Issiki Collection is not unusually large, numbering 16,236 
moths, but the rich representation of Eastern Palearctic and Oriental species makes 
the collection an extremely valuable research tool. Probably 95% of the known 


160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Japanese microlepidopterous fauna is represented in addition to nearly all of the 
described Formosan species. The collection is particularly strong in the primitive 
families, an area in which Dr. Issiki specialized. Seventy-eight holotypes are present 
and nearly 200 secondary types, several of the latter having been contributed in 
recent years by Prof. Issiki’s former students. In addition to Microlepidoptera, the 
order Mecoptera was also an early research interest of Dr. Issiki. As a result, his 
collection also contains over 1,000 specimens of this order which, likewise, exhibits 
excellent coverage of both Formosa and Japan. 

The Formosan collection represents the only serious attempt to survey the Micro- 
lepidoptera of that country and was accumulated during Dr. Issikis tenure as 
Professor of Agriculture at the Imperial University at Taipei from 1920 to 1948. 
Examples of this material were sent to Edward Meyrick and, thus, formed the basis 
of Meyrick’s studies on Formosan moths. Unfortunately, the collection was con- 
siderably larger than at present but was partially destroyed due to neglect following 
Prof. Issiki’s permanent departure to Japan in 1948. The only material known to 
survive of the original Formosan collection, other than those duplicates retained by 
specialists such as Meyrick, is the synoptic representation which Issiki was able to 
transport at the time of his departure. During his residence in Formosa, Prof. Issiki 
also collected for a brief interval in New Guinea, assembling a small but select 
representation of Microlepidoptera from that area. 

In 1949 Dr. Issiki was appointed Professor of Agriculture at the Univeristy of Osaka 
Prefecture, a post he held until his retirement in 1961. It was during this period 
that the major portion of his Japanese material was acquired. His influence as a 
teacher was also impressive, as several of the present generation of Japanese micro- 
lepidopterists studied under him during this period. Dr. Issiki now resides with his 
family in Ito City where he still actively pursues his interests on the biology of 
Japanese Microlepidoptera. 


Donatp Ray Davis, Curator, Department of Entomology, Smithsonian Institution, 
Washington, D.C. 20560. 


A NEW GENERIC NAME IN MIDILINAE (PYRALIDAE) 
Eupastranaia Becker, new genus 


Pastranaia Munroe, 1970, p. 63-64; preocc. by Pastranaia Orfila, 1955, p. 30-31. 

The genus Pastranaia was erected by Orfila (1955) for a new species of Nemopteri- 
dae (Neuroptera). Unfortunately Munroe (1970) chose the same name in his 
excellent work “Revision of the subfamily Midilinae (Lepidoptera: Pyralidae),” for 
the fenestrata group of this subfamily. 

This homonymy was communicated to Dr. Munroe who recommended that the 
author propose a new name and that it continue to honor Dr. J. A. Pastrana. 

The author is very grateful to Dr. Munroe for recommending that he publish the 
new name. 


References 
Munrog, E. 1970. Revision of the subfamily Midilinae (Lepidoptera: Pyralidae). 
Men. Entomol. Soc. Can. 74, 94 p. 


Orrita, R. N. 1955. Un nuevo Nemopteridae (Neuropt.) americano con una 
sinopsis de la familia. Rev. Soc. Entomol. Argent. 17: 29-32. 


Vitor OsMAR Becker, InterAmerican Institute of Agricultural Sciences, Organiza- 
tion of the American States, Turrialba, Costa Rica. 


aS ee eee eS 


Mea ee He ee ee 


ee ee ee ee Se ee ee ee ee ee 


Natl 


ee ee ae 


re Ee ee PT ee a Ee ee ee ee Ey ee yee 
i 


EDITORIAL COMMITTEE OF THE JOURNAL 
Editor: THEoporE D. SarcEnt, Department of Zoology, 
University of Massachusetts, Amherst, Massachusetts 01002 
K. S. Brown, J. M. Burns, R. H. Carcasson, J. P. DoNnAHUE 
J. F. Gates Criarxe, C. D. Ferris, R. O. KENDALL, J. H. MASTERS 
_L. D. Micter, A. P. Puart, J. R. G. Turner 


> 


> 


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 CITeED, 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: A!] 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% 
>< 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 article’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 approx- 
imate 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. 

Page Charges: Authors with grant or institutional funds are requested to pay a 
charge of $24.00 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 
case of authors who have no grant or institutional funding, as it is not intended that 
any author should pay this charge from personal funds. However, all authors will 
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: Dr. C. V. Covell, Department of Biology, University of 
Louisville, Louisville, Kentucky 40208. 


ALLEN PRESS, INC. aes LAWRENCE, KANSAS 
use 


CONTENTS 


LEPIDOPTERA FEEDING AT PUDDLE-MARGINS, DUNG, AND CARRION. 
To Ae DD OUEST Ng rr Rui 


A New GeENuSs AND SPECIES OF OECOPHORIDAE FROM TROPICAL 


AMERICA? #'\J.: F. Gates Clarke 0 ae 


RECORD AND ILLUSTRATION OF SOME INTERESTING Motus FLYING 
In Texas (SPHINGAE, CTENUCHIDAE, NoctumAgE, Norto- 
DONTIDAE, (GEOMETRIDAE, PyYRALIDAE, CossmpAE). André 
Blanchard). 8g 


CHROMOSOME NUMBERS FOR PLEBEJUS (ICARICIA) ACMON, P. LUPINI, 


AND P. NEURONA (LycAENDAE). Carll Goodpasture —_ 


CALLOPHRYS (INCISALIA) POLIOS (LYCAENIDAE): DISTRIBUTION IN 


NortH AMERICA AND DESCRIPTION OF A NEW SUBSPECIES. 
Clifford D: Ferris and Michael S. Fisher _. 


THE Larva or LOXAGROTIS KYUNE (Barnes) (NocrumaE). George 
bh. Godjrey 2 
METHODS FOR EXTERNALLY SEXING MATurRE LARVAE AND PUPAE OF 
LimMENITIs (NYMPHALIDAE). Philip J. Kean and Austin 
Fo Phage: note ob A NSIOL ECON AMCOMNMOANMRIINRMCUNO |S 


ILLUSTRATIONS OF HELICONIUS (NYMPHALIDAE): SOME RARE AND 


IMPORTANT SPECIMENS. John R. G. Turner : 
POPULATIONS.; OF PAPILIO ANDRAEMON BONHOTEI SHARPE AND 


PAPILIO ARISTODEMUS PONCEANUS SCHAUS (PAPILIONIDAE) IN 
BiscAYNE NATIONAL MONUMENT, FLorma. Larry N. Brown 


A NEw SPECIES OF THE GENUS GLENOIDES McDuNNoucH ( GEOMET- 


RIDAE). André Blanchard 


Notes ON VIRGINIA BUTTERFLIES, WITH Two New STATE RECORDS. 
Charles V. Covell, Jr. and Gerald B. Straley 


GENERAL NOTES 


A massive migration of Kricogonia (Pieridae) in Campeche, Mexico. 
Eduardo C. Wellg M. 


Midges (Diptera: Ceratopogonidae) sucking blood of caterpillars. D. 


G. /Sevastonulo iw OE is 
An alternative cause of dimorphism in Papilio pupae (Papilionidae). D. 
G. Sevastopubo: 2 Oi Os 
Abnormalities and heredity.. D. G. Sevastopulo 
On Glenoides texanaria (Geometridae) with designation of the lectotype. 
Frederick: Hy Rindge ao ne es oN 
Host records for Brephidium exilis (Lycaenidae). Arthur M. Shapiro __ 
An attempted interfamilial mating (Lycaenidae, Nymphalidae). Arthur 
Me SREB ee eee a SAN 


Issiki collection of microlepidoptera to the Smithsonian Institution. 
Donald: Ray Dapisicn gu MU OE CE 


A new generic name in Midilinae (Pyratidaeh) Vitor Osmar Becker 


103 


109 


112 


119 


122 


130 


136 


ms el i ee ee le 


a te 


tea. 


Stal pe 


Volume 27 1973 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 


10 August 1973 


THE LEPIDOPTERISTS’ SOCIETY 
EXECUTIVE COUNCIL 


J. F. Gates Ciarxe (Washington, D.C.) President 

Harry K. Crencu (Pittsburgh, Penn.) President-elect 
ALEXANDER B. Kiors (New York, N.Y.) 1st Vice President 
C. F. Cowan (Berkhamsted, England) Vice President 

E. G. Munroe (Ottawa, Ontario) Vice President 

S. S. Nicouay (Virginia Beach, Va.) Treasurer 

Lee D. Mixxer (Sarasota, Fla.) Secretary 


Members at large (three year term): M. C. Nuevsen (Lansing, Mich.) 1974 
A. BLANCHARD (Houston, Texas) 1973 D. C. Fercuson (Washington, D.C.) 
R. B. Dominick (McClellanville, S.C.) 1975 

1973 R. O. Kenpatt (San Antonio, Texas) 
J. P. DonanveE (Los Angeles, Calif.) 1973 1975 
J. M. Burns (Cambridge, Mass.) 1974 J. A. Powext (Berkeley, Calif.) 1975 
R. H. Carcasson (Vancouver, B.C.) 1974 


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 $10.00 
Student members—annual dues $7.50 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $15.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia 23455. 


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


by Cyru. F. pos Passos 


Price, postpaid: Society members—$5.00, others—$7.50; uncut, unbound signatures 
available for interleaving and private binding, same prices; hard cover bound, mem- 
bers—$8.00, others—$10.00. Revised lists of the Melitaeinae and Lycaenidae will be 
distributed to purchasers free (separately with paper covered copies and unbound 
signatures, bound in with hard covered copies). 


The Lepidopterists’ Society is a non-profit, scientific organization. The office of 
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520. 
Second class postage paid at Lawrence, Kansas, U.S.A. 66044. 


ate “Gg eee gee me ee eer aR a GT i sitet 


ee 2 oat 


maar 


JOURNAL OF 


Tue LEPIDOPTERISTS’ SOCIETY 


Volume 27 1973 Number 3 


TAXONOMIC SIGNIFICANCE OF REFLECTIVE PATTERNS 
IN THE COMPOUND EYE OF LIVE BUTTERFLIES: A 
SYNTHESIS OF OBSERVATIONS MADE ON SPECIES 
FROM JAPAN, TAIWAN, PAPUA NEW GUINEA 
AND AUSTRALIA 


ATUHIRO SIBATANI 
30 Owen St., Lindfield, New South Wales 2070, Australia 


During observations in the field of New South Wales, I came to notice 
that some Australian Lycaenidae had unusual semi-transparent and some- 
times brightly coloured eyes which I had not come across before in some 
other parts of the world, including temperate and tropical Eurasia and 
America. This character could be observed only in live or recently killed 
butterflies. The regular occurrence of this type of eye in certain lycaenid 
groups strongly suggested its taxonomic usefulness. Upon extending 
my observation to other butterfly families, I soon realised that in such 
semi-transparent eyes there were usually certain reflective spots which 
changed their position according to the direction of observation, and 
that these spots were observed almost invariably in Pieridae and Nym- 
phalidae (s.str.), but not in Papilionidae and Hesperiidae, and variably 
in Satyridae, Danaidae and Lycaenidae. Moreover, the pattern of these 
spots also appeared to be of taxonomic significance. During the past 
two years I have thus accumulated records of my own observations on 
the superficial feature of the eye in butterfly species occurring in New 
South Wales and Papua New Guinea. 

Meanwhile, my attention was drawn to the extensive monograph, “The 
Compound Eye of Lepidoptera,” by Yagi and Koyama (1963). In this 


work the authors not only recorded the patter of reflective spots in 
fresh eyes for the majority of butterfly species in Japan and many species 
from Taiwan, but also correlated them to the histologic structure o! the 
ommatidium and thus clarified the optical basis of the appearance 0! 


these spots. Although they gave a number of suggestions about the 


162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


taxonomy and phylogeny of butterflies as well as other Lepidoptera 
families based on their studies on the compound eye of Lepidoptera 
at large, I feel that this extensive and important work treating the subject 
in depth has so far failed to have an impact on the taxonomy of but- 
terflies. 

In order to pay due tribute to the work, and to fill the lacuna of 
knowledge and interest in this aspect of butterfly morphology and 
taxonomy among lepidopterists, I wish to briefly redescribe the super- 
ficial (but not histologic) features of the butterfly eye only to the extent 
useful to careful field workers, on the basis of records compiled by 
Yagi & Koyama (1963) as well as my own observations. The material 
so collated covers species from Japan, Taiwan, Papua New Guinea and 
Australia, thus representing some of the major butterfly groups in the 
Palaearctic, Indo-Malayan and Australian Regions. The limitation of 
this character as a taxonomic criterion is the fact that the eye must be 
observed soon after an insect is captured, but this is certainly much less 
demanding than is the knowledge of foodplants and early stages for 
inclusion in taxonomic investigations. It is desired that, through the 
cooperation of field workers in various countries, patterns of live but- 
terfly eyes will soon be recorded for the groups not included in this 


paper. 


General Description and Terminclogy 


The superficial appearances of butterfly eyes may be divided into two 
major types. One is the eye looking totally dark; the other is the eye 
looking semi-transparent, lightly or brightly coloured and bearing at 
least one, and usually seven or more, dark spots which change their 
position and sometimes their shape when observed from different di- 
rections. 

The first type is observed in Papilionidae and Hesperiidae in which 
the eye may reflect strong incident lights in the form of a hexagonal 
bright central area. The images of such an eye are well recorded in 
some photographs of butterflies taken in the field, especially those of 
papilionid species: Papilio aegeus aegeus Donovan (Deger & Eden, 
1970, p. 5, fig. 4); Papilio protenor demetrius Cramer (Kohiyama, 
Takase & Fujioka, 1971, fig. 5); Papilio demoleus sthenelus Macleay 
(D’Abrera, 1971, p. 41). The hexagonal spot may be observed in the 
eye of dead and dried specimens, suggesting that it is mainly due to 
surface reflection, and thus is not the subject of this paper. In some 
Satyridae and most Lycaenidae, the eye looks completely dark without 
any hexagonal bright spot. An example may be seen in the photograph 
of Jalmenus eichorni (sic) Staudinger given on p. 3 of Deger & Eden 


VOLUME 27, NUMBER 3 163 


Fig. 1. A live eye of Pieris rapae Linnaeus, enlarged from a part of the original 
photograph of Deger & Eden (1970), and reproduced through the courtesy of the 
authors and publisher. 


(1970), and of Taraka hamada Druce given in fig. 47 of Kohiyama et al. 
CRD 

A good illustration of the second type of eye may be found in the 
photograph of Pieris rapae (Linnaeus) shown in fig. 3 on p. 9 of Deger 
& Eden (1970), of which I reproduce a part as Fig. 1 by courtesy of 
these authors. As seen in the original photograph printed in colour, the 
eye itself is pale greenish grey with a prominent dark central spot which 
is surrounded by six conspicuous primary side spots arranged in a hexa- 
gonal array. Around these primary side spots, there are vague dark 
shadows in the form of some discrete patches, which represent the 
secondary side spots, which are rather ill-defined in this species. | 

These spots were called by Yagi & Koyama (1963) the “pseudopupils. 
The name should have been phrased the ~ aise pupils” to contorm with 
a uniform Latin derivation rather than a mixture of Greek and Latin 


ones. However, the word “pupil” even with the adjective “false” sounds 

inadequate because of its possible an eanenareae bias. The more 
> | $4 “> "ht =) fy ‘ : a @ 

descriptive expression “reflective spots’ is being u inost as a jargon 


among Drosophia geneticists for a pair of appar similar spots in 


164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the eye of the wild-type individuals of Drosophila melanogaster Meigen. 
I want to borrow this expression in place of the “pseudopupil,” since 
the latter has not yet been established among lepidopterists, and will 
abbreviate it as RS in the rest of this paper. 


Main Types and Subtypes of the Compound Eye 


According to Yagi & Koyama (1963) the eye with RS may be divided 
into two major types. One has a large central spot only. In the other the 
central spot is smaller and usually accompanied by primary side spots 
which however may become very faint, and frequently by the secondary 
side spots which may surround strongly marked primary side spots. 

The nature of the appearance of these RS has not been formulated 
in exact physical or mathematical terms, but is correlated by Yagi & 
Koyama (1963) with an at least partial absence of pigment around the 
distal half of the ommatidium, allowing the oblique incident light to 
pass through one ommatidium into another. The distribution and extent 
of pigmentation seem to be related to the variation of the patterns of 
RS, but I will not deal with this aspect here. When the butterfly dies, 
the appearance of RS and the semi-transparency of the eye are progres- 
sively lost hand-in-hand, suggesting a change in the refractive index of 
the substance filling the ommatidium. In the dark eye without RS, the 
ommatidia appear to be optically isolated from one another by a pre- 
dominant pigmentation along the septa of each ommatidium. 

In the terminology of Yagi & Koyama (1963) the dark eye without RS, 
the light eye having only one (large) central RS, and the light eye 
having basically seven or more RS were called the non-pseudopupil 
type, mono-pseudopupil type, and multi-pseudopupil type, respectively. 
In this paper I will simply call them Type I, II and III eye, respectively. 
If necessary, they may be called the eye with no RS, single RS and 
multiple RS. I have never seen a Type II eye myself and its superficial 
character is somewhat obscure to me. In Fig. 2 I have assembled 
diagrammatic representations of these three basic types, and modifica- 
tions (or subtypes) of the Type III. 

In the subtype IIa, the side spots are very obscure. The primary side 
spots are conspicuous in IIIb; their position may be close to the central 
spot or to the periphery of the eye. This last example may be seen 
in the photograph of Polyura pyrrhus sempronius (Fabricius) on p. 
25 of D’Abrera (1971). In IIIc, the secondary side spots are apparent. 
Sometimes the intensity of all the spots becomes more uniform, giving the 
eye a mesh-like appearance. In IIId, radial striations appear, especially 
around the central spot, and each RS may take a hexagonal shape. In 
Ilfe, individual RS’s take a rectangular shape and tend to be confluent. 


VoLUME 27, NUMBER 3 165 


I | II Ia 


Id We ITl f III g Ih 


Fig. 2. Diagrammatic representation of types and subtypes of the reflective eye 
pattern in butterflies. Modified from Yagi & Koyama (1963) except for IIIe and h 
which are original, through the courtesy of Prof. N. Koyama. 


IIIf is characterised by dark areas intercalating the RS and tending to 
connect one another to circumscribe the latter. III[g represents a very 
conspicuous type in which the RS’s are longitudinally united to form a 
striated pattern in the eye. IIIh is the most ill-defined pattern; here the 
eye looks dark and the various modifications described in IIIc—g seem 
to appear to various extents in combination. 

Table 1 lists the distribution of the major types and some less fre- 
quently encountered types in each of the butterfly families' occurring 
in the areas covered, except for Riodinidae for which no observation 
has been available. Assignments of species to individual types and sub- 
types are compiled in the last section of this paper. 


Intraspecific Variation 


Repeated observations on different individuals of the same species 
have given consistent results in most cases, but there does seem to be 
some intraspecific variation in the appearance of the live eye. The 


variation may fall into several categories: 1. individual variation; 2. 
1The higher classification of butterflies is in a state of confusion at the moment Almost 
every author seems to have his or her own system of classification. The most logical approach 
to the problem may be that of Ehrlich (1958) but his system has not been followed by_ the 
majority of contemporary workers in the world. There is too little room for intermediate 
classification in some of the Ehrlich’s families which lumped together a number of distinct 
taxonomic groups. Here I have followed the more conventional classification for that reason. 
However, I admit that the standard of my recognising individual families is nO ae man 
1968) tor Satyridae, Fox 


scientific. For definition of individual families I followed Miller 
(1956) for Ithomiidae, and Eliot (pers. comm.) for Lycaenidae. Also 
splitting genera into smaller units is more (or too) conspicuous for Huropean 
fauna than Australian and New Guinean ones, and this necessarily caused 
recognising individual genera. 


the recent trend olf 
and Japanese 
unevenness in 


166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TasLeE 1. Distribution of eye types among various families of butterflies (O, 
common, predominant types; and @, unusual types, in individual families). 


Family I Il Ill 


Hesperiidae 
Papilionidae 
Pieridae 
Danaidae 
lithomiidae 
Amathusiidae 
Satyridae 
Nymphalidae 
Acraeidae 
Libytheidae 
Lycaenidae 


Cer OO 


O O80O0O 8800 
Oe 
O 
@ 


developmental change taking place soon after emergence of the adult; 
3. sexual dimorphism; 4. subspecific or seasonal difference; and 5. 
genetic polymorphism in one population. At the present stage of in- 
vestigation it is still difficult to make a distinction in this sense among 
some of the observed examples of intraspecific variation. In the following 
I give some examples worth noting. 

1. Danaus chrysippus petilia Stoll (Danaidae) males caught at one 
place (New South Wales) at the same time showed two subtypes of the 
Type III eye: e and h. Slight variation between a and b, b and ¢, or c 
and g may be encountered among a number of species having the Type 
mmeyc 

2. Yagi & Koyama (1963) noted that the eye of Aporia Hiibner 
(Pieridae ) turns from Type IIIf to Type I soon after emergence, obviously 
owing to a colour development around ommatidia during that period. 

3. A difference in the eye pattern between the two sexes was observed 
in two lowland species of Heteronympha Wallengren (Satyridae) in 
New South Wales: H. merope merope Fabricius and H. mirifica Butler, 
in which males showed IIIb or IIIb-g eyes and females typical IlIlg 
eyes. Another example was recognised in Ogyris amaryllis amaryllis 
Hewitson (Lycaenidae) in which males showed IIIg eyes while a single 
female specimen had IIIa eyes. Males of some other species of Ogyris 
Westwood have IIIg eyes like the male of O. amaryllis, but I have not 
yet examined females of these other species. My record includes a 
somewhat doubtful case of Hypolimnas alimena eremita Butler (Nym- 
phalidae): male, Ile; female, I1[d—but this needs confirmation. In 
most of the observations I have made, I have not recorded the sex of 
the observed specimens explicitly, and it is possible that there are sexual 


VOLUME 27, NUMBER 3 167 


dimorphisms among the species recorded without reference to sex in 
this paper. 

4. So far I have not come across any definite proof that subspecies 
or seasonal forms of one and the same species can differ in the reflective 
pattern of the eye. Nor do I know of any evidence for a genetic poly- 
morphism related to this character among butterflies. However, in 
Drosophila melanogaster (Diptera) the somewhat similar RS’s are char- 
acteristic of wild-type eyes and are absent from all the known eye-colour 
mutants. 


A Brief Description of the Eye Pattern in Various Taxonomic 
Groups and Evaluation of its Taxonomic Significance 


Hesperiidae. All the examined species covering Coeliadinae, Pyrginae, 
Trapezitinae and Hesperiinae have similar Type I eyes, so that this 
eye pattern can be regarded as an invariable character of the family. The 
pyrgine genus Chaetocneme Felder has red instead of dark brown or 
black eyes whose colour is stable post mortem. It would be of interest 
to know whether or not the eye of this genus follows the general pattern 
of the family. 

Papilionidae. All the genera so far examined and reported, covering 
Parnassiinae (Zerynthiini and Parnassiini) and Papilioninae (Graphiini, 
Papilionini and Troidini) show Type I eyes, so that this eye pattern 
can be taken as a universal character of the family. 

Pieridae. The family is characterised by the Type HI eye. A remark- 
able exception is the genus Aporia Hiibner having Type I eyes. Another 
interesting point is the appearance of IIIf eyes in Gonepteryx Leach and 
Ixias Hiibner (Yagi & Koyama, 1963). The colour of the pierine eye 
ranges from greenish yellow (Colias Fabricius and Eurema Hubner) 
to bluish white (Prioneris Wallace), via the commonest pale greenish 
or whitish grey of most of the genera including Catopsilia Hubner, 
Pieris Schrank and Delias Hiibner. The major subtypes of the eye are 
IIIa, b and c. | 

Danaidae. This family present certain difficult problems. Eyes ol 
many species are dark but still usually some dark and light patterns are 
discernible. The diagram shown in Fig. 2 IIIh is an interpretation of 
the complicated pattern encountered frequently in this family. Yag! 
& Koyama (1963) noted that certain species of Euploea Fabricius im 
Taiwan had eyes without RS. I also noted the same with several Euploed 
species I collected in New Guinea. However, some specimens of certain 
species in New Guinea or New South Wales had patterns like Ig 
eyes. I am not certain at the moment whether the variability ol the 


eye pattern in this group is at least partly « limorphism 


168 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Distribution of Type I and III eyes among some higher taxa of 
Satyridae. 


Taxa Type I Type III 
BIINAE Melanitis 
ELYMNIINAE 
Lethini 
Lethe-Series Lethe europa, L. chandica, Lethe marginalis, L. diana, L. 
Ninguta, Neope sicelis, Harima 
Pararge-Series Lopinga Lasiommata, Kirinia 
Mycalesini Mycalesis, Orsotriaena 
SATYRINAE 
Hypocystini 
Hypocysta-Series | Hypocysta Harsiesis, Platypthima, 
Pieridopsis 
Tisiphone-Series* Argynnina, Oreixenica, Geitoneura, Heteronympha 
Heteronympha penelope, merope, H. mirifica 


H. paradelpha, H. banksii, 
H. cordace, Tisiphone 


Coenonymphini Coenonympha 
Erebiini Erebia 
Satyrini 
Satyrus-Series Minois 
Oeneis-Series Oeneis 


* Miller (1968) called this the Xenica-Series, but Xenica Westwood, 1851, is an objective 
synonym of Tisiphone Hiibner, 1816-1826 (Hemming, 1967). 


Working in the field it was not always possible to compare live males 
and females of the same species. Also the specific identification of in- 
dividual specimens of Euploea in the field posed some problems in New 
Guinea. Obviously much more careful and extensive observations on 
many individuals are needed before something more definite can be 
stated about the eye of this difficult family. 

Ithomiidae. The only known example (Tellervo Kirby) has sub- 
type IIIb eyes with a vivid yellow ground colour. The eye of this family 
therefore seems to be at variance with that of Danaidae or Satyridae. 

Amathusiidae. The only known example (Taenaris Hibner) showed 
Type I eyes. 

Satyridae. The majority of the species of this family have eyes of 
Type IlIg with a grey or grey-brown, but sometimes yellow, ground 
colour, but there occur a number of exceptions. A glance at Table 2 
reveals that the difference is not apparently related to any taxonomic 
groupings, because both Type I and III eyes are found side by side in 
many recognised higher taxa, and even within a single genus, almost 
throughout the family. However, in most cases the eye pattern may be 
regarded as a good character of individual genera, especially in Hy- 
pocystini. 


VoLUME 27, NUMBER 3 169 


Also noteworthy is a consistent difference between the two sexes as 
exemplified by the two lowland species of Heteronymphe (sce above). 
One theory might be that the difference in the eye pattern is more adap- 
tive than phylogenetic, but nothing is known about the possible cor- 
relation of certain eye types with the habitat or some other ecological 
or behavioral variables. 

Nymphalidae. Most species of this family have eyes of Type III, 
but there is an odd example of Type I (Clossiana thore jezoensis Mat- 
sumura). I have a record of a live female of Hypolimnas deois divina 
Fruhstorfer having Type I eyes, while another record of mine indicates 
that the same species (male ?) had subtype IIIb eyes. Some species 
(Dichorragia nesimachus nesiotes Fruhstorfer and Hestina assimilis 
formosana Moore) are assigned to Type II by Yagi & Koyama (1963). 
Generally, the subtype IIIc predominates, but subtype [Id eyes are 
widely seen among members of Nymphalinae (s.str.). The subtype IIa 
appears in Apaturinae. Many species have brightly coloured eyes: 
yellow or orange predominates in Argynninae; grey or brown in Nym- 
phalinae; bluish grey in Limenitinae; light orange or yellow-brown in 
Apaturinae; green in some species of Cyrestis Boisduval, and bluish black 
in Dichorragia Butler. In Charaxinae, Charaxes latona papuensis Butler 
has bright orange eyes, while Polyura pyrrhus sempronius Fabricius 
shows Type IIIb eyes which look almost as dark as Type I eyes. 

Aecraeidae. As far as known, the eye belongs to Type I. Perhaps this 
character may justify the family status of this group. 

Libytheidae. The only known case is the subtype IIIf. 

Lycaenidae. As noted by Yagi & Koyama (1963), the majority of the 
species have Type I eyes. However, there are some genera with Type 
III eyes, which are for some unknown reason frequently met with in the 
Australian region. This latter type is found in various subfamilies, but 
its occurrence generally serves as a criterion at least for a genus, some- 
times for even a higher taxon. 

Thus, in Polyommatinae, Zizula Chapman, Zizina Chapman and 
Zizeeria Chapman (including the very anomalous Z. alsulus Herrich- 
Schaeffer), which are somewhat remotely related, have yellowish grey 
eyes of the subtype IIIa—b. Theclinesthes Rober and Catopyrops 
Toxopeus stand out with similar yellowish grey eyes among the genera 
related to Nacaduba Moore. Two species currently placed in Neolucia 


Waterhouse and Tumer: serpentata Herrich-Schaeffer and sulpitius 
Miskin also have eyes similar to those of Theclinesthes, while Neolucia 
agricola Westwood and N. mathewi Miskin have Type | eyes. | have 
found that other features such as male genitalia and antennae of ser- 


pentata and sulpitius showed a very strong affinity to those ol 


170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Theclinesthes and were unlike any other groups of Lycaenidae I know of, 
let alone Neolucia agricola and N. mathewi. Obviously there is a need 
of generic rearrangement for some species hitherto placed in Theclinesthes 
and Neolucia, and the eye pattern is consistent with many other characters 
with respect to this point. Among genera allied to Candalides Hubner 
with Type I eyes, Erina Swainson is peculiar with Type III eyes, which 
seems to point to a good generic status of Erina. 

In Theclinae, Hypochrysops Felder & Felder and Philiris Rober have 
yellow eyes belonging to subtype Illa or b-g. Contrary to previous 
treatments by authors as recent as Common (1964), Tite (1963) or 
D’Abrera (1971), the two genera are taxonomically closely related as 
is seen from the wing shape, antennae and male genitalia. Eliot (pers. 
comm.) has reached the same conclusion independently. Paralucia 
Waterhouse & Turner and Pseudodipsas Felder & Felder have grey sub- 
type IIa or b-c eyes and seem to stand fairly close to the preceding two 
genera. More peculiar is Ogyris Westwood, males having characteristic 
Type IlIg eyes with a grey-brown ground colour, like those of many 
satyrid species, and the only female ever examined having subtype 
IIla eyes. Two species of Hypochlorosis Rober in New Guinea had 
yellowish Type III eyes. According to a photograph shown by Kohiyama 
et al. (1971), Spindasis takanonis Matsumura may have a brown Type 
III eye, but this is not certain. 

Curetinae seems to have disparate eye types in the single genus 
Curetis Hiibner. It would be of interest to see whether or not there 
occur some clusters of genera having Type III eyes in areas other than 
the Australian region. 


Assignments of Various Species of Butterflies to Individual 
Types and Subtypes of the Compound Eye 


Assignments taken from Yagi & Koyama (1963) are marked with 
an asterisk, and those inferred from the figures of Kohiyama et al. (1971) 
with two asterisks. 


SPECIES WITH TYPE I EYES 


Hesperiidae. Chaospes benjamini japonica Murray*; Bibasis aquilina chrysaeglia 
Butler*; Erynnis montanus Bremer*; Daimio tethys Ménétriés; Trapezites eliena 
éliena Hewitson, T. iacchoides Waterhouse, T. phigalia phigalia Hewitson; Toxidia 
peron Latreille; Signeta flammeata Butler; Mesodina halyzia halyzia Hewitson; 
Leptalina unicolor Bremer & Grey*; Aeromachus inachus Ménétriés*; Isoteinon 
lamprospilus formosanus Fruhstorfer*; Notocrypta curvifascia C. & R. Felder*, N. 
waigensis waigensis Plotz; Thymelicus leoninus Butler*, T. sylvaticus Bremer*; 
Ochlodes venata herculea Butler*, O. ochracea rikuchina Bulter*; Hesperia florinda 
Butler*; Potanthus flavus Murray*; Telicota eurotas eurychlora Lower; Cephrenes 
augiades sperthias Felder; Polytremis pellucida Murray*; Pelopidas jansonis Butler*, 
P. mathias oberthiiri Evans*; Parnara guttata Bremer & Grey*. 


VoLUME 27, NUMBER 3 lyal 


Papilionidae. Parnassius eversmanni daisetsuzanus Matsumura*, P. stubbendorfii 
hoenei Schweitzer*, P. glacialis Butler*: Luehdorfia puziloi inexpecta Sheljuzhko*, 
L. japonica Leech*; Pachliopta aristolochiae interpositus Fruhstorfer*; Troides 
aeacus kaguya Nakahara & Esaki*; Ornithoptera priamus richmondits Gray; Byasa 
febanus Fruhstorfer*, B. polyeuctes termessus Fruhstorfer*, B. alcinous Klug*; 
Papilio machaon hippocrates C & R. Felder*, P. xuthus Linnaeus*, P. macilentus 
Janson*, P. protenor amaura Jordan*, P. protenor demetrius Cramer*, P. memnon 
heromus Fruhstorfer*, P. castor formosanus Fruhstorfer*, P. nephelus chaonulus 
Fruhstorfer*, P. aegeus aegeus Donovan, P. aegeus ormenus Guérin, P. ambrax 
ambrax Boisduval, P. euchenor euchenor Guérin, P. helenus fortunius Fruhstorfer*, 
P. thaiwanus Rothschild*, P. polytes pasikrates Fruhstorfer*, P. bianor takasago 
Nakahara & Esaki*, P. bianor dehaani C. & R. Felder*, P. ulysses autolycus Felder, 
P. laglaizei Depuiset, P. anactus Macleay; Princips demoleus libanius F ruhstorfer*, P. 
demoleus sthenelus Macleay; Chilasa agestor matsumurae Fruhstorfer*: Graphium 
cloanthus kuge Fruhstorfer*, G. sarpedon nipponum Fruhstorfer*, G. sarpedon con- 
nectens Fruhstorfer*, G. sarpedon choredon Felder, G. doson positanus Fruhstorfer*, G. 
eurypylus lycaonides Rothschild, G. agamemnon ligatus Rothschild, G. wallacei 
wallacei Hewitson, G. macleayanus macleayanus Leach, G. weiskei Ribbe. 

Pieridae. Aporia hippia japonica Matsumura*, A. crataegi adherbal Fruhstorfer*. 

Danaidae. Idea leuconoé clara Butler*; Euploea leucostictos hobsoni Butler’, 
E. sylvestor swinhoei Wallace*, E. tulliolus koxinga Fruhstorfer*. 

Amaihusiidae. Taenaris myops kirschii Staudinger. 

Satyridae. Lethe chandica rathnacri Fruhstorfer*, L. europa pavida Fruhstorfer*; 
Ninguta schrenckii menalcas Fruhstorfer*; Neope goschkevitschii Ménétriés (or 
N. niphonica Butle:?)*; Lopinga achine achinoides Butler*; Hypocysta adiante 
adiante Hubner, H. pseudirius Butler, H. metirius Butler, H. aroa aroa Bethune- 
Baker (?, from Madang, New Guinea), H. euphemia Westwood; Argynnina cyrila 
Waterhouse & Lyell, A. tasmanica Lyell; Heteronympha paradelpha paradelpha 
Lower, H. penelope penelope Waterhouse, H. banksii banksii Leach, H. cordace 
cordace Geyer; Oreixenica lathoniella herceus Waterhouse & Lyell; Tisiphone abeona 
abeona Donovan; Erebia niphonica Janson*, E. ligea takanonis Matsumura*; Oeneis 
daisetsuzana Matsumura*. 

Nymphalidae. Clossiana thore jezoensis Matsumura*; Hypolimnas deois divina 
Fruhstorfer @. 

Acraeidae. Acraea issoria formosana Fruhstorfer*; Miyana meyeri Kirsch. 

Lycaenidae. Taraka hamada Druce*; Curetis brunnea Wileman*; Arthopoétes 
pryeri Murray*; Ussuriana stygiana Butler*; Japnoica lutea Hewitson*, J. saepestriata 
Hewitson*; Shirozua jonasi Janson*; Antigius attilia Bremer*, A. butleri Fenton*; 
Araragi enthea Janson*; Ravenna nivea Nire*; Wagimo signatus quercivorus 
Staudinger*; Iratsume orsedice Butler*; Favonius saphirinus Staudinger*, F. orientalis 
Murray*, F. yuasai Shirozu*, F. cognaius Staudinger*, F. jezoensis Matsumura*, 
F. ultramarinus hayashii Shirozu*; Neozephyrus taxila japonicus Murray*; Chryso- 
zephyrus smaragdinus Bremer*, C. aurorinus Oberthiir (Koyama, pers. comm. ), 
C. ataxus kirishimaensis Okajima*; Narathura japonica Murray*, N. bazalus turbata 
Butler*, N. alkisthenes Fruhstorfer, N. micale centra Evans; Jalmenus ictinus Hewit- 
son; Hypolycaena phorbas phorbas Fruhstorfer, H. danis deripha Hewitson; Deudorix 
epijarbas diovis Hewitson; Rapala varuna simsoni Miskin, R. varuna formosana 
Fruhstorfer*, R. arata Bremer*; Strymonidia w-album fentoni Butler®, ». mera 
Janson*; Ahlbergia ferrea Butler*; Lycaena phlaeas daimio Seitz; Heliophorus ila 
matsumurae Fruhstorfer*; Anthene lycaenoides Felder, subsp. (New Guinea); 
Holochila neuropacuna Bethune-Baker, H. ardosiacea Tite, H. absimilis Felder, H. 
consimilis Waterhouse; Cyprotides cyprotus Olliff; Candalides xanthospilos Hubner; 
Microscene heathi heathi Cox; Niphanda fusca shijimia Fruhstorfer*; Prosotas dubiosa 


ee ee, » Byenhctorfer® ) 
dubiosa Semper, P. nora nora Felder, P. nora formos ihstorfer*, P. papuana 


172 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Tite, P. felderi Murray; Nacaduba biocellata biocellata Felder, N. ruficirca Tite, 
N. kurava parma Waterhouse & Lyell, N. berenice Herrich-Schaffer; Paraduba 
owgarra Bethune-Baker, P. metriodes Bethune-Baker; Ionolyce helicon Felder, subsp. 
(New Guinea); Erysichton lineata meiranganus Rober; Neolucia agricola agricola 
Westwood, N. mathewi Miskin; “Castalius’ mindarus mindarus Felder; Pistoria 
nigropunctatus nigropunctatus Bethune-Baker; Callictita cyara Bethune-Baker; Danis 
hymetus taygetus Felder, D. hymetus hymetus Felder, D. albula Grose-Smith, D. 
hebes Druce; Pepliophorus inops piluma Druce; Jamides purpurata purpurata Grose- 
Smith, J. aratus Cramer, subsp. (?), J. nemophila electus Grose-Smith, J. croitus 
pseudeuchylas Strand, J. aleuas nitidus Tite, J. celeno suwndana Fruhstorfer; Cato- 
chrysops panormus papuana Tite, C. amasea Waterhouse & Lyell; Lampides boeticus 
Linnaeus; Syntarucus plinius pseudocassius Murray; Everes argiades hellotia 
Ménétriés*, E. lacturnus Hiibner, subsp. (New Guinea); Tongeia fischeri Evers- 
mann*; Pithecops nihana urai Bethune-Baker*; Celastrina argiolus ladonides de 
YOrza*, C. sugitanii Matsumura*, C. limbata himlicon Fruhstorfer*, C. albocaerulea 
sauteri Fruhstorfer*, C. philippina nedda Grose-Smith, C. argioloides Rothschild, 
C. drucei drucei Bethune-Baker, C. leucothalia Jordan, C. meeki meeki Bethune- 
Baker; C. owgarra Bethune-Baker; Udara rona Grose-Smith, U. dardia owgarra 
Bethune-Baker; Ewpsychellus dionisius Boisduval; Maculinea teleius kazamoto Druce*; 
Sinia divina barine Leech*; Euchrysops cnejus cnidus Waterhouse & Lyell; Plebejus 
argus micrargus Butler*; Lycaeides argyrognomon ~praeterinsularis Verity*, L. 
subsolana yagina Strand*, L. subsolana yarigatakeana Matsumura”. 


SPECIES WITH TYPE II EYES 


Nymphalidae. Dichorragia nesimachus nesiotes Fruhstorfer*; Hestina assimilis 
formosana Moore”. 


SPECIES WITH TYPE III EYES 


Subtypes are shown in parentheses after each names wherever adequate records 
are available. 

Pieridae. Colias erate poliographus Motschulsky (c)*, C. palaeno aias Fruhstorfer 
(c)*, C. palaeno sugitanii Esaki (c)*; Eurema hecabe mandarina de YOrza (b)*, E. 
hecabe hobsoni Butler (b)*, E. laeta bethesba Janson (b)*, E. esakii Shirozu (b)*, 
E. blanda arsakia Fruhstorfer (b)*, E. smilax Donovan (b), E. candida puella 
Boisduval (b); Catopsilia pomona Fruhstorfer (c)*, C. crocale Cramer (c), C. 
pyranthe pyranthe Linnaeus (c)*, C. pyranthe crokera Macleay (b); Gonepteryx 
mahaguru niphonica Verity (£)*; Appias indra aristoxenus Fruhstorfer (c)*; Delias 
nysa nysa Fabricius (c); Cepora nandina eunama Fruhstorfer (c)*, C. perimele 
latilimbata Butler (c), C. euryxantha Honrath (c), C. abnormis Wallace (c); 
Elodina egnatia angulipennis Lucas (b), E. parthia Hewitson (b), E. padusa 
Hewitson (a); Prioneris thestylis formosana Fruhstorfer (c)*; Ixias pyrene insignis 
Butler (£)*; Anthocharis scolymus Butler (c)*, A. cardamines isshikii Matsumura 
(b)*; Pieris rapae rapae Linnaeus (c), P. rapae curcivora Boisduval (c)*, P. 
canidia canidia Linnaeus (c)*, P. melete Ménétriés (c)*, P. napi nesis Fruhstorfer 
(c)*, P. napi japonica Shirozu (c)*; Leptosia nina niobe Wallace (c)*; Leptidea 
amurensis Ménétries (a)*. 

Danaidae. Danaus plexippus Linnaeus (I-IIIh); Limnas chrysippus petilia Stoll é 
(g-h, e), @ (d or e-h); Tirumala hamata septentrionis Butler (g)*; Radena similis 
similis Linnaeus ( g)*; Parantica aglea maghaba Fruhstorfer (g¢)*, P. melaneus swinhoei 
Moore (g)*, P. sita niphonica Moore (g)*, (h?)**; Euploea althaea juvia Fruhstorfer 
(h?)*, E. mulciber barsine Fruhstorfer (h?), E. core corinna Macleay 2 (g). 

Ithomiidae. Tellervo zoilus hempsal Fruhstorfer (b). 

Satyridae. Penthema formosanus Rothschild (b?); Melanitis leda leda Linnaeus 
(f)*, M. leda destitans Fruhstorfer (g), M. phedima oitensis Matsumura (f)*, M. 


VOLUME 27, NUMBER 3 173 


phedima polishana Fruhstorfer (f)*, M. amabilis Boisduval subsp. (New Guinea) 
(g); Mycalesis gotama fulginia Fruhstorfer (g)*, M. francisca formosana Fruhstorfer 
(g)*, M. terminus kyllenion Fruhstorfer (g); Lethe marginalis Motschulsky (¢)*, 
L. diana Butler (g)*, L. sicelis Hewitson (g)*; Harima callipteris Butler (b-c)*: 
Kirinia epaminondas Staudinger (g)*; Lasiommata deidamia interrupta Fruhstorter 
(g)*; Harsiesis hygea hygea Hewitson (g); Pieridopsis virgo Rothschild & Jordan 
(g); Platypthima homochroa Rothschild & Jordan (g), P. decolor Rothschild & 
Jordan (g); Geitoneura klugi klugi Guérin (b), G. acantha acantha Donovan 2 
(b-g); Heteronympha merope merope Fabricius ¢ (b), 2 (g), H. mirifica Butler 
6 (b-g), 2 (g); Ypthima arctous papuana Fruhstorfer (g), Y. arctous arctous 
Fabricius (g), Y. argus Butler (g)*, Y. yamanakai Sonan (g)*, Y. multistriata 
Butler (g)*, Y. esakii Shirozu (g)*; Coenonympha oedippus annulifer Butler (g)*; 
Minois dryas bipunctatus Motschulsky (g)*. 

Nymphalidae. Cupha prosope oderca Fruhstorfer (b); Mellicta ambigua niphona 
Butler (b)*; Melitaea scotosia Butler*; Clossiana freija asahidakeana Matsumura 
(b-c)*; Brenthis daphne rabdia Butler (c)*, B. ino tigroides Fruhstorfer (c)*; 
Argynnis paphia geisha Hemming (c)*, A. anadyomene midas Butler (c)*; Damora 
sagana liane Fruhstorfer (c)*; Fabriciana adippe pallescens Butler (c)*, F. nerippe 
C. & R. Felder (c)*; Mesoacidalia charlotta fortuna Janson (c)*; Argyronome laodice 
japonica Ménétriés (c)*, A. ruslana lysippe Janson (c); Argyreus hyperbius hyperbius 
Linnaeus (c)*, A. hyperbius inconstans Butler (b); Vindula arsinoe rebeli Fruhstorfer 
(ec); Vagrans egista offaka Fruhstorfer (c-d); Araschnia burejana strigosa Butler 
(b)*, A. levana obscura Fenton (b)*; Polygonia c-aureum Linnaeus (d)*; P. c-album 
hamigera Butler (d)*, P. vau-album samurai Fruhstorfer (d)*; Kaniska canace no- 
japonicum von Siebold (d)*, K. canace drillon Fruhstorfer (d)*; Nymphalis antiopa 
asopos Fruhstorfer (d)*, N. xanthomelas japonica Stichel (d)*; Inachis io geisha 
Stichel (d)*; Aglais connexa Butler (d)*; Cynthia cardui Linnaeus (c)*; Vanessa 
indica Herbst (e)*; Precis almana almana Linnaeus*, P. lemonias lemonias Linnaeus”, 
P. iphita iphita Cramer*, P. erigone tristis Miskin (b); Yoma algina etonia Fruhstorter 
(c); Symbrenthia hippocle formosana Fruhstorfer*, S$. hippocle hippocrates Staud- 
inger; Hypolimnas misippus Linnaeus (RS round and H-shaped)*, H. bolina Lin- 
naeus*, H. bolina nerina Fabricius (e), H. alimena eremita Butler 6 (e), 2 (d), 
H. deois divina Fruhstorfer (b); Doleschallia bisaltide guralca Grose-Smith (c); 
Kallima inachus formosana Fruhstorfer*; Mynes geoffroyi ogulina Fruhstorfer (b); 
Cyrestis achates achates Butler (b), C. thyodamas mabella Fruhstorfer*; Phaedyma 
shepherdi damia Fruhstorfer (b); Neptis praslini meridionalis Talbot (b), N. 
sappho intermedia Pryer (c)*, N. hylas luculenta Fruhstorfer*, N. nata lutatia 
Fruhstorfer*, N. philyra excellens Butler*, N. rivularis insularum Fruhstorter*, N. 
pryeri jucundita Fruhstorfer*, N. alwina kaempferi de YOrza*; Athyma selenophora 
laela Fruhstorfer (b-d)*, A. perius perius Linnaeus*; Ladoga camilla japonica 
Ménétriés (b-d)*, L. glorifica Fruhstorfer (b-d)*; Helcyra chionippe thyiada Fruh- 
storfer (c); Apatura ilia substituta Butler (a-b)*; Hestina japonica C. & R. Felder 
(a-b)*; Sasakia charonda Hewitson (a-b)*; Charaxes latona papuensis Butler (c); 
Polyura pyrrhus sempronius Fabricius (I-IIIb). 

Libytheidae. Libythea celtis celtoides Fruhstorfer (f)*, L. celtis formosana 
Fruhstorfer (f)*. . . 

Lycaenidae. Curetis acuta paracuta de Niceville (a)*; Paralucia aurifer Blanchard 
8 (b-c), P. aenea aenea Miskin 8 (b-c); Pseudodipsas brisbanensis brisbanensis 
Miskin ¢ (b), P. cuprea Sands ¢ (a); Hypochrysops delicia delicia Hewitson 
(b), H. ignita ignita Leach ¢ (a), H. cyane Waterhouse & Lyell (b-g), H. 
epicurus Miskin, H. byzos Boisduval, H. pythias pythias Felder ¢ (b), H. rufinus 
Grose-Smith @ (b), H. polycletus rex Boisduval (b-g); H. argyriorufa van Eecke 
(a), H. chrysargyra Grose-Smith ¢ (b-g); Philiris innotatus Miskin $, P. albi- 
humerata Tite ¢ (a), P. unipunctata Bethune-Baker ¢ ), F. griseldis griseldis 
Staudinger (b), P. moira moira Grose-Smith (b); Ogy ‘va gela Waterhouse 


174 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


6 (g), O. ianthis Waterhouse ¢ (g), O. abrota Westwood ¢ (g), O. amaryllis 
amaryllis Hewitson 6 (g), @ (a); Hypochlorosis lorquini metilia Fruhstorfer (a), 
H. humboldti Druce (a); Spindasis takanonis Matsumura ?**; Erina acasta Cox 
6 (a), E. hyacinthina hyacinthina Semper 6 (a); Catopyrops ancyra mysia Water- 
house & Lyell ¢ (a-b), C. florinda estrella Waterhouse & Lyell @ (a-b); The- 
clinesthes miskini Lucas 6 (a), T. onycha onycha Hewitson 6 (a-b); “Neolucia’ 
sulpitius sulpitius Miskin (a), “N.” serpentata serpentata Herrich-Schaffer (a); 
Zizeeria maha argia Ménétriés (b)*, Z. maha okinawana Matsumura (a)*, Z. knysna 
karsandra Moore (a), “Z.” alsulus alsulus Herrich-Schaffer (a); Zizina otis aruensis 
Swinhoe (a); Zizula hylax dampierensis Rothschild (a). 


SUMMARY 


The observations of Yagi & Koyama (1963) on the superficial reflective 
pattern of fresh compound eyes based on butterflies from Japan and 
Taiwan are extended to those from Papua New Guinea and Australia. 
Three basic patterns of the eye are recognised: Type I, with no reflective 
spots (RS); Type II, with single RS; and Type III, with multiple RS. 
The last one is divided into several subtypes according to the number, 
shape, distribution and modification of RS. A number of higher taxa 
may be well characterised by the eye patterns. Records of assignments 
of individual species to various eye types are compiled. 


ACKNOWLEDGMENTS 


Thanks are due to Prof. N. Koyama, Shinshu University, Ueda, Japan, 
for his permission to utilise some of the original figures and valuable 
suggestions; to Mrs. Dorothy Deger and Mr. Anthony Healy, Sydney, for 
their supply of a photograph and permission to reproduce a part of a 
figure in the book of which Mrs. Deger is a co-author; to Mr. D. P. 
Sands, DASF, Lee, Papua New Guinea, for his critical reading of the 
manuscript; and to Prof. T. Shirozu, Kyushu University, Fukuoka, Japan, 
for drawing my attention to the work by Yagi & Koyama (1963). I am 
also indebted to Lt. Col. J. Eliot, Taunton, England, for information on 
his recently completed work on the higher classification of Lycaenidae 
which is now in press. I also acknowledge the permissions granted by 
the National Parks and Wildlife Service, Government of New South 
Wales, and the Department of Agriculture, Stock and Fisheries, Papua 
New Guinea, to collect specimens used in this work in Ku-ring-gai Chase 
National Park, New South Wales, and in Papua New Guinea. I also 
express my deep appreciation to Messers T. L. Fenner, DASF, Konedobu; 
A. Hutton, DASF, Garaina; R. Straatman, Sogeri; and Dr. J. L. Glessitt 
and his staff at the Bernice P. Bishop Museum, New Guinea Field Station, 
Wau (now Wau Ecology Institute), for their generous help to my col- 
lecting activities in Papua New Guinea. 


VOLUME 27, NUMBER 3 175 


LITERATURE CITED 


Common, I. F. B. 1964. Australian Butterflies. Jacaranda, Bristbane. 132 p. 
D’Asrera, B. 1971. Butterflies of the Australian Region. Landsdowne, Melbourne. 


AON SD. 
Decer, D. & R. Even. 1970. Collecting Australian Butterflies. Horwitz, North 
Sydney. 52 p. 


EueuicH, P. R. 1958. The comparative morphology, phylogeny and higher clas- 
sification of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci. 
Bull. 39: 305-370. 

Fox, R. M. 1956. A monograph of the Ithomiidae (Lepidoptera), Part 1. Bull. 
Amer. Mus. Nat. Hist. 111: 1-76. 

HemMmMinc, F. 1967. The generic names of the butterflies and their type-species 
(Lepidoptera: Rhopalocera). Bull. Brit. Museum (Nat. Hist.): Entomol. 
Suppl, 9: I—510. 

KontyAMA, K., T. TaxasE & T. Fujioka. 1971. Butterflies of Japan. Yama-to- 
Keikcku, Tokyo. 204 p. (in Japanese). 

Minter, L. D. 1968. The higher classification, phylogeny and zoogeography of 
the Satyridae (Lepidoptera). Mem. Amer. Entom. Soc. 24: 1-174. 

Tire, G. E. 1963. A revision of the genus Candalides and allied genera (Lepidop- 
tera: Lycaenidae). Bull. Brit. Museum (Nat. Hist.): Entomol. 14: 199-260. 

Yaci, N. & N. Koyama. 1963. The Compound Eye of Lepidoptera: Approach 
from Organic Evolution. Shinkyo, Tokyo. 320 p. 


STUDIES ON THE CATOCALA (NOCTUIDAE) OF SOUTHERN 
NEW ENGLAND. IV. A PRELIMINARY ANALYSIS OF 
BEAK-DAMAGED SPECIMENS, WITH DISCUSSION OF 

ANOMALY AS A POTENTIAL ANTI-PREDATOR FUNCTION 
OF HINDWING DIVERSITY 


THEODORE D. SARGENT 


Department of Zoology, University of Massachusetts, 
Amherst, Massachusetts 01002 


An intensive study of the biology of the Catocala moths is presently 
being conducted in southem New England (Sargent & Hessel, 1970; 
Kellogg & Sargent, 1972; Sargent, 1972a). During the course of these 
investigations, substantial numbers of beak-damaged specimens have 
been collected. The present study was undertaken in hopes that an 
analysis of such specimens might shed some light on various aspects of 
the predator-prey relationships involving birds and these moths. 


Prior Studies 


Beak-damaged Lepidoptera have attracted considerable attention in 
the literature, though most prior studies have been concerned almost 


176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


exclusively with butterfly examples. A number of interpretations of 
various damage patterns have been advanced, and some of these seem 
generally accepted. Thus, V-shaped tears in the wings, which suggest 
that the specimens involved have torn free from bird attacks, have been 
regarded as evidence of palatability in their bearers. On the other hand, 
disabling injuries (i.e. those sufficient to prevent flight) have been 
regarded as evidence that their bearers are unpalatable or otherwise 
noxious (e.g. Poulton, 1913). Unpalatability has also been advanced as 
an explanation for crisp beak-imprints on the wings, the presumption 
being that specimens exhibiting such marks have been purposely released 
by birds (e.g. Collenette & Talbot, 1928). This last explanation receives 
support from studies showing a relatively high frequency of crisp beak- 
imprints on the wings of aposematic species (e.g. Carpenter, 1941). 
Beak-damage patterns have also provided evidence that the small, eye- 
spot markings on the margins of butterfly wings direct the attacks of 
birds (e.g. Marshall & Poulton, 1902; Swynnerton, 1926; Blest, 1957), 
and so function as deflective devices. 

Although beak-damage patterns in moths have received little attention, 
the prior work with butterflies encouraged the present study and raised 
hopes that some insight could be gained into predator-prey relationships 
involving the Catocala moths. 


Anti-Predator Functions of Catocala Wings 


Catocala moths appear to rely upon the bark-like crypsis of their fore- 
wings as their primary defense against predation. A number of studies 
reveal the extent to which this crypsis is enhanced by means of behavioral 
adaptations, e.g. the selection of appropriate backgrounds (Sargent, 
1966, 1968, 1969a, 1973; Keiper, 1968; Sargent & Keiper, 1969), and the 
adoption of appropriate resting attitudes (Sargent, 1969b). It seems 
likely that forewing variations, particularly the dramatic polymorphisms 
exhibited by many species, function to augment the effectiveness of 
crypsis by foiling predator tendencies to form “specific searching images” 
(Tinbergen, 1960) for particular cryptic patterns (Sargent, 1972b). 

Hindwing function in Catocala moths has not been extensively studied. 
It is generally assumed that these boldly patterned, and often colorful, 
structures are examples of “flash coloration,’ being revealed when 
crypsis fails to deter attack and a moth takes flight, only to be concealed 
when the moth again alights. Such “flash and cover” sequences are 
presumed to confuse predators as to the whereabouts of the moths 
(Cott, 1940; Ford, 1967). 

It is sometimes assumed that the flash of the hindwings is itself the 
important anti-predator device, functioning to startle predators and 


VoLUME 27, NUMBER 3 177 


thereby adversely affecting the efficiency of their attacks. In this event, 
Catocala hindwings would have a function similar to that suggested for 
the large eye-spots of saturniid moths (Blest, 1957: Coppinger, 1970). 

Catocala hindwings might also serve a deflective function. In this 
case, predators would direct their attacks toward these prominent 
structures, thereby being directed away from more vulnerable body parts. 
Catocala hindwing patterns would then be functionally related to the 
small eye-spots found along the margins of some butterfly wings, which 
are known to direct predator attacks (Poulton, 1890; Blest, 1957). 

The interspecific diversity found in hindwing colors and patterns in 
the genus Catocala has also been suggested as an anti-predator adapta- 
tion (Sargent, 1969c). In this view, hindwing variation introduces the 
potential of novelty (unfamiliar stimuli) or anomaly (unexpected stimuli ) 
as a startle mechanism into predator-prey relationships involving birds 
and several Catocala species. For example, a bird, after several suc- 
cessive encounters with species possessing a particular hindwing pattem 
(e.g. yellow and black bands), might become habituated to that pattern; 
but this same bird, upon encountering a distinctly different hindwing 
pattern (e.g. pink and black bands), might be effectively startled. If 
the level of startle exhibited by a bird to a particular hindwing pattern 
were a function of the extent and recency of its experience with that 
pattern, then considerable advantage might accrue to an assemblage of 
rather similar cryptic species which evolved a variety of hindwing pat- 
terns (see discussion of schizomimicry; Sargent, 1969c). 

While it appears likely that Catocala hindwings serve one or more of 
the preceding anti-predator functions, it is also quite possible that they 
play an important role in other aspects of the biology of these moths. 
The rather surprising lack of intraspecific hindwing variation in the 
genus suggests that these structures may function as releasers and anti- 
hybridization devices in courtship and mating behaviors. Little is known 
of these behaviors, but in one species, C. relicta Walker, mating can oc- 
cur under conditions of essentially complete darkness (Sargent, 1972a). 
However, until much more is known of Catocala courtship and mating 
behaviors, the question of a sexual role for the hindwings must remain 
open. 


METHODS 
The primary aims of the present study were to describe and interpret 
. R - : Ve . ] 
the beak-damage patterns found on the wings of wild-caught Catocata 
moths. In order to achieve these aims, two procedures were followed: 
(1) recording the behavioral interactions between captive birds and 


Catocala moths in aviaries, and collecting t! ) which escaped 


178 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


under these circumstances for later analysis of their beak-damage pat- 
terns; and (2) collecting all Catocala specimens showing suspected beak- 
damage from a large field sample being obtained in connection with 
other studies. 


Aviary Study 


The Catocala used in this study were 50 fresh, undamaged specimens 
which were taken in a Robinson trap in Leverett, Massachusetts during 
July and August of 1971. These moths were released to birds in two 
aviaries (each 8 X 5 X 10 ft.) which were located on the roof of the 
Morrill Science Center on the University of Massachusetts (Amherst ) 
campus. The aviaries were lined with fine-mesh hardware cloth which 
prevented escape of the moths. 

Each aviary housed seven blue jays (Cyanocitta cristata) which were 
1-2 years old, and which had been hand-reared from approximately 14 
days of age by Dr. Alan C. Kamil and his graduate students in the 
Psychology department. These birds had been tested in discrimination 
and learning-set experiments (e.g. Hunter & Kamil, 1971), but their prior 
experience with insects was limited to the mealworms (Tenebrio larvae ) 
which were used as reinforcements in these experiments, and to various 
flies, etc. which frequented their aviaries. The birds were normally 
maintained on a free-feeding schedule of food (mynah pellets) and 
water, but prior to the release of Catocala into the aviaries, the food was 
withdrawn for a period of 16-24 hours. Under these circumstances, 
the moths were quickly attacked by the birds. 

The observations of moth-bird interactions included careful attention 
to such matters as: (1) whether a moth was attacked while flying or 
resting; (2) if attacked while resting, the direction from which attack 
occurred (rear, side, etc.); and (3) if escaping from attack, the duration 
of retention in the bird’s beak (instantaneous, several seconds, etc.). 

Of the 50 Catocala specimens released to the birds, 23 were recovered 
for analysis of their beak-damage patterns. Every effort was made to 
recover each moth which escaped its initial attack, but some specimens 
suffered second or third attacks before they could be captured. The birds 
became progressively more efficient at taking the moths, losing 10 of 
the first 12 released, but successfully capturing each of the last 15 speci- 
mens. Moths that were eaten by the birds were initially ingested wings 
and all, but after some experience the birds invariably attempted to 
remove the wings by holding the moths in their feet and pecking with 
their beaks. None of the birds developed the clean, efficient shearing-off 
of the wings, without use of the feet, that I have witnessed in blue jays 
in the field. 


VoLUME 27, NUMBER 3 179 


Field Sample 


A total of 2047 Catocala specimens of 33 species were recorded at 
lights (incandescent, black light fluorescent, and mercury vapor) and 
bait (brown sugar—beer mixture) in Leverett, Massachusetts during the 
summer of 1971. Of this total, 1623 specimens were carefully examined 
in glass jars (primarily for sexing, by means of the frenulum), and if 
suspected beak-damage was discovered during this examination, the 
specimen was retained for later comparison with specimens obtained 
in the aviary study. Of the 75 wild-caught specimens that were retained, 
65 were later judged to exhibit clear beak-damage, with the remainder 
exhibiting bat-inflicted damage patterns. . 


RESULTS 


The behavioral observations in the aviaries, coupled with subsequent 
analyses of specimens, permitted a classification of beak-damage patterns, 
as follows: 


Type I. Attack: while moth in flight 
_ Characteristic damage: unilateral; 
tear from one wing only 
a. hindwing tear (Figs. la; 4a,b,c; 7a) 
b. forewing tear (Figs. 1b; 4d) 


Type II. Attack: while moth resting; grasp of 
beak not including costa of either forewing 
Characteristic damage: corresponding 
tears from ipsilateral forewing and hindwing 
a. forewing and hindwing tears 
overlapping when wings fully 
closed; unilateral (Figs. 2a; 5a,b) 
b. forewing and hindwing tears over- 
lapping when wings partially 
closed; unilateral (Figs. 2b; 5c; 7b) 
ce. forewing and hindwing tears over- 
lapping on both sides when 
wings fully closed; bilateral (Figs. 2c; 5d) 


Type III. Attack: while moth resting; grasp of beak 
including costa of one forewing 
Characteristic damage: crisp beak-imprint 
on one forewing; tear from ipsilatera! 
hindwing 


180 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


a. apex of beak-imprint directed 

toward (but not across) inner mar- 

gin of forewing; unilateral (Figs. 3a; 6a,b; 7a,b ) 
b. apex of beak-imprint directed 

toward (and across) costal margin 

of forewing; unilateral or bilateral (Figs. 3b; 6c) 
e. apex of beak-imprint directed 

toward (and across) inner margin 

of forewing; bilateral (Figs. 3c; 6d) 


This classification of beak-damage patterns, based on specimens obtained 
in the aviary study, also proved adequate for classifying the field sample 
specimens. The number of examples of each damage pattern obtained 
in both the aviary and field samples is given in Table 1. Note that a 
number of specimens (6 in the aviary sample, 8 in the field sample) 
exhibited two damage patterns (e.g. Fig. 7). 

Most of the field specimens were easily assigned to categories within 
the preceding classification, but occasional difficulties were encountered. 
For example, three specimens were apparently grasped from the rear 
while resting such that the apex of the bird’s beak almost, but not quite, 
reached the forewing costa. The resulting damage, which technically 
had to be classified as Type Ila, included at least a portion of a crisp 
beak-imprint, and this is a characteristic of Type II] damage. Another 
problem was posed by four specimens which exhibited smeared, rather 
than crisp, beak-imprints. However, since these specimens otherwise 
met the criteria of Type III damage, they were so classified. 

The only damage pattern which was not obtained in the field sample 


Taste 1. Distribution of damage patterns among Catocala in the aviary and 
field samples. 


Number of Individual Examples 


Damage Patterns Aviary Sample Field Sample 


12 33 
1 
16 
3 


Type I 


Type II 


Type III 13 


qaT7TrnMna 729 Of 
ed Oe WM Ole 
bo 


Totals 29 73 


VOLUME 27, NuMBER 3 18] 


i: 


b 


Fig. 1. Type I damage patterns. Diagrammatic representations of bird attacks 
on flying moths (left), and the resulting specimens (right). Damage is usually 
confined to one hindwing (a), rarely to one forewing (b). 


was Type IIIc. The attack which leads to this damage pattern (Fig. 3c) 
was the one that blue jays in the aviaries used almost exclusively after 
they had had some experience with Catocala. A moth grasped in this 
fashion rarely escaped, as its wings were securely held in place. How- 
ever, on one occasion a bird apparently loosened its grip when attempting 
to transfer a moth from its beak to its feet, and the escaped specimen 
was recovered (Fig. 6d). Presumably such “carelessness’ would be very 
rare in nature. 


182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Cc 


Fig. 2. Type Il damage patterns. Diagrammatic representations of bird attacks 
on resting moths (left), and the resulting specimens (right). Damage may be 
inflicted when the wings are fully closed (a) or partially spread (b), and while 
usually unilateral, may be bilateral (c) on occasion. 


VoLUME 27, NUMBER 3 183 


TT. 


representations of bird attacks 
t). The beak-mark is 


usually confined to one forewing, including an imprint o! he apex (a) or not (b), 


but may extend across both forewings (c). 


Fig. 3. Type III damage patterns. Diagrammatic 
on resting moths (left), and the resulting specimens (righ 


184 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 


tm! DD 


Fig. 4. Specimens exhibiting Type I damage patterns: A. C. antinympha, Ia 
(8 August 1970, Leverett, Mass., Robinson trap); B. C. unijuga, Ia (30 August 
1971, Leverett, Mass., bait); €C. C. palaeogama, Ia (28 August 1971, Leverett, 
Mass., Robinson trap); D. C. residua, Ib (8 July 1972, Fontana, No. Carolina, 
black light—specimen courtesy D. F. Schweitzer). 


As mentioned previously, a number of specimens which were initially 
retained in the field sample were later judged to be bat-damaged rather 
than bird-damaged. On one occasion a moth was observed to escape a 
bat attack near the Robinson trap, and this specimen was immediately 
recovered (Fig. 8b). Other specimens exhibiting the same distinctively 
frazzled or tattered outer wing margins (e.g. Fig. 8a) were presumed 
to be bat-damaged as well. 


TABLE 2. Distribution of damage patterns among field-caught Catocala of two 
hindwing types. 


Number of Individual Examples 


Damage Patterns 


Saat Total 
Hindwing Types I II III Sample 
Chromatic 28 14 10 1228 
Achromatic 6 7 8 395 
Percent Achromatic 17.6 33.3 44.4 24.3 


Note: The achromatic hindwing totals include one specimen of C. relicta (banded, as 
opposed to uniformly black, upper hindwing surface) under damage patterns I and III, and 
there were 28 specimens of this species in the total sample. 


VOLUME 27, NUMBER 3 185 


tm D 


Fig. 5. Specimens exhibiting Type II damage patterns: A. C. ilia, Ila (5 
August 1971, bait); B. C. epione, Ila (10 August 1971, black light); C. C. flebilis, 
IIb (4 September 1971, Robinson trap); D. C. retecta, IIc (12 September 1971, 
Robinson trap). All specimens from Leverett, Mass. 


It seemed of some interest to ascertain whether the various types of 
beak-damage were distributed in the same way among moths having 
very different types of hindwings. Accordingly, the major beak-damage 
patterns were tabulated separately for moths having chromatic and 
achromatic hindwings (Table 2). (Chromatic hindwings include those 
characterized by color, this color generally providing a ground for 
prominent black bands on both the upper and lower wing surfaces. 
Achromatic hindwings include those characterized by the absence of 
color, with the upper wing surface generally solid black, and the lower 
wing surface banded with black and white. Both hindwing types may 
have a more-or-less prominent white fringe. ) 

Analysis of the data in Table 2 revealed that significantly more speci- 
mens with achromatic hindwings were found with Types II and Il] 
damage (escaping attacks while resting) than were found with Type 
I damage (escaping attacks while flying), or than were present in the 
total field sample (chi-square 2 x 2 contingency tests; P’s < 0.09). 


DiIscussION 


In the present sample of wild-caught Catocala moths, 4% of the in- 


dividuals exhibited clear evidence of at leas! bird attack. These 
collected moths had successfully escaped their attacks, but it may be 


186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


‘ont DS 


Fig. 6. Specimens exhibiting Type III damage patterns: A. C. ultronia, [ila (ex- 
perimental moth 2); B. C. retecta, Illa (experimental moth 6); C. C. ultronia, 
IIIb (11 August 1970, Robinson trap); D. C. concumbens, IIc (experimental moth 
10). All specimens from Leverett, Mass. 


assumed that many other individuals were not so successful. Thus it 
appears that avian predation on Catocala moths is substantial. In such 
a circumstance, one would expect the prey to have some highly evolved 
defensive strategies. The bark-like crypsis of Catocala forewings, coupled 
with appropriate behaviors, seems a clear example of such a strategy. 
The anti-predator functions of Catocala hindwings have not been clearly 
established, but an analysis of the present beak-damaged specimens 
provides evidence for both deflective and startle functions. 

Nearly half of the beak-damaged specimens were apparently attacked 
while in flight, and all but one of these exhibited only hindwing damage. 
It appears that bird attacks are frequently directed towards these 
structures when Catocala moths are flying, and that such attacks may 
result in damage which is not highly detrimental to the moths (some 
specimens have been taken at lights which are missing virtually all of 
one hindwing). Thus Catocala hindwings apparently function on some 
occasions in a fashion similar to the colorful, but expendable, tails of 
certain lizards (Cott, 1940). 

The remaining half of the beak-damaged specimens were apparently 
attacked while resting. Escape from these attacks seemed to be of two 
sorts: (1) a pulling-free of the moth while being tightly gripped by a 
bird, resulting in a tearing of the wings around the region of beak con- 


VOLUME 27, NUMBER 3 187 


A eae | 6B 


Fig. 7. Specimens exhibiting two damage patterns: A. C. ultronia, Ua on the 
left side, Ia on the right side (11 August 1971, black light); B. C. wltronia, IIb on 
the left side, IIIa on the right side (19 August 1971, Robinson trap). Both specimens 
from Leverett, Mass. 


tact (Type II damage); and (2), a release of the moth ajter being 
tightly gripped by a bird, resulting in a clear beak-imprint along some 
of the lines of beak contact (Type III damage). Escapes of the first 
sort apparently occurred when the size, speed, and strength of the moth 
enabled it to break free from the grip of a bird. It is postulated that 
escapes of the second sort were at least in part a result of startle responses 
of the predators to the sudden appearance of the contralateral hindwing 
(i.e. opposite the side being gripped). The startle response is viewed 
as effecting a momentary relaxation of a bird’s grip, this relaxation 
enabling the moth to make its escape, and leaving a crisp beak-imprint 
on some part of its wings. (It seems likely that some of the specimens 
exhibiting Type II damage could have startled their predators, but 
that tears normally occurred in the wings, rather than beak-imprints, 
because a bird’s grip in such cases included only the more fragile por- 
tions of the forewing. Thus, the normal swift escape reactions of the 
moths might have resulted in significant tearing of the wings before 


A gd Fae 


Fig. 8. Specimens exhibiting bat-inflicted damage: A. C. ili@ (5 August L971] 


bait); B. C. habilis (18 September 1971, Robinson trap). Both specimens 
Leverett, Mass. 


188 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the birds relaxed their grip, with further tearing perhaps resulting from 
subsequent wear on the wings. ) 

There appears to be a contradiction between the functions of deflection 
and startle for the same structures with respect to the same predators. 
However, the resolution of this apparent contradiction may lie in an 
understanding of the nature of startle in this situation. It is suggested 
that the degree of startle exhibited by a bird in any encounter with 
a Catocala hindwing pattern is not so closely related to that hindwing 
pattern per se, as it is to the degree of expectation of that pattern which 
the bird brings to the encounter on the basis of its past experience. Thus, 
anomaly, defined in terms of departure from expectation, is regarded 
as the critical factor in determining whether a bird will react by releasing 
a Catocala moth which suddenly displays a particular hindwing pattern. 
This instantaneous reaction might not interfere in any substantial way 
with the ability of a bird to attack a flying moth, when time to adjust 
to the appearance of the hindwings might be available. 

An “anomalous stimulus” (defined in terms of its departure from ex- 
pectation and the momentary startle it elicits) would be clearly distin- 
guishable from a “frightening stimulus” (which is usually defined in terms 
of its absolute properties and the innate rejection it elicits), and could 
often be distinguished from a “novel stimulus” (which is usually defined 
in terms of its unfamiliarity and the avoidance it elicits) (see Brower, 
1971). Anomaly and novelty are obviously closely related, and although 
an anomalous stimulus need not be a novel stimulus (as these are here 
defined), the two phenomena may produce similar physiological effects 
in predators (e.g. high arousal (Coppinger, 1970), or an “orienting 
reflex” (Sokolov, 1960) ). 

The primary evidence that Catocala hindwings function as anomalous 
stimuli is provided by the specimens which exhibit crisp beak-imprints 
on their wings (Type III damage). Such beak-marks on lepidopteran 
wings have long been regarded as evidence that the specimens involved 
were captured and subsequently released by birds. Since beak-marks 
of this sort are most often found on aposematic species, it has been 
assumed that release of the specimens followed predator recognition 
of some noxious quality (usually odor or taste) of the captured prey. 
However, Catocala moths, as far as is known, are entirely palatable 
(many species are readily eaten by cage-birds (pers. obs.), and the 
work of Jones (1932) supports the assumption of their palatability). 
Why should these moths be released after capture by birds? 

The answer to this question may be related to some sort of startle 
response on the part of predators, and it is suggested that interspecific 
hindwing variation provides the key to understanding the situation. This 


VOLUME 27, NUMBER 3 189 


hindwing variation introduces the potential of anomalous stimuli into 
predator-prey relationships involving these moths. Thus. a predator 
is seen as building up expectations regarding future hindwing patterns 
on the basis of its past experiences with these patterns, reacting more 
and more efficiently if these expectations are met, but inefficiently if 
they are countered. Inefficiency presumably results from some involun- 
tary response (e.g. gaping) which is elicited by an unexpected stimulus, 
and which interferes with the completion of normal attack, allowing a 
moth to escape. 

Some indirect evidence for this view is provided by an analysis ot 
the distribution of beak-damage patterns on moths having different 
hindwing types. The most obvious discontinuity among Catocala hind- 
wings occurs between the chromatic and achromatic patterns. As in- 
dividuals possessing chromatic hindwings are more common than those 
possessing achromatic hindwings (the latter comprising less than 25% 
of the Catocala taken in Leverett each year), it might be predicted that 
achromatic hindwings would be less often encountered, and therefore 
more often anomalous, than chromatic hindwings. Analysis of the data 
in Table 2 reveals that beak-damage patterns II and III are more com- 
monly found on individuals possessing achromatic hindwings than would 
be expected on the basis of chance. This finding suggests that achromatic 
hindwings are particularly effective as startle devices, and anomaly may 
provide an explanation for this effectiveness. 

Predators exhibiting a tendency to react inefficiently to anomalous 
stimuli would provide strong selection pressures favoring diversity in 
their prey. Diversity would result in an increased number of potential 
predator expectations, and a corresponding increased number of po- 
tential anomalous stimuli. Anomaly, within the limits of the advantage 
it provides, would then favor the origin and maintenance of considerable 
diversity in sympatric assemblages of closely related species. Hindwing 
diversity among moths in the genus Catocala may represent a response 
to pressures of this sort. . 

Whenever one suggests that selective advantages result from inter- 
specific hindwing diversity in Catocala moths, then the apparent problem 
posed by the lack of intraspecific hindwing diversity must be faced. 
If selection has favored diversity on the one hand, why has it opposed 
diversity on the other? This problem would be easily solved if the 
hindwings were involved in specific recognition; serving, for example, as 
releasers in mating behaviors and therefore as anti-hybridization devices. 
Thus far there is no evidence that the hindwings serve any such function, 
but the possibility must remain open until Catocala mating behaviors are 
thoroughly studied. 


190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Another explanation for the lack of intraspecific hindwing variation 
may be related to the notion that parsimony would prevail with respect 
to the genetic bases of adaptive diversity in nature. Adaptive diversity 
in the case of Catocala hindwings is spread over an assemblage of species 
and seems to involve a large, but finite, number of categories, with each 
category being limited to a more-or-less fixed percentage of the total 
assemblage. The origin and maintenance of such diversity would seem 
to be more easily achieved if monomorphism, rather than polymorphism, 
characterized the species involved. 


SUMMARY 


An attempt is made to describe and interpret beak-damage patterns 
found on the wings of Catocala moths. Two samples of beak-damaged 
specimens were studied: (1) 23 individuals recovered after being at- 
tacked by blue jays in aviaries; and (2) 65 individuals retained from 
a large sample of these moths taken at bait and lights in the field. 
Analyses of these moths resulted in a classification of beak-damage pat- 
terns into three major types: I (attacked while flying, tear from one 
wing); II (attacked while resting, tears from ipsilateral forewing and 
hindwing); and III (attacked while resting, beak-imprint on at least 
one wing). About half of the specimens in both the aviary and field 
samples were attacked while flying, the other half being attacked while 
resting. 

The damage patterns obtained provide evidence that Catocala hind- 
wings serve both deflective and startle functions. The probable nature 
of the startle function is discussed in detail, and it is suggested that a 
sudden, unexpected display of hindwings results in startle (and con- 
sequent unsuccessful completion of attack) in predators. The evidence 
for this view is provided by the crisp beak-imprints found on the wings 
of many Catocala specimens, and by the distribution of these beak- 
imprints among moths having different types of hindwings. Anomaly 
(defined as departure of prey from predator expectation, with resultant 
startle in the predator) is proposed as a functional and adaptive basis 
for hindwing diversity in Catocala moths. 


ACKNOWLEDGMENTS 


I thank Dr. Alan C. Kamil for providing the blue jays used in the 
aviary study, and for critically reading the manuscript. Dale F. Schweitzer 
kindly provided a number of Catocala specimens for study. My wife, 
Katherine, ably assisted in the preparation of the figures, and patiently 
helped in many other ways. 


VoLUME 27, NUMBER 3 191 


LITERATURE CITED 


Biest, A. D. 1957. The evolution of eyespot patterns in the Lenidoptera. Be- 
haviour 11: 209-256. 

Brower, L. P. 1971. Prey coloration and predator behavior, in Topics in the 
Study of Life: The BIO Source Book. Harper & Row, New York. p. 360-370. 

Carpenter, G. D. H. 1941. The relative frequency of beak-marks on butterflies 
of different edibility to birds. Proc. Zool. Soc. London A 111: 223-231. 

CoLLENETTE, C. L. & G. Tartpor. 1928. Observations on the bionomics of the 
Lepidoptera of Matto Grosso, Brazil. Trans. Entomol. Soc. London 76: 391- 
Al6. 

Coppincer, R. P. 1970. The effect of experience and novelty on avian feeding 
behavior with reference to the evolution of warning coloration. in butterflies. 
II. Reactions of naive birds to novel insects. Amer. Nat. 104: 323-335. 

Cort, H. B. 1940. Adaptive Coloration in Animals. Methuen, London. 508 p. 

Forp, E. B. 1967. Moths. 2nd. ed. Collins, London. 266 p. 

Hunter, M. W. III & A. C. Kamin. 1971. Object-discrimination learning set and 
hypothesis behavior in the northern bluejay (Cyanocitta cristata). Psychon. 
Ser. 22; 271-273. 

Jones, F. M. 1932. Insect coloration and the relative acceptability of insects to 
birds. Trans. Roy. Entomol. Soc. London 82: 443-453. 

Kerer, R. R. 1968. Field studies of Catocala behavior. J. Res. Lepid. 7: 113- 
ie 

Keiutocc, C. G. & T. D. Sarcenr. 1972. Studies on the Catocala (Noctuidae ) 
of southern New England. II. Comparison of collecting procedures. J. Lepid. 
Soc. 26: 35-49. 

MarsHatL, G. A. K. & E. B. Poutton. 1902. Five year’s observations and ex- 
periments (1896-1901) on the bionomics of South African insects, chiefly directed 
to the investigation of mimicry and warning colours. Trans. Entomol. Soc. 
London: 287-584. 15 pls. 

Poutton, E. B. 1890. The Colours of Animals. Intern. Sci. Ser. 68, London. 
360 p. 

1913. Disabling and other injuries found in the Lepidoptera and their 
interpretation. Proc. Entomol. Soc. London: xix—xxii. 

SARGENT, T. D. 1966. Background selections of geometrid and noctuid moths. 
Science 154: 1674-1675. 

1968. Cryptic moths: effects on background selections of painting the 

circumocular scales. Science 159: 100-101. 

1969a. Behavioral adaptations of cryptic moths. I. Experimental studies 

on bark-like species. J. N. Y. Entomol. Soc. 77: 75-79 

—. 1969b. Behavioral adaptations of cryptic moths. III. Resting attitudes 

of two bark-like species, Melanolophia canadaria and Catocala ultronia. Anim. 

Behav. 17: 670-672. | 

1968c. A suggestion regarding hindwing diversity among moths of the 

genus Catocala (Noctuidae). J. Lepid. Soc. 23: 261-264. 

1972a. Studies on the Catocala of southern New England. II. Mating 

results with C. relicta Walker. J. Lepid. Soc. 26: 94-104. 

1972b. Sketches of New England moths. 5. Polymorphisms. Man & 

Nature (September): 25. 


1973. Behavioral adaptations of cryptic moths. VJ. Further experimental 
studies on bark-like species. J. Lepid. Soc. 27; 8-12 | 

& S. A. Hesset. 1970. Studies on the Catocala (Noctui lae of southern 

New England. I. Abundance and seasonal occurrence of the species, L961- 


1969. J. Lepid. Soc. 24: 105-117. 


192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


& R. R. Kereper. 1969. Behavioral adaptations of cryptic moths, I. 
Preliminary studies on bark-like species. J. Lepid. Soc. 23: 1-9. 

SoxoLtov, E. N. 1960. Neuronal models and the orienting reflex, in M. A. B. 
Brazier, ed., The Central Nervous System and Behavior. Joseph Macy Jr. 
Found., New York. p. 187-276. 

SWYNNERTON, C. F. M. 1926. An investigation into the defences of butterflies 
of the genus Charaxes. Proc. III Intern. Entomol. Congr., Zurich (1925) 2: 
478-506. 

TINBERGEN, L. 1960. The natural control of insects in pinewoods. I. Factors 
influencing the intensity of predation by songbirds. Arch. Neer!. Zool. 13: 
265-343. 


AN ORCHID ATTRACTANT FOR MONARCH BUTTERFLIES 
(DANAIDAE) 


W. H. WAcNER, Jr. 
Department of Botany, University of Michigan, Ann Arbor, Michigan 48104 


In view of the recent spurt of research on insect-flower relationships, 
and in particular the studies of Dodson and his co-workers on biologically 
active compounds in orchid fragrances (Dodson et al., 1969), it is 
especially interesting to discover an orchid which seems to have a 
practically “irresistible” attraction for monarch butterflies. 

The monarch butterfly is unquestionably the best known butterfly 
species in the United States, and its biology has been the subject of a 
great deal of attention (Urquhart, 1960). In the Great Lakes states, 
as elsewhere, the monarch is famous for its swarming behavior prepara- 
tory to migration southward to the Gulf Coast and Mexico. Swarming 
is observed in the latter part of September and early October as a rule, 
although Moore (1960) reported a swarm in the middle of August, 
“, . . thousands of individuals on Seul Choix Point on the north shore of 
Lake Michigan,” and Urquhart points out that migration actually has its 
beginnings in July. 

During the first two weeks of autumn, monarch butterflies are fre- 
quent everywhere in the vicinity of Ann Arbor in southeastern Michigan. 
Their behavior is languid, and they soar slowly across fields and along 
roadsides, feeding especially upon flowers of various species of asters 
(e.g. Aster azureus, A. ericoides, A. laevis, A. novae-angliae). In culti- 
vated legume fields the monarchs visit red clover (Trifolium pratense ) 
and alfalfa (Medicago sativa) primarily. In general, butterfly diversity 
is low at this time during most years—a few sulphurs (Colias philodice, 
C. eurytheme), some worn swallowtails (Papilio polyxenes especially), 


VOLUME 27, NUMBER 3 193 


buckeyes (Precis lavinia), painted ladies (Vanessa spp.), and occasional 
skippers (esp. Epargyreus clarus, Hesperia leonardus, and rarely southern 
migrants such as Atalopedes campestris). Angle-wings (Polygonia 
comma, P. interrogationis) and mourning-cloaks (Nymphalis antiopa) 
are seen along woods edges and in old orchards. Although Milbert’s 
tortoise shell (Nymphalis milberti) may become common in some years, 
monarchs and sulphurs dominate the picture. Wherever a lepidopterist 
drives on a warm, sunny day in early fall he finds Danaus and Colias 
on roadside asters, indicating a “good day” for butterflies. 

On just such a day, 19 September 1970, we encountered an extra- 
ordinary circumstance. The greenhouses at the University of Michigan 
Gardens, located at Dixboro, Washtenaw Co., Michigan (approximately 
five airline miles from the center of Ann Arbor), were discovered to 
be “full of monarchs,” as reported by one of our students. Looking into 
the matter we found that more accurately one part of one greenhouse 
was occupied by the butterflies. But the scene was remarkable; there 
were at least 200 individuals, as best we could count them. They were 
flying slowly to-and-fro, banging against the windows. At one corner 
of the house were five dozen butterflies settled on the glass panes. 
Something within this unit of the greenhouse was obviously attracting 
the monarchs. They would fly to the top of the greenhouse from the 
grounds nearby, and then fly down through the ventilators into the 
house. On 22 September 1970, the ANN ARBOR NEWS carried a 
story entitled “A Butterfly Invasion” describing the situation as follows, 
“More than 100 of the large butterflies are now gliding around the 
orchid greenhouse, beating their black-striped, orange-brown wings 
against the glass panes. . . . For some reason the butterflies have 
neglected adjoining greenhouses with their different plant varieties.” 
The reporter had consulted entomologists for an explanation. One of 
them commented, “At this point it is anybody’s guess why the monarchs 
have invaded this particular greenhouse. It could be something dif- 
ferent about the temperature, or humidity of the air, or some special 
odor from the greenhouse.” It was this last suggestion that we acted 
upon. 

The so-called “orchid greenhouse” actually contained many other kinds 
of plants besides orchids, including bromeliads, ferns, and certain aroids. 
We made the assumption that some plant—either its vegetative parts or 
its flowers—was generating a substance that was so powerful as to 
attract the insect from the neighboring fields and woods to the roof of 
the greenhouse and then down through the ventilators. Of all the plants 
growing there, only one was visited selectively by the monarchs—an 


orchid. However, the butterflies would on! by casually, stop 


194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


briefly, then leave, not to return. This particular orchid species has 
numerous flowers, but a monarch would come for its brief stop only 
to a single flower, and then depart. In spite of this seemingly almost 
random interest in the flower, we moved the orchid plant into another 
greenhouse in order to observe what effect this might have. We soon 
learned that there was no question about it: this was the plant. Very 
soon the monarchs now floated down into this greenhouse, as before 
they had in the other. And no new monarchs flew into the house where 
they had previously been attracted. 

The orchid has been identified as Epidendrum paniculatum Ruiz & 
Pavon, which is a widespread species (or species-complex) ranging 
from Central America to Peru. The plant is commonly and readily 
grown in horticulture for its somewhat fragrant flowers, these about 
2 cm. across, brownish to rose-white in color, which are borne in 
spreading, many-flowered panicles a foot or more in length. Historically 
this is one of the best-known members of its genus. Over three-quarters 
of a century ago, it was written (Veitch & Sons, 1887-1894) that this 
plant is, “One of the first epiphytal orchids known to science, and one 
of the most widely distributed of the genus. It was discovered more 
than a century ago by the Spanish botanists Ruiz and Pavon, near 
Huayaquil (Guayaquil?) in Peru, and has since been gathered by vari- 
ous collectors in many localities in tropical South America widely remote 
from each other, but always at a considerable elevation on the Cordilleras 
from Bolivia northwards to Venezuela. As a species it is very variable, 
due doubtless to diversity of station and its wide distribution... .” 
Dodson has written me of his personal observations in Costa Rica and 
Ecuador that this species or closely related ones will attract ithomiid 
butterflies. He suggests that perhaps the substances involved in the 
attraction of monarchs are pyrolizidine alkaloids. The orchid is, as 
indicated, widespread in cultivation and therefore readily obtainable 
for experimental work. 

Morrell (1960) reports that heliotrope (Heliotropium indicum, Bo- 
raginaceae) is attractive to danaids. Plants dried from 48 hours to a 
fortnight may be used. It should be interesting to determine whether 
there is a common factor in the substances produced by the orchid 
described here and the heliotrope. 

We were especially curious to learn whether the Epidendrum orchid 
attracted the sexes differentially, so we kept records for two years on 
the numbers that came into the greenhouses. We also set up what 
might be called “an obstacle course” to see whether we could entice the 
monarchs into the court of a large building, forcing them to fly up and 
over the sides, using the orchid as an attractant. 


VOLUME 27, NUMBER 3 195 


Both in 1970 and 1971 we missed the main flights. In 1970, on 21 
and 22 September there were still numerous butterflies being attracted 
into the greenhouse (a total of 119 for both days), but the weather 
became cloudy and the temperatures dropped, so they were considerably 
less common, the numbers dropping to 0-16 per day. Observations 
in the field after 22 September indicated that monarchs were either ex- 
tremely rare or absent; nevertheless, a few kept showing up at the 
greenhouses. Males were in slight excess—66 males to 51 females from 22 
to 30 September inclusive. In 1971, the orchid plant came into bloom 
rather late, but we kept records for the period 29 September through 
3 October inclusive, and counted a total of 47 males and 50 females. The 
sex ratio of attracted monarchs thus appears to be approximately 1:1. 

In 1972 we experimented to find out how much “trouble” the butter- 
flies might undergo to reach the flowers of this orchid by bringing it into 
the Natural Sciences Building on the Main Campus of the University 
of Michigan, Ann Arbor. Here in the center of the city, in a building 
four stories high, the plant was placed in a third floor office near a 
window opening into the central courtyard. In order for a butterfly to 
reach the plant it would have to fly over the walls of the building and 
then select the right window out of over 100 facing the court. So that 
the butterfly would not see the flowers, the venetian blind was lowered but 
the window left open. In spite of these seeming obstacles, three males 
did appear at and entered through the window, indicating that the 
attractive forces of this orchid for monarchs must be very strong indeed. 
The period of our “obstacle course” test was 15-17 September, and the 
butterflies came from 0955-1235. One butterfly appeared each day. 

From the evidence that we have seen, Epidendrum paniculatum is a 
powerful attractant for monarch butterflies. We have made no attempt 
to identify the compound or compounds which are active in the attrac- 
tion, but various persons have described their subjective interpretation 
of the scent of the flowers as “lemony” or “slightly spicy.” It is to be 
hoped that further studies of this attractant can be made, and also that 
students of monarch butterfly biology may be able to use the orchid in 
their research, e.g., for luring migrating specimens into buildings or 
other enclosures for purposes of marking. 

I wish to acknowledge the help of Caloway Dodson, Edward L. 
McWilliams, and L. J. Melton in making this study. I especially thank 
Louis Ludwig for his interest and help in gathering data. 


LITERATURE CITED 


Dopson, C. H., R. L. Dresster, H. G. Hits, R. M. ADAMS 6 N. H. Wi IAMS. 
1969. Biologically active compounds in orchid fragra Science 164: 1243 
1249, 


196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Moorg, S. 1960. A revised annotated list of the butterflies of Michigan. Occ. 
Papers Mus. Zool., Univ. Michigan. No. 617. 39 p. 

Morre.xi, R. 1960. Common Malayan Butterflies. Farrold & Sons, Norwich. 64 p. 

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

VeiTcu, J. & Sons. 1887-1894. A Manual of Orchidaceous Plants cultivated under 
Glass in Great Britain. Pollett, London. 108 p. 


ON ORNITHOPTERA PRIAMUS CAELESTIS ROTHSCHILD, 
DEMOPHANES FRUHSTORFER AND BOISDUVALI 
MONTROUZIER (PAPILIONIDAE ) 


H. Borcu 


Rabaul, New-Britain, Territory of Papua and New Guinea 
and 


F. ScHmMID 


Entomology Research Institute, 
Canada Department of Agriculture, Ottawa, Ontario 


In October and November of 1970 and 1971, the senior author under- 
took two trips by boat to Trobriand Islands, Louisiades Archipelago and 
Woodlark Island, to study the life histories of and collect Ornithoptera 
priamus caelestis Rothschild, O. p. demophanes Fruhstorfer and O. p. 
boisduvali Montrouzier. This article consists of field notes, descriptions 
of immature stages and a selection of photographs taken in the field 
by H. Borch. The young stages are described here for the first time. 
The taxonomic and phyletic considerations on the imagines are by F. 
Schmid. The eggs and the first three larval instars are identical in the 
three forms and likely are very similar to those of other subspecies of 
priamus; therefore, they are not described here. 


Ornithoptera priamus caelestis Rothschild 
General Observations 


Imagines and immatures were seen and collected on Misima Island 
(Misima, Liag and Larama), Nimoa Island (Nimoa), Sudest Island, 
Hemenahei Island and Moturina Island. The food-plant is Aristolochia 
tagala. The ova are always laid singly on the food-plant or on any 
nearby object or plant. An ant, Oecophylla smaragdina (native name, 
Kura Kum), has been observed preying on the eggs, sometimes sucking 
dry dozens of them. The newly emerged larvae eat their egg-shells and 
sometimes turn also to other unhatched eggs and completely devour them. 


VoLUME 27, NUMBER 3 197 


The larvae are always found on the food-plant, but when they are 
moulting they crawl for shelter into nearby shrubs. We noted a number 
of cases of cannibalism. This usually occurs just after ecdysis when 
larvae will attack and devour newly formed pupae. We once observed 
three larvae completely devour a pupa. 

The pupae are usually found on the underside of the leaves, or on 
nearby shrubs, or on grass or even sometimes on trees as high as 20 
ft. from the ground. They are usually found in shady situations. 

The imagines were seen mostly in the hills, as high as 300 ft. elevation, 
and very rarely at sea-level. Often, early in the morning, males were 
observed flying directly to female pupae to check if they were emerging. 
They hovered over the pupae for some minutes and finally flew away. 
They were observed doing this day after day. When a female finally 
emerged, it mated before its wings were fully dried. 


Taxonomy 


Fowth and fifth-instar larvae (Fig. 1). Basic colour intense velvety black. All 
tubercles partially red, none cream towards the base, except the dorsal tubercles 
on 4th abdominal segment. Cream saddle-marks small, not extending far laterally. 
Length at maturity: 67-75 mm. 

Pupa (Fig. 2). Basic colour rather dark brown. All markings very pale and 
contrasting. Two bright yellow areas: a small one on the pronotum and a large one 
on the metanotum and all abdominal segments, limited laterally by the wing-cases 
and abdominal tubercles. Lenght 48-50 mm. Pupal period: 22-26 days. 

Imagines. This small blue form is well known taxonomically. We shall not 
describe it again, but simply indicate variations that we observed on 36 ¢ é and 
30 © @ at our disposal. The anal blue band of the anterior wing is always nar- 
rower than the radial band and usually regular; in half of our specimens, it is 
interrupted in the middle of the anal margin. The number of the black discal 
spots on the posterior wing varies from 5 to 0, the usual number being 3 or 4. 
Almost all specimens show a black shadow on the discal cell and some have the 
whole disc powdered with black up to the discal spots. The extension of the black 
area is proportional to the size of the discal spots. We have seen only two speci- 
mens with uniformly blue posterior wings. Three specimens out of four have a 
yellow translucent spot in the cell Sc + RI — Rs. Specimens with yellow marginal 
dots have been reported and named flavopunctata Rousseau-Decelle, but only one 
of our males shows a pair of these dots. 

The female is very constant. The basic colour of both wings is deep brown, 
Both wings are very spotted, with a large oval mark in the discal cell of the anterior 
wings. The spots are uniformly rather dark. 


Phyletic Position 


Along with urvilleana, caelestis is the only known blue subspecies of 
priamus. An interesting point would be to know \ it Is derived from 
urvilleana (or from a common blue ancesto) evolved from a 
green form like poseidon and acquirec. © 


olour in a parallel 


198 JouURNAL OF THE LEPIDOPTERISTS SOCIETY 


Vy 


WY 
Wy 


Yj 


yy Yj ty 
UY 


Yj 
YY 


Figs. 1-2. O. p. caelestis: 1, fifth-instar larva; 2, pupa. Figs. 34. O. p. 
demophanes: 3, fifth-instar larva; 4, fourth-instar larva (other specimen). 


VOLUME 27, NUMBER 3 199 


manner to urvilleana, as did other subspecies like hecuba and bornemanni 
which show a tendency to be partly tinted with blue. 

The absence of the coloured cubital band on the anterior wing separates 
caelestis from the complex of poseidon, demophanes, boisduwali, borne- 
mann, hecuba, arruana and pronomus and readily places it in the line 
of urvilleana, priamus, admiralitatis, richmundia and euphorion. The 
regularity of the anal band on the anterior wing and its frequent inter- 
ruption along the anal margin separates it also from the four last named 
subspecies and relates it to urvilleana. The posterior wing is less rounded 
than in priamus, admiralitatis, richmundia and euphorion and has its 
anterior apical angle slightly protruding, as in urvilleana. Furthermore, 
the disc of the same wing is always more-or-less clouded with black, 
again as in urvilleana. Below, the posterior wing is blue with a yellowish- 
green marginal border in both forms. This shows rather convincingly 
that caelestis is the nearest relative of urvilleana and that their blue 
sheen is a monophyletic character inherited from an already blue common 
ancestor. 


O. priamus demophanes Fruhstorfer 


Since the original description by Fruhstorfer in 1913, from the 
Trobriand Islands, no mention of this form has been recorded in the 
literature. Its status is universally considered as doubtful. However, 
specimens are not rare in collections. The question we shall try to 
resolve here is if this form is a good subspecies or indeed a simple 
variation of poseidon, as are so many of Fruhstorfer’s so-called aberra- 
tions. 


Taxonomy 


Fourth and fifth instar larvae (Figs. 3-4). Basic colour velvety black, in some 
cases grey-black. Tubercles on thoracic segment 1 and all lateral tubercles black. Dor- 
sal and laterodorsal tubercles on thoracic segments 2 and 3 and all abdominal segments 
partially red, without cream colour. Cream saddle-marks on abdominal segment 
A broad, extending laterally to the spiracles and almost contiguous on the dorsum. 
Two additional cream saddle-marks usually on segments 5 and 6, decreasing in size 
(Fig. 3). A few specimens with these three marks on segments 3, 4, 5 (Fig. 4) or 
only on segments 4 and 5. One large larva measured 90 mm in length. Duration of 
larval period 25-29 days. 

Pupa (Figs. 5-6). Basic colour dull yellowish-brown, in some cases tinted with 
pinkish. Dorsal saddle-mark bright yellow. A fine brown line middorsally through 
the saddle-mark to last abdominal segment. A heavy dark brown line laterally below 
the wing cases. Tenulae yellow. Two short and sharp black-tipped processes on 
abdominal segments 5 to 8. Average length 60 mm. Pupae period: 24-28 days. 

Imagines (Figs. 10-12). From a dozen pairs that we have studied, we 
three specimens showing the extremes of variation, 


» illustrate 


JourRNAL OF THE LEPIDOPTERISTS SOCIETY 


200 


8, 


? 


7, larva 


1 


Figs. 7-8. O. p. boisduval 


pupae. 


demophanes: 


’ 


. 0-6. O. p 


gs 


i 


pupa 


VOLUME 27, NUMBER 3 201 


Fig. 9. O. p. poseidon, 6. Figs. 10-12. O. p. demophanes, ‘ variation of three 
specimens. 


202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Phyletic Position 


To assess the status of demophanes, we shall compare it with poseidon 
to see if the differences are important and constant enough to be con- 
sidered of subspecific importance. Poseidon is a very variable subspecies 
in size as well as in pattern. One of the patterns rather frequently 
observed is characterized by the cell Sc + RI — Rs on the posterior wings 
being entirely black and with 2 or 3 small black discal spots sharply 
decreasing in size. A specimen of this form is shown on Fig. 9, but it 
is clear that many intermediates exist. This type of coloration is not 
peculiar to poseidon, since we know it for admiralitatis and bornemanni 
also. It is to this form of poseidon only that we shall refer below. 

The ¢ of demophanes is remarkably constant in size as well as in 
pattern. Likely, it is genetically much more homogenous than poseidon, 
which can be expected in a form geographically restricted to an island. 
The green colour of demophanes is more often tinted with blue than is 
poseidon, though not frequently. In poseidon, the radial and anal bands 
are broad, regular and of equal width. In demophanes, the radial band 
is slightly narrower than in poseidon, the anal band is constantly and 
definitely narrower than the radial one, being sometimes reduced to a 
simple line (Fig. 11) and somewhat irregular. The reduction occurs from 
the posterior and outer sides with the result that the black margin is 
broader than in poseidon. 

The cell Sc+RI-—Rs of the posterior wing is sometimes entirely 
black, sometimes green, but always with a certain amount of black. 
The number of the black discal spots on the wings varies from 1 to 5, 
the first one often merging with the black area. The second one is the 
largest and the size of the following ones decreases more rapidly than 
in poseidon. Under the posterior wing, the discal spots are always clearly 
bigger than in poseidon and with frequent black indentations from the 
black margin in the cells Rs - M1 and M1 — M2. The yellow translucent 
spot on the cell Sc + RI — Rs is of course always absent in demophanes as 
well as in the form of poseidon considered here, but on one specimen of the 
latter we have noted two pairs of yellow marginal dots in the cells Rs — 
RI and M1 — M2; these dots always seem to be absent in demophanes. 

The 2 of demophanes is also very stable in size and coloration. We 
doubt that it can be separated from the highly variable 2 of poseidon 
but it can be described as follows: basic colour of the anterior wing 
light brown, except on the three margins that are dark brown. Spots 
numerous, dark gray, always present in the discal and the marginal cells. 

Basic colour of the posterior wing dark brown, with the spots as in 
poseidon. 


eee 


VoLUuME 27, NUMBER 3 203 


Figs. 13-17. O. p. boisduvali: 13-15, ¢ 6, variation of three specimens; 16-17, 
@ ©, variation of two specimens. 


204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The typical imaginal characters of demophanes are neither numerous 
nor important. But their constancy plus the remarkable fact that the 
caterpillar has three saddle-marks where poseidon shows only one, plus 
its insular isolation lead us to think that it is genetically distinct enough 
from poseidon to deserve a full subspecific status. It certainly evolved 
from specimens of the form of poseidon we considered above which 
populated the Trobriand Islands not long ago and became isolated there. 


O. priamus boisduvali Montrouzier 


Since its original description by Montrouzier, from Woodlark Island 
in 1852, only very few references have been made of this subspecies in 
the literature. They are all brief expressions of doubt on its validity, since 
Montrouzier’s description is insufficient to distinguish it. No specimens 
seem to be present in collections. We are very glad to have rediscovered 
this interesting subspecies that was forgotten or ignored for 120 years 
and to make it known adequately to the lepidopterological world. 


Taxonomy 


Fourth and fifth-instar larvae (Fig. 7). Basic colour velvety black. All tubercles 
partially red, those on abdominal segments 5 to 10 red and cream near the bases. 
Cream saddle-marks on abdominal segment 4 extending from the bases of dorsal 
tubercles and narrowing to the spiracles. Length 90 mm. Duration of larval period 
25-29 days. Feeding on local race of Aristolochia tagala with typical flowers and 
seed-pods but with a slight pink colouring on the stems. 

Pupa (Fig. 8). Basic colour dull yellowish-brown as in demophanes, but dorsal 
half sometimes darker than ventral half. Seen dorsally the outer edge of wing 
cases seeming to protrude more laterally. 

Imagines (Figs. 13-17). This subspecies shows a remarkable stability in all of 
its characters because of the small number of the populations and its isolation on a 
small island. Since it is closely related to poseidon we shall quote here only the 
characters that distinguish the two subspecies and shall again refer only to the 
form of poseidon we discussed above and illustrated in Fig. 3. Of the 20 pairs we 
studied, we selected four specimens showing the extremes of variation. Both sexes 
are small. Expanse of ¢: 8-12 mm; 9: 13-15 mm. 


Phyletic Position 


The green colour of the ¢ is distinctly more acid and bluish than in 
poseidon and remarkably constant. The outer margin of the anterior 
wing is slightly more tilted and the anal angle rounded. The posterior wing 
is comparatively small, very rounded, even more so than in euphorion 
and the outer margin shows only a very slight crenulation. On the 
anterior wing, both the radial and anal bands are slightly narrower than 
in poseidon, of equal width and the latter is slightly irregular. The cubital 
band is always present but never very well developed. On the same wing 
below, there is always a small coloured spot in the discal cell, the other 
spots being slightly reduced. 


VOLUME 27, NUMBER 3 205 


On the posterior wing there is a black area, usually large, extending 
not only into the cell Sc + RI — Rs as in poseidon, but also into the 
discal cell and into the anterior angle of the cell M1 — M2, which is 
never the case in poseidon. The apex of the first mentioned cell is 
always green. The number of the black discal spots varies from 1 to 5, 
the first one merging often with the black area. The black marginal 
border is well developed. The underside of the posterior wings is 
powdered with black at the extreme base. The discal spots are slightly 
larger than in poseidon and there are occasionally short black indenta- 
tions from the black margin. Yellow spot and dots are always lacking. 

The 2 is the darkest of all the subspecies of priamus known to us. 
The basic colour of the anterior wing is brown-black but generally washed 
with grey in all the marginal cells from the apex of the discal one. The 
only constantly present spot is in the cell Cul — Cu2 and is divided into 
two; it is present below only, though it is visible by transparency from 
above. There might also be two tiny spots in the cells M3 — Cul and 
Cu2 -— 2A, but they are not visible from above. 

The posterior wing is as shown in Figs. 16-17. Above, the spots are 
usually very dark. but are lightened to cream yellow in the cell Sc + 
RI — Sr. Beneath, the spots are distinctly lighter. They can be pure 
yellow, especially the two anterior ones, but the three posterior ones 
remain clouded with grey. 

Montrouzier’s type very likely disappeared long ago, but we do not 
find it necessary to designate a neotype, since there is no doubt about 
the identity of the subspecies. Boisduvali is obviously derived from the 
same form of poseidon as demophanes. It has evolved in the same direc- 
tion but has gone further, probably because its original population was 
isolated at an earlier period or because Woodlark Islands are more re- 
motely distant from the New-Guinean mainland than are Trobriand 
Islands. 


LITERATURE CITED 


FruustorFer, H. 1912. Neue Indo-Australische Rhopaloceren. Deutsch. Entomol. 
Zeit. Iris. 27: 130-139. 

MontrovuzieER, FATHER. 1852. Suite de la Faune de ile de Woodlark ou Moiou. 
An. Soc. Phys. Nat. Lyon: 393-395. 

RorTHscHitpD, W. 1898. New Lepidoptera from the East. Nov. Zool. 5: 216-219. 


206 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


PROJECT PONCEANUS: A REPORT ON FIRST EFFORTS TO 
SURVEY AND PRESERVE THE SCHAUS SWALLOWTAIL 
(PAPILIONIDAE ) IN SOUTHERN FLORIDA? 


Cuarces V. COVELL, Jr. 
Department of Biology, University of Louisville, Kentucky 40208 


AND 


GrorceE W. RAwson 
10425 Amherst Avenue, Silver Spring, Maryland 20902 


In this time of environmental awareness the subject of butterfly con- 
servation is receiving increasing attention in the United States. Lepi- 
dopterists have long been concerned with decreasing numbers of certain 
species; but only recently have efforts been undertaken to protect habitats 
and rare butterflies that populate them. Although the English have ac- 
complished much in this field (such as the reintroduction of the Large 
Copper, Lycaena dispar Haworth), Americans have made only small 
endeavors in the past. Rawson (1961) attempted unsuccessfully to intro- 
duce a small population of Eumaeus atala florida (Rober) into the Ever- 
glades National Park; but with more groundwork and support, future 
efforts along these lines may prove successful. 

The Schaus Swallowtail, Papilio aristodemus ponceanus, was described 
by Schaus (1911) from specimens he collected in the Miami area in 
May, 1898. Originally considered a separate species, and considered so 
by Holland (1930, 1931), it was given subspecific rank by Bames and 
McDunnough (1917). Bates (1934) and subsequent workers have con- 
tinued referring to it as a subspecies of aristodemus Esper. 

The popular and rather sensationalized report by Grimshawe (1940) 
of her discovery and rearing of ponceanus, with statement of its extinction 
on Lower Matecumbe Key (a paper considered by some to be the “last 
word” on the butterfly) did much to add to the impression among 
lepidopterists that ponceanus is a rare insect in Florida. The three 
articles by Henderson (1945a, b; 1946) summarized known information, 
and proved that the butterfly had not been rendered extinct by the 
September 1935 hurricane, as Grimshawe had claimed. Although some 
specimens were no doubt missed, Henderson’s papers listed collecting 
localities, dates, and owners of known specimens taken through 1945. 
Additional records since then are apparently not recorded is the literature. 
Klots (1951) and Kimball (1965) discussed ponceanus as rare, and urged 


1 University of Louisville Contributions in Biology No. 157 (New Series). 


VOLUME 27, NUMBER 3 20 


~l 


collectors to help preserve it. Young (1955) mentioned it as a butterfly 
one would be happy just to see, much less capture, and also recorded 
that the specimen figured in Holland (1931) was from Key West. C. V. 
Covell, Jr. took a ragged female on Key Largo on 31 March 1961 (re- 
ported in the Field Season Summary of the Lepidopterists’ Society for 
that year)—an unusual date, since all other records apparently fall 
between mid-April and late June. Collections of ponceanus during the 
1960°s seem few, and may indeed reflect some degree of scarcity. We 
believe, however, that it was merely not sought much by collectors. 

Rutkowski (1971) gave good new ecological and biological informa- 
tion, based on his 1970 experience with ponceanus on Key Largo. This 
article, plus a letter circulated by Kent H. Wilson urging that lepidop- 
terists take a hand in preserving what he thought was the last population 
in the U.S. of ponceanus, stimulated the authors to undertake such a 
project. 


The 1972 Survey Trip 


We first decided to find out if ponceanus is really as threatened as 
people seemed to think. We planned to visit southern Florida in early 
May 1972, and look for the swallowtail in lands protected by the U.S. 
Government from both collectors and developers. Through personal 
communication with Dr. William B. Robertson, Research Biologist at 
Everglades National Park, we learned that Torchwood (Amyris elemifera 
L.), the primary foodplant of ponceanus, is not known to be common 
anywhere in the Everglades National Park, but is well established on 
some of the keys making up the Biscayne National Monument. A second- 
ary foodplant, Wild Lime (Xanthoxylum fagara L.), while present in 
the Everglades National Park, was not thought to be very common. 
Robertson agreed to arrange a trip for us to Biscayne National Monument 
on 1] May. 

Through a travel grant from the Tom Wallace Conservation Fund at 
the University of Louisville, Covell and student assistant Gregory Florence 
drove to Florida City, where a rendezvous with Rawson was accomplished 
on 10 May. After a preliminary planning session with Robertson and 
Dr. William Hendrickson, Ecologist, at Everglades National Park head- 
quarters, the rest of the day was spent making camp at Long Pine Key 
campground, and seeking ponceanus in the Flamingo area of the Park 
(where X. fagara was reported to be growing). Papilio cresphontes 
Cramer was common, but no ponceanus were seen. 

Early on 11 May we met Dale Engquist, Superintendent of Biscayne 
National Monument, and Ranger George Sites, at the Monument head- 
quarters at Homestead Bayfront Park. Ranger Sites was to be our guide 


208 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


and boatman on the expedition, and botanist George Avery from Miami's 
Fairchild Garden came along to seek two rare plant species. A few 
light showers preceded hot, humid weather as we motored toward the 
first of the keys we were to visit. 

We tied the boat to the tangle of Red Mangrove (Rhizophora mangle 
L.) that completely surrounded the first island, and picked our way 
some distance through this dense vegetation. When we reached the 
higher hardwood hammock, George Avery pointed out some Torchwood 
trees. A swallowtail appearing to be ponceanus flew past at some 
distance; and another was netted by Covell as it flew beside him. The 
presence of the species there was verified; we saw about 15 more in as 
many minutes before we worked our way back to the boat. The second 
island was more easily penetrated, and ponceanus were seen (two taken) 
flying along or across paths which honeycombed the hammock. None 
were seen on a smaller, third key. Upon our return to the mainland, we 
were assured that Monument officials would be on the lookout for 
unauthorized collectors on these islands. 

On 12 May we looked in other parts of the Everglades National Park 
for ponceanus, but found none. The following day took us to Key Largo, 
where we surveyed the ponceanus population along paths from Rt. 905 
toward the Atlantic with Terry Dickel of Homestead. We took several 
specimens, all males, varying from fresh to slightly worn in condition. 
None were observed visiting blossoms, and they were not really abundant. 
Dickel informed us that the weekend before he had seen six collectors 
on Key Largo, apparently seeking ponceanus. We found very few eggs 
and larvae on the foodplants, and later learned that one collecting group 
had taken a large number of ova from foodplants there through systematic 
examination of the leaves. The Key is still not developed to any great 
extent; but we did find that the spot where Rutkowski had made his 
observations in 1970 is apparently gone now, bulldozed to form a marina- 
community called “Worlds Beyond.” We determined this from careful 
directions kindly given us by Rutkowski through personal communication. 

Other butterfly species recorded on 11 May in Biscayne National 
Monument included: Epargyreus zestos Geyer, Battus polydamas (Lin- 
naeus ), Papilio cresphontes Cramer, Ascia monuste (Linnaeus), Phoebis 
agarithe maxima (Neumoegen), Hemiargus ammon bethunebakeri Com- 
stock and Huntington, Ewunica tatila tatilista Kaye, Phyciodes frisia 
(Poey ), Heliconius charitonius tuckeri Comstock and Brown, Dryas julia 
cillene (Cramer), and Danaus gilippus berenice (Cramer). On 13 May 
on Key Largo we recorded the same (except B. polydamas, A. monuste, 
P. frisia and D. gilippus berenice) plus the following: Wallengrenia otho 


VOLUME 27, NUMBER 3 209 


otho (Smith), Polygonus leo (Gmelin), Appias drusilla (Cramer), Mar- 
pesia petreus (Cramer), and Agraulis vanillae (Linnaeus ). 


DIscussiION 


We felt that we had accomplished our mission, in that P. aristodemus 
ponceanus seems to be well established on at least two of the islands in 
the Biscayne National Monument. Except for the uncontrollable phe- 
nomenon of climatic traumas (freezes and hurricanes), the Schaus 
Swallowtail seems safe from real or imagined threats of extinction via 
development, pesticides and overcollection. On Key Largo, developers 
do pose some threat, but probably not for some years to come. There 
are Over nine miles of relatively undisturbed hardwood hammock along 
Rt. 905 from Rt. U.S. 1 to Ocean Reef at the northern tip. We feel that 
ponceanus is probably established in some other pockets in the keys, and 
hope to continue our survey activities. No efforts to introduce ponceanus 
to other sites seem to be necessary to the survival of the U.S. ponceanus 
population. 

This butterfly does, however, have a tenuous foothold in the United 
States, and we urge collectors to give this species a conservationist's 
concern when collection or purchase of specimens are considered. 


ACKNOWLEDGMENTS 


As “Project Ponceanus” progressed, many people contributed to its im- 
plementation in various ways. We are grateful to all for their help, and 
especially to the following: Dr. William B. Robertson, Research Biologist, 
Everglades National Park, for making our visit possible; Superintendent 
Dale Engquist and Ranger George Sites of Biscayne National Monument; 
Mr. Terhune S. Dickel, Homestead, Fla.; Mr. Gregory G. Florence, 
Bardstown, Ky.; Dr. W. Hendrickson, Ecologist, Everglades National 
Park; and Mr. Frank Rutkowski, New York City. We are also indebted 
to Drs. William Clay and Burt Monroe, Jr. of the Biology Dept., Univer- 
sity of Louisville, for making available to us funds from the Tom Wallace 
Conservation Fund. 


LITERATURE CITED 


Barnes, W. & J. H. McDunnoucu. 1917. Check List of the Lepidoptera of 
Boreal America. Herald Press, Decatur, Ill. 392 p. 


Bates, M. 1934. Notes on some tropical Florida butterflies (Lepid.: Rhopalo- 
cera). Entomol. News 45: 166-169. ae 
GrimsHAWwE, F. M. 1940. Place of sorrow: The world’s rarest butterfly and 

Matecumbe Key. Nature Mag. 33: 565-567, 611 | 
HenveERSon, W. F. 1945a. Papilio aristodemus ponceana Schaus (Lepidoptera: 


Papilionidae). Entomol. News 56; 29-32 


210 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1945b. Additional notes on Papilio aristodemus ponceana Schaus (Lepi- 

doptera: Papilionidae). Entomol. News 56: 187-188. 

1946. Papilio aristodemus ponceana Schaus (Lepidoptera: Papilionidae) 
notes. Entomol. News 57: 100-101. 

Houianpb, W. J. 1930. Notes on some American butterflies, mainly relating to 
their classification and nomenclature. Part 1, Papilionidae, Pieridae, Nymphalidae 
(Danainae). Ann. Carnegie Mus. 19: 185-204. 

1931. The Butterfly Book, Revised ed. Doubleday, Garden City, N.Y. 
A494 p. 

KimBaALiL, C. P. 1965. Arthropods of Florida and Neighboring Land Areas. Vol. 
1. Lepidoptera of Florida. Fla. Div. Plant Industry, Gainesville. 363 p. 
Kuots, A. B. 1951. A Field Guide to the Butterflies. Houghton Mifflin, Boston. 

349 p. 

Rawson, G. W. 1961. The recent rediscovery of Eumaeus atala (Lycaenidae) in 
Florida. J. Lepid. Soc. 15: 237-244. 

Rutkowski, F. 1971. Observations on Papilio aristodemus ponceanus ( Papilioni- 
dae). J. Lepid. Soc. 25: 126-136. 

Scuaus, W. 1911. A new Papilio from Florida, and one from Mexico (Lepid.). 
Entomol. News 22: 438—439. 

Younc, F. N. 1955. Notes on collecting Lepidoptera in southern Florida. Lepid. 
News 9: 204-212. 


NOTES ON THE LIFE CYCLE AND NATURAL HISTORY 
OF BUTTERFLIES OF EL SALVADOR. I. PREPONA 
OMPHALE OCTAVIA (NYMPHALIDAE) 


ALBERTO MuyYSHONDT 
101 Avenida Norte #322, San Salvador, E] Salvador 


For a number of years my sons and I have been collecting and breeding 
butterflies in the vicinity of San Salvador (600-900 m. altitude), capital 
city of El Salvador. Since the life cycle of many neotropical butterflies 
is not completely known, many species have been classified solely on 
the morphological characteristics of the adults. It seems desirable there- 
fore to place on record the various facts that we have found. This we 
intend to do in a series of articles dealing with the life cycle, host plants, 
and general natural history of the species we have been able to breed. 

A major difficulty has been the identification of the species described, 
as we are dependent on A. Seitz (ed.) (1924, Macrolepidoptera of the 
World, Vol. 5. The American Rhopalocera 1907-14), that is, according 
to many modern authors, “. . . replete with errors which cause much con- 
fusion.” (Klots, 1960). To partially overcome this handicap, Drs. F. H. 
Rindge and A. B. Klots of the American Museum of Natural History, and 
L. D. Miller of the Allyn Museum of Entomolgy, have made at least 


VOLUME 27, NUMBER 3 a1) 


tentative determinations of the material. Specimens of adults and of 
their early stages have been placed in these museums, so as to be avail- 
able for students of the groups. 

To give a clear idea of the habitats of the species described, it seems 
appropriate to make a rough description of the country. El] Salvador is 
the smallest country of Central America, and the most densely populated 
of continental America: 21393 km, with 3,500,000 hab. It has the shape 
of a rough parallelogram that lies between 87° 40’ and 90° 13’ east, and 
13° 11’ and 14° 40’ north, having Guatemala to the west, Honduras 
to the north, and the Golfo de Fonseca to the east. It has shores only 
on the Pacific Ocean (south). Being separated from the Caribbean by 
the high mountains of Sierra Madre, its climate is not affected by the 
monsoon conditions. There are six months of dry season (November 
to April) and six of rainy season (May to October). This fact influences 
greatly the flora and thence, the fauna. According to Serrano & Serrano 
(1972), only 348 species of Rhopalocera have been reported from EI 
Salvador. 

There are four main climatic zones: hot tropical lowlands (0-800 m. 
altitude), warm tropical plains (800-1200 m.), cool tropical highlands 
(1200-1800 m.) and cold tropical highlands (1800-2700 m.). In the 
lowlands, cotton, rice, sugar cane, corn and cattle are grown. In the warm 
plains, there are sugar cane, corm, cattle, some coffee, fruits and vege- 
tables. In the steeper cool highlands, there is mostly coffee, with some 
vegetables, fruits and flowers. The cold highlands are located in the 
NW part of the country, and there some forests are left. 

Due to the country’s dense population, most of the land is under intense 
cultivation, so what is left of wild vegetation consists of heavily disturbed 
second-growth plant communities, localized mostly along rivers and 
ravines. An advantage for the collector of insects in general and but- 
terflies in particular is that when being based in the capital city, San 
Salvador, he can find a wide range of altitudes and habitats within a 
range of 50 km, making it possible to find about 90% of the local species 
without the need of long traveling. 

It is within this range that we have collected eggs and/or larvae ot 
about 130 species of butterflies in order to study their early stages and 
developmental time. Among these species is Prepond omphale octavia 
Fruhstorfer, a rather scarce and elusive species of the family Nymphalidae. 
Some authors, e.g. Brues, Melander & Carpenter (1954), place the genus 
Prepona in the subfamily Nymphalinae, but prefer to use the more widely 
accepted subfamily Charaxinae. In this article we relate what we have 
found about the life cycle, behavior of immature stages and adults, host 
plant and habitat of Prepona o. octavia in El Sal) ador. 


bo 
fad 
bo 


JourNAL OF THE LEPIDOPTERISTS SOCIETY 


Since 1968 we have observed adults of P. 0. octavia at different al- 
titudes, ranging from sea level to about 2000 m., mostly in the neighbor- 
hood of coffee plantations (which can be considered man-made forests, 
due to the local technique of planting the coffee under shade trees, 
mostly Inga spp.), or near rivers and ravines. Yet, until December 1971, 
we had been unable to obtain eggs. At that time we found a female in 
the process of oviposition right in town, and three eggs were collected. 
Once the food plant was identified, a two-month search was made in 
an area of 20 blocks, and 21 larvae in different stadia were found. 
Fifteen were collected and six left on the plants. 

The eggs were photographed and put in individual plastic bags, as were 
the 15 larvae. The larvae were supplied with fresh leaves of the host 
plant every two days. Attempts were made to determine the develop- 
mental time under laboratory conditions, to photograph and measure the 
different stadia, and to keep material of the early stages preserved in 
alcohol. The transparent bags were kept under ambient temperature 


and lighting conditions until the emergence of the adults. No moisture 
control was made. 


Life Cycle Stages 


Egg. Pure white, spherical with slightly flattened base and depression at 
mycropyle, which is surrounded by a tiny ridge. No visible sculptures at 10x 
magnification. Diameter, 2.5 mm. All hatched in 7 days. 

First instar larva. Light brown all over. Head naked and roundish, slightly 
thicker than thoracic segments. Body naked, thickening gradually to 2nd abdominal 
segment, which is made prominent by two warts located one at each side, at sub- 
dorsal area, lighter brown than the general color. Body tapering to 6th abdominal 
segment then keeping about the same thickness to the 10th, which ends in two 
short and stubby “tails.” Anal prolegs slightly smaller than the other prolegs. 
Lateral ridge starting at thoracic segments subspiracularly, ending low at side of 
first abdominal segment. Another lateral ridge originates at 2nd abdominal seg- 
ment between the wart and spiraculum, which is placed much higher than the 
rest, and terminates at the “tails” of the 10th. Spiracula of subsequent abdominal 
segments are placed under this ridge, except on the 8th abdominal segment where 
it is above, and so out of line with the others. Larvae at emergence 3.5 to 4 mm, 
growing to 1.3 cm before moulting. Time, 12-13 days. 

Second instar larva. Darker brown on upper surface of body, where the two 
warts on 2nd abdominal segment are conspicuous due to lighter shade. Head, under- 
surface of body, and caudal zone all around, light brown. Head pyramid-shaped, with 
two fused epicranial horns, projecting higher than humped abdominal segments 
(1st and 2nd). Distinct “neck” formed by narrow Ist and 2nd thoracic segments. 
True legs tiny on Ist, slightly bigger on 2nd, and still bigger on 3rd thoracic seg- 
‘ments. Anal prolegs much reduced as compared to the other prolegs. Tails on 
tenth abdominal segment elongated. Head and body always naked, but tiny 
grainings most apparent around upper thoracic segments. Ridges lighter colored than 
rest of body. Reaching about 2.5 cm in 8-10 days. 

Third instar larva. Lighter brown than 2nd instar. Thin black lines dorsally at 
thoracic segments. Grains on head more noticeable and fused horns with tip slightly 


VOLUME 27, NUMBER 3 913 


Figs. 1-7. Propona omphale octavia Fruhstorfer: 1, egg (2.5 mm); 2, first 
instar larva 10 days after hatching (1 cm); 3, second instar larva 2 days after moulting 
(1.7 em); 4, third instar larva 10 days after moulting (3.8 cm); 5, fourth instar larva 
10 days after moulting (5.2 cm); 6, fifth instar larva 12 days after moulting (7.2 
cm); 7, prepupa. 


curved back. Anal prolegs still more reduced, and tails more elongated and slightly 
crooked. Reaching about 3.8 cm in 9-12 days. 

Fourth instar larva. Base of true legs on 3rd thoracic segment and prolegs on 
3rd, 4th, 5th and 6th abdominal segments much thickened, seeming to start just 
below spiracula. Body develops an indefinite and faint pattern of darker brown 
shade dorsally. Anal prolegs still more reduced. Tails longer and quite crooked. 
Reaching about 5.2 cm in 11-14 days. 

Fifth instar larva. Same aspect as 4th instar, but much longer and thicker, so 
that head appears disproportionately small. Very much reduced anal prolegs. The 
day before entering pre-pupal stage, color becomes “transluscent” light brown. 
Reaching 7 or 7.5 cm in 19-26 days. . 

Pre-pupa. Thick and incurvated, so touching with the horns the twig from which 
it hangs. General color transluscent brown, the warts now being darker than the 
body. Anal tails positioned at either side of twig. Time, one day. 


A214 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Ve, 
— 


Figs. 8-14. Prepona omphale octavia Fruhstorfer; 8-10, pupa—lateral, dorsal 
and ventral views (3.2 cm long); 11-12, adult male—dorsal and ventral view (7 
cm); 13-14, adult female—dorsal and ventral view (8 cm). 


VOLUME 2/, NUMBER 3 BALLS 


Pupa. Leaf green. Very humped dorsally across the first abdominal segments 
tapering to cremaster and to head. Spiracula brown, with prominent orange see 
around first abdominal one. Occasional orange stains in wing cases near antennae. 
Cremaster light brown and bifid head points light orange. Becoming dark gray 
shortly before adult emergence. Measures 3 to 3.3 cm long, 1.3 io 1.7 cm dorso- 
ey at widest point and 1.2 to 1.6 cm laterally at widest point. Time, 12-14 

ays. 

Adult. Dorsally, basic color is dark gray to dark grayish brown. Forewing with 
iridescent light blue band that starts near costa subapically, with outer border 
parallel to wing’s outer margin, inner border of band sinuose and diverging from 
outer border, so as to make band at inner margin of wing much wider than at 
costal margin. Hindwing with similarly colored band not reaching the costal margin, 
nor the outer angle, extending itself down parallel to the outer margin, but not 
reaching the anal angle, covering discal area without touching the inner margin or the 
basal area. Males, in addition to the light blue band in the forewing, have a dark 
blue area that covers the space between the band and the base of the wing. In the 
hindwing males have basally, under the cubital vein, a tuft of honey-colored “hairs.” 
The females lack the dark blue area in the forewing, and have an “eye” near the 
anal angle in the hindwing. 

Ventrally, wings are pearly gray basally, darker gray distally; the forewing with 
thin black lines forming an elaborate pattern, that in the hindwing is less elaborate. 
The hindwing has two medium-sized “eyes,” dark colored and surrounded by a 
clear gray ring; one near the outer angle, the second corresponding to the eye seen 
dorsally in the female, near the anal angle. Basic color and markings are darker in 
males than in females. 

Both sexes have the proboscis pink, the antennae black. Females have a wing 
span of 8 cm, while males are only 7 cm. Total developmental time: from 79 to 
97 days, females being slower than males. 


Natural History 


The foodplant, Andira inermis (Wright) Urban, is a robust leguminous 
tree that has very thick, dark green, imparipinate foliage, with leaves 
ranging from 6 to 10 cm. long, and a profuse lilac inflorescence from early 
March to mid-April. It has a rounded tree-top, and has been favored 
as an ornamental tree alongside sidewalks and parks in towns. In natural 
conditions it usually grows near rivers, thence its local name: Almendro 
de Rio (River Almond). These trees shed all their leaves from late 
December to late January, stay bare for about two weeks before growing 
new leaves that take another two weeks ito reach maturity. After that, 
the trees flower. 

Soon after the first instar larvae come out of the egg, they eat the 
egg shell completely, and stay under the leaf without further eating for 
one day. They move then to the tip of the leaf and eat around the 
central vein, leaving it bare. With frass stuck with silk they build a 
continuation to the bare vein, that then appears to project beyond the 
leaf limits, and the larvae keep perched on it while not feeding, usually 
head pointing outwards. This behavior is kept all through the first, 
second and third instars. Commonly during the third instar a larva 


216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


moves to another leaf because it has eaten the first one, and builds another 
“perch” on it with the now heavier frass. During the fourth stadium 
the larvae start wandering about the tree for feeding purposes, and keep 
motionless for long periods of time when not feeding, striking two 
characteristic poses: (1) head and thorax hanging at side of the twig 
where larvae hold with just the central prolegs, with the rear section of 
the abdomen either hanging also, or slightly raised from the seventh 
abdominal segment caudad; (2) head and thorax raised from second 
thoracic segment anteriad and last abdominal segments from seventh 
also raised. During the fifth stadium the larvae behave similarly. When 
newly hatched, and more markedly during subsequent stadia, the larvae 
walk with a balancing movement from side to side, due possibly to the 
very humped back and the fact that the reduced anal prolegs are not 
used for holding to the twigs while walking, having thence little leverage. 
During the prepupal stage, larvae weave a narrow girdle around a twig 
and then hang from it with the much reduced anal prolegs, positioning 
the crooked anal tails one at either side of the twig. Before hanging, the 
larvae clean the digestive tract by expelling a considerable amount of 
liquid with excrement. When becoming a pupa the larval skin splits 
from the head, breaking the head capsule in half and separating the 
fused horns. The larval attitude is all along very passive, being slow 
moving. No defense movements have been noticed during the pupal 
stage either, even when handled. Adults emerge very rapidly from the 
pupa shell, eject an amount of brown meconium, and are ready to fly 
in about 20 minutes. 

The adults of P. o. octavia, like most Charaxinae, have a swift and 
powerful flight that produces a rustling noise, somewhat like Hesperidae. 
This flight has been described by Lichy (1962) “. . . algo ruidoso y 
con movimientos de alas perceptibles, aunque rapidisimos,” (somewhat 
noisy and with perceptible wing movements, even though extremely 
swift). Males are very belligerent and chase any flying animal or 
object near their perching site. Females are larger than males and when 
ovipositing, circle the host tree several times at different levels before 
alighting in a cluster of mature leaves. There, sitting on the lower sur- 
face of the leaf, she deposits one egg. The female repeats the process 
several times before moving to another tree. The eggs we saw being 
laid were between 2.5 and 3.5 m. above the ground, the tree being about 
6 m. tall. The hour, 13:15. 

Both sexes are assiduous visitors of fermenting fruits, and some in- 
dividuals have been observed feeding in specific spots on trees, halfway 
up the trunks. Both sexes favor shaded coffee plantations, ravines, and 


VOLUME 27, NUMBER 3 AW 


rivers with thick second growth forests and patches of thick vegetation 
along shorelines. 

It was noticed that when newly emerged males and females were 
pressed on the thorax, they emitted a light green liquid from the base 
of the wings, and that they produced a mouldy odor when handled. 

We did not find a single case of parasitism in the 24 individuals 
observed. We found one instance of predation, a Chrysopa larva attacking 
a 2nd instar larva of Prepona o. octavia, which was killed without making 
any defense effort. The remains of a pupa were found still attached to 
a twig, only the abdominal segments affixed to the cremaster, with some 
body tissues left inside. One larva died while moulting to 2nd instar 
and one 4th instar larva died of a disease that softened the body. 

Three adult females were dissected, one each on days one, two and 
three after emergence. No eggs were found in any of them. A bright 
green fluid was found inside the abdomen. 


DiIscussION 


As this is the first time the life cycle of P. 0. octavia is fully described, 
some interesting facts have been found. The egg shape is very similar 
to the egg shape of the various species of Anaea found in E] Salvador, 
even though its size is much bigger. The shape of the larvae from the 
2nd stadium on resembles the shape of the larvae of Anaea (Zaretis ) 
itys Cramer (ms in prep.). The behavior of the three initial instars of 
P. o. octavia is very much like the behavior of those instars in various 
Anaea spp. except for the balancing gait of Prepona while walking. The 
spiracula of the 2nd abdominal segment in P. o. octavia is located very 
high, and this would explain why in the drawing of another Prepona 
(P. amphimacus Fabricius ) that appears in Comstock (1961, p. 174, fig. 
234), this spiraculum is lacking. This same characteristic, not so drastic, 
is found in the various Anaea spp. larvae we have observed, as is the 
slightly out-of-line spiraculum on the 8th abdominal segment. All of 
these factors seem to confirm the grouping of the two genera under one 
subfamily: the Charaxinae. 

The absense of parasitism found while breeding this species is striking 
considering the long developmental cycle (two and a half to three 
months), and the apparent lack of chemical and mechanical defenses. 
The larvae are very slow moving and passive, and the plant family 
Papilionaceae in general is not reputed for having poisonous components. 
Thus it appears that the immature stages of this species rely solely upon 
mimicry for protection: the larvae look like fragments of dry leaves, and 
the pupae are very inconspicuous in green foliage. It should be empha- 


218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


sized that while we studied the life cycle of P. 0. octavia, the life cycles 
of two other species which were feeding on the same trees were being 
observed: Panthiades bitias bitias (Cramer) and Theritas lisus (Stoll) 
(both Lycaenidae). These two species produced braconid wasps most 
of the time. This fact excludes the explanation that the lack of parasitism 
in P. 0. octavia was due to the absense of parasites at the time the life 
cycle was being studied. 

A possible cause of massive mortality of young larvae during the dry 
season could be the characteristic of the host plant, Andira inermis, of 
shedding all of its leaves rather abruptly during the period between late 
December and late January (not all the trees shed their leaves at the 
same time). The trees remain bare of leaves for a couple of weeks before 
growing new ones, and it is another two weeks before the new leaves 
reach the maturity the larvae require for feeding. In natural conditions 
A. inermis trees grow too far away from one other to allow larvae to move 
from one tree to another. Even when planted as ornamentals along 
sidewalks and parks in towns, they are placed from 6 to 10 m. apart. 

It is possible that P. 0. octavia uses other plants of the same group 
as a food plant, although we have not observed this. We can say in 
any case, it is one of the few species of Rhopalocera that have profited 
by man-made changes in natural ecology. 

According to the time it took to develop under laboratory conditions, 
we can assume that there are four generations a year of P. 0. octavia 
in E] Salvador. We can expect also that the females are slow in reaching 
sexual maturity, and that even if they produce eggs for a considerably 
long period of time (as do some other Lepidoptera with slow develop- 
mental time), they would not produce a very large number of them, 
because individuals of this species are rather scarce in spite of the little, 
if any, parasitism and predation the species is subject to, on one hand, and 
the abundance of the foodplant on the other. These factors would tend 
to make the species very abundant if the females were highly fertile. 

Besides three other species of Prepona found in El Salvador, that 
resemble very much P. o. octavia, there is an Apaturinae, Doxocopa 
cherubina Felder, that is superficially very similar to it. Whether or 
not these two species form a mimetic complex, we do not know. If 
they do, which is the model and which the mimic? The fact is that 
both species are very scarce locally and that circumstance would go 
against the tenet of accepted Batesian mimicry theory that requires the 
protected model to be more abundant than its unprotected mimic. The 
fact that P. o. octavia feeds on a tree that is not reputed to have poisonous 
properties would tend to eliminate the possibility of this resemblance 
being a Muellerian mimicry case. 


VoLUME 27, NUMBER 3 219 


ACKNOWLEDGMENTS 


We are grateful for the kind assistance of Dr. Lee D. Miller (Allyn 
Museum of Entomology) who identified the species mentioned, and 
made constructive criticism on the manuscript. We alsa thank Dr. 
Alexander B. Klots, for his encouragement to present the results of our 
work, and Drs. Theodore D. Sargent and Allen M. Young, who gave many 
valuable suggestions. My younger son, Pierre, first observed oviposition 
in P. o. octavia. Specimens of early stages and adults are deposited with 
the Allyn Museum of Entomology. 


LITERATURE CITED 


Brurs, C. T., A. L. MetanperR & F. M. Carpenter. 1954. Classification of 
Insects. Bulletin of the Museum of Comparative Zoology, at Harvard College. 
Vol. 108. Cambridge, Mass. 

Comstock, W. P. 1961. Butterflies of the American Tropics, The Genus Anaea, 
(Lepidoptera, Nymphalidae). 

Kiots, A. B. 1960. A Field Guide to the Butterflies. Houghton Mifflin, Boston. 

Licuy, R. 1962. Apuntes sobre los Agrias Doubleday. (Nymphalidae, Charaxidi- 
nae). Revista de la Facultad de Agronomia. Vol. II, No. 4. Maracay. Venezuela. 

SERRANO, FRANCISCO Y SERRANO, MicurL E. 1972. Las Mariposas de El Salvador. 
Primera parte: Papilionidae. Comunicaciones, 2a. Epoca. Vol. 1 #1. Uni- 
versidad de El Salvador. 


TWO NEW SPECIES OF PHYCITINAE FROM TEXAS, WITH 
DESCRIPTION OF TWO NEW GENERA (PYRALIDAE) 


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


Triozosneura A. Blanchard, new genus 


Tongue well developed. Antenna (Fig. 12) simple; finely pubescent in male. 
Labial palpus (Fig. 4) upcurved, rough scaled, reaching level of vertex, third 
segment very short (on denuded palpus (Fig. 13) it appears less than %s the length 
of second segment). Maxillary palpus squamous. Vestiture entirely of scales. 

Forewing (Fig. 11): Smooth, eleven veins, Ri absent, cell longer than half the 
length of the wing; discocellular vein weak curved; vein Cuz from near lower angle 
of cell; Cu: from the angle, slightly separated at base from stalk of Mes; Ms and 
M:; stalked for about 1% their lengths; Re contiguous or partly fused, for about +s 
its length, with the stalk of Rss; Rs and Rs stalked for about 34 their lengths; vein 
R: from cell. 

Hindwing (Fig. 11): With veins Cu: and Ms both present; cell at lower angle 
about half as long as wing; discocellular vein deeply concave; vein Cuz from near 
lower angle of cell; vein Cu: shortly united with the stalk of Mes; Ms and Msg stalked 


220 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-7. Holotypes and paratypes. Figs. 1-4. Triozosneura dorsonotata: 1, 
holotype ¢, Mt. Locke, Davis Mts., 27 August 1970; 2, paratype ¢, Big Bend Nat. 
Park, Green Gulch, 31 March 1971; 3, paratype @, Big Bend Nat. Park, Green 
Gulch, 6 May 1972; 4, head of holotype. Figs. 5-7. Glyphocystis viridivallis: 5, 
holotype ¢, Big Bend Nat. Park, Green Gulch, 23 March 1971; 6, paratype 2, Big 
Bend Nat. Park, Green Gulch 12 May 1972; 7, paratype ¢, Big Bend Nat. Park, 
Green Gulch, 28 March 1971. 


for well over half their lengths; S. and Rs shortly, and weakly anastomosed beyond 
cell. 

Male genitalia (Figs. 8, 8a): Uncus hoodlike subtriangular; tegumen strongly 
sclerotized laterally from its junction with vinculum to midheight where it supports 
the lateral arms of the gnathos; gnathos hoodlike, with acute apex, deeply cleft 
dorsally, fused apically, with semi-elliptical ventral opening; valve narrow, unarmed, 
with strongly sclerotized costa; transtilla absent; juxta narrow embracing *% of 
circumference of aedeagus; aedeagus half as long as combined height of vinculum, 
tegumen and uncus; vesica armed with a bunch of numerous cornuti, and what 


VOLUME 27, NuMBER 3 


bo 
bo 
— 


12 


Figs. 8-13. Triozosneura dorsonotata: 8, genitalia of ¢ holotype, aedeagus 
omitted (slide A.B. 2395); 8a, aedeagus; 9, genitalia of 2 paratype (slide A.B. 
3034); 10, bursa of 2 paratype (slide A.B. 2909); 11, venation; 12, basal segments 
of antenna; 13, denuded left labial palp (inner aspect) of a damaged, discarded 
2 specimen (slides A.B. 2647A, 2647B). 


222 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


appears as a small, thin, sclerotized membrane, bent as a half cylinder; vinculum 
about as broad as long. 

Female genitalia (Figs. 9, 10): Genital opening simple, narrowly sclerotized 
ventrally; ductus seminalis from ductus bursae, midway between genital opening 
and bursa; bursa with two signa, each consisting of a disk armed with hollow 
spines. Figs. 9 and 10 represent the bursae of two different specimens. It is believed 
that the bursa of the former is over-inflated, whereas that of the latter, which is 
depressed al] around the ductus bursae, thus forming a circular groove and a ridge, 
is more nearly normal. In Fig. 10 one signum only was represented for the sake of 
clarity: it is on the inner wall of the ridge and seen through the membrane of 
the bursa. 


If and when other species of this genus are discovered some of the 
characters of this description will appear only of specific value, but it 
would be awkward to guess which ones at the present time. 

The venation of the hindwing makes Triozosneura a member of 
Heinrich’s Group I and venetional division B. Using Heinrich’s key for 
division B, we are led through couplets 1, 3, 8, 9, 16, 21, 22 and 24 to cou- 
plet 25 which offers two alternatives, none of which applies satisfactorily 
to the case at hand: 26 is eliminated because the transtilla is absent from 
the male genitalia, 27 because the ductus seminalis is from the ductus 
bursae in the female genitalia. Thus we are led to a dead end in which, 
by chance, there are only four genera, namely: Coptarthria, Anadelosemia, 
Gabinius and Ceracanthia. It is enough to examine the detailed descrip- 
tion of these four genera, and the figures of the male and female genitalia 
of all species belonging to them, to convince oneself of the necessity of 
a new genus. 


Triozosneura dorsonotata A. Blanchard, new species 
(dies, I, we, B, 4 ch, ce, BD, IO, IL ile, ils) 


Head and thorax covered with gray or black scales tipped with white, more 
contrastingly mixed on the palpus. Abdomen yellowish gray. Forewing above gray, 
darker along costa, variably suffused with whitish on each side of the antemedial 
line, in the cell, and in the fold. Antemedial line white, starting on costa % 
distance from base to apex, reaching dorsal margin 24 distance from base to tornus, 
outwardly bilobed between (often indistinctly); contrastingly borderd with black 
outwardly, between costa and radius, where it is aimed toward tornus; widely, 
roundly excurved between radial and anal veins, where the black outer line almost 
disappears except for a conspicuous black spot on cubital vein, marking the notch 
between the two lobes; contrastingly bordered with black on both sides between 
anal vein and inner margin. Subterminal line mostly indistinct. Discal dots obsolete. 
On some specimens a reddish or yellowish diffuse spot covers the discocellular vein. 
Some veins are marked in black, particularly Rs and M; and the cubital in the basal 
space. A black blotch beyond the middle of the inner margin is generally separated 
from the antemedial band by a paler spot. Terminal dots and line absent. Fringe 
gray, somewhat lighter than the background of the wing. Hindwing yellowish-white, 
a little darker along the outermargin, mainly in females; fringe concolorous with 
disk of wing. Beneath: forewing brownish gray, a little darker along costa, hind- 
wing as above. Wing expanse: male 25 to 28 mm.; female 25.5 to 29 mm. 

Holotype: Male, Davis Mountains, Mount Locke, McDonald Observatory 


VOLUME 27, NUMBER 3 22:3 


Figs. 14-18. Glyphocystis viridivallis: 14, genitalia of ¢ holotype, aedeagus 
omitted (slide A.B. 3030); 14a, aedeagus; 15, genitalia of 2 paratype (slide A.B. 
3028); 16, venation; 17, basal segments of antenna; 18, denuded left labial palp 
(inner aspect) of damaged discarded specimen (slide A.B. 2649). 


Grounds, Texas 27 August 1970, genitalia on slide A.B.2395, deposited in the Na- 
tional Museum of Natural History (No. 72379). 

Paratypes: Big Bend Nat. Park, Basin, 7 April 1967, 1¢, (A.B.627); Green 
Gulch, 25 March 1971, 19, (A.B.2909); 31 March 1971. 18; 6 May 1972, 19, 
(A.B.3034); 12 May 1972, 16. Davis Mts., 5 miles SE of Mt. Livermore, 29 August 
1970; 19, (A.B.2396). 


224 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Glyphocystis A. Blanchard, new genus 


Tongue well developed. Antenna finely pubescent; on male with a shallow sinus 
at base of shaft, containing a row of minute black teeth half hidden between two 
rows of scales (Fig. 17). Labial palpus (Fig. 18) oblique, not grooved, smooth 
scaled, reaching level of vertex; third segment about % length of second, not 
projected forward. Maxillary palpus squamous, vestigial. 

Forewing (Fig. 16): Smooth, with eleven veins; cell about % length of wing; 
discocellular vein concave and weak between veins Mi and M,; vein Cuz, from before 
the lower outer angle of cell; Cu: from the angle, closer to Ms than to Cu, M> 
and Ms separated at base; M: from below upper outer angle of cell, straight; R: 
and R; stalked for a little over half their lengths; R: and R» separately from cell. 

Hindwing (Fig. 16): With veins Mz and Cu, both present. Cell at lower angle 
about half as long as wing; discocellular vein deeply concave, considerably extended 
at lower angle; vein Cuz from before angle; Cu: from angle, connate to the stalk 
of M2s; Mz and Ms anastomosed for about half their lengths; M1: connate to the 
contiguous parts of Rs; and Sc, which is about the same length as the free portion 
of Sc. 

Male genitalia (Figs. 14, 14a): Apical margin of uncus subrhomboidal; gnathos 
terminating in a spatulate hook; valve with strongly sclerotized costa, markedly 
produced at apex, about as long as combined length of tegumen and uncus; transtilla 
absent; juxta V-shaped; vinculum not quite as wide as long, becoming narrower and 
truncate anteriorly; aedeagus almost as long as combined length of vinculum, 
teugmen and uncus, stout; vesica armed with one stout cornutus, almost as long 
as aedeagus. 

Female genitalia (Fig. 15): Genital opening simple; ductus bursae somewhat 
contracted and membranous at junction with bursa; bursa wide and _ sclerotized 
posteriorly, bulbous and membranous anteriorly, provided on its right side with 
a large lobe; most of the inner surface of the lobe, a collar around ductus bursae, 
and some of the dorsal surface of the bursa covered with an inner, densely spinose 
mat. Ductus seminalis from the anterior end of the lateral lobe. 


In spite of the absence of a transtilla, and of substantial differences 
in the gnathos of the male genitalia, the following species could have 
been described in Catastia, but in view of considerable differences in 
the female genitalia this course of action was deemed unadvisable. 


Glyphocystis viridivallis A. Blanchard, new species 


(Ciies, &, 6 7% 14, I4e, 5, 16, iy, 8) 

Head, collar, first and second segments of palpus clothed with dark gray, white- 
tipped scales; gray and white more contrasting on second segment of palpus, third 
segment acute, black. Thorax concolorous with background of forewing. Forewing 
background, a nearly uniform slate-gray produced by dark gray, white tipped scales; 
generally a little darker in basal space. Antemedial line whitish, outwardly bordered 
by a black line which is generally much wider between costa and cubitus than in 
fold; varying from almost straight to a little sigmoid (outwardly convex above Cu, 
concave in fold) starting on costa 14 distance from base to apex, reaching inner 
margin %4 distance from base to tornus; a dark blotch of variable extent adnate 
to and basad of am line, along inner border. Subterminal line whitish, inwardly 
bordered by a thin black line (thicker and darker near costa), roundly retracted 
on vein Mi, and again, but less so in fold, reaching tornus at base of some very 
long fringe scales. A black triangular blotch distad of, and adnate to s.t. line near 
apex. Fine terminal black line. Fringe long, light gray, consisting of scales of three 


VOLUME 27, NUMBER 3 225 


different lengths, tipped with darker gray. Hindwing pale yellowish-white, darkened 
near apex and, in the female, along outer margin. Beneath: forewing yellowish-gray, 
hindwing yellowish-white. Wing expanse: 21 to 24 mm. 

Holotype: Male, Big Bend National Park, Green Gulch, Texas, 28 March 1971, 
gen. prep. A.B. 3030, deposited in the National Museum of Natural History, (No. 
72380 ). 

Paratypes: All from Big Bend Nat. Park, Green Gulch: 9 October 1969, 14, 
(Geb 0t®) 25) March 1971, 56 ¢, (A.B. 3030, 2649, 2650); 3 May 1972, 19, 
(A.B. 3029); 12 May 1972, 14, 12, (A.B. 3028). 


ACKNOWLEDGMENTS 


I am deeply grateful to Dr. D. C. Ferguson of the Entomolgy Research 
Division, U.S.D.A., for revising the manuscript and making several very 
helpful suggestions. The authorizations given me by Mr. C. D. Laughlin 
to collect on the McDonald Observatory grounds, and by the Administra- 
tion of Big Bend National Park to collect around and in the Chisos 
Mountains are also gratefully acknowledged. 


LITERATURE CITED 


Herricu, C. 1956. American Moths of the subfamily Phycitinae. U.S. Nat. 
Mus. Bull. 207. 


NOTES ON THE TAXONOMIC STATUS OF 
HYALOPHORA COLUMBIA (SATURNIIDAE ) 


MicHaEL M. CoL.iins 
924 Mendocino Avenue, Berkeley, California 94707 


The work of many authors (Sweadner, 1937; Weast, 1959; Collins & 
Weast, 1961; Wright, 1971) has shown that the various forms of 
Hyalophora are not reproductively isolated from one another. Females 
of any form of Hyalophora (in the restricted sense of Ferguson, 1972) 
will attract and mate indiscriminately with males of any other form. 
Generally ova laid by cross-mated females are viable and usually produce 
fertile F,; males and sterile females. Backcrossing the F; male with a 
female of either parental form or even a third form again produces fertile 
males and generally sterile females. Occasionally these females may lay 
some fertile ova. 

Intergrades and hybrids occur in nature. A population currently des- 
ignated “kasloensis” (Cockerell) exists between H. gloveri (Strecker) 
and H. euryalus (Boisduval) in Idaho, western Montana, and British 


226 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Columbia and is believed to be an intergrade between these two forms. 
H. cecropia (Linnaeus) produces occasional hybrids with gloveri. I 
obtained such an individual when a hybrid-like cocoon collected along 
with typical gloveri in the Black Hills of South Dakota produced a female 
very similar to laboratory hybrids. Mr. Duke Downey (pers. comm.) has 
collected supposed hybrids in Sheridan, Wyoming, but only rarely. The 
contact between gloveri and cecropia may be the recent result of a west- 
ward invasion of the latter species (Cockerell, 1929; Peterson & Worden, 
1962). 

Wild hybrids between H. columbia (Smith) and cecropia have been 
cited by Sweadner (1937). I collected a hybrid male along with typical 
columbia one mile west of Whiteshell Provincial Park along Highway 
One in Manitoba on 18 June 1963. This specimen will be described in 
a later paper. No true blending occurs between cecropia and the other 
forms but introgression is indicated by columbia and gloveri specimens 
with red scales in the extradiscal band. About half the specimens of 
gloveri and columbia sampled by Sweadner in areas of contact with 
cecropia exhibited this trait. The density of recognizable hybrids in 
nature is lower than predicted from the results of laboratory crosses. In 
addition, the mobility of the males during the mating flight and of the 
females during oviposition assures a potentially high rate of gene flow. 
Selection must act strongly against hybrids in some way. Hybrids may 
be ill-adapted such as in oviposition habits and consequent foodplant 
acceptance. H. columbia spins a small cryptically colored cocoon on 
exposed tamarack branches. When cecropia breeds with this form the 
larger non-cryptic hybrid cocoon may be more easily found by predators. 

Given the ability of the Hyalophora forms to intergrade, the origin 
and present status of columbia seemed to be an excellent subject for 
study. In the United States gloveri is quite distinct phenotypically and 
is separated geographically from the small dark form in Maine, Wisconsin, 
and Michigan designated columbia. In western Canada the gloveri popu- 
lation exhibits a phenotype nearer to columbia in size but often even more 
brightly colored than the Rocky Mountain phenotype. Consequently, 
gloveri and columbia have up to now been considered separate species. 
On the basis of geographic distribution, wing pattern, and identical 
genitalic structure, Sweadner believed columbia arose from gloveri at 
the end of the last period of glaciation. To account for the apparent lack 
of intergradation, he believed the two became geographically isolated 
by foodplant preference. The gloveri population in the prairies of Alberta, 
Saskatchewan, and Manitoba, which has been given the subspecies name 
nokomis, feeds on wolf willow (Shepherdia) and to a lesser extent on 
willow (Salix). The Hyalophora in the swamps east of Winnipeg, 


VoLUME 27, NUMBER 3 227 


recognized as columbia, is thought to feed entirely on tamarark (Larix). 
These food plants are not isolated, however. The Shepherdia of the plains 
meets to the north a broad band of spruce-tamarack forests ranging from 
British Columbia across Alberta and Saskatchewan to Manitoba where 
the conifers dip south to the east of Winnipeg (Harlow & Harrar, 1949). 
Thus, a foodplant bridge connects the two populations but at a more 
northern point than was sampled by Sweadner. 


Field Work 


I conducted field research in Canada in 1963, 1964, and 1966 to better 
sample these populations. Tied females and females in specially designed 
moth traps were placed at intervals throughout selected areas. The moth 
trap consists of a nine inch metal funnel mounted at one end of a screen 
cylinder two to three feet high. A small cage is suspended above the 
mouth of the funnel and confines a Hyalophora female. Attracted males, 
in their frenzy to reach the female, eventually collide with the funnel 
and slide through its enlarged opening. Thus, several males can be 
taken at each site and the female can be used as bait for up to 4 or 5 
days. 

Three population areas were sampled. The general trap line was along 
Highway One east and west of Winnipeg. The small dark phenotype 
was first taken just outside of the tamarack bog 30 miles east of Winnipeg 
near Richer. Thirty-two specimens were taken at regular intervals up to 
the Manitoba-Ontario Border beyond which no traps were placed. No 
males of any form were collected beyond the western extent of the bog 
30 miles east of Winnipeg to a point 90 miles west of Winnipeg. Shep- 
herdia does not seem to occur in this immediate area and extensive 
farming must further isolate the two Hyalophora forms. Stands of Shep- 
herdia occur west of here along fence rows and in untillable terrain. 
To the west the first male of the brightly colored phenotype was taken 
96 miles east of Brandon, Manitoba. A total of 17 specimens was taken 
westward along Highway One to Swift Current, Saskatchewan. The 
only cecropia collected was trapped 5 miles west of Virden, Manitoba 
on 21 June, 1963. 

In an attempt to collect a predicted intermediate form between gloveri 
and columbia, I collected north along Highway Ten to Flin Flon, Mani- 
toba and then west to Prince Albert Park, Saskatchewan. The terrain 
near Flin Flon is unusual in that the tamarack bog is interrupted by 
granite outcroppings and the flora is consequently more varied. Shep- 
herdia was not seen near Flin Flon. The possible implications of this 
environmental change was discussed below. Frequent cold weather 
limited collecting but 12 males were taken from 17 to 22 June, 1964. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


228 


XX 


RS 


RCC 


VoLUME 27, NUMBER 3 229 


Analysis of Specimens 


When the population samples collected were compared superficially, 
it was clear that not all specimens could be placed into one of two distinct 
phenotypes. Furthermore, the light and dark moths were not always 
associated with the expected foodplant community. One of the males 
from the tamarack bogs east of Winnipeg was as brightly colored as the 
average Shepherdia-feeding phenotype and at least one of the specimens 
from the latter population was quite as dark as those from east of Win- 
nipeg. Sweadner reported similar specimens from the large samples he 
secured. 

The moths taken from the north were quite variable; some were as 
dark as those from east of Winnipeg and others were as bright as the 
prairie phenotype. Most were intermediate in coloration (Fig. 1). To 
present a more objective comparison, I quantified this color variance and 
analyzed it statistically. While the more northern sample was somewhat 
small, I believe the results are significant. The ground color of the dark 
phenotype is usually accompanied by a darkening of the color band distal 
to the white wing band. Under the microscope one sees that this dark- 
ening is the result of an increase in the number of black scales relative 
to the number of white scales. Using a magnification of 40x with a 
grid reticle I measured the relative scale density in this band contained 
in the cell formed by veins Cu; and M3. The total numbers were con- 
verted to percentages and then plotted as a distribution with the mean 
and standard deviation. Fig. 2 shows that the specimens from northern 
Manitoba are more variable than those from east of Winnipeg and are 
very intermediate for this character. 

Surprisingly, the moths from northern locales were larger on the 
whole in both average and maximum size than specimens taken further 
south. Males collected east of Winnipeg (32 specimens) averaged 50.3 
mm measured from point of attachment of the forewing to the apex. 
These specimens ranged from 45.0 to 54.0 mm. Moths collected from 
250 to 375 miles NW of Winnipeg (12 specimens) averaged 55.8 mm 
and ranged from 50.0 to 61.0 mm. The prairie sample (17 specimens) 
averaged 52.2 mm and ranged from 48.0 to 55.0 mm. 


eS 


Fig. 1. Extreme light and dark specimens for each population area: H. columbia: 
1, Manitoba, 8 mi. e. Richer, 18 June 1966; 2, Manitoba, 17 mi. e. Richer, 18 June 
1966. Intermediate Hyalophora: 3, Manitoba, 3 mi. s. Baker's Narrows, 21 June 
1964; 4, Saskatchewan, 80 mi. w. Flin Flon, Ballantyne Bay, 22 June 1964. H. gloveri 
nokomis: 5, Manitoba, 3 mi. e. Brandon, 17 June 1966; 6, Manitoba, 5 mi. w. 
Virden, 21 June 1963. 


230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


x=80 S.D.=8.4 HYALOPHORA COLUMBIA 


© eee a, 
30-75 MILES EAST OF WINNIPEG 


Y% BLACK SCALES 
100 90 80 70 60 50 40 30 


INTERMEDIATE HYALOPHORA 
S.D.= 11.0 x=71 


= 2 a 250 - 375 MILES NORTH-WEST OF WINNIPEG 


H. GLOVERI NOKOMIS 


er 90-500 MILES WEST OF WINNIPEG 


Pn Am BO 


Fig. 2. Comparative scale counts of specimens collected in the three population 
areas. Scales were counted in a cell of the grey band of the FW formed by veins 
Cu: and Mz. 


After a thorough examination, I am in agreement with Ferguson 
(1972) that there are no significant structural differences, including 
genitalic characters, between gloveri nokomis and columbia adults. As 
with the adults, the larvae of these two forms differ mainly in size and 
coloration. I have reared over 100 gloveri larvae from north central 
Montana and found 25% to exhibit up to the fifth instar the same black 
dorsal and lateral scoli seen in immature columbia larvae. In the last 
instar the columbia larva differs from gloveri mainly in the color of the 
enlarged dorsal tubercules on segments 2, 3, and 4; these scoli vary from 
dull red-brown to a brighter red and are encircled at their bases by black. 
In gloveri these same scoli are usually yellow with a reduced black basal 
ring. I have seen a larva from Sheridan, Wyoming which possessed dull 
orange thoracic dorsal scoli. The remaining dorsal and lateral scoli are 
nearly identical in both columbia and gloveri; those on the sides vary 
from light blue tipped with white to nearly all white while the dorsal 
scoli on the abdominal segments are pale yellow. I have reared a few 
Montana (Toole Co.) gloveri which possessed yellowish orange dorsal 
scoli. Published descriptions of fifth instar gloveri larvae are varied. 
Sweadner (1937) states that all the dorsal scoli are straw yellow, “but 
in a few exceptions those on segments two, three, and four are slightly 
deeper in shade.” He further mentions a larva from Glacier National 


VOLUME 27, NUMBER 3 231 


Park, Montana with the dorsal scoli on segments 2 and 3 colored a 
“dull burnt orange.” Cooley (1908) reared gloveri larvae from Bozeman 
(Gallatin Co.), Montana and described the dorsal scoli on segments 
2 and 3 as “coral red” in color. Few descriptions of the gieveri nokomis 
larva have been published but Freedley (1908) describes the dorsal 
scoli as all yellow in Alberta specimens. Thus it appears that while 
columbia larvae are rather constant in coloration, a certain percentage 
of fifth instar gloveri larvae possess reddish dorsal scoli on segments 2 
and 3 and the immature larvae of both forms may be nearly identical 
except for size. Mature hybrid larvae of columbia x gloveri may exhibit 
either all yellow or all red dorsal scoli on segments 2, 3, and 4. The 
cocoons of gloveri nokomis and columbia are said to be indistinguishable. 


Laboratory Breeding 


The results of interbreeding were limited by the high rate of disease 
present in all my Hyalophora larvae, including pure strains. Neverthe- 
less, two female adults were reared on Larix from a mating between 
a female from Montana (Toole Co.), and a male taken 58 miles east 
of Winnipeg. Both of these females appeared fully fertile and laid a 
normal compliment of ova when bred to Montana gloveri. No exact 
count was made of the number of ova laid or the percentage of hatch 
but approximately 90% of the ova produced larvae. The resulting larvae 
were unfortunately lost to disease. More recently, hybrids between 
columbia (Livingston Co., Michigan) and gloveri from western Utah 
were reared by Mr. James Tuttle and Robert Weast. Two pairings of 
a male columbia X female gloveri yielded 92% and 90% hatch of ova 
laid. The reciprocal cross produced an 89% and 61% hatch (James 
Tuttle, pers. comm.). Robert Weast (pers. comm.) succeeded in in- 
breeding the F; hybrids of gloveri x columbia; the three females laid 
an average of 150 ova each. This is a normal number for the smaller 
Hyalophora forms. Fertility was over 90%. The results of this and future 
crossbreeding should be quantitatively compared to the degree of fertility 
seen in crosses involving other forms, such as gloveri X euryalus. Further- 
more, it should be determined if the fertility between gloveri and columbia 
changes as a function of geographic (and ecologic) separation of the 
populations studied. 

Some new information is available on the conifer feeding habits in 
the Hyalophora. In 1963 I reared to maturity on Larix two normal gloveri 
from a Russian Olive feeding population in Montana. Two dozen larvae 
were initially started successfully on Larix but again disease disrupted 
the experiment. In England gloveri is reported to be commonly reared 
on European Larch (Crotch, 1956). Dr. Thomas Koerber of the U.S. 


SOCIETY 


2 


JOURNAL OF THE LEPIDOPTERISTS 


232 


SIWOYON 1IY¥dAOTS “H 


VYOHdOIVAH JLVIGAWaAFLNI 


VIGWNIOD VYOHdOTVAH 


O 
3 


VOLUME 27, NUMBER 3 Dio 


Forest Service has collected wild euryalus larvae and cocoons on Douglas 
Fir (Pseudotsuga) at several locales in the Klamath and Cascade Moun- 
tain ranges of northern California. We have both reared gloveri and 
euryalus on Douglas Fir in captivity. A more detailed discussion of 
this work will be published at a later time. I am at present rearing 
thirteen columbia larvae which I have found will switch in the fifth in- 
star from Larix to Pseudotsuga after 30 to 45 minutes hesitation. During 
the first instar five larvae of a total of 24 initially began feeding on 
Pseudotsuga but eventually wandered to nearby Larix twigs. I did 
not force any of the first instar larvae to feed on Pseudotsuga due to the 
present scarcity of columbia stock. I feel that the ability of the various 
Hyalophora to feed in captivity on conifers, even reluctantly, is especially 
significant when one considers that many rather monotypic species, such 
as Callosamia promethea (Drury), may be regionally quite food plant 
specific (Collins & Weast, 1961). One would expect the acceptance of 
conifers to decrease outside the normal range of these food plants. 


Conclusion 


Gene flow between the brightly colored prairie population and the 
population of dark moths in the tamarack bogs east of Winnipeg produces 
in each population occasional phenotypes similar to the opposite form. 
The variability in the northern population of intermediates is also the 
result of gene flow from the two extreme populations. This clinal varia- 
tion seems not to be any more sudden in the sense of a “step cline” than 
is the transition from the Rocky Mt. phenotype into the bright prairie 
form. Both clines involve phenotypic and foodplant changes. 

Selection must favor the dark phenotype in the tamarack bogs and 
the lighter form in the prairie region since on the average the two popula- 
tions have rather distinct phenotypes. North of Winnipeg the environ- 
ment undergoes a transition from plains to conifer forest. Perhaps here 
where the intermediate form occurs, selective forces do not clearly favor 
either the light or dark phenotype, thus promoting the rate of gene ex- 
change. See Fig. 3 for food plant distribution and collecting sites. 

Melanism in other Lepidoptera has been shown to be controlled by 
a relatively few alleles. Thus, while the dark phenotype described as 
typical of columbia may appear to be qualitatively different from gloveri, 
the genetic basis for the apparent differences is probably minor. Whether 
this melanism is directly adaptive or is somehow linked with other 
adaptive gene systems is unknown. 

The conifer feeding habit has been shown to not be restricted to 
columbia and may even prove to be widespread in the western forms. 
Workers should attempt to determine if “kasloensis” and gloveri feed 


234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


naturally on either Douglas Fir (Pseudotsuga) or Western Larch (Larix 
occidentalis) in the Bitteroot and Rocky Mountain ranges. 

The selective basis for the melanism seen in columbia may operate 
during the larval stage. Perhaps the darker columbia larva is more crypti- 
cally colored in comparison to the lighter gloveri larva when feeding on 
the sparsely needled Larix. The immature columbia are especially dit- 
ficult to locate when reared in sleeves on larch. If the genes controlling 
melanin production in the larva are somehow linked with those controlling 
scale pigmentation in the adult, predator selection would quickly produce 
dark adults while acting upon the larval stage. Admittedly, such a gene 
system would be exceptional. Larvae are subject to much greater preda- 
tion pressure than the short-lived nocturnal adults, however. The “eco- 
logical image” of the adults is the underside of their wings as they rest 
with their wings folded over their backs. Furthermore, euryalus, gloveri, 
and columbia are much more similar when viewed in this aspect. Thus, 
the visual variance in dorsal wing coloration may not be the result of 
selection for this characteristic per s2. Selective breeding for light and 
dark larvae and adults may reveal a correlation. 

In summary, the obvious similarity of the two forms in all stages, the 
clinal variation of the melanization of the adults, and the demonstrated 
lack of isolation mechanisms coupled with the fertility seen in hybrid 
females all suggest columbia is not a separate species but is rather a 
melanic subspecies of gloveri. It is hoped that this paper will encourage 
and aid further investigation. 


ACKNOWLEDGMENTS 


I would like to sincerely thank Dr. C. Don MacNeill for continued 
help in the conception and analysis of the research project and in pre- 
paring this paper. I am also grateful for the living material and breeding 
information kindly provided by James Tuttle and Robert Weast. 


LITERATURE CITED 


CockERELL, T. D. A. 1929. The spread of Samia cecropia. J. Econ. Entomol. 
22: 97-98. 

Coxuins, M. M. & R. D. Weasr. 1961. Wild Silk Moths of the United States. 
Collins Radio Corp. 138 p. 

Cootey, R. A. 1908. Sixth annual report of the state entomologist of Montana. 
Bull. 75, Mont. Agric. College Exp. Sta. 

Crotcu, W. F. B. 1956. <A Silk Moth Rearer’s Handbook. The Amateur Entomol. 
Soc., England. p. 143. 

Frercuson, D. C. in Dominick, R. B. et al. 1972. The Moths of America North 
of Mexico. Fasc. 20.2B, Bombycoidea (in part). p. 252, 256, 259. 

FREEDLEY, W. J. 1908. The early stages of Samia columbia nokomis. Can. 
Entomol. 40: 350-354. 


VOLUME 27, NUMBER 3 235 


Hartow, W. M. & E. S. Harrar. 1949. Textbook of Dendrology. McGraw-Hill. 
p. 120-125. | 
Peterson, L. O. T. & H. A. Worpen. 1962. The cecropia moth in the prairie 
provinces. Contr. 20 Forest Nursery Sta. Research branch, Can. Agric., Indian 
Head, Sask. 

SWEADNER, W. R. 1937. Hybridization and the phylogeny of the genus Platysamia. 
Ann. Carnegie Mus. 25: 163-242. 

WeastT, R. D. 1959. Isolation mechanisms in populations of Hyalophora (Saturni- 
idae). J. Lepid. Soc. 13: 212-216. 

Wricut, D. A. 1971. Hybrids among species of Hyalophora. J. Lepid. Soc. 
25: 66-73. 


NOTES ON SIPROETA AND METAMORPHA WITH FIGURES 
OF SIPROETA EPAPHUS GADOUI MASTERS (NYMPHALIDAE) 


Joun H. MAsTERs 
5211 Southern Avenue, South Gate, California 90280 


Siproeta epaphus gadoui Masters (1967) was described from a series 
of specimens collected by Albert and Mary Lou Gadou at El Pao, Bolivar, 
Venezuela. The subspecies is of particular interest because it is inter- 
mediate in many respects between S. epaphus (Latreille) and S. trayja 
Hubner, which have been considered to be distinct species in the past. 
S. gadoui appears to be derived from trayja stock rather than from the 
geographically adjacent epaphus, although the entire Amazon Valley now 
separates trayja and gadoui. The description of gadoui was hurried in 
order to make the name available for a genus revision by Richard M. Fox 
and Alden C. Forbes and the new subspecies was not figured. The 
original description did promise figures in the coming revision of Fox & 
Forbes. The senior author of this revision, R. M. Fox, died on 25 April 
1968 with the manuscript partially completed. F. Martin Brown com- 
pleted the manuscript which was finally published on 24 December 
1971, albeit without figuring gadoui. I feel that it is important to figure 
this interesting butterfly and figures are hereby presented (Fig. 1). 

The Fox & Forbes revision (1971) may not be readily available to all 
lepidopterists and a summary may be useful. The genus Metamorpha was 
divided into two genera: Metamorpha Hubner (type species elissa 
Hubner ), and Siproeta Hubner (type species trayja Hubner ). Differences 
were cited in venation (in Metamorpha a vestige of the posterior tip of the 
third discocellular is present on the forewing), male forelegs (in Siproeta 
the tarsus is little more than half the length of the tibia, in Metamorpha 


236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Siproeta epaphus gadoui Masters: A. holotype male, El Pao, Bolivar, 
Venezuela; B. same, underside; C. allotype female, El Pao, Bolivar, Venezuela, 
upperside. All specimens natural size. 


it is one-fourth the length), and male genitalia (in Siproeta the tegumen 
and uncas are still separated by a suture, but in Metamorpha the suture 


has been lost and they are fused). A checklist of species and subspecies 
follows: 


VOLUME 27, NUMBER 3 237 


1.1 Siproeta epaphus 


a. S. e. epaphus (Latreille) 1811 Mexico to Bolivia, eastward 
into northern Venezuela. 

b. S. e. gadoui Masters 1967 Only known from E] Pao, 
Bolivar, Venezuela. 

c. S. e. trayja Hubner 1823 Southern Brazil and Paraguay. 

1.2 Siproeta superba 
a. S. s. superba Bates 1864 Southern Mexico, Guatemala 
and Honduras. 
b. S. s. euoe Fox & Forbes 1971 Costa Rica. 


1.3 Siproeta stelenes 

a. S. s. biplagiata (Fruhstorfer) 1907 All of Central America, 
northern Colombia, southern 
Texas and Cuba. 

b. S. s. stelenes (Linnaeus) 1758 Jamaica, Hispaniola, Cayman 
Islands, Virgin Islands, 
Leeward Islands, etc. 

c. S. s. meridionalis (Fruhstorfer) 1909 Trinidad and all of South 
America excepting northern 
Colombia and western Ecuador. 


d. S. s. sophene (Fruhstorfer ) 1907 Western Ecuador. 
2.1 Metamorpha elissa 
a. M. e. elissa Hubner 1919 Colombia to Peru and throughout 
the Amazon Valley. 
b. M. e. pulsitia Fox & Forbes 1971 Eastern Bolivia. 


The onl: effect upon the Nearctic fauna is that Metamorpha stelenes, 
species number 524 in the Dos Passos (1964) checklist, should be placed 
in the genus Siproeta. 


LITERATURE CITED 


Dos Passos, C. F. 1964. A Synonymic List of the Nearctic Rhopalocera. Mem. 
Lepid. Soc. 1. 145 p. 

Fox, R. M. & A. C. Forses. 1971. The butterflies of the genera Siproeta and 
Metamorpha (Lepidoptera: Nymphalidae). Ann. Carnegie Mus. 43: 223-247. 

Masters, J. H. 1967. A new Siproeta (Lepidoptera: Nymphalidae) from Vene- 
zuela. Ann. Carnegie Mus. 39: 193-194. 


NEW FOODPLANT RECORDS FOR PAPILIO POLYXENES F. 
(PAPILIONIDAE ) 


That the larvae of Papilio polyxenes F., the eastern black swallowtail butterfly, 
feed solely on plants of the carrot family, the Umbelliferae, is well documented (e.g., 
Scudder 1889, The Butterflies of the Eastern United States and Canada II; Holland 
1931, The Butterfly Book; Dethier 1941, Amer. Nat. 75: 61-73; Forbes 1960, 
Lepidoptera of New York and Neighboring States IV). The following three umbelli- 
fers have not previously been reported as larval food plants for this species. 

On 1 June 1970, at the summit of Shaw’s Ridge, Highland County, Virginia, | 
found two P. polyxenes larvae feeding on a meadow parsnip, Thaspium barbinode 
(Michx.) Nutt., which was growing in company with golden alexanders, Zizia aptera 
(Gray) Fern., on a dry roadside bank along Route 250. I reared the larvae to 


238 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


pupation on T. barbinode and Z. aptera collected at that site and on Pseudotaenidia 
montana Mackenz., mountain pimpernell, gathered at the summit of Shenandoah 
Mountain along the same route. Both larvae had been parasitized, and wasps, 
Trogus pennator (F.), emerged from the chrysalids on 18 June 1970. 

Identifications of the plants were kindly verified by Dr. Peter Hyypio, L. H. 
Bailey Hortorium, Cornell University, and plant specimens have been deposited in 
the Herbarium. The identity of the chrysalids was verified by Dr. John G. Francle- 
mont, Department of Entomology, Cornell University, and the parasites were identi- 
fied by Dr. Robert W. Carlson, Systematic Entomology Laboratory, USDA, Wash- 
ington, D.C. Insect specimens have been deposited in the Cornell University 
Collection, Lot 1023, Sublot 16a. 


SHERRY S. ReHR, Department of Entomology, Cornell University, Ithaca, New 
York. 14850. 


A YELLOW ABERRATION OF LERODEA EUFALA (HESPERIIDAE) 


Four of the five commonest, most widespread Hesperiinae in the Central Valley 
of California are golden-yellow in color: Hylephila phylaeus (Drury), Atalopedes 
campestris (Bdv.), Polites sabuleti (Bdv.) and Ochlodes sylvuanoides (Bdv.). The 
fifth, Lerodea eufala (Edw.), is dark brown with a few small whitish spots on the 
forewing above and below. On 4 September 1972, a male L. eufala was taken at 
Willow Slough, Yolo County, California, in which the dark brown is completely 
replaced by golden-yellow of nearly the exact shade prevalent dorsally in O. 
sylvanoides. The replacement extends to the body, appendages, and both wing 
surfaces. The whitish spots are normal. Many Hesperiine genera include both 
golden-yellow and brown species, and in some cases one sex will be yellow, the 
other brown. The yellow L. eufala suggests that the change from one color to the 
other is biochemically “easy.” If there is a mimetic or other advantage in golden- 
yellow pigmentation in California grassland skippers, the evolutionary opportunity 
for L. eufala to partake of it seems to be available. 


Artuur M. SuHapirno, Department of Zoology, University of California, Davis, 
California 95616 


A POPULATION OF LETHE APPALACHIA (SATRYRIDAE) 
FROM WEST CENTRAL FLORIDA 


The southern limit in the geographical range of the eyed-brown satyr, Lethe 
appalachia Chermock, is reported by Kimball (1965, Lepidoptera of Florida, Fla. 
Dept. Agr.) to be northern Florida. Klots (1951, Field Guide to the Butterflies, 
Houghton Mifflin Co.) indicates that the species ranges southward in the Ap- 
plachian mountains into Georgia and to a few swampy locations in northern Florida. 
In July and August 1972, a sizeable population of this species was discovered in 
a swampy forest located two miles south of Zephyrhills, Pasco Co., Florida. The 
swamp is adjacent to Crystal Springs, the large spring contributing to the head- 
waters of the Hillsborough River in west central Florida. This locality is approxi- 
mately 200 air miles south of those areas on the Florida-Georgia border where 
appalachia previously has been taken. 


VOLUME 27, NUMBER 3 239 


The initial collection was made on 28 July 1972, by members of my entomology 
class at the University of South Florida. Seven specimens were collected and many 
others observed flying about the swamp. Nearly all of them were in perfect 
condition indicating a recent emergence of adults. The dominant trees in this 
swamp are: water oak, Quercus nigra L.; bald cypress, Toxodium distichum (L.) 
Rich; sweet gum, Liquidambar styraciflua L.; blue beech, Carpinus carolina Walt.; 
red maple, Acer rubrum L.; water hickory, Carya aquatica (Michx.) Nutt.; and water 
ash, Fraxinus caroliniana Mill. Within the swamp, eyed browns were closely as- 
sociated with specific areas having a ground cover of giant sedge, Rhynchospora 
inundata (Oakes) Fernald. In August, several larvae of L. appalachia were found 
feeding on this sedge and adults continued to exhibit a distinct preference for flying 
about and resting in the sedge patches. 

The swamp was revisited on 7 October 1972, and several adult L. appalachia 
were still present and actively flying during the fall season. However, most of 
these individuals exhibited rather worn and tattered wings. 

I have directed considerable effort toward collecting this species in other ap- 
parently suitable localities along the west-central coast of Florida without success. 
It appears that the colony at Crystal Springs is very local in distribution and perhaps 
represents a southern disjunct population which is more or less isolated from those 
in northern Florida. 

Other species of butterflies collected within the Crystal Springs swamp in 1972 
include the following: Euptychia gemma (Hubner), E. hermes sosybia (Fabricius), 
Battus philenor (Linnaeus), B. polydamus lucayus (Rothschild and Jordan), Papilio 
polyxenes asterius Stoll, P. cresphontes Cramer, P. glaucus Linnaeus, Graphium 
marcellus (Cramer), Danaus gillippus berenice (Cramer), Heliconius charitonius 
tuckeri Comstock and Brown, Agraulis vanillae nigrior Michener, Phyciodes tharos 
(Drury), Polygonia interrogationis (Fabricius), Vanessa atalanta (Linnaeus), Lim- 
enitis archippus (Cramer), Asterocampa clyton (Boisduval and Leconte), A. 
celtis (Boisduval and Leconte), Saiyrium calanus (Hiibner), Urbanus proteus 
(Linnaeus), Pyrgus oileus (Linnaeus), Erynnis zarucco (Lucas), Wallengrenia otho 
(Abbot and Smith), and Lerema accius (Abbot and Smith). 


Larry N. Brown, Department of Biology, University of South Florida, Tampa, 
Florida 33620. 


NOTES ON THE OCCURRENCE OF HESPERIA PAHASKA MARTINI 
(HESPERIIDAE) IN COLORADO 


In August of 1967, one somewhat worn pair of specimens of a species of 
Hesperia were collected near Gateway, Mesa Co., Colorado. In May of 1968, 1969 
and 1970 additional specimens of this same species were collected at Black Ridge 
Breaks, also in Mesa Co., Colorado. Upon first examination, these specimens were 
identified as Hesperia viridis (Edwards), but a closer examination of the short 
series, including genitalic dissections and comparison to long series of both Hesperia 
viridis and Hesperia pahaska pahaska Leussler from various sections of Colorado, 
established them as members of the Hesperia pahaska complex. Further comparison 
with descripitions of members of this complex in MacNeil (1964, Univ. Calif. Publ. 
Zool. 35: 136, 142-151, Pl. 1) established the specimens to be Hesperia pahaska 
martini MacNeill, the first reported specimens of this subspecies for Colorado, rhe 
most proximate published localities (MacNeill, op. cit. ) are in Arizona (15 mi. 
WNW of Kayenta, Navajo Co.) and Utah (Beaver, Beaver Co.). Callaghan (1970, 
News Lepid. Soc. 3: 9) reported martini from the LaSal Mts., San Juan Co., Utah, 


240 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


collected in early June of 1969. It may be noted that the former two listed by 
MacNeill are several hundred miles from both Colorado localities but that reported 
by Callaghan is within 50 air miles of the Gateway locality in Mesa Co., Colorado. 
MacNeill gives no indication of the possibility of martini occurring in Colorado by 
his distribution map for the Hesperia pahaska complex. It is now noted that the 
range of this subspecies has been extended eastward and that it has a much closer 
association with H. pahaska pahaska than previously known. 

Although substantial series of this new discovery in Colorado are not available, 
it seems to depart from MacNeill’s description of martini in two distinct characters. 
First, the macular band on the undersurface of the hindwings is composed of small 
rather than large spots, comparable with those of eastern Colorado pahaska pahaska. 
Second, in several specimens, the amount of fulvous suffusion in the borders of 
the fore and hindwings above is not as extensive and therefore, the insect is not 
as brightly marked as specimens from westward in its range. Intermediacy to H. p. 
pahaska is suggested by these two characters; however, the differences between 
these specimens and eastern slope pahaska are too substantial to consider them 
as intermediates and they should be referred to as martini. Colorado martini are 
comparable in size to that given by MacNeill with a FW range of 15-16 mm for 
the males and 16-18 mm for the females. . 

MacNeill suggests that this subspecies is double brooded, flying in the spring 
and again in September. Colorado specimens have been recorded from both of 
these times (May and August), thus confirming his suspicion. The larval food- 
plant is unknown but may be Bouteloua gracilis (H.B.K.) Lag.; Steud., the host 
for pahaska pahaska in Douglas Co., Colorado (J. Scott, in litt.). 


MicuaEL S. Fisner. 1200 Summit Road, Route 2, Parker, Colorado 80134. 


RECENT SMITHSONIAN LEPIDOPTERA ACCESSIONS 
The Wilbur S. McAlpine Collection 


Through the kindness and generosity of Mr. Wilbur S. McAlpine, Union Lake, 
Michigan, the bulk of his collection of Lepidoptera has come to the Smithsonian 
Institution. 

This collection consists of over 12,000 specimens and is rich in material from the 
state of Michigan. In addition there is a series of Alaska butterflies collected by 
him in 1906 and again in 1911 and 1912 when he was Assistant Surveyor in a 
survey of coal claims at Homer, Alaska. Mr. McAlpine also was interested in the 
saturniid Hyalophora columbia and its hybrid with H. cecropia in Michigan and 
acquired a long series of these moths. 

The most important part of his collection, however, consists of the genus 
Calephelis which was the subject of his, “A Revision of the Butterfly Genus Calephelis 
(Riodinidae)” (J. Res. Lepid. 10(1): 1-125, 1971). All described species of 
Calephelis are represented with the following types of new species and subspecies 
coming to the Smithsonian: Calephelis sixola, C. perditalis donahuei, C. muticum, 
C. rawsoni, C. freemani, C. arizonensis, C. sinaloensis nuevoleon, C. dreisbacki, C. 
stallingsi, C. huasteca, C. montezuma, C. acapulcoensis, C. azteca, C. yucatana, C. 
maya, C. wellingi, C. wellingi baleuensis, C. clenchi and C. schausi. Paratypes of 
the above and those of other species are included in the collection. 

Other museums received holotypes and paratypes of some of McAlpine’s new 
species and subspecies. The museums and the holotypes they received are: American 
Museum of Natural History: Calephelis laverna trinidadensis, C. mexicana, C. 


VOLUME 27, NUMBER 3 241 


sacapulas, C. browni and C. tapuyo. The British Museum (Natural History) received 
Calephelis guatemala, C. braziliensis and C. burgeri. The following were sent to 
the Carnegie Museum: Calephelis nemesis bajaensis, C. costaricicola (neotype), C. 
sinaloensis, C. matheri, C. inca and C. aymaran. Calephelis nemesis dammersi and 
C. n. californica are in the Los Angeles County Museum. 


The E. J. Newcomer Collection 


Mr. E. J. Newcomer, formerly of Yakima, Washington, concentrated his collecting 
on Lepidoptera of that state and divided his collection between Washington State 
University and the Smithsonian Institution, the latter receiving nearly 3,000 speci- 
mens. This block of material, from a restricted geographical area, forms a very 
representative collection of that fauna, and is particularly strong in the genus 
Speyeria. 


The F. T. Vallins Collection 


This excellent assemblage of material consists of more than 22,000 blues and 
coppers of the family Lycaenidae. All of the specimens are from the Palaearctic 
Region, and although. there are no holotypes, there are long series from throughout 
the ranges of the species. These demonstrate all known forms of intraspecific 
variation that occur in the various localities. This is an extremely valuable collection 
for the study of variation and comparison of parallel development in American 
species. 


J. F. Gates Ciarke, Senior Entomologist, Department of Entomology, Smithsonian 
Institution, Washington, D.C. 20560. 


NOTES AND NEWS 


PROPOSED AMENDMENTS TO THE CONSTITUTION 
OF THE LEPIDOPTERISTS’ SOCIETY 


The following changes in the Constitution of the Society have been proposed 
in order to: 

1. add the immediate Past President to the Executive Council; and 

2. change the inauguration date of the President, Vice-Presidents, and the 
Members-at-Large of the Executive Council from 1 January to the date of the 
annual meeting. 

Notice is hereby given, in accordance with Article XII, Section 1, that these 
proposed amendments to the Constitution will be sent to the members with the 
ballots in November, 1973. Each section containing a proposed change is reproduced 
below in full, with new language in CAPITAL LETTERS; there are no proposed 
deletions. 

Article IV, Section 2: The business and affairs of the Society, not otherwise 
provided for, shall be controlled by an Executive Council, consisting of the President, 
President-elect, THE MOST RECENT AVAILABLE PAST PRESIDENT, three 
Vice-Presidents, the Secretary, the Secretary-elect, the Treasurer, the Treasurer-elect, 
and nine other members of the Society. Action on all amendments to the By-Laws 
and all appointments and elections by the Executive Council shall be obtained by 
a canvass by the Secretary of all members of the Council. 


242 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Article V, Section 2: Election of Officers. All officers shall be elected by ballot. 
The President and all Vice-Presidents shall be elected for the term of one year, 
and shall be eligible to succeed themselves once. The Secretary and Treasurer shall 
be elected for the term of three years and shall be eligible to succeed themselves 
twice. The nine other elective members of the Executive Council shall be elected 
for the term of three years; three of them shall be replaced each year; these members 
shall not be eligible to succeed themselves. The President, the Secretary and the 
Treasurer shall be elected a year prior to the time they take office. For each office, 
the nominee receiving the highest number of ballots shall be elected. THE 
PRESIDENT, VICE-PRESIDENTS, AND NEWLY-ELECTED MEMBERS-AT- 
LARGE OF THE EXECUTIVE COUNCIL SHALL TAKE) OF iChat tHe 
BUSINESS MEETING OF THE ANNUAL MEETING FOLLOWING THEIR 
ELECTION; IN THE EVENT THAT NO ANNUAL MEETING IS HELD IN 
A GIVEN YEAR, THESE OFFICERS SHALL ASSUME OFFICE ON THE 
ANNIVERSARY OF THE LAST ANNUAL MEETING. ALL OTHER officers shall 
take office at the beginning of the calendar year for which they are elected. 


¥ 


~ sz 


EDITORIAL COMMITTEE OF THE JOURNAL 
Editor: THEoporE D. SarcEnt, Department of Zoology, 

University of Massachusetts, Amherst, Massachusetts 01002 
K. S. Brown, J. M. Burns, R. H. Canrcasson, J. P. DoNAHUE 
J. F. Gates Ciarke, C. D. Ferris, R. O. KENDALL, J. H. MASTERS 
fe D. Miairer, A.’P. Pratt, J. R. G. Turner 


> 


> 


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, 814 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 Crtep, 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 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 article’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 FIGURES, 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 approx- 
imate 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. 

Page Charges: Authors with grant or institutional funds are requested to pay a 
charge of $24.00 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 
case of authors who have no grant or institutional funding, as it is not intended that 
any author should pay this charge from personal funds. However, all authors will 
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: Dr. C. V. Covell, Department of Biology, University of 
Louisville, Louisville, Kentucky 40208. 


ALLEN PRESS, INC. eerrea LAWRENCE, KANSAS 
ugar 


CONTENTS 


TAXONOMIC SIGNIFICANCE OF REFLECTIVE PATTERNS IN THE Com- 
POUND EYE OF LivE BUTTERFLIES: A SYNTHESIS OF OBSERVA- 
TIONS MADE ON SPECIES FROM JAPAN, TAIWAN, PapuA NEW 
Guinea anp Ausrrauia. Atuhiro Sibatani 


STUDIES ON THE C4TocALA (NocrumaE) oF SOUTHERN NEW 
ENcLAND. IV. A PRELIMINARY ANALYSIS OF BEAK-DAMAGED 
SPECIMENS, WITH DIsCUSSION OF ANOMALY AS A POTENTIAL 
ANTI-PREDATOR FUNCTION OF Hinpwinc Diversity. Theodore 

“Dy Sargent ee a 


An OrcHip: ATTRACTANT FOR MONARCH BUTTERFLIES (DANAIDAE). 
W. A. Wagner, Jeep ee 


On ORNITHOPTERA PRIAMUS CAELESTIS ROTHSCHILD, DEMOPHANES 


FRUHSTORFER AND BOISDUVALI MONTROUZIER (PAPILIONIDAE ). 
H. Borch and FF. Schmid 2.) 05.) 


PRroyEcT PoNcEANUS: A REPORT ON FirsT EFFORTS TO SURVEY AND 
PRESERVE THE SCHAUS SWALLOWTAIL (PAPILIONIDAE) IN 
SOUTHERN Fioriwa. Charles V. Covell, Jr. and George W. 
Rawson 


NOTEs ON THE LirE CycLE AND NATURAL History OF BUTTERFLIES 
oF Ex Satvapor. I. PREPONA OMPHALE OCTAVIA (NYM- 
PHALIDAE). Alberto Muyshondt 


Two New SPECIES OF PHYCITINAE FROM TEXAS, WITH DESCRIPTION 
oF Two New GENERA (PyRALIDAE). André Blanchard 
NOTES ON THE TAXONOMIC STATUS OF HYALOPHORA COLUMBIA 
(SaTurnmpAE). Michael’ M. Collins 
NOTES ON SIPROETA AND METAMORPHA WITH FIGURES OF SIPROETA 
EPAPHUS GADOUI Masters (NYMPHALIDAE). John H. Masters 


GENERAL NOTES 


New foodplant records for Papilio polyxenes F. (Papilionidae). Sherry 
S.. Fama so aie NG i A rr 
A yellow aberration of Lerodea eufala (Hesperiidae). Arthur M. Shapiro — 
A population: of Lethe appalachia (Satyridae) from west central Florida. 
Larry: INA GPOLON gl ea oa 


Notes on the occurrence of Hesperia pahaska martini (Hesperiidae) in 
Colorado.) Michael §.;\ Fisher 000 


Recent Smithsonian Lepidoptera accessions. J. F. Gates Clarke 


NoTEs AND NEwsS ______.--------- RO IN IE Uk Oa 


161 


192 


196 


206 


210 


219 


Volume 27 1973 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 


30 November 1973 


‘THE LEPIDOPTERISTS’ SOCIETY 
EXECUTIVE COUNCIL 


J. F. Gates Ciarxe (Washington, D.C.) President 

Harry K. Ciencu (Pittsburgh, Penn.) President-elect 
ALEXANDER B. Kiots (New York, N.Y.) Ist Vice President 
C. F. Cowan. (Berkhamsted, England) Vice President 

E. G. Munroe (Ottawa, Ontario) Vice President 

S. S. Nico.ay (Virginia Beach, Va.) Treasurer 

LEE D. MILER (Sarasota, Fla.) Secretary 


Members at large (three year term): M. C. Nievsen (Lansing, Mich.) 1974 
A. BLANCHARD (Houston, Texas) 1973 D. C. Fercuson (Washington, D.C.) 
R. B. Dominick (McClellanville, S.C.) 1975 

1973 R. O. KenpatL (San Antonio, Texas) 
J. P. DonanueE (Los Angeles, Calif.) 1973 1975 
J. M. Burns (Cambridge, Mass.) 1974 J. A. Powexu (Berkeley, Calif.) 1975 
R. H. Carcasson (Vancouver, B.C.) 1974 


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 $10.00 
Student members—annual dues $7.50 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $15.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia 23455. 


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


by Cyr. F. pos Passos 


Price, postpaid: Society members—$5.00, others—$7.50; uncut, unbound signatures 
available for interleaving and private binding, same prices; hard cover bound, mem- 
bers—$8.00, others—$10.00. Revised lists of the Melitaeinae and Lycaenidae will be 
distributed to purchasers free (separately with paper covered copies and unbound 
signatures, bound in with hard covered copies). 


The Lepidopterists’ Society is a non-profit, scientific organization. The office of 
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520. 
Second class postage paid at New Haven, Connecticut 06520 and at additional mail- 
ing office. 


JOURNAL OF 


tae LEPIDOPTERISTS’ SOCIETY 


Volume 27 1973 Number 4 


DESCRIPTIONS OF NEW NEOTROPICAL HESPERITIDAE 


S. S. NIcOoLAY 
1500 Wakefield Drive, Virginia Beach, Virginia 23455 


Throughout the past 20 years, I have by collecting, trade and limited 
purchase, acquired a number of specimens of neotropical Hesperiidae 
which I believe are unnamed. A new subspecies of the Pyrrhopyginae 
was described previously (Nicolay & Small, 1969); eight species of the 
Pyrginae and Hesperiinae are herein described as new. 

The descriptions utilize the English system of numbered veins and 
interspace identification found in Evans’ works. Wing measurements 
are from base to apex. Unless of specific importance, generic characters 
are omitted from the descriptions of new species. Lengthy descriptions 
of the male genitalia are omitted, for each is shown in lateral view with the 
left valva removed, the aedeagus in place and the inner surface of the 
right valva figured. Notable differences from other species in the genus 
are discussed where appropriate and helpful. All line drawings were 
made by the author. 


Ouleus dilla baru Nicolay, new subspecies 
Figs. 1, 2, 9, 9a 
Male: Length of forewing, 15 mm. Upperside: both wings very dark brown, 


with faint, vague post discal band on forewing; a post basal and post discal band on 
the hindwing. Underside: dark brown; faint bands of upperside repeated on both 
wings. Female: Unknown. 

Holotype male, Potrerillos, 1100 m., Chiriqui Province, Republic of Panama, 
14 February 1970, S. S. Nicolay, collector. Holotype will be deposited in the 
American Museum of Natural History, New York, New York. 


O. dilla was recently named by Evans (1953) from two specimens 
collected in Ecuador in 1896. The subspecies baru differs from the 
nominate form in having a completely dark brown hindwing beneath; 
dilla is white on the dorsal half of this wing. O. d. baru looks almost 
exactly like O. fridericus salvina and some dark specimens of O. f. 


244 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-8. New Neotropical Hesperiidae (Pyrginae and Hesperiinae): (1, 2) 
Ouleus dilla baru Nicolay, upper and underside, holotype male, Potrerillos, Chiriqui 
Province, Panama, 1100 m., 14 February 1970; (3, 4) Tosta sapasoa Nicolay, 
upper and underside, holotype male, Sapasoa, Rio Huallaga, 500 m., San Martin, 


VOLUME 27, NUMBER 4 945 


fridericus and the very dark, unmarked O. calavius. The male genitalia 
of all are quite distinct, yet it is only with the most careful scrutiny that 
adults may be separated. Field collecting provides additional challenges, 
for many of the species of Ouleus look exactly like species of the genus 


Staphylus that occupy the same habitats, and fly in much the same 
manner. 


Tosta sapasoa Nicolay, new species 
Figs. 3, 4, 10, 10a 


Male: Length of forewing, 20 mm. Upperside: forewing brown, shaded and 
vaguely formed into areas of different intensity; basal two-thirds dark chocolate 
brown, outer third a paler brown; with a pronounced white-centered costal fold. 
Hindwing lighter brown, base darkened by heavy concentration of dark brown hairs: 
vague, narrow, brown discal and central bands, fading into ground color at tornus. 
All wings with a very faint purple cast; fringes brown. Underside: forewing base 
dark brown from costal margin to interspace lb, the color formed by a heavy con- 
centration of dense, short hairs; remainder of wing paler brown; fringes brown. 
Hindwing fulvous-brown, base darker; irregular discal and central bands of upper 
surface repeated, appearing more obvious due to lighter ground color; basal half of 
interspace lc and adjoining cell area densely covered by long, grey hairs; interspace 
1 and lb heavily scaled in a mixture of light brown and grey scales, paler along 
inner margin and tornus; a light submarginal line of grey scaling continued from 
tornus along outer margin through interspace 3; fringes brown. Palpi below with 
a mixture of grey and brown hairs; above head and palpi black, thorax and abdomen 
dark brown; below abdomen light grey, with a single ventral black stripe. Antennae 
uniformly shining black above and below; nudum 21. Female: Unknown. 

Holotype male, Sapasoa, 500 m., Rio Huallaga, San Martin, Peru, May 1954, 
Felix Wytkowski, collector. The holotype will be deposited in the American Museum 
of Natural History, New York, New York. 


Evans (1953) erected the genus Tosta for a group of species allied 
to the genera Anastrus and Achlyodes, yet with marked differences from 
both. Certainly, the male genitalia of T. sapasoa bear a superficial 
resemblance to some Achlyodes and Anastrus species. Unlike the latter, 
sapasoa has a well developed costal fold in the male; it does not have 
a hair tuft on the hind tibia, nor the associated thoracic pouch. In wing 
pattern sapasoa resembles some species of Anastrus, yet the wide, heavy 
thorax, short, stout abdomen and very short forewing cell, place this 
species in the genus Tosta. 

Based on the very brief description of Evans’ T. taurus (1953), sapasoa 
closely resembles this species but is somewhat larger and is obviously 
separated by the genitalic differences. The male genitalia in sapasoa 


< 
Peru, May 1954; (5, 6) Damas immacula Nicolay, upper and underside, holotype 
vials. Colon (Sta. Rita) 300 m., Panama, 4 January 1969; (7, 5) Damas immacula 


Nicolay, upper and underside, allotype female, same locality as male, 1 January 1969. 


246 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


VOLUME 27, NuMBER 4 247 


form a very densely chitinous organ, replete with an extraordinarily wide, 
“horned” uncus and heavily spined cuillar. Both species are currently 
known only from single male types. 


Virga paraiba Nicolay, new species 
Figs. 12, 12a, 13, 14, 15, 16, 29 


Male: Length of forewing, 9 mm. Upperside: forewing dark brown; a discal 
yellow spot in each of spaces lb, 2 and 3, the largest in space 2; a tiny yellow upper 
cell spot, a straight line of three small yellow apical spots in spaces 6-8; sparse 
yellow scaling over the basal half of costal margin, the base of spaces la and Ib. 
Two small, dark brands over and under the origin of vein 2 (Fig. 29) their outer 
margins coincident with the inner edge of spot in space 2. Fringes sordid yellow, 
darkened at each vein end producing a faint checkered pattern. Hindwing dark 
brown; a discal row of fused yellow elongated spots in spaces 2-5; a few long 
yellow hairs over base of each wing; fringes light yellow with darkened vein ends 
producing a checkered pattern. Underside: forewing black; vein 12 and veins 
at apex yellow-scaled; also the medium veinlet from base to cell-end; a scattering of 
light scales across the discal area of spaces la and 1b with discal yellow spots in 
spaces 2 and 3; two pale violet spots near termen in spaces 4 and 5; a straight 
row of 3 apical spots; cell spot yellow. A yellow-scaled sub-terminal line from 
vein 1 to apex; a narrow post-terminal black line adjoins the wider dark brown base 
of grey fringes, darkened at each vein end. Hindwing dark brown, almost obscured 
by large areas of light violet spots and yellow-scaled veins; at the base of each wing, 
an arc of light violet conjoined spots from space lb through cell and space 7; a 
broad irregular light violet discal band from space 1b through 7, most obvious in 
spaces Ic, 4, 5, and 7. All veins yellow and a yellow sub-terminal line; a heavy 
post-terminal dark line formed by the dark bases of yellow fringes, themselves 
darkened at each vein end. Head, palpi and thorax with mixed yellow and black 
scales; abdomen dark brown dorsally, yellow striped at the side, cream colored 
below with 2 dark ventro-lateral stripes. Antennae yellow below, lightly scaled with 
black at each joint, the color pattern reversed on the upper side. Female: Length 
of forewing, 8 mm. All wing maculation above and below, and body coloring 
repeated as in the male. 

Holotype male, Joao Pessoa, Paraiba, Brazil, 31 March 1954, Jorg Kesselring, 
collector. Allotype female, same locality and collector, 8 February 1953. Para- 
types: 15 male and 12 female paratypes, same locality and collector with dates 
recorded in the months of December, January, February, March, April and May in 
the years 1953 and 1954. The holotype will be deposited in the American Museum 
of Natural History, New York, New York. Paratypes will be deposited as follows: U.S. 
National Museum, Washington, D.C.; Carnegie Museum, Pittsburgh, Pennsylvania; 
Allyn Museum of Entomology, Sarasota, Florida; British Museum of Natural History, 
London, England and with Olaf H. H. Mielke, Departamento de Zoologia, Uni- 
versidade Federal do Parana, Curitiba, Parana, Brazil. 


< 

Figs. 9-12. Male genitalia of new Neotropical Hesperiidae ( Pyrginae and 
Hesperiinae): (9, 9a) Ouleus dilla baru Nicolay, lateral view of genitalia and 
ventral view of gnathos and uncus (9a); (10, 10a) Tosta sapasoa Nicolay, lateral 
view of genitalia and ventral view of gnathos and uncus (10a); (11, lla) Pamba 
boyaca Nicolay, lateral view of genitalia and ventral view of gnathos and uncus 
(lla); (12, 12a) Virga paraiba Nicolay, lateral view of genitalia and ventral view 
of gnathos and uncus (12a). 


248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 13-16. Virga paraiba Nicolay: (13, 14) holotype male, upper and under- 
side, Joao Pessoa, Paraiba, Brazil, 30 May 1953; (15, 16) allotype female, upper 
and underside, same locality as male, 8 February 1953. (Approximately 2 life size ) 


Evans (1955) writes that the species virginius Moschler is variable 
and could possibly be divided into subspecies with more material. Wing 
markings on both surfaces of paraiba are similar in many respects to 
those of Moschler’s species, but the male genitalia are very different. In 
paraiba the uncus and gnathos are of equal length, the uncus with a 
distinct high crown; the gnathos of virginius is much shorter (almost 
non-existent) than the uncus and the uncus has no high crown. The 
valvae of paraiba are tapered evenly to a single curved point throughout 
their length; those of virginius are broad and sharply compressed to a 
point only at the very end. 

Evans (1955) gives as a generic character, “8 upf with a small, black, 
rounded brand over the origin of vein 2.” In V. paraiba, without bleaching 
and removal of all but the brand scales, this indeed is what appears under 
the microscope; the actual brand-scales (Fig. 29) are covered rather 
effectively by a layer of other specialized, larger, black scales. Hayward 
(1951) figures this brand as one of three examples included in the genus 
Callimormus Scudder where he placed the Virga species austrinus. His 
key to the Callimormus species contains a brand (estigma) description 
for austrinus which is the same as that figured for paraiba. Critical 
examination of other Virga species may well determine this brand form 
to be the correct generic character. 


VOLUME 27, NUMBER 4 249 


The upper cell spot on the forewing appears to be a variable feature 
and is present in 9 of 16 males and in 8 of 13 females examined. 

All specimens in the type series were taken by Mr. and Mrs. Jorge 
Kesselring at the type locality near their home. I have seen no specimens 
of paraiba from other localities. 


Pamba boyaca Nicolay, new species 
Kies tl Wa, 17, 18, 31 


Male: Length of forewing, 14 mm. Upperside: unmarked dark chocolate brown 
with sparse green hairs and scales on the collar and thorax. Forewing with a narrow, 
almost invisible tri-partite stigma from base of vein 3 along cubitus nearly to base 
of vein 2, thence directly in two short segments to middle of vein 1 (Fig. 31). 
Underside: forewing unmarked dark brown with heavier black scaling at base; hind- 
wing dark chocolate brown with faint small cream colored postmedian spots in 
spaces 3 and 6; mixed green and brown hairs clothing thorax and base of legs; 
palpi thickly covered with intermixed grey and brown scales; antennae black, sparse 
yellow scaling under club; nudum 11. Female: Unknown. 

Holotype male, Arcabuco, 2200 m., Department of Boyaca, Colombia, 31 January 
1971, S. S. Nicolay, collector. Paratype: One male, same data as the holotype. 
The holotype will be placed in the American Museum of Natural History, New York, 
New York. The single male paratype remains in the author’s collection. 


The genus Pamba was erected by Evans (1955) to accommodate a 
single unnamed species from Ecuador. The generic characters that 
establish this grouping are the shape of the long, narrow tri-partite 
stigma and the relatively long antenna with an obtuse apiculus. Boyaca 
differs from the sole previously known species pamba by its general lack 
of maculation on both wing surfaces; pamba has a narrow sub-tornal 
yellow area on the upperside hindwing, small silvery apical spots on the 
forewing underside and a pale yellow discal band on the hindwing 
underside. The valvae of the male genitalia are similar, but the uncus 
of boyaca is one of the most unusual forms I have encountered in any 
skipper. A dorsal plate or shield rises vertically from the horizontal 
plane of the uncus and dominates the entire lateral aspect of the genitalia. 
The chitinous sleeve or ring through which the aedeagus is articulated 
is neither mentioned nor shown by Evans in his illustration or description 
of pamba; it is however, a very obvious part of the male genitalia of 
boyaca (Fig. 11). 


Papias trimacula Nicolay, new species 
Figs. 19, 20, 25, 25a 


Male: Length of forewing, 15 mm. Upperside: both wings pale brown, the 


forewing with three small pale yellow discal spots in interspaces lb, 2 and 3, the 
largest in interspace 2. Underside: both wings brown with pale fulvous overscaling 
along costal area of forewing and on all of hindwing; yellow spots in interspaces 


2 and 3 on forewing; base and disc dark brown. Palpi missing; pale yellow scaling 


250 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 17-24. New species of Neotropical Hesperiidae (Hesperiinae): (17, 18) 
Pamba boyaca Nicolay, upper and underside, holotype male, Arcabuco, 2200 m., 
Dept. of Boyaca, Colombia, 31 January 1971. (19, 20) Papias trimacula Nicolay, 
upper and underside, holotype male, Ft. Clayton, Canal Zone, 21 September 1963; 
(21, 22) Vettius chagres Nicolay, upper and underside, holotype male, Colon 
(Santa Rita), 300 m., Panama, 29 January 1972; (23, 24) Vettius chagres Nicolay, 
upper and underside, allotype female, Colon, Pifa, Panama, 200 m., 24 November 
WPA, 


VOLUME 27, NUMBER 4 Ie 


around eyes and collar; head, thorax and abdomen brown above, paler below with 
abdomen pale yellow, almost white. Antennae with a faint checkered pattern above, 
more pronounced below, yellow under club, nudum red-brown, 3/10. Female: 
Unknown. 

Holotype male, Ft. Clayton, Panama Canal Zone, 21 September 1983, G. B. Small, 
collector. The holotype will be deposited in the American Museum of Natural 
History, New York, New York. 


Trimacula is undoubtedly the lightest colored species in the genus, 
rivaling Evans’ smaller species quigua from Venezuela. Although some 
others of the genus are marked with spots on the underside, trimacula 
is the only species with obvious, albeit pale and inconspicuous, markings 
on the upper surface. In this respect as well as the single added segment 
of the nudum on the apiculus (10 instead of 9), this species. does not fit 
the exact generic criteria set forth by Evans (1955). However, the male 
genitalia with a quadrifid uncus, very long saccus and equally long 


aedeagus places trimacula in the genus Papias as defined by Godman 
(1900). 


Vettius chagres Nicolay, new species 
Figs. 21, 22, 23, 24, 27, 27a, 32 


Male: Length of forewing, 17 mm. Upperside: forewing black, white hyaline 
spots in spaces 2, 3 lower cell and small sub-apical spots in spaces 6 and 7, the 
larger in space 6; a dense layer of blue-white hairs from the base along vein 1 in 
space la to midwing; short black brands above and below the origin of vein 2. Sides 
of collar and tegulae dark orange. Hindwing black with a prominent blue-white 
streak from base the length of space 1b, completely separate from the triangular 
white discal band running from spaces 2—5, the white spots hyaline in spaces 3-5, 
widest in space 2 consisting of long white hairs, all spots divided by dark veins. 
Fringes narrowly white at tornus. Underside: forewing dull black; a wide, short 
yellow pre-apical band cut by dark veins in interspaces 4 thru 8; hyaline white spots 
as above, with added small semi-hyaline white spot in lb. Hindwing, space la 
dull orange; space 1b and distal half of 1c black; basal half of spaces le through 4 
and cell orange; a tapered discal band of white semi-opaque spots in spaces 3-5, 
spaces 3 and 4 yellow from end of spots to termen; spaces 5 and 6 black from base 
to termen; spaces 7 and 8 clear yellow except bases and costal margin narrowly 
black; fringes paler at tornus. Sides of thorax at base of wings, orange. Female: 
Length of forewing, 19 mm. Upperside: same as in male, but forewing longer, 
narrower with a white spot mid space lb; hindwing wider. Underside: same as in 
male. 

Holotype male, Colon (Santa Rita), Republic of Panama, 300 m., 29 January 
1972, S. S. Nicolay, collector. Allotype female, Panama, Colon, Pina 200 m., 24 
November 1972, H. L. King, collector. Paratypes: 1 male, same locality as holotype, 
4 February 1970; 1 male, Gatun, Canal Zone, 10 January 1972, S. S. Nicolay, collector; 
3 males, same locality as holotype, 5 January 1969, 7 February 1969; 4 males, Gatun, 
Canal Zone, 9 December 1969; 2 females, same locality, 26 June 1970, 2 December 
1972, G. B. Small, collector; 1 male, same locality as holotype, 19 February 1969; 
16 males and 3 females, Pifia, Colon, Republic of Panama, 200 m., H. L. King, 
collector. The holotype will be placed in the American Museum of Natural History, 
New York, New York. Paratypes will be deposited as follows: U.S. National Museum, 
Washington, D.C.; Carnegie Museum, Pittsburgh, Pennsylvania; Allyn Museum of 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


252, 


Figs. 25-27. Male genitalia of new Neotropical Hesperiidae (Hesperiinae) : 
(25, 25a) Papias trimacula Nicolay, lateral view of genitalia and ventral view of 
gnathos and uncus (25a); (26, 26a) Damas immacula Nicolay, lateral view of 
genitalia and ventral view of gnathos and uncus (26a); (27, 27a) Vettius chagres 
Nicolay, lateral view of genitalia and ventral view of gnathos and uncus (27a). 


VOLUME 27, NUMBER 4 pas ys: 


Entomology, Sarasota, Florida; and the British Museum of Natural History, London, 
England. The allotype and remaining paratypes will remain in the collections of 
the author and Mr. G. B. Small. 

The male genitalia of chagres are almost identical to those of V. phyllus 
Cramer. Evans (1955) lists four subspecies for phyllus and I originally 
considered chagres to be a fifth taxon. But, with additional collecting and 
further careful study it became rather obvious that chagres and phyllus 
were two distinct, separate species. Both species have been collected in 
the forest-clothed hills and mountains on the Atlantic coast side of the 
Isthmus in the Gatun/Pifa and Colon area. I have found no intergrades 
in either sex between the two species in a careful study of rather sub- 
stantial series of both species, taken within a few miles of one another 
and, in some instances, in the same locality. 

There are a number of consistent, distinct differences between the two 
species: on the upperside, chagres lacks the orange scaling on the fore- 
wing costa, the spot in space 1b on the forewing is missing or tiny, and 
the white discal band on the hindwing is triangular in shape, with a 
definite, long concave dorsal edge; phyllus has the basal half of the costa 
on the forewing, dull orange, the white spot in space 1b of the forewing 
is always present, and the white discal band on the hindwing is rectangu- 
lar (almost ovoid), and about the same width throughout. The underside 
of the hindwing in chagres is strikingly different from typical phyllus 
or its subspecies; the large orange basal area and the appearance of the 
white discal spots of the upperside are the most notable differences. The 
obvious white streak mid space lb in phyllus is missing in the chagres 
male, and marked by a few white scales in the female. 

The known range of chagres thus far appears to be the forested hills 
on the Atlantic side of the Isthmus in the Canal Zone, and the adjoining 
areas of the Republic of Panama. 


Damas immacula Nicolay, new species 
Figs. 5, 6, 7, 8, 26, 26a, 30 


Male: Length of forewing, 23 mm. Upperside: all wings unmarked, dark, 
chocolate brown; forewing with a broad, grey bipartite stigma divided by vein 2. 
Hindwing fringes yellow from inner margin through tornus to vein 6. Underside: 
forewing unmarked dark brown; hindwing unmarked dark chocolate brown with a 


faint reddish tinge; fringes yellow-orange from inner margin through tornus to 
vein 6. Female: Length of forewing, 24 mm. Upperside: all wings unmarked dark 
chocolate brown. Underside: forewing brown, with a vague pale sub-apical band 
from the costal margin to mid-termen dividing wing into a vaguely darker apical 
area and lighter proximal area; space 1b paler with a wash ol yellow overscaling; 
hindwing dark brown from the costal margin through cell and portion of space 3, 
ventral half clear yellow; a thin dark brown marginal line to vein 2; yellow fringes 


from inner margin through tornus to vein ©. oe 
Holotype male, Colon (Santa Rita), 300 m., Republic of Panama, 4 January 


254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 28-28a. Male genitalia of Thespius inez Nicolay, lateral view and ventral 
view of gnathos and uncus (28a). 


1969, collector, S. S. Nicolay. Allotype female, same locality and collector, 1 
January 1969. Paratypes: 2 males with the same data as the holotype; 2 males, 
same locality and collector, 1 January 1969; 1 female, same data as allotype; 1 female, 
same locality and collector, 5 January 1969; 2 females, same locality, 1 January 
1969, 16 February 1969, collector, G. B. Small; 1 female, Farfan, Canal Zone, 2 
February 1968, collector, S. S. Nicolay. The holotype will be deposited in the 
American Museum of Natural History, New York, New York. Paratypes will be 
deposited in the U.S. National Museum, Washington, D.C., and in the Allyn 
Museum of Entomology, Sarasota, Florida. The allotype and remaining paratypes 
will remain in the author's collection and that of Mr. G. B. Small. 


Evans (1955) in his discussion of Damas clavus Herrich-Schaffer, the 
only species in this genus, refers to the rather wide variability in wing 
markings to be found in both sexes. It has become apparent to me, after 
an exhaustive search of the literature, and study of extensive material in 
the American Museum and Smithsonian collections, that this variability 
does not include the unmarked, spotless species herein described as 
immacula. In spite of the very close similarity in the male genitalia of 
the two, it would be difficult to place immacula in the category of a sub- 
species; I have taken both sexes of “typical” clavus in the same area and 
at the same time with immacula. This fact would also tend to negate 
the possibility that it might be only a seasonal form. Although Evans 
mentions 3 males and 1 female in the long series of clavus in the British 
Museum collection as being without spots on the forewings, none is 
from Panama. Of the two sexes, females of immacula offer the most 
compelling and obvious characters of a specific value. Yet, the yellow 


Ol 


VOLUME 27, NUMBER 4 25 


30 


Figs. 29-33. Forewing male stigmatal patterns: (29) Virga paraiba Nicolay; 
(30) Damas immacula Nicolay; (31) Pamba boyaca Nicolay; (32) Vettius chagres 
Nicolay; (33) Thespius inez Nicolay. 


hindwing beneath is not mentioned in previous literature, nor does Evans 
discuss this feature in combination with his single spotless female. The 
wide yellow ventral half of the underside of the hindwing is a color 
pattern also found in other large skipper species found in this same 
region—Tromba xanthura Godman, Astraptes anaphus annetta Evans, 
and Achlyodes busirus heros Ehrmann. 

D. immacula is found in the tropical forested hills on the Atlantic and 
Pacific sides of the Isthmus of Panama. Both sexes are particularly 
attracted to the large purple and white flower of a “morning glory” vine 
of the genus Ipomaea in the Family Convolvucidae that covered fallen 
timbers and areas laid waste by wood-cutters. Both sexes are very wary 
and powerful fliers. With extreme care, they could be netted while 
feeding on the flowers, but once missed, offered no second chance for 
capture. 


Thespius inez Nicolay, new species 
Figs. 28, 28a, 33, 34, 35 


Male: Length of forewing, 24 mm. Upperside: forewing dark brown, base of 
interspaces lb and 1 thinly clothed with long blue hairs; a faint white spot mid 
interspace lb, a narrow rectangular spot in space 2, a small square spot in space 3, 
a small triangular spot in space 4; 3 small sub-apical square spots in-line from 
spaces 6-8; a very small crescent-shaped upper cell spot; all spots white and 
hyaline. A narrow, pale, broken stigma from base of vein 3 to vein 1 (Fig. 32). 
Fringes at tornus yellow to vein 2. Hindwing dark brown, base clothed in long 
blue hairs, with a pale yellow tornus 2 mm at the widest point centered at vein i. 
extending to vein 2 and opposite through tornus to inner margin; tornal fringes 
yellow. Underside: forewing red-brown along costa with lilacine scaling on outer 
margin from apex widening to include spot in space 4, terminating at a point just 
inside space 2; dark brown in disc, hyaline spots as on upperside; spot in space 
lb enlarged by white scaling. Hindwing red-brown with a wide bar of heavy 
lilacine scaling from vein 6 through end of cell to mid-point of vein L: abdominal 
fold dark brown, remainder of wing with intermixed lilacine scaling, at some points 


256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 34 & 35. Thespius inez Nicolay, upper and underside, holotype male, 
Arcabuco, 2200 m., Dept. of Boyaca, Colombia, 31 January 1971. 


quite heavy; tornus pale yellow, 4 mm wide from vein 2 to inner margin; fringes 
yellow from vein 5 through tornus. Head, thorax and tegumen dark red-brown, 
mixed with long blue hairs on the thorax. Abdomen dark brown, anterior clothed 
with long blue hairs, posterior tip yellow. Palpi with mixed brown and light grey 
hairs and scaling. Below, thorax and legs clothed with red-brown hairs, abdomen 
with orange-yellow hairs; Antennae brick-red at bend of apiculus, yellow on the 
club beneath. Female: Unknown. 

Holotype male, Arcabuco, 2200 m., Department of Boyaca, Colombia, 31 January 
1971, S. S. Nicolay, collector. The holotype will be placed in the American Museum 
of Natural History, New York, New York. 


This is a large hesperiid by any standard, and the largest of this genus 
I have yet seen. The reduced hyaline markings of the forewings, complete 
lack of markings on the hindwings above, and the yellow tornal marking 
on both hindwing surfaces make inez something other than a typical 
species of the genus Thespius. Wing maculation and the male genitalia 
relate it closely to T. pinda Evans, known only from the type which I 
have not seen, and the subspecies ovallei Bell of T. tihoneta Weeks. The 
illustration of tihoneta Weeks (1905) bears little resemblance to inez. 
My vivid impression of inez is that, like many of the genus, it is very wary 
and an incredibly fast and powerful flyer. It is also, like its closest rela- 
tives, rather rare; each of the three is currently known only from a single 
male in each species. 

It is a pleasure to name this interesting species after Inez Schmidt- 
Mumm, the wife of Dr. Ernesto W. Schmidt-Mumm of Bogota, Colombia. 


ACKNOWLEDGMENTS 


I want to thank Mr. Gordon B. Small of the Canal Zone and Dr. 
Ernesto W. Schmidt-Mumm, Bogota, Colombia for their help with 
extensive field work, and invaluable logistic support for my own collect- 
ing efforts; their enthusiasm and assistance make such a study as this 
both a possibility and a pleasure. I very much appreciate the help of 


VOLUME 27, NUMBER 4 257 


Mr. and Mrs. H. L. King who collected an extensive series of the new 
species of Vettius for this study. Additionally my thanks to Gordon 
Small and Gerald Straley for helpful suggestions on the manuscript and 
to Dr. C. Don MacNeill of the Oakland Museum for his critical re- 
view of the final draft. To Dr. Ronald W. Hodges of the Smithsonian 
Institution, my thanks for help in solving a rather interesting and puzzling 
problem of subspeciation. Wm. D. Field, U.S. National Museum and Dr. 
F. H. Rindge, American Museum of Natural History provided the as- 
sistance and cooperation that allowed me to examine the material in 
the collections in their care. 

Most of the photographs were made with the help of WO G. G. 
Williams, SSgt. Richard E. Banzal and Pfc. Robert Egner of the U.S. 
Marine Corps. Their technical help, capable assistance, and enthusiastic 
support are deeply appreciated. Photographs of the female of the new 
Vettius species were made by the Smithsonian Institution. 

Determination of plant specimens was made by Dr. Robert Dressler 
of the Smithsonian Institution in the Canal Zone. 


LITERATURE CITED 


BELL, E.L. 1938. A New Genus and Five New Species of Neotropical Hesperiidae 
(Lepidoptera-Rhopalocera). Amer. Mus. Novitates, No. 1013. 11 p. 

1959. Descriptions of Some New Species of Neotropical Hesperiidae 
(Lepidoptera-Rhopalocera). Amer. Mus. Novitates, No. 1962. 16 p. 

Evans, W. H. 1953. A catalogue of the American Hesperiidae. Part III, Pyrginae, 
Section 2. British Museum of Natural History, London. 246 p. pl. 26-53. 
1955. A catalogue of the American Hesperiidae. Part IV, Hesperiinae 

and Megathyminae. British Museum of Natural History, London. 449 p. pl. 
54-88. 
GopMan, F. C. & O. Satvin. 1887-1901. Biologia Centrali-Americana. Insecta. 
Lepidoptera-Rhopalocera. London. Vol. 2: 244-637; Vol. 3: Tabs. 73-106. 
Haywarp, K. I. 1950. Genera Et Species Animalium Argentinorum. Hesperiidae, 
Tomus Secundus, Hesperiinae. Tucuman, Argentina. 347 p. 26 pl. 

Mretke, O. H. H. 1968. Lepidoptera of The Central Brazil Plateau. II New 
Genera, Species and Subspecies of Hesperiidae. J. Lepid. Soc. 22: 1-20. 
Nicotay, S. S. & G. B. Smarty. 1969. A New Subspecies of Pyrrhopyge creon 

(Hesperiidae) from Panama. J. Lepid. Soc. 23: 127-130. 
Weexs, A. G., Jr. 1905. Illustrations of Diurnal Lepidoptera, with Descriptions. 
University Press, Boston. 117 p. 45 pl. 


258 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


THE LIFE CYCLE OF DIRCENNA RELATA (ITHOMIIDAE) 
IN COSTA RICA 


ALLEN M. YOUNG 
Department of Biology, Lawrence University, Appleton, Wisconsin 54911 


This paper summarizes a variety of observations on the life cycle and 
natural history of the neotropical butterfly, Dircenna relata Butler & 
Druce, as studied in Costa Rica. While such studies of Brazilian members 
of the tribe Dircennini have been conducted (Brown & D’Almeida, 
1970), the Central American ithomiid fauna generally remains to be 
studied in this respect. The excellent systematic studies of Central Ameri- 
can Ithomiidae (Fox, 1968) provide a good basis for comparative studies 
on the biology of these butterflies and forms the foundation for examina- 
tion of phylogenetic trends with respect to foodplant exploitation, be- 
havior patterns, and a wealth of other factors underlying population 
biology. This paper represents an effort to study the biology of Costa 
Rican Ithomiidae, and supplements other similar reports (Young, 1972, 
in press). 

METHODS 


Field observations were carried out intermittently during June— 
September 1971 at Bajo la Hondura (San Jose Province), a montaine 
tropical forest locality (800-1000 m. elev.) in central Costa Rica. Ob- 
servations were confined to a small breeding population of D. relata 
located at the bottom of the steep ravine at Bajo la Hondura; this popula- 
tion was discovered in exposed second-growth brush bordering the Rio 
Claro in the ravine. 

Field studies or observations consisted of: (1) description of the life 
cycle, including estimation of developmental time and studies of larval 
hostplant specificity, (2) analysis of oviposition behavior, and (3) notes 
on larval behavior. Developmental time and larval hostplant specificity 
were studied in the laboratory. For these purposes, larvae at low densities 
were confined in tightly sealed plastic bags containing cuttings of host- 
plants. 


RESULTS 
Life Cycle and Developmental Time 


The egg (Fig. 1,4) is barrel-shaped and truncated at the base. It 
bears several deep vertical ribs and many less distinct horizontal ribs. 
The egg (0.7 mm X 0.6 mm) is cream-colored when first laid but becomes 
deep yellow within a day. The vertical ribs remain cream-colored as the 


VOLUME 27, NUMBER 4 259 


egg changes color. The first instar larva is light green with three dorsal 
rows of yellow spots. The two outer rows appear continuous and the 
head is uniformly light green. The larva measures about 3.5 mm by the 
first molt. The second instar (Fig. 1,B) is remarkably similar to the 
first but the central row of yellow spots now becomes interrupted with a 
complex patchwork of yellow and black spots. It is difficult to describe 
the body color pattern in terms of segments because these have become 
strongly subdivided in the Dircennini. The spots in the central row be- 
come square-shaped while the two outer rows continue to remain less 
distinct. The second instar is about 8 mm long by the second molt. 

The third instar (Fig. 1,C) is very similar to the previous one, although 
now the general body color becomes dark green. The yellow patchwork 
and mottling of previous instars is retained in this instar. By the third 
molt the larva is about 13 mm long. The fourth instar (Fig. 1,D) re- 
sembles the previous instar and measures about 19 mm long by the 
fourth molt. The fifth and final instar (Fig. 1,E) is lighter green (like 
the second instar) and the yellow and black patchwork has become 
more diffuse. This instar attains a size of about 24 mm in length by 5 mm 
in width before changing into a truly mobile prepupa. 

The prepupa is uniformly translucent light green with the yellow mark- 
ings barely visible. It is an extremely active form but seldom crawls off 
the hostplant for pupation. The pupa measures about 11 mm in length 
and 7 mm in width (dorsoventrally in the thoracic region) and its colora- 
tion is somewhat variable; on light backgrounds the pupa is light green 
with gold markings (Fig. 1.F) while on dark backgrounds the pupa is 
reddish-brown with gold markings (Fig. 1G). The gold markings in 
both forms are confined to rings around abdominal segments, legs, thorax, 
and wing veins. The cremaster in both cases is pink. To the human 
observer the pupae of this butterfly are very dazzling and reflective and 
when several are found together in the field they give the appearance of 
large drops of water reflecting sunlight. 

Adults of both sexes are similar in size, having a forewing radius of 
about 31 mm (N= 23). There is very little sexual dimorphism in the 
delicate translucent orange ground coloration of the wings (Fig. 1,H). 
Descriptions of wing coloration are given by R. Haensch in Seitz (1924) 
and more recently in greater detail by Fox (1968). 

Developmental time from egg to adult is summarized in Table 1. 
Despite a mean developmental time of 25 days, females take about a day 
longer to complete ontogeny. In a total of 50 emergences, the sex ratio 
did not deviate significantly from unity, there being 28 males and 22 


females. 


260 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


7 


Fig. 1. Life cycle of Dircenna relata Butler & Druce: (A) eggs; (B—-E) second, 
third, fourth, and fifth instars, respectively; (F-—G) light and dark forms of pupa, 
respectively; and (H) adult female (above) and male (below). 


VoLUME 27, NUMBER 4 261 


TaBLE 1. The developmental time* of Dircenna relata Butler & Druce (Ithomiinae ) 
on its larval foodplant, Solanum hispidum (Solanaceae ). 


Instars 
Statistics Egg 1 2 3 4 5? Pupa 
Mean (days) 5 2 2 2, 3 5 6 
ae S15. 0.3 0.1 0.4 0.2 0.2 0.82 0.8 
No. individuals studied 42, Si 37 37 34 34 34 


Died 0 6° 0 0 $ 0 0 
a Measured in the laboratory conditions (20—24°C). 
b Includes an active prepupa which lasts about one day. 


¢ This mortality apparently due to handling effects. 
d Females longer in development, by about 1 day, occurring in the fifth instar. 


Larval Host Plant Specificity 


At Bajo la Hondura the major larval hostplant is Solanum hispidum 
Pers. (Solanaceae) which grows along the edges of the Rio Claro (Fig. 
2). This plant (see Standley, 1937 for description) is common in thickets 
of second-growth vegetation in central Costa Rica between 1300-2000 m 
elevation, and occurs as an armed shrub or small tree (1.5-3.0 m tall) 
(D.C. Wasshausen, pers. comm.). Geographically this species occurs 
northward to Mexico and perhaps southward into Panama. At the study 
site individual plants are highly scattered along the river edge (Fig. 2) 
and seldom occur in homogeneous patches. 

Although this was the only hostplant I found for D. relata at Bajo la 
Hondura, caterpillars complete development successfully (with same 
developmental time) in the laboratory on several other species of 
Solanum. I tested the following species from various parts of Costa Rica: 
(1) S. laurefolium (from Vara Blanca, Heredia Province, 600 m elev.), 
and (2) S. orchraceo-ferrugineum (from San Miguel, Heredia Province, 
100 m eleyv.). Both of these species appear very similar to S. hispidum 
both in life form and distribution of spines on leaves and stems. Neither 
one occurs at Bajo la Hondura, however. 


Larval Behavior 


Larvae devour their empty egg shells and always remain on the ventral 
surface of the leaf. Feeding may begin virtually anywhere on a leaf (i.e. 
first instars do not necessarily feed at the edge of a leaf). Although 
several caterpillars may be found on a single leat of S. hispidum, they 
are never gregarious. This is in sharp contrast to the larvae of Mechanitis 
isthmia isthmia Bates (Ithomiidae), in which both feeding and resting 
are highly gregarious. But nongregarious larval behavior also occurs in 
other species of Costa Rican ithomiids such as Godyris caesiopicta 


262 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Naas 


Fig. 2. Habitat and hostplant of Dircenna relata Butler & Druce at Bajo la 
Hondura, Costa Rica. Both adult D. relata and individuals of Solanum hispidum (large 
inset to the left), the major larval host plant, are distributed sparsely along the rocky 
edge of the Rio Claro (remainder of photo). The hostplant does not occur in the 
primary-growth forest lining the Rio Claro further back. 


Niepett* (Young, in prep.), Napeogenes tolosa amara Godman (Young, 
in prep.), Hymenitis nero (Hewitson) (Young, 1972), and Pteronymia 
notilla Butler & Druce (Young, in press). 

Larvae appear to be cryptically colored when on the undersides of the 
very hairy leaves of S. hispidum, and younger instars are especially diffi- 
cult to find in the field. All instars are generally diurnal feeders, resting 
at night near major veins of leaves. In the field, larvae of all instars are 
most frequently encountered on the lowest leaves of S. hispidum, and 
mixed age groups are frequently seen. Larvae have never been found on 
apical leaves. 

Neither predation nor parasitism on larvae have been observed in the 
field. Of a total of 37 additional larvae (of all instars) collected in the 
field and reared in the laboratory, none gave rise to parasitic flies or 
wasps. 

* This is the old name for this subspecies, and it was brought to my attention by Keith S. 


Brown, Jr., who encountered it in Lepidoptera Niepettiana (Vol. 2); it was renamed by R. M. 
Fox as G. z. sorites. 


VOLUME 27, NUMBER 4 263 


Oviposition Behavior 


Oviposition sequences were witnessed on several occasions, first on 
11 July 1971 at 1220 (CST). A female patrols around the hostplant, lights 
on the ventral surface of a large, older leaf near the ground, and walks 
around, depositing several eggs. Eggs are loosely clustered (Fig. 1,A) 
and many are only loosely attached to the leaf by entwining hairs (i.e. 
there is no actual adhesion of eggs to the leaf). Eggs are not clustered 
in an orderly fashion as seen in other species of ithomiids (e.g. Mechanitis 
isthmia) but rather are scattered haphazardly on the leaf. It is possible 
that some eggs may become jarred loose, perhaps resulting in substantial 
mortality. However, it is equally possible that eggs are held securely by 
the very dense leaf hairs (Fig. 1,A). 

Eggs frequently are laid on leaves where eggs had been desposited 
previously. The same female will often return to the same leaf to lay 
additional eggs over several successive oviposition sequences. Eggs 
are not laid on the younger apical leaves of the hostplant. An individual 
female, on the average lays between 1 and 14 eggs during a single visit 
to a leaf. Females always “drum” a leaf with the forelegs before laying 
an egg. Oviposition at Bajo la Hondura probably occurs throughout the 
day, although it is most frequent between 1130 and 1430. 

Undoubtedly ovipositing females are quite vagile owing to the highly 
dispersed distribution of S. hispidum. A female ovipositing on a given 
individual hostplant will fly away after several oviposition sequences 
(i.e. after about 10 minutes of interrupted patrolling) and may search 
for other individual plants on which to oviposit. 

Courtship activity has never been observed in the immediate vicinity 
of hostplants. This behavior may occur in the understory of the primary 
growth forest that covers the sides of the ravine at Bajo la Hondura. 


DISCUSSION | 


Although the Ithomiidae are generally solanaceous-feeding butterflies 
in the caterpillar stage, rendering at least one species unpalatable to 
some predators (Brower & Brower, 1964), it is apparent that this group 
has experienced several lines of phylogenetic diversification into several 
tribes differing substantially in morphology of both adults and imma- 
tures (Fox, 1940, 1967, 1965; Brown & D’Almeida, 1970). One of these 
tribes is the Dircennini, which contains several genera including Cera- 
tinia, Dircenna, and Pteronymia (see Brown & D’Almeida, 1970, p. 14-15). 
Since these and other genera within this tribe presumably share some 
portion of their evolutionary history, it is interesting to discuss the life 
cycle and natural history data for D. relata in terms of similar studies of 


264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


other genera in the tribe, and genera in other tribes. It is realized, how- 
ever, that contemporary differences in the biological properties of dif- 
ferent species and genera in any animal group must not only be molded 
by evolutionary history, but also by simultaneous ecological factors 
operating on breeding populations (Birch & Ehrlich, 1967). The follow- 
ing comparisons are limited to a few illustrative examples that emphasize 
both similarities and divergences in life cycle and natural history. 

Young (in press) found that Pteronymia notilla at Cuesta Angel in 
Costa Rica has an egg-adult developmental time of about 30 days on 
Cestrum megalophyllum (Solanaceae), which is about five days longer 
than the developmental time of D. relata on Solanum hispidum, under 
similar conditions. Brown & D’Almeida (1970) report a developmental 
time of about 45 days for Ceratiscada canaria on Solanum caavurana 
under presumably laboratory conditions in Rio de Janeiro. Young (1972) 
found the developmental time of Hymenitis nero (not in the Dircennini) 
on Cestrum standleyi (eggs from a Cuesta Angel population) to be about 
30 days in the laboratory. Based on these very fragmentary records in- 
volving different foodplants, it is still not possible to assign generic or 
tribal differences in developmental time for these ithomiids. With 
respect to the duration of the egg stage, if we assume equal environ- 
mental conditions for the forementioned species, the genera Dircenna, 
Pteronymia, and Hymenitis (as represented by the species discussed) all 
have an incubation period of 4-5 days; but the incubation period of 
Ceratiscada is 9-10 days (Brown & D’Almeida 1970). It is very likely 
that there exists a wealth of unpublished measurements of egg-adult 
developmental in many genera and species of ithomiids on various food- 
plants by workers in Central and South America; I would like to make a 
plea to bring all of these data together and look for a phylogenetic 
pattern of developmental time (within tribes), especially where similar 
foodplants are involved. If more information is obtained regarding leaf 
toughness and secondary compounds for different genera of Solanaceae, 
such data would also shed light on developmental time differences re- 
sulting from foodplant differences rather than evolutionary history among 
the ithomiids. Since incubation period may be only indirectly influenced 
by foodplant (in cases where the female’s foodplant affects egg dura- 
tion), it may be best to examine differences in duration of the egg stage 
among genera and tribes. 

Rather pronounced differences in external morphology of immatures 
are also evident among different genera of ithomiids. For example, the 
egg of D. relata is very different in appearance from that of Hymenitis 
nero, Pteronymia notilla, and Ceratiscada canaria. First, the egg of D. 


VOLUME 27, NUMBER 4 265 


relata is deep yellow, essentially spherical, and has a system of external 
horizontal and vertical ribs; the egg of P. notilla is white and more 
oblong in shape than the egg of D. relata (Young, in press). Further- 
more, it lacks very definite horizontal ribs. While the ege of H. nero 
is also uniformly white, it’s shape approaches that of D. relata and it 
has a similar array of vertical ribs as seen in P. notilla. Although the egg 
of C. canaria is white, the shape and distribution of vertical and horizontal 
ribs (grooves) are similar to that of D. relata (as determined from the 
egg description of C. canaria given in Brown & D’Almeida, 1970). The 
size of the egg of D. relata is closer to that of C. canaria while those of 
the other two genera (Pteronymia and Hymenitis) are smaller. 

The coloration and extent of hair on body segments varies very 
strikingly when Dircenna (as seen by D. relata) caterpillars are compared 
to caterpillars of other genera such as Hymenitis, Pteronymia, and 
Ceratiscada. While it is not necessary to go into a detailed account of 
these differences here, the caterpillars of D. relata are hairy while those 
of H. nero, P. notilla, and C. canaria generally lack hairs on body seg- 
ments in all instars (see larval descriptions in Brown & D’Almeida, 1970; 
Young, 1972, in press, and present report). Furthermore, the color 
patterns of the body in caterpillars of H. nero, P. notilla, and C. canaria 
consist of a dark green background color with lighter lateral stripes, 
usually yellow or light blue; the color pattern of the caterpillars of D. 
relata is very different, being a linear patchwork of yellow spots on a 
light green and speckled background color (Fig. 2, B-E). 

The pupae of these genera are also different. The major difference 
is in the extent of reflective coloration on the external surfaces. The 
pupa of P. notilla is generally devoid of gold or silver markings, with 
the color being translucent yellowish-green (Young, in press). The 
pupae of both H. nero and C. canaria are remarkably similar in terms of 
the highly reflective silver cover on the wings (Young, 1972; Brown & 
D’Almeida, 1970). The network of reflective gold coloration on the pupa 
of D. relata (Fig. 2,F,G) is different from the previous two color patterns. 
It is interesting that there occurs considerable similarity between the 
pupae of Hymenitis and Ceratiscada since these genera belong to dif- 
ferent tribes (Fox, 1940), while considerable divergence in appearance 
of the pupa occurs within the Dircennini, as typified by D. relata, P. 
notilla, and C. canaria. 

Finally, it is interesting to consider some differences in larval and 
adult behavior among different genera. The ithomiid Mechanitis isthmia 
(see Fox, 1967) lays its eggs in tight clusters on the food plant (pers. 
obs., Costa Rica and El Salvador), but most other ithomiids in other 


266 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


genera do not show this type of oviposition behavior. For example, H. 
nero, P. notilla, C. canaria generally lay their eggs singly but D. relata 
exhibits loose clustering of eggs (Fig. 2,A). The size and orderliness of 
egg clusters in D. relata are very different from M. isthmia. From recent 
considerations (Labine, 1968) it would seem that marked differences in 
oviposition behavior accompany differences in egg productivity of in- 
dividual females in different species, and differences in the spatial 
distribution of preferred foodplants (for egg-laying) in different species. 
It is therefore difficult to obtain phylogenetic correlations for egg-laying 
behavior in butterfly groups such as the Ithomiidae. 

It follows from the oviposition behavior that different genera have 
different degrees of larval gregariousness on the foodplant. The cater- 
pillars of M. isthmia form very tight and coordinated groups on leaves 
of the foodplant (pers. obs.) in which there is group feeding and resting 
periods. This is very different from group organization in D. relata 
where each caterpillar is very individualistic for feeding, resting, and 
general locomotor activities. The more usually encountered situation, as 
seen in H. nero, P. notilla, and C. canaria, is where caterpillars occur 
singly on leaves and there is little or no evidence of group interaction of 
any sort when more than one individual is on the same leaf. For the 
ithomiids, it is clear that the degree of group interaction in caterpillars 
is a direct consequence of the type of oviposition. 


SUMMARY 


(1) The life cycle of the itomiid butterfly, Dircenna relata Butler & 
Druce, is described for the first time. 

(2) The egg-adult developmental time on a natural foodplant, Solanum 
hispidum Pers. (Solanaceae) is about 25 days under constant laboratory 
conditions. But caterpillars are capable of successfully completing de- 
velopment on several other species of Solanum. 

(3) The eggs are laid in loose clusters on leaves of the foodplant, and 
the number of eggs in each cluster is very variable. Eggs are generally 
laid on lower, older leaves and the caterpillars subsequently form loose, 
disorganized aggregations. There is no evidence of coordinated activity 
patterns (feeding, resting, etc.) among individuals within a “group.” 
The loose association of the caterpillars is interpreted as a direct result 
of the loose clustering of eggs. 

(4) An attempt is made to compare morphology, developmental time, 
and behavior patterns among different genera of Ithomiidae within the 
tribe Dircennini and outside it. In the absence of further data on these 
species and others, it is difficult to separate phylogenetic from contempo- 


VOLUME 27, NUMBER 4 26 


~l 


rary ecological selective forces for the characteristics studied. A plea is 
made for the bringing together of unpublished data on developmental 
time and foodplants as gathered by different workers in Central and 
South America, so that a more grandiose attempt to look for phylogenetic 
trends in these characteristics underlying population biology and habitat 
selection can be made. 


ACKNOWLEDGMENTS 


This is a contribution from a College Science Improvement Grant 
(COSIP, GY-4711) awarded to Lawrence University. Research facilities 
in Costa Rica were provided by Dr. J. Robert Hunter of the Associated 
Colleges of the Midwest. Technical assistance in the field and laboratory 
was given by Patrick Eagan. Dr. D. C. Wasshausen (Smithsonian) 
identified all Solanaceae. Dr. Keith S. Brown, Jr. (Rio de Janeiro) also 
identified the species studied, and as a reviewer, made very helpful sug- 
gestions on the manuscript. 


LITERATURE CITED 


Bircy, L. C. & P. R. Emruicu. 1967. Evolutionary history and population biology. 
Nature (London) 214: 349-352. 

Brower, L. P. & J. V. Z. BRowER. 1964. Birds, butterflies, and plant poisons: a 
study in ecological chemistry. Zoologica 49: 137-159. 

Brown, K. S., Jn. & R. F. D’Atmema. 1970. The Ithomiinae of Brazil (Lepidop- 
tera: Nymphalidae). II. A new genus and species of Ithomiinae with comments 
on the tribe Dircennini D’Almeida. Trans. Amer. Entomol. Soc. 96: 1-17. 

Fox, R. M. 1940. A generic review of the Ithomiinae. Trans. Amer. Entomol. 
Soc. 66: 161-207. - 

1967. A monograph of the Ithomiidae (Lepidoptera). Part II. The 

tribe Mechanitini Fox. Mem. Amer. Entomol. Soc. 22. 190 p. 

1968. Ithomiidae (Lepidoptera: Nymphaloidea) of Central America. 
Trans. Amer. Entomol. Soc. 94: 155-208. 

Lapine, P. A. 1968. The population biology of the butterfly, Euphydras editha. 
VIII. Oviposition and its relation to patterns of oviposition in other butterflies. 
Evolution 22: 799-805. 

Serrz, A. 1924 (ed.). The Macrolepidoptera of the World. Vol. 5. The American 
Rhopalocera. Stuttgart, A. Kernan Verlag. 1139 p. 

Younc, A. M. 1972. On the life cycle and natural history of Hymenitis nero 
(Lepidoptera: Ithomiinae) in Costa Rica. Psyche 79: 254-294. 

Notes on the biology of Pteronymia notilla (Ithomiidae) in a Costa Rican 

mountain forest. J. Lepid. Soc. (in press). 


268 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


THE GENUS EUMAROZIA HEINRICH (OLETHREUTIDAE ) 


J. F. Gates CLARKE 
National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 


The genus Eumarozia was proposed by Heinrich (1926: 110) for the 
single species Grapholitha (Poecilochroma) malachitana Zeller, and 
untill now, has remained monobasic. 

The second species, described below, first came to our attention in 
1970 and again in 1972, when it was submitted for determination. The 
third species which I now include in this genus was described by Meyrick. 
These species are of particular interest because they greatly extend the 
range of Eumarozia. 


Eumarozia malachitana (Zeller ) 
Fig. 2 


Grapholitha (Poecilochroma) malachitana Zeller, 1875, Verh. zool.—bot. Ges. Wien 
M5)p DSP. 

Penthina malachitana (Zeller), Fernald, 1882, Trans. Amer. Ent. Soc. 10: 33 (no. 
200); Fernald, 1891, in Smith, List Lepid. Bor. Amer., 91 (no. 4818). 

Olethreutes malachitana (Zeller), Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull. 52: 
452 (no. 5044); Kearfott, 1903, in Smith, Check List Lepid. Bor. Amer., 101 
(no. 5447); Walsingham, 1914, in Godman and Salvin, Biol. Centr. Amer. 42: 
(Lepid. Heter. 4): 252; Forbes, 1923, Mem. 68, Cornell Univ. Agr. Exp. Sta., 
457; Bottimer, 1926, J. Agric. Res. 33(9): 817. 

Argyroploce malachitana (Zeller), Barnes & McDunnough, 1917, Check List Lepid. 
North America, 168( no. 6854). 

Eumarozia malachitana (Zeller), Heinrich, 1926, U.S. Nat. Mus. Bull. 132: 111, 
figs. 60, 194, 413; McDunnough, 1939, Mem. So. Calif. Acad. Sci. 2(1): 
40(no. 6634); Jones, 1943, Lepid. Nantucket and Martha’s Vinyard Islands, 
Mass., 149, 206; McKay, 1959, Can. Entomol. 91(Suppl. 10): 159, fig. 155; 
Kimball, 1965, Arthropods of Florida and Neighboring Land Areas. Lepid. 
Florida, 256(no. 6634). 


Type: British Museum (Natural History ). 

Type locality: Missouri. 

Distribution: The species malachitana is widely distributed in the eastern and 
southern United States and its range extends into Mexico. According to specimens 
in the U.S. National collection and the collection of Dr. Annette F. Braun, Cincinnati, 
Ohio, the distribution is, alphabetically by states, as follows: ARKANSAS: Devil's 
Den State Park, Fayetteville (June); pisrricr OF COLUMBIA: Washington (Sept.); 
FLORIDA: Lake Placid, St. Petersburg, Sarasota (Feb., Apr.); GEORGIA: Savannah 
(Aug.); mLLINoIs: Quincy (Sept.); INDIANA: Bedford (Sept.); KANSAS: Eureka, 
Pittsburg (June); MARYLAND: Hyattsville, Plummer’s Island (Aug.); MuissouURI: 
Joplin, “Mtn. Grove,” “C. Mo.” (June); NorTH CAROLINA: Highlands, Knotts Is., 
Southern Pines (June, July); oH10: Cincinnati (Sept.); okKLAHOMA: Oklahoma City 
(Sept.); TExAs: Brownsville, Kerrville, Lk. Charlotte, San Benito (Apr.); VIRGINIA: 
Falls Church, Wicomico Ch. (Aug., Sept.). Heinrich (1926: p. 111) also recorded 
the species from “. . . Central America and South America.” There are two speci- 
mens before me from Mexico from Salina Cruz, Oaxaca, and Orizaba (Wm. Schaus 


VOLUME 27, NUMBER 4 269 


coll.). I have seen none from South America. Presumably Heinrich accepted Zeller’s 
original record “Sudamerica.” 

Foodplants: Diospyros virginiana L. (persimmon); Ostrya virginiana ( Mill.) K. 
Koch (hop hornbeam); Philoxerus sp.; Pyrus communis L. (pear); Cassia sp. 
“black sapota” (Achras? sp.). 


Eumarozia beckeri Clarke, new species 
Figs. 1, 3, 4 


Alar expanse 13-15 mm. Labial palpus creamy white; third segment and outer 
side of second, light ochraceous buff. Antenna gray; cilia short, whitish. Head with 
face and vertex buff; posteriorly gray with white-tipped scales. Thorax grayish 
fuscous, tegula grayish fuscous, some scales white-tipped. Forewing ground color 
grayish fuscous; beyond basal fifth of ground color a shining buff transverse fascia 
extends from costa to dorsal margin, the dorsal third of fascia overlaid with leaden 
metallic scales; on costa, preceding the pale fascia, a triangular spot of grayish olive; 
the pale fascia is followed on its outer side by a transverse band of grayish olive, 
the latter edged outwardly with a narrow fascia of leaden-metallic scales; both 
grayish olive areas show a golden sheen in certain lights; outer half of wing deep 
hellebore red variously marked with fuscous; from end of cell at about vein 5, a 
crescentic fuscous mark ending on termen; on tornus a quadrate fuscous spot and a 
fuscous blotch between the latter and the crescentic mark; subterminally a curved line 
of small leaden-metallic spots; around termen a series of 5 or 6 fuscous dots; cilia 
fuscous with some red scales mixed. Hindwing grayish fuscous, slightly darker 
toward termen; extreme costal edge of male buff, followed inwardly with a longitudi- 
nal band of-black scales; cilia grayish with a darker subbasal line. Foreleg gray 
strongly suffused fuscous on outer side; tarsal segments spotted with white; mid- 
and hindlegs leaden gray. Abdomen grayish fuscous to leaden gray with a few 
scattered buff scales ventrally and in tuft. 

Male genitalia (USNM 24095): Harpe with deeply incised neck; cucullus 
rounded, broad; Spc! a single long seta; Spc’ absent; base of cucullus with series of 
strong setae; ventral edge of sacculus with cluster of long setae and inner surface 
setaceous. Gnathos a moderately broad band. Uncus reduced, rounded. Socius a 
setaceous lobe at end of flattened stalk. Vinculum broadly rounded. Tegumen oval. 
Anellus subtriangular strongly attached posteriorly to a scoop-shaped member 
through which the aedeagus articulates. Aedeagus slender, curved; vesica armed 
with a cluster of long cornuti. 

Female genitalia (USNM 24096, 24097): Ostium produced dorsally, nearly 
rectangular. Antrum not differentiated. Inception of ductus seminalis from anterior 
end of ductus bursae, slightly posterior to junction with bursa copulatrix. Ductus 
bursae sclerotized for most of its length, slender. Bursa copulatrix membranous. 
Signa two triangular plates with serrate edges. 

Holotype: USNM No. 72436. 

Type locality: Costa Rica, Turrialba. 

Distribution: Costa Rica. 

Foodplant: Juglans olanchanum Standl. & L. Wms. 

Described from the holotype ¢, 55 ¢¢ and 55 92°@Q paratypes from the type 
locality dated 29 August to 3 October 1972, V. O. Becker; and 2 ¢ ¢ paratypes Costa 
Rica, Turrialba (12 January 1970), no collector, rf. Juglans. Paratypes are distributed 
as follows: 5 24,5 22 (25-30 September 1972) in Museu de Entomologia de la 
Universidad de Costa Rica; 48 6 6, 48 22 (9 September—3 October 1972) Becker 
Collection in the Department de Zoologia da Universidade Federal do Parana, Brasil; 
266,292 (12 January 1970 and 29 August 1972), U.S. National Museum. 


The two species treated here are closely allied but may be distinguished 


270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Vy 


Yi) 


Fig. 1. Eumarozia beckeri, new species. Turrialba, Costa Rica. Fig. 2. Eumarozia 
malachitana (Zeller). Devil’s Den State Park, Arkansas. 


VOLUME 27, NuMBER 4 21 


Fig. 3. Eumarozia beckeri, new species. Ventral view of male genitalia with left 
harpe omitted. 


easily on pattern and genitalia. In E. malachitana the olive marking near 
center of forewing is nearly oval in shape, does not reach the dorsal edge 
and is bordered by a slender buff or white line. In the case of beckeri, 
however, the olive marking is in the form of a transverse fascia which 
reaches from costa to dorsum. The male genitalia are similar; Spc' 
(Heinrich, 1926: fig. 60) being prominent in both species; but Spc? is 
absent in beckeri. The only substantial difference between the female 
genitalia lies in the size and shape of the ostium which is much narrower 
and longer in beckeri than in malachitana. 

It gives me great pleasure to name this species in honor of Victor Omar 
Becker who collected and reared the major part of the type series. 


Eumarozia elaeanthes (Meyrick), new combination 


Argyroploce elaeanthes Meyrick, 1927, Exotic Microlepid. 3: 340. 

Olethreutes elaeanthes (Meyrick), Clarke, 1958, Catalogue of the Type Specimens 
of Microlepidoptera in the British Museum (Natural History) described by 
Edward Meyrick, 3: 507, pl. 252, figs. 1—la. 


Type: British Museum (Natural History). 
Type locality: Bolivia, Andes, 10,000 feet. 
Distribution: Known only from the type locality. 


212 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


VOLUME 27, NUMBER 4 273 


Foodplant: Unknown. 


Meyrick described elaeanthes from the unique type, and as far as I 
know, the type is the only known specimen. The genitalia of the male 
elaeanthes, which I figured (1958: pl. 252, figs. 1-la), are typical for 
the genus except Spc? ( Heinrich, 1926: fig. 60) is missing, as in beckeri. 
Inside costa, at basal third of harpe, is a cluster of strong setae, and from 
the ventral edge of sacculus is a series of long, hairlike setae in elaeanthes 
which are absent in malachitana. In the forewing of elaeanthes there 
is no white line delineating the olive green area as in malachitana. 1 
have not figured elaeanthes here; the figure cited is adequate for 
recognition. 


. é 
e 
e e re) 
) Or 
ES 
a BES oh, 
e £ malachitan a (Zeller) ae 
x E. beckerl, new species he! po pat | 
E. elaeanthes (Meyrick) - Bolivian p 5 


Fig.5. Eumarozia distribution. 


ACKNOWLEDGMENTS 


The photographs for this paper were made by Victor E. Krantz and 
the drawings were made by George Venable, both on the staff of the 
Smithsonian Institution. 


< 


Fig. 4. Eumarozia beckeri, new species. Ventral view of female genitalia. 


274 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


REFERENCES 


CLARKE, J. F. Gates. 1958. Catalogue of the Type Specimens of Microlepidoptera 
in the British Museum (Natural History) described by Edward Meyrick 3: 
1-600, pl. 1-298. London. 

Hernricu, C. 1926. Revision of the North American Moths of the Subfamilies 
Laspeyresiinae and Olethreutinae. U.S. Nat. Mus. Bull. 132. 216 p., 76 pl. Wash- 
ington, D.C. 

Kimpatut, C. P. 1965. Arthropods of Florida and Neighboring Lands Areas. 
Lepidoptera of Florida. An Annotated Check List. v + 363 p. 26 pl. Gainesville, 
Florida. 

WALSINGHAM (LORD THOMAS DE Grey). 1909-1915. In Godman and Salvin, 
Biologia Centrali-Americana, 42 (Lepidoptera-Heterocera) 4: i-xii, 1-24(1909); 
25-40( 1910); 41-112( 1911); 113-168(1912); 169-224( 1913); 225-392(1914); 
393-482( 1915); figs. 1-30, pls. 1-10. London. 


THREE NATURAL HYBRIDS OF VANESSA ATALANTA RUBRIA 
<x CYNTHIA ANNABELLA (NYMPHALIDAE) 


THomas FE. Dimock 
464 Fairfax Avenue, Ventura, California 93003 


In an effort to better understand the relationship and variation of two 
local butterflies, Vanessa atalanta rubria (Fruhstorfer) and Cynthia an- 
nabella Field, not only as adults but also as larvae, I have been rearing 
to maturity all vanessid larvae encountered on their various foodplants. 
In the vicinity of northeastern Thousand Oaks, California, foodplants for 
C. annabella are Althaea rosea (L.) Cav. (Hollyhock), Malva parviflora 
L. (Cheeseweed), both Malvaceae; and Urtica holosericea Nutt. (Sting- 
ing Nettle), Urticaceae. Urtica holosericea grows abundantly along an 
intermittent creek, which flows through the Lang Ranch property, and 
is the only local foodplant for V. a. rubria. The surrounding floral com- 
munities are coastal sage scrub, chaparral, southern oak woodland (Munz, 
1968), and annual pasture grassland. The area is but a half mile from 
city subdivisions. 

On 3 April 1972 I collected two second instar larvae on two Urtica 
plants occupying a position further downstream than any other Urtica. 
Several leaves were taken as a food supply. In later instars both larvae 
appeared at a glance to be C. annabella, so when all the Urtica leaves 
were consumed the larvae were given leaves of Althaea rosea, which 
they readily devoured to maturity. At this time one larva hung and 
pupated, and later emerged as C. annabella. The other continued growth 
and surpassed in size all other C. annabella reared to date. Finally it 
pupated, and when the hardened pupa was observed, characteristics of 


VOLUME 27, NUMBER 4 NES, 


both V. a. rubria and C. annabella were displayed. I then decided it 
would be wise to take some notes of description in case the specimen 
proved to be a hybrid, which it was. It is regrettable that closer attention 
was not given to the larva, but as it fed and simply appeared to be C. 
annabella, it was assumed to be merely a giant individual. Therefore, 
the following larval description is written from memory and is very 
general. 


Hybrid #1. Mature larva. Coloration within usual range of variation shown by 
C. annabella and V. a. rubria. A dark form, ground color dark grey, with a series of 
indistinct subdorsal black patches (as in some V. a. rubria). Supralateral rusty- 
orange coloration, typical of many C. annabella, very reduced. Lateral line dull 
flesh-colored. Spines black. Shape as in C. annabella. Pupa. Shape as in V. a. 
rubria, abdominal segments flattened laterally but not to extent shown by V. a. 
rubria. Ground color tan or light brown. Most obvious features are subdorsal pair 
of white spots on metathorax, first, and second abdominal segments (saddle 
spots’) (these are bright gold in V. a. rubria and white in C. annabella). Also a 
subdorsal pair of smaller white spots on mesothorax, and middorsal point on this 
segment lighter than ground color. The pair of subdorsal points, and the single 
row of middorsal points on abdominal segments 3 through 7, dull orange. Supra- 
lateral gold spot which occurs on many V. a. rubria on the fourth abdominal segment 
here present only as yellowish spot lighter than ground color. Grey lateral line on 
the abdominal segments. Only marking on wing case a thin, black vein line on discal 
cell at base of M:-Cu:. Pupal measurements: length 21 mm, width 8 mm, depths: 
thorax 8 mm, saddle 6 mm, abdomen 8 mm. (For comparison, pupal measurements 
of the parent species (average of five specimens each) are: V. a. rubria: length 
21.7 mm, width 8 mm, depths: thorax 8 mm, saddle 6.3 mm, abdomen 8 mm. C. an- 
nabella: length 19 mm, width 7 mm, depths: thorax 6.25 mm, saddle 5.25 mm, 
abdomen 7.25 mm.) Pupal duration, 9 days at room temperature. V. a. rubria and 
C. annabella both have pupal durations of 8 to 9 days at room temperature. 


Following the pupation of this first individual, with the realization that 
it was a hybrid, I returned to the foodplant locality and collected all 
remaining larvae. Eleven were found: two hybrids and nine V. a. rubria. 
Again, however, the hybrids were not suspected until after the final 
moult. These two individuals differed from the previous hybrid, so brief 
descriptions are given for both. 


Hybrid #2. Mature larva. Ground color black as in some V. a. rubria. Lateral 
line pale yellow. Head entirely black. Spines black. Otherwise like V. a. rubria. 
Pupa. As in hybrid #1. Pupal measurements: length 21.3 mm, width 7.5 mm, 
depth: thorax 7.5 mm, saddle 6 mm, abdomen 7.5 mm. Pupal duration 8 days. 

Hybrid #3. Mature larva. Ground color dark brown with numerous tiny yellow 
spots laterally and dorsally except for narrow middorsal line of solid ground color. 
A rusty-orange spot on each abdominal segment between subdorsal and supralateral 
spines. Lateral line large and yellow, interrupted at each segment by a lateral spine 
base of the ground color. Head entirely black. Spines black. Shape as in C. annabella. 
Pupa. As in hybrid #1. Pupal measurements: length 20 mm, width 7 mm, depths: 


we 


thorax 7 mm, saddle 5.5 mm, abdomen 7 mm. Pupal duration 8 days. 


Adult hybrids #1 and #2 emerged without problems. Hybrid #3 had 


276 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 1-5. Vanessa atalanta rubria, Cynthia annabella, and three natural hybrids: 
(1) V. a. rubria, male, ex-larva on Urtica holosericea, northeast Thousand Oaks, 
Ventura Co., Calif., elev. 980 ft., 17 April 1972; (2) C. annabella, male, ex-larva 
on U. holosericea plus Althaea rosea, northeast Thousand Oaks, Ventura Co., Calif., 
3 April 1972; (3) hybrid #1, male, same data as C. annabella; (4) hybrid #2, male, 
same data as V. a. rubria; (5) hybrid #3, male, same data as V. a. rubria. 


to be assisted by the author, at the expense of the pupal shell, from the 
terminal four pupal shell segments, as these failed to separate from the 
body of the adult. During the last day of pupal duration the abdomen 
failed to darken while coloring developed normally over the remainder 
of the pupa. Probably as a result of this trouble, the butterfly, after 
extraction from the pupal shell, failed to expand its hindwings to their 
full extent. 


VOLUME 27, NUMBER 4 


Figs. 6-10. Undersides of corresponding specimens in Figs. 1-5. 


Figs. 1-10 depict the parent species and the three hybrid specimens. 
The color of the subapical bar on the forewing upperside is white in V. a. 
rubria and orange in C. annabella. In the hybrids this bar is only slightly 
lighter orange than in C. annabella. An interesting feature on the under- 
side of the left forewing of hybrid #3 is a streak of orange in the apical 
area. All three hybrids are males. 

It is remarkable that hybrids of V. a. rubria and C. annabella occur in 
nature, and I can only speculate on the circumstances responsible for 
their production. Both species are hilltoppers, and it has been shown 
by Shields (1967) that fertilization of the females takes place on hilltops. 


278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


It seems safe to assume that when males of both species are on a given 
hilltop they court and fertilize females of their own species. If, however, 
a virgin female of one species hilltopped and found the summit occupied 
only by one or more males of the other species, interspecific courtship, 
copulation, and fertilization might then occur. There must certainly be 
many behavioral and environmental factors opposing this process, other- 
wise one might expect hybrids to be more common. 

Field (1971) has removed annabella from the genus Vanessa and 
placed it in Cynthia. In view of the natural hybridization of V. atalanta 
rubria and C. annabella, it would seem that the validity of such a 
separation is doubtful. Until further studies of the hybrids can be made, 
including laboratory crosses and backcrosses, together with comparative 
morphological studies of the early stages, it seems best to respect for the 
present Field’s revision. 

The adult hybrid specimens, their mature larval cast skins, and the 
pupal shells of hybrids #1 and #2 are retained in the collection of the 
author. 


ACKNOWLEDGMENTS 


I am most grateful to Mr. Herman G. Real of San Mateo, California, 
for his review and criticism of the manuscript, and to Dr. Theodore D. 
Sargent for his editorial suggestions. 


LITERATURE CITED 


Fretp, W. D. 1971. Butterflies of the Genus Vanessa and of the Resurrected 
Genera Bassaris and Cynthia (Lepidoptera: Nymphalidae). Smithsonian Contr. 
Zool., No. 84. Smithsonian Institution Press, Washington, D.C. 75 p., 160 figs. 

Munz, P. A. 1968. A California Flora. University of California Press, Berkeley 
and Los Angeles. 1681 p. 

SHIELDS, O. 1967. Hilltopping. J. Res. Lepid. 6: 69-178. 


NOTES AND NEWS 


Erratum 


In my paper, “Observations on some Phycitinae (Pyralidae) of Texas with descrip- 
tions of two new species,” (J. Lepid. Soc. 24: 249-255, 1970), the species 
Dioryctria Auranticella (Grote) is reported in error. Dr. E. G. Munroe, who saw 
the three specimens, when he examined my collection in early 1973, identified them 
as Dioryctria rossi Munroe. 


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


VOLUME 27, NUMBER 4 279 


LIFE HISTORY OF CALLOPHRYS S. SHERIDANII (LYCAENIDAE) 
AND NOTES ON OTHER SPECIES! 


CuirForD D. FErris? 
College of Engineering, University of Wyoming, Laramie, Wyoming 82070 


Although Callophrys sheridanii sheridanii (Edwards) (see Brown & 
Opler, 1970, concerning authorship) was described in 1877, little was 
known until recently about its life history. Eff (in Brown et al., 1957) 
reported the foodplant as Eriogonum umbellatum Torr. The type locality 
for this butterfly is “Big Horn Mountains,” near Sheridan, Wyoming. 
The present study is based upon live material collected in the vicinity of 
Pole Mountain (East of Laramie), Medicine Bow National Forest, ca. 
8500’, Albany Co., Wyoming, April-May, 1971-72. 

A few ova of C. sheridanii were secured in the field. Live females were 
captured and caged over the hostplant. Very few additional ova were 
obtained in this manner, however, as the females were very reluctant to 
Oviposit in captivity, even when placed with the foodplant and various 
nectar sources in a large breeding cage out of doors. 

Females appear to oviposit selectively upon E. umbellatum. No ova 
or signs of larval feeding were observed upon two other species of 
Eriogonum, E. flavum Nutt. and E. subalpinum Greene, which are 
sympatric with E. umbellatum. In the field, ova were found on the 
largest of the plants only. Only one or two eggs were found per plant. 

In April and May, when the ova are deposited, E. umbellatum is a low- 
growing trailing plant. Some plants spread out to a foot or more in 
diameter. The rosettes are not yet well developed, and the foot-high 
flower stalks do not appear until June and July. The plant is quite 
common on open hillsides in the Transition Zone in various areas of 
Wyoming. 


Description of Immature Stages 


Ovum: Eggs a very pale green, oblate spheroids approximately 0.8 mm in diam- 
eter; laid singly at the base of a leaf rosette, on the upper surface of the stem; color 
blends with color of new buds on the plant. Depending upon temperature, eggs may 
remain up to a week before hatching. Prior to emergence of larva, ova turns a 
milky white. 

First Instar: First instar larvae pale greenish-white and covered with long fine 
black hairs; head light green. Larvae do not eat egg shells. Stadium one lasted 2-3 
days and larvae increased in length from about 1-3 mm. 


1 Published with the approval of the Director, \Wyoming Agricultural Experiment Station, as 
Journal Article JA 560. : A 
2 Research Associate, Allyn Museum of Entomology, Sarasota, Florida. 


280 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


g A 

Fig. 1. Callophrys s. sheridanii and its foodplant: (a) E. umbellatum (in situ) 
as it appears in April; (b) leaf damage (in situ) produced by sheridanii larva; 
(c) larva-damaged leaves; (d) ovum of C. s. sheridanii; (e) early 4th instar larva; 


(£) pupa; (g) mature (4th instar) larva, dorsal view; (h) same, lateral view, scale 
divisions are 1 mm; (i) dorsal and lateral views of late third instar larva. 


Second Instar: Stadium two larvae a uniform green, covered with short, stiff 
black-tipped spines; black dots visible at bases of spines. This stage lasted from 
3-4 days and larvae increased in length to 5 mm. 

Third Instar: Third instar larvae similar to second instar, but stiffer spines, pink 
at base. As larvae matured, two rows of luminous white spots appeared (Fig. 1, i); 


VoLUME 27, NuMBER 4 2981 


sad (b) 


(G7 


Fig. 2. Callophrys s. sheridanii: (a) 1st instar larva feeding, and cross-section 
through a middle segment; (b) lateral and dorsal views of 2nd instar larva, and 
cross-section through a middle segment; (c) dorsal view of 3rd instar larva, and 
cross-section through a middle segment, also lateral view of head (black) tucked 
under frontal segment. 


not surface spots, but appearing to be well below skin surface. Stadium three lasted 
approximately two weeks and larvae increased in length to 0.9-1 cm. 

Fourth Instar: Fourth instar larvae similar to third, but white spots attenuated 
and gradually fading as larvae matured. Green color approximately matched that 


282 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of fresh leaves of foodplant. Including prepupal stage, stadium four lasted about 
three weeks, and larvae increased in size to about 1.5 cm. 

Pupa: Pupae medium brown color and covered with short brown hairs. No other 
distinguishing characters. Measured 0.8-0.9 cm. Pupation occurred on the earth 
in debris at bottom of rearing container. 


The larvae exhibited rather peculiar feeding habits. Rather than con- 
suming the leaves of the foodplant from the edges or tips, they ate holes 
from the under or upper surfaces of the leaves into the leaf centers. 
On the untouched sides, the leaves appeared to have been attacked by 
leaf mining insects, but when turned over, the larvae-produced holes 
could be seen. Fig. 1 (b,c) illustrates leaf damage. This characteristic 
feeding pattern, which persisted through all larval instars, aided in dis- 
covering larvae in the field. 

Larvae were not observed to spin silken mats prior to moulting, as do 
some other Theclinae larvae, nor did they eat the cast-off skins. 

During rearing, larvae were maintained at 20° C, which is a somewhat 
higher temperature than that of the natural habitat. 


Notes on Other Species 


It was observed that two other genera of Lepidoptera use E. umbel- 
latum as a larval host. While collecting ova of C. sheridanii in May 
1971, a mature lycaenid larva was found feeding on E. umbellatum. It 
was a medium green color with a lateral pink and white stripe. At first 
this was thought to be a mature sheridanii larvae, as ova can be found 
throughout the month-long flight period of the adults. The larva pupated 
and a male Plebejus (Icaricia) acmon lutzi Klots emerged in July 1971. 
Apparently this species must overwinter in the third or fourth instar, as 
adults are not on the wing until June in the Pole Mountain area, and very 
cold weather prior to collection of the larva precluded development 
from an egg. 

As mentioned previously, larvae of C. sheridanii were field-collected in 
1972 by looking for the characteristic leaf damage they produce. It was 
also found that a moth larva produces the same sort of damage. Efforts 
to obtain adults from these larvae were unsuccessful. They were leaf- 
rollers and spun a light silken network in their feeding area. They had 
black heads and were about the same color as sheridanii larvae. 

On 11 July 1971, females of Callophrys sheridanii neoperplexa ( Barnes 
and Benjamin) were observed ovipositing on Eriogonum umbellatum var. 
dichrocephalum Gand. on the rocky hillsides about 1-2 miles southeast 
of Galena Summit, ca. 8200’, Blaine Co., Idaho. The ova were similar to 
those of C. s. sheridanii. Rearing was not attempted. 


VOLUME 27, NuMBER 4 283 


ACKNOWLEDGMENTS 


Appreciation is expressed to Dr. John M. Reeder, University of 
Wyoming, Laramie, and to Dr. James L. Reveal, University of Maryland, 
College Park for determining the species of Eriogonum cited, and to Mr. 
Michael M. Ellsbury, Ft. Collins, Colorado, who assisted in the care of 
the larvae and provided the photographs shown in Fig. 1 (e and i). 


LITERATURE CITED 


Brown, F. M., Err, D. & B. Rorcer. 1957. Colorado Butterflies. Denver Museum. 
Denver, Colorado. 

Brown, F. M. & P. A. OpLer. 1970. The types of the lycaenid butterflies described 
by William Henry Edwards. Part 1I—Theclinae and Strymoninae. Trans. Amer. 
Entomol. Soc. 96: 19-77. 


DOWN-VALLEY FLIGHT OF ADULT THECLINI (LYCAENIDAE) 
IN SEARCH OF NOURISHMENT 


JAmes A. Scott 
60 Estes Street, Lakewood, Colorado 80226 


Down-valley movement of adult Theclini was reported by MacNeill 
(1967) for Satyrium saepium Boisduval. In this paper I will report 
down-valley flight of four additional species and will show the presumed 
flight pattern (Fig. 1) and the purpose of the down-valley flight. 

There are two types of mate-locating behavior among males of butter- 
flies (Scott, in press). In perching species, males sit at characteristic sites 
and dart out at passing objects in search of females, often returning to or 
near the previous spot after an investigative flight. Virgin females fly 
to these mating sites after emergence. In patrolling species, males fly 
almost constantly in search of females. 

Down-valley flight is defined as continuous rapid flight predominantly 
in a down-valley direction. Such flights can occur for several reasons, 
including migration. In patrolling species, male patrolling may appear 
similar to down-valley flight, but differs in that males investigate fe- 
males, flight upstream occurs about as frequently as downstream, and 
males usually patrol on hillsides in addition to valleys. Females may 
have different flight behavior in the two situations. In this paper I show 
that down-valley flight is part of a feeding response (Fig. 1). 


284 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fig. 1. Postulated flight pattern of two adult Theclini in search of nourishment. 


MetTHODS AND RESULTS 


Down-valley flight of Callophrys johnsoni Skinner, Callophrys au- 
gustinus Westwood, and Atlides halesus Cramer was observed 18, 19, 20, 
and 21 March 1972 at Thompson Canyon, Yolo County, California. Be- 
havior of the three species was identical. C. augustinus was only about 
two-thirds as abundant as C. johnsoni, and A. halesus was uncommon. 
The interval between the appearance of individuals passing the stationary 
observer averaged about 10 minutes for C. johnsoni on 18 March, and in- 
creased to about 25 minutes on 21 March, as abundance declined. These 
intervals for C. augustinus were about 15 minutes, increasing to about 30 
minutes. Because observations were made near the end of the brood, 
both species declined in numbers with time. Weather was similar on all 
four days, and so did not affect the observed numbers. Only about 
five A. halesus were seen per day. Individuals flew down-valley about 
a meter about the ground at a rapid rate of perhaps 3 meters per second 
for both Callophrys species; Atlides flew slightly faster. Very few in- 


VOLUME 27, NUMBER 4 985 


dividuals of these species flew up-valley. Individuals were observed fly- 
ing far enough apart so that they could seldom see one another in flight. 
They flew down-valley from about 0930 to 1600 on all days, but most 
individuals were observed between 1200 and 1400, the warmest part of 
the day. Where the road along the stream bottom ran onto the hillside, 
individuals flew along the stream bottom rather than the road. 

Down-valley flight of Erora laeta quaderna Hewitson was observed on 
6 April 1966 near Stewart Campground, Cochise County, Arizona. Down- 
valley flight of both sexes was the same as that of the Callophrys 
species except that individuals flew down-valley mainly along a road 
in the canyon bottom. Down-valley flight also occurred mainly in warm 
midday hours. . 

C. johnsoni males and females fed in the valley bottom on mud and on 
flowers of Rhus trilobata Nutt., and (rarely) on Cercis occidentalis Torr. 
C. augustinus fed there on mud and on Rhus. A. halesus and E. laeta 
Edwards fed on mud. 

Females outnumbered males in the two Callophrys species. Observed 
sex ratios based on collections by several persons during the 4 days 
were: C. johnsoni, 336, 592; C. augustinus, 15¢, 452; A. halesus, 22, 
22. The sex-ratio of E. laeta was 326, 312. 


DISCUSSION AND CONCLUSIONS 


Down-valley flight among Theclini has been recorded previously only 
at high population density. MacNeill (1967) observed down-valley flight 
of Satyrium saepium in a small dry ravine. About 5-15 individuals per 
minute flew down-valley about a meter above the ground, from 1130 to 
1200; very few individuals flew up-valley. The present observations 
indicate that the phenomenon may be widespread at fairly low density 
in Theclini, but is seldom noticed except at high density. 

The continuous down-valley flight contrasts markedly with the short 
flights characteristic of mate-locating behavior. All five species are 
perching species. Details of mate-locating behavior of these species 
follow: 

1) Oakley Shields (pers. comm.) observed mate-locating behavior of 
C. johnsoni on a hilltop next to Thompson Canyon, where many males 
perched on the tops of tall trees, from which they often chased each 
other. I observed the same behavior in the closely related species C. 
spinetorum Hewitson occurring on small pines near hilltops in Grand 
Canyon, Arizona, in the afternoon. 

2) Powell (1968a) described mate-locating behavior of C. augustinus. 
Males perched on a small tree in a clearing from 1030 to dusk. Mating 


286 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


was observed only in the late afternoon. However, I observed copulation 
at 1110 on a tree on a hillside at Alpine Lake, Marin County, California, 
4 April 1970. 

3) I have observed many A. halesus males perching on the tops of 
small trees on hilltops at midday and in the afternoon at four localities 
in New Mexico and Arizona. 

4) Many E. laeta males were observed perching on the tops of juniper 
trees on hilltops in the afternoon near Emory Pass, Sierra County, New 
Mexico, 5 April 1966, and near Onion Saddle, Cochise County, Arizona, 
6 April 1966 and 24 April 1973. 

5) I observed mate-locating behavior of S. saepium at numerous lo- 
calities in Jefferson County, Colorado, where males perched about one 
meter above the ground on the sides of small pine and juniper trees on 
ridgetops and hilltops. Males perched mostly on the eastern side of trees 
in the morning, and on the western side in late afternoon. The preferred 
side was less predictable at midday, but males often chose the south 
side. The perching males darted after passing objects (mainly other 
males) and usually returned to the same tree after an investigative flight. 
Such behavior occurred from 0715 to 1600. A copulating pair was found 
at 1155 on one of the trees where males were perching. 

The presumed flight pattern is shown in Fig. 1. Individuals fly from 
hillsides to the valley bottom, then downstream to a feeding site. Having 
fed, they depart to the hillsides again. The following points support 
these contentions: 1) individuals were most abundant in a small area in 
the middle of Thompson Canyon; only one individual was seen at the 
mouth of the canyon, and only one (a female showing preoviposition be- 
havior toward the larval hostplant) was seen near the head of the canyon; 
(MacNeill (1967) also noted that down-valley flight was confined to a 
small area); 2) study on four consecutive days showed that individuals 
do not move down-valley one day and up-valley the next. More than 20 
individuals of each of the two Callophrys species were seen to fly down- 
valley each day, whereas less than 4 per day flew up-valley. 

Down-valley flight may be advantageous to these species because 
flowers and water are often more abundant in the central portions of 
small canyons in arid areas of the western United States. Down-valley 
flight may prove to be limited to arid areas, since it involves movement 
toward regions of high moisture. Individuals need to seek nourishment 
in the valley bottom because flowers are much less common on hillsides. 
Also, larval hostplants grow on hillsides, and do not serve as nectar 
sources for the adult butterflies. The known larval hostplants are as 
follows: Arceuthobium campylopodum Engelm. for C. johnsoni, a plant 


VOLUME 27, NUMBER 4 287 


parasitic on Pinus sabiniana Doug]. in Thompson Canyon (Shields, 1965); 
many plants for California C. augustinus (Powell, 1968b); Phoradendron 
for A. halesus, a plant parasitic on live oak; and Ceanothus sp. for S. 
saepium (Clench, 1961). 

Net down-valley movement was not observed for either sex in any of 
the species. The flight pattern postulated in Fig. 1 does not result in net 
down-valley movement, because individuals of at least C. johnsoni, A. 
halesus, E. laeta, and S. saepium apparently move upslope to seek mates 
on hilltops. Scott (1973) and Shields (1967) have shown that hilltops 
serve as mating sites for some butterflies. 

Finally, down-valley flight also probably occurs in other butterfly 
taxa. I have observed what appears to be down-valley flight in Oeneis 
chryxus Doubleday and O. uhleri Reakirt (both Nymphalidae, Satyrinae ) 
in southern Colorado. However, about 20-30 percent of the individuals 
of these species seen were flying upstream; individuals of these two 
species also were observed feeding on mud in the gully bottom. 


SUMMARY 


Down-valley flight (continuous rapid flight predominantly in a down- 
valley direction) was observed in four species of Theclini and appears 
to be part of a behavior pattern in which individuals seeking nourishment 
fly to the valley bottom, fly downstream until mud or flowers are found, 
feed, then fly to hillsides. Mate-locating behavior, which occurs else- 
where (usually on hilltops), is very different. 


LITERATURE CITED 


Ciency, H. K. 1961. Tribe Theclini, in P. R. & A. H. Ehrlich, How to Know 
the Butterflies. Brown, Dubuque, Iowa. p. 177-220. 

MacNem., C. D. 1967. A unidirectional mass movement by Satyrium saepium. 
J. Lepid. Soc. 21: 204. 

PoweELt, J. A. 1968a. A study of area occupation and mating behavior in Incisalia 
irioides (Lepidoptera; Lycaenidae). J. N. Y. Entomol. Soc. 76: 47-07. 

1968b. Foodplants of Callophrys irioides. J. Lepid. Soc. 22: 225-226. 

Scorr, J. A. 1973. Mate-locating behavior of butterflies. Amer. Midl. Nat. (in 
press ). 

SureLps, O. 1965. Callophrys (Mitoura) spinetorum and C. (M.) johnsoni; their 
known range, habits, variation, and history. J. Res. Lepid. 4: 233-250. 

1967. Hilltopping. J. Res. Lepid. 6: 69-1785. 


288 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


THE IDENTITY OF MACARIA INAPTATA WALKER AND 
ITAME VARADARIA (WALKER) (GEOMETRIDAE) 


DouctLas C. FERGUSON 


Systematic Entomology Laboratory, ARS, USDA 
c/o U.S. National Museum, Smithsonian Institution, Washington, D.C. 20560 


Macaria inaptata Walker has been consistently misidentified since the 
time of its original description. In North American lists it was treated as 
a variety of Episemasia solitaria (Walker) by early authors following 
Hulst (1894: 306). More recently the name has stood as that of a distinct 
species in the Semiothisa minorata-distribuaria group, having been 
arbitrarily transferred to Semiothisa when Macaria was recognized as a 
junior synonym (e.g. McDunnough, 1938: 158). In the collection at the 
U.S. National Museum I found a finely executed colored drawing of a 
moth resembling Semiothisa aemulataria (Walker), but mislabelled as 
the type of Macaria inaptata. This illustration appears to represent an 
unnamed species that I was preparing to describe, and it thus led to the 
present investigation. When I recently examined the type of inaptata, I 
found that it is actually a female of Itame varadaria; thus Macaria 
inaptata Walker, 1861: 886, becomes a synonym of Caberodes ? varadaria 
Walker, 1860: 251. 

At least three other names have been proposed for this species, and 
its synonymy and type data are as follows: 


Itame varadaria (Walker) 


Caberodes ? varadaria Walker, 1860, List of the Specimens of Lepidopterous Insects 
in the Collection of the British Museum 20: 251. 
Type locality: Georgia (From the Milne collection and probably collected by 


Abbot). 
Types: A male holotype in the collection of the British Museum (Natural 


History ). 
Macaria inaptata Walker, 1861, ibid. 23: 886. NEW SYNONYM. 

Type locality: United States. As the species is known only from the extreme 
Southeast, and the donor was stated to be Doubleday, I hereby restrict the type 
locality to St. Johns Bluff, Duval Co., the site of Doubleday’s Florida collecting. 

Types: A female holotype in the collection of the British Museum (Natural 
History ). 

Aspilates abbreviata Walker, “1862”[1863], ibid. 26: 1673. 

Type locality: Georgia (from the Milne collection and probably collected by 
Abbot). 

Types: A male holotype in the collection of the British Museum (Natural 
History ). 

Diastictis florida Hulst, 1896, Trans. Amer. Entomol. Soc. 23: 334. 

Type locality: Florida. 


VOLUME 27, NUMBER 4 289 


Figs. 1-3. Itame varadaria (Wlk.): (1) summer brood male, Wedge Plantation, 
McClellanville, South Carolina, 19 August 1971; (2) spring brood male, same 
locality, 31 March 1967; (3) spring brood female, University Conservation Reserve, 
Welaka, Putnam Co., Florida, 11 March 1962. (All collected by the author.) Photos 
by photographic laboratory of the Smithsonian Institution. 


Types: The number of types was not stated although the description reads as 
though Hulst may have had only one. However, there is one male type in the collec- 
tion of the American Museum of Natural History, and there are two labelled as types 
in the U.S. National Museum. Of the latter, one is labelled “N.Y.” (probably in- 
correctly) and is assumed to be a spurious type. The other is labelled as from 
Florida and the Hulst collection, and may be a syntype. All three specimens repre- 
sent the same species as varadaria. 


Itame varadaria is a species with which few lepidopterists have been 
familiar; it is limited to the Deep South and has not been common in 
collections. It is known to occur from McClellanville, South Carolina, to 
southern Florida and along the Gulf Coast to the vicinity of Houston, 
Texas. It has at least two broods, the spring specimens (March-April) 
being slightly larger, darker and less brightly marked than summer ones 
(July-August). Otherwise nothing is known of its life history. 

As the species has not been illustrated, I figure a summer brood male 
(Fig. 1) and spring brood male and female (Figs. 2-3). This female 
was compared with the holotype of inaptata and agrees in almost every 
detail. I. varadaria varies from dull yellowish to grayish brown with a 
darker brown pattern, and the male has rather wide bipectinate antennae. 
No very closely related species are known, but the male genitalia of 
varadaria are nevertheless typical of the genus in every respect, being 
hardly distinguishable from those of the Itame argillacearia-occiduaria 
group. I have examined 34 specimens. 

Aspilates donataria Walker (“1862”[1863]: 1673), listed in the genus 
Itame as number 4787 by McDunnough (1938: 161), is a nomen dubium. 
It may also be the same as varadaria but has never been identified as any 
known species. The type was reported by Dyar (“1902"[1903]: 315) 
to be lost, and I am advised by Mr. D. S. Fletcher of the British Museum 
(Natural History) that it still has not been found. The original descrip- 
tion could fit varadaria except that the size seems too large. The type 
locality is Georgia. 


290 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


I am indebted to Mr. D. S. Fletcher for kindly responding to my in- 
quiries about the Walker types and for lending specimens, and to Drs. 
C. W. Sabrosky and J. F. G. Clarke for conveying specimens to and from 
London. 


LITERATURE CITED 


Dyan, H. G. “1902”[1903]. A List of North American Lepidoptera and Key to 
the Literature of this Order of Insects. Bull. U.S. Natl. Mus. 52. 723 p. 

Huxst, G. D. 1894. Notes on types of North American Geometrina in European 
collections.—I. Entomol. News 5: 302-306. 

McDunnoucu, J. H. 1938. Check List of the Lepidoptera of Canada and the 
United States of America, 1, Macrolepidoptera. Mem. So. Calif. Acad. Sci. 1. 
DZ ie 

WALKER, F. “1862’[1863]. List of the Specimens of Lepidopterous Insects in the 
Collection of the British Museum 26: 1479-1796. 


TRANSFER OF CYMORIZA ABROTALIS WALKER, 1859, FROM 
NYMPHULA SCHRANK TO DISMILILA DYAR 
(PYRALIDAE: NYMPHULINAE, MIDILINAE) 


EUGENE MUNROE 


Biosystematics Research Institute, Canada Department of Agriculture, Ottawa, Ontario 


Cymoriza abrotalis Walker (1859: 956) was described from a single 
male from Rio de Janeiro “in Mr. Fry’s collection.” Hampson (1897: 
140) transferred the species to Nymphula, without comment. This place- 
ment was followed by Klima (1937: 92). 

The holotype of C. abrotalis is in the Oxford University Museum. A 
photograph made by myself in 1958 (Fig. 1) shows clearly that the 
species is a midiline not a nymphuline, and that it belongs to the genus 
Dismidila Dyar. Although I had seen and photographed the type, I had 
not noticed the relationship and I omitted the name from my Revision 
of the Midilinae (Munroe, 1970). The maculation and wing shape show 
that the species is very close to Dismidila similis Munroe, type-locality 
Buenavista, East Bolivia, and to D. obscura Munroe, type-locality 
Petropolis, Rio de Janeiro State, Brazil. In colour it resembles D. similis 
more than D. obscura, but it has the hyaline spot of the forewing much 
larger. In the absence of series and not having compared the types 
directly I think it better not to establish synonyms. I content myself with 
transferring the species to Dismidila as Dismidila abrotalis (Walker), 
new combination. 


VOLUME 27, NUMBER 4 291 


Fig. 1. Cymoriza abrotalis Walker, holotype, 6. Oxford University Museum. 


LITERATURE CITED 


Hampson, Sir Georcre F. 1897. On the classification of two subfamilies of moths 
of the family Pyralidae: the Hydrocampinae and Scoparianae. Trans. Entomol. 
Soc. London. 127-240. 

Kima, A. 1937. Lepidopterorum catalogus. Pars 84. 226 p. ’s-Gravenhage. 

Munroe, E. 1970. Revision of the subfamily Midilinae (Lepidoptera: Pyralidae). 
Mem. Entomol. Soc. Canada 74. 94 p. 

WALKER, F. 1859. List of the specimens of lepidopterous insects in the collection 
of the British Museum. Part XIX.-Pyralides. p. 799-1036. London. 


UNUSUAL COPULATORY BEHAVIOR IN THE 
NYMPHALIDAE AND SATYRIDAE 


EDWIN M. PERKINS, JR. 


Department of Biological Sciences, University of Southern California, 
Los Angeles, California 90007 


Interspecific, heterosexual coupling among Rhopalocera in the wild is 
relatively uncommon enough to warrant being recorded in the literature 
(Hovanitz, 1949; Shigeru, 1956; Stallings et al., 1959; Downey, 1962; 
Shapiro & Biggs, 1968; Perkins & Gage, 1970; Priestaf, 1970; Platt & 
Greenfield, 1971). Even more rare are occurrences of intergeneric, 
heterosexual coupling (Frechin, 1969; Jae, 1972). 


292 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


To date, however, a search of the literature confirms that instances of 
neither intraspecific, pleoheterosexual coupling nor intergeneric, homo- 
sexual coupling are known. The following account cites examples of 
each. 

While visiting the Bentsen State Park National Wildlife Refuge, 
situated thirteen miles northwest of McAllen, Hidalgo County, Texas on 
13 November 1962, I came upon a small collateral tributary of the Rio 
Grande, whose moist sand bar was nearly covered with imbibing butter- 
flies. Bewildered by this spectacle, I cautiously approached and noted 
several hundred individuals, including examples of Libytheana bachmanni 
larvata Strecker, Phyciodes phaon Edwards, Zerene cessonia gen. aut. 
rosa M’Neill, Danaus gilippus strigosus Bates, Papilio cresphontes Cramer, 
Kricogonia lyside castalia Fabricius, Eurema lisa Boisduval, Hemiargus 
ceranus zachaeina Butler & Druce, Atlides halesus estesi Clench, and 
Myscelia ethusa Boisduval. 

Of particular interest were three specimens that appeared to be in 
copula. Despite their efforts to escape my net, all three were genitalically 
attached and remained so until placed in a cyanide bottle. Examination 
of the unlikely threesome revealed 2 ¢ 6 and 1 2 P. phaon (Fig. 1). 

Twenty-two months later, on 11 August 1964, a similarly surprising 
event took place at Camp Sherman, Jefferson County, Oregon. This day, 
while collecting in the lush meadows that border the Metolius River, I 
had already taken several copulating pairs of Speyeria cybele leto Behr, 
S. atlantis dodgei Gunder, and S. mormonia erinna Edwards. 

Beside a willow-bordered drainage ditch, I noted a tall blade of grass 
on which two, in copula butterflies were at rest: affixed by their claspers 
were 1 ¢ S. m. erinna and 1 6 Cercyonis pegala ariane Boisduval. When 
disturbed, the S. m. erinna labored upward—in a forward direction— 
under the weight of the attached C. p. ariane, which it pulled after it. 
The pair (Fig. 2) was captured in flight at 1400 PDT, placed in a separate 
cyanide bottle, spread and genitalically examined at the same time, and 
given cross-reference data labels in order to minimize the considerable 
confusion that already prevailed! All specimens are currently housed in 
the Allyn Museum of Entomology, Sarasota, Florida. 

It is tempting to speculate about the overall importance of these ob- 
servations regarding the breakdown of behavioral and/or mechanical 
isolating mechanisms within or between species. Certainly, neither of the 
two particular unions described could definitively subserve any reproduc- 
tive end. Behavioral aberrations such as these, and that described by 
Heitzman (1964), not only challenge the “lock and key” hypothesis, but 
also suggest the possibility of a parallel development in precopulatory 


VOLUME 27, NUMBER 4 293 


2 


Figs. 1-2. Specimens taken in copula: (1) Phyciodes phaon males and female 
(center), dorsal surfaces (Bentsen State Park, Hidalgo Co., Texas, 13 November 
1962, E. M. Perkins, collector); (2) Speyeria mormonia erinna male (above), and 
Cercyonis pegala ariane male (below), ventral surfaces (Camp Sherman, Jefferson 
Co., Oregon, 11 August 1964, E. M. Perkins, collector). 


behavior and/or chemical configuration of pheromones in certain, dis- 
similar lepidopterous taxa. It is interesting to note that capture time of 
specimens in the latter event coincides with frequency polygons of ob- 
served mating times for both Nymphalidae and Satyridae (Miller & 
Clench, 1968). Although the significance of both records is subject to 
debate, if not entirely enigmatic, these data are placed on permanent 
record in the hope that they might in some way shed additional light 
on our knowledge regarding insect behavior, pheromones, and/or 
evolution. 


LITERATURE CITED 


Downey, J. C. 1962. Inter-specific pairing in Lycaenidae. J. Lepid. Soc. 16: 
235-237. 

FrecHIN, D. 1969. A notable intergeneric mating (Lycaenidae). J. Lepid. Soc. 
Joe 115. 

HeirzMan, R. 1964. The story of a “mixed up” Thorybes pylades (Hesperiidae ). 
J. Lepid. Soc. 18: 169-170. 

Hovanirz, W. 1949. Interspecific matings between Colias eurytheme and Colias 
philodice in wild populations. Evolution 3: 170-173. 

Jaz, R. J. 1972. Natural inter-breeding of close nymphalid groups. J. Lepid. Soc. 
DOI 28: 


294 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Miter, L. D. & H. K. CLtencu. 1968. Some aspects of mating behavior in butter- 
flies. J. Lepid. Soc. 22: 125-132. 

PERKINS, E. M. & E. V. Gace. 1970. On the occurrence of Limenitis archippus 
L. lorquini hybrids. J. Res. Lepid. 9: 223-226. 

Piatt, A. P. & J. C. GREENFIELD, Jr. 1971. Inter-specific hybridization between 
Limenitis arthemis astyanax and L. archippus (Nymphalidae). J. Lepid. Soc. 
25: 278-284. 

PriestaF, R. C. 1970. Courtship and mating between Chlosyne neumogeni and 
Chlosyne californica (Nymphalidae). J. Lepid. Soc. 24: 226. 

SHaprro, A. M. & J. D. Biccs. 1968. A hybrid Limenitis from New York. J. Res. 
Lepid. 7; 149-152. 

SuicEru, A. A. 1956. Hybrids between Colias eurytheme and C. interior (Pieridae). 
J. Lepid. Soc. 10: 9-14. 


NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF 
BUTTERFLIES OF EL SALVADOR. 


Il. ANAEA (ZARETIS) ITYS (NYMPHALIDAE) 


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


This is the second article of a series dealing with what my sons and I 
have discovered about the life cycle and natural history of butterflies 
found in the neighborhood of San Salvador, capital city of El Salvador. 
The life cycles of many neotropical butterflies are apparently incom- 
pletely known, and therefore classification has been solely on the basis 
of the adult morphological characteristics. 

The determination of the species mentioned herein has been done by 
Dr. Lee D. Miller of the Allyn Museum of Entomology. Adults and at 
least some specimens of the early stages have been placed in that museum, 
so as to be available for students of the groups. 

In the introduction of our prior article (1973), a rough description of 
the country, its climatic zones and other pertinent information was given, 
so as to make an understandable picture of the habitats of the species 
described in these articles. 

Anaea (Zaretis) itys Cramer, belonging to the subfamily Charaxinae, 
has been placed at different times in several genera: Papilio (by Cramer, 
1777), Siderone (by Westwood, 1850), Zaretes (Frihstorfer, 1909); 
and has been described several times, due perhaps to the great geographi- 
cal, seasonal and individual variation shown. Comstock (1961) uses the 
name Anaea (Zaretis) itys, leaving open the possibility that valid sub- 
species might be found later. 


VOLUME 27, NUMBER 4 295 


We had captured a few specimens of this elusive butterfly since 1968, 
always at around 1000 m, usually while feeding on decaying fruits. 
But until August 1970 we had never seen the larvae and did not know 
the foodplant. At that time a larva of medium size, but of unusual shape, 
was captured on a small tree later identified as Casearia nitida (L.). 
The larva produced an adult male in mid-September. Once the foodplant 
was known, we made several extensive searchs through the different 
zones of the country where the plant is found, at altitudes ranging from 
sea level to about 1200 m. Many larvae in different stadia were found 
and adults were obtained from them. The first time we found eggs was 
during November 1971, when a female was observed ovipositing on a 
medium-sized tree in the neighborhood of San Salvador. This female 
laid about 17 eggs, five of which were collected and put in individual 
transparent plastic bags. Black-and-white photos were taken of the 
eggs, and of the larvae at various stages; and measurements and records 
of developmental time were kept. Head capsules, and larval specimens 
of each instar were kept in alcohol. The bags in which the process took 
place were kept at ambient temperature and lighting conditions. Since 
that time we have been able to obtain eggs at different times of the year 
- and have repeated the process several times with about the same results. 


Life cycle stages 


Egg. Nearly spherical, about 1 mm diameter, with flattened base and slight depres- 
sion at micropyle. Color yellowish. Surface smooth (nothing noticeable at 10x 
magnification). All hatched in 6 days. 

First instar larva. Head naked, brown, roundish, with slight depressions be- 
tween epicrania and cervical triangle. Body grayish, naked, wedge-shaped. Changes 
color and shape after feeding, becoming light brown, and thickening across 2nd 
abdominal segment. Annulets appear in the segments. Grows from 2.5 mm to 5.2 
mm in 7 days. 

Second instar larva. Head and body dull dark brown. Short and thick horns on 
the apex of epicrania. Anal segments with short and flattened “tails.” Spiracula of 
lighter shade than body. Spiraculum on 2nd abdominal segment located much 
higher than the rest, and the one on the 8th abdominal segment slightly out of line 
with the preceding ones also. Larvae grow from 9.5 mm—1l.1 cm in 6 days. 

Third instar larva. Head dark gray with black, blunt, thick and short horns on 
epicrania. Tiny graining apparent on head surface. Body dark grayish brown, with 
lateral light ridge starting at first thoracic segment running to first abdominal seg- 
ment supraspiracularly. Another light brown ridge originating high on second abdomi- 
nal segment under a lateral pyramidal projection located in the subdorsal area, going 
through remaining abdominal segments, supraspiracularly, ending at side of the 
short anal tail. Lateral projections on the second abdominal segment united 
postero-dorsally by thick and dark brown ridge bordered by clear margin, and each 
one terminated by a scolum with minute spines. Spiraculum immediately below 
the projected scolum, completely out of line with rest of spiracula. 3rd and 4th 
abdominal segments each crossed dorsally by a clear brown line, parallel to the 
clear margin on 2nd abdominal segment. A large rhomboid of light brown color 
covering the dorsal area of the 5th, 6th and 7th abdominal segments and _ partly 


296 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the 8th. A scattering of tiny spines all over the body. Larvae grow to from 1.6—2.0 
cm in 8 days. 

Fourth instar larva. Same general shape and coloration as third instar, but has 
developed some spines on epicranium, below the horns and at sides of them. Whitish 
graining on thoracic segments. Light rhomboid on last abdominal segments bordered 
now by black triangles, and many minute spines noticeable on these segments. Larvae 
grow to from 2.3—-3.1 cm in 9-11 days. 

Fifth instar larva. Lighter color than 4th instar, hump at 2nd abdominal segment 
very exaggerated. Dorsal area of abdominal segments with lighter brown rhomboid 
markings. Spines more noticeable now, mostly at lateral zones of last abdominal 
segments. Spines now prominent at sides of head and at base of and between 
epicranial horns. Some of these spines have a white tip. Tails look very flattened 
and united at the base. Larvae grow to from 4.3-5 cm in 11-19 days. 

Pre-pupa. Considerably shorter and thicker than 5th stadium and of lighter 
color. Stays incurvated laterally with anal prolegs affixed to silken mat for one day. 

Pupa. Can be light green or light brown. Cremaster very elaborated and reddish- 
brown. Abdominal segments tapering abruptly from wing case to cremaster, and 
thoracic segments gradually to slightly bifid head. Yellowish ridge dorsally separating 
tapering abdominal segments from the rest, and from border of wing cases. Spiracula 
inconspicuous light brown. Measurements vary from 1.5-1.9 cm long, 1.0-1.2 cm 
laterally at widest point and 0.9 to 1.1 cm dorsoventrally at widest point. Dura- 
tion 8-10 days. . 

Adults. Adults show a marked sexual dimorphism, and minor individual differ- 
ences between specimens of the same sex. Males. Both fore and hindwing orange 
dorsally, with darker apical markings on forewing, and darker submarginal broken 
line on hind wing, with small black dots in variable number between the dark line 
and outer margin. Apex more or less acute, outer margin undulate, more or less 
convex above tornus and inner margin more or less concave at side of tornus. 
Usually with two transparent roundish “windows” located on cells M3 and Cu 1, 
near discal cell, but lacking in some individuals. Hindwing with a humeral lobe 
and emargination at inner angle. Outer margin of hindwing concave, more or less 
sinuose, with blunt and projected anal angle, and anal lobe. 

Ventrally both wings brownish orange with darker line that goes from anal angle 
to mid-costal area on hindwing, continuing from mid-inner margin to apex on fore- 
wing. This line imitates the central vein of a leaf, and there are besides faint shadings 
imitating secondary veins, mostly on hindwings. Females. Shape of wings as in 
male, with apex generally more acute. Forewing dorsally, light yellow with brown 
markings at apex. Hindwing with orange shading, more pronounced at anal angle. 
Both wings ventrally dirty yellow with brown markings imitating, as in male, the 
venation of a leaf. “Windows” usually present on forewing, but may be lacking. 

In both sexes body same color as dorsal wings. Palpi, eyes and proboscis, brown. 
Antennae orange. Individual differences have been noticed on specimens emerging 
during the same month. Size of individuals of the same sex is variable; in general 
females are larger than males. Average measurement is 4.2 cm for males and 
5.3 em for females (tip-to-tip of spread forewings). Total development time, under 
laboratory conditions, ranges from 56-68 days, females usually taking more time 
than males. 


Natural History 


We have found eggs and larvae of A. (Z.) itys on only one species of 
the Flacourtiaceae family (also known as Samydaceae): Casearia nitida 
(L.) Jacq. This plant grows to become a small tree about 5 m high and 
is fairly abundant from sea level to 1200 m of altitude, in heavily 
disturbed second-growth plant communities, usually along ravines or 


297 


VoLUME 27, NUMBER 4 


\ 


r 


al 


c 


st 


in 


t 
4) th 


irs 
( 
ifth 


Jon 


bout 1 em 
3 


a 
bout 3 cm 
al \ 


a 
eC 


( 


> 


egg, about 1 mm 
ral 


) 


1 


tys Cramer 


is) 4 
, about 5 mm 


Zaret 


( 


Anaea 


]-9. 
larvae ready to moult 


Figs. 


ird 
instar 
9x 1 cm. 


6) £ 
5x 0. 


( 


1 


> 


a 
1eW 


> 


I'v 
r 


a 
rent 


> 


sta 
a 
and \ 


in 
ary 


. 


tare 
rsal 


second 
do 


ins 
> 


) 
ral 


3 


( 
) fourth 


(5) fo 
8, 9) pupa, late 


. 
? 


? 


8 cm 
7 


( 


about 1 
larva, about 4.5 cm; 


tar larva, 


Ins 


298 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Figs. 10-14. Anaea (Zaretis) itys Cramer: (10) adult male with “windows,” 
dorsal view; (11) adult female with “windows,” dorsal view; (12) adult male 
without “windows,” ventral view; (13) adult female with “windows,” ventral view. 
(14) adult male without “windows,” dorsal view. The black bars measure 1 cm. 


VoLUME 27, NuMBER 4 299 


unused land. The plant sheds all of its leaves prior to the rainy season, 
during March or April. When bare of leaves, the plant flowers, and some 
time later starts to grow the new leaves. By July-August the coffee-like 
fruits are green and become yellowish-orange by October. The leaves 
of this species are attacked by a disease that curls the edges and turns 
the rolled portion dark brown and dries it. 

Recently-emerged larvae of A. (Zaretis) itys, like all the larvae of 
Charaxinae we have observed, completely eat the egg shell, and stay 
under the leaf for a full day without further eating. They then move 
to the edge of the leaf, usually to the tip, eat around a vein until it is 
bare and prolong it with excreta affixed with silk. This formation is used 
as a resting place through the first, second and third stadia, the larvae 
usually keeping the head pointing outwards. During the fourth stadium 
the larvae leave this resting place and wander about the plant. While 
not feeding they stay motionless at the edge of a leaf imitating to perfec- 
tion a dryed portion of it. This behavior is kept also through the fifth 
stadium. When ready to pupate the larva shortens and weaves a silken 
mat under a leaf or on a twig with many leaves and affixes the anal 
prolegs to it. Then the prepupal larva positions itself in the peculiar 
fashion of the genus Anaea, incurvated sideways, not hanging. 

The pupae of Anaea (Z.) itys, like some other Anaea spp. can be either 
light green or light brown, regardless of environment, and their shape 
is very similar to A. (Consul) fabius Cramer and A. (Memphis) eurypyle 
confusa Hall, (MSS in prep.), being quite hard to tell from these. The 
pupae are rather rigid and will only occasionally react when handled 
with short lateral movements. 

The adult of A. (Z.) itys, like most Charaxinae, are swift fliers, what 
makes them hard to see in the field unless they are feeding, which they 
do on fermented fruits and from open cavities of tree trunks. The fe- 
males when ready to oviposit, fly rapidly around the foodplant and 
alight on a group of mature leaves. Sitting under one of them, they 
deposit one egg on the underside and immediately resume their flying. 
They repeat this action about five times before taking a rest on a 
neighboring tree for some ten minutes, and then go back to flying around 
the foodplant, depositing some more eggs, until a total of about 30 are 
deposited before the female flies away. Both sexes often sit on tree 
trunks, head pointing downwards. Males chase other butterflies that 
cross their territory. We have observed males attacking falling leaves. 


DISCUSSION 


According to Comstock (1961), there is at least some information 
about the life cycle of Anaea (Zaretis) itys in the works of Sepp (1828) 


300 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


and Muller (1886) but apparently this is the first complete life cycle 
description with photographs. 

The eggs of this species are very similar to the eggs of Anaea (Consul) 
fabius, A. (C.) electra Westwood, A. (Memphis) eurypyle confusa and 
A. (M.) pithyusa R. Felder, except for the yellowish, rather than greenish, 
color, as in the bigger egg of A. (Siderone) marthesia Cramer. First 
instar larvae are very similar also to the larvae of the species mentioned, 
except for color. From second stadium on, they resemble more the larvae 
of Prepona spp. in shape and behavior, even though the Anaea spp. 
mentioned have similar habits up to the third instar. The pupae again 
are very much like the pupae of A. (C.) fabius, A. (C.) electra and A. 
(M.) eurypyle confusa, even in the characteristic of being at times light 
brown and at times light green. We thought at first that this duality of 
coloration was caused by “environmental conditions during pupation” as 
Harrison (1963) assumed for the same phenomenon in Opsiphanes 
tamarinidi Felder (Brassolidae). But on several occasions groups of 
larvae of A. (Z.) itys, A. (C.) fabius, A. (M.) e. confusa and even Opsi- 
phanes tamarindi and O. cassina fabricii Bdv. kept under similar condi- 
tions all through their developmental stages have produced simultanously 
and indiscriminately green and brown pupae, invalidating this explana- 
tion. The duality of coloration in the pupae of these species is not de- 
pendent on the sex of the individuals either: we have had males and 
females come out of brown and green pupae. Whether or not this 
phenomenon is hereditary, we have not yet been able to find out. 

The close affinity between the genus Prepona and the genus Anaea 
is evident: apparently this species, A. (Zaretis) itys, is intermediate be- 
tween Prepona and “typical” Anaea. As is the case in Prepona spp., the 
spiraculum on the second abdominal segment of A. (Z.) itys larvae is 
located very high on the subdorsal area, being not so high in the other 
Anaea spp. Yet the spiraculum on the eighth abdominal segment is 
slightly higher in the three cases. 

The larvae of A. (Z.) itys are very slow moving and apparently have 
no chemical or mechanical defenses, relying solely on mimicry for 
protection against predation, as in Prepona spp. During the initial three 
instars Prepona spp., A. (Zaretis) itys, and the other species of Anaea 
we have studied use the same strategy: the bared vein prolonged with 
frass that they use for resting, imitating to perfection dried portions of 
leaf tissue still attached to the vein. But from the fourth instar on the 
larvae of A. (Z.) itys behave more like Prepona larvae, staying motionless 
usually on the edge of a diseased leaf, mimicking its curled and dried 
edge. Moving about for feeding purposes is usually done at dawn or 


VOLUME 27, NUMBER 4 301 


dusk, minimizing the chances of diurnal bird predation. The pupae, 
green or brown, are very hard to spot among the foliage. The adults, 
besides their rapid and vigorous flight, mimic decaying leaves when at 
rest giving them a near perfect concealment among vegetation. 

The mimicry exhibited, while reducing predation, does not protect 
from parasitism caused by Tachinidae of the kind that deposit the eggs 
on the surface of the leaves where the larvae are feeding. Quite often 
we have collected larvae that have been killed during the last larval 
stage or shortly after pupation by tachinid larvae, not yet determined. 
The larvae of A. (Z.) itys are also very prone to a disease that softens 
their bodies, making them burst and die. | 

As in the case of Prepona o. octavia (Muyshondt, 1973), a possible 
cause of massive larval mortality is the characteristic of the foodplant, 
Casearia nitida, of shedding all the leaves in a short period of time, just 
prior to the rainy season, (late March and April). The shrubs or small 
trees then remain bare of leaves for a period of two to three weeks. 

It is possible that the larvae of A. (Z.) itys feed on other related species 
of the Flacourtiaceae family, as is the case with A. (Siderone) marthesia, 
although we have not found evidence to support this. 

Under laboratory conditions, this species took an average of 62 days 
to complete development from egg to adult. Therefore there could be 
some six generations a year. Since the species is not common, we suggest 
that the females of A. (Z.) itys are not particularly prolific and that 
parasites kill many larvae. This suggestion seems to be supported by the 
fact that adults of A. (Zaretis) itys are mostly found during the dry season, 
and, “during the dry season the population of small insects,” (including 
parasites ), “is distinctly reduced seemingly by desiccation associated with 
small body size” (Janzen and Schoener, 1968). In addition to the paucity 
of parasites during the dry season, their efficiency in laying eggs on the 
leaves might be affected by the heavy dust layer deposited by the sea- 
sonal northern winds. 


ACKNOWLEDGMENTS 


We are grateful for the valuable assistance of Stephen R. Steinhauser, 
who besides sharing his own observations on the adult behavior of this 
species, gave us free access to his technical library. We are particularly 
indebted to Dr. Lee D. Miller who identified the specimens and to Dr. 
T. D. Sargent who made invaluable criticisms and suggestions regarding 
the manuscript. Specimens of early stages and adults have been placed 
in the Allyn Museum of Entomology, Sarasota, Florida. 


302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


LITERATURE CITED 


Comstock, W. P. 1961. Butterflies of the American Tropics, the Genus Anaea, 
(Lepidoptera, Nymphalidae). Amer. Mus. Nat. Hist., N.Y. p. 30. 

CrAMER, P. 1777. Papillons exotiques des trois parties du Monde: L/Asie, 
L’Afrique et L’Amérique. Amsterdam. Vol. 2, p. 34. 

FRUustToRFER, H. 1909. Entomol. Zeitschr. Stuttgart. Vol. 23, p. 166. 

Janzen, D. H. & T. W. ScHOENER. 1968. Differences in insect abundance and 
diversity between wetter and drier sites during a tropical dry season. Ecology 
49: 96-110. 

Harrison, J. O. 1963. On the biology of three banana pests in Costa Rica. 
(Lepidoptera: Limacodidae, Nymphalidae). Ann. Entomol. Soc. Amer. 56: 
87-92. 

MUu.ier, W. 1886. Zool. Jahrb. Zeitschr. Syst. Geogr. Biolog. der Tiere, Jena. 
Voleiep 497 

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

Sepp, J. 1928. Surinaamische Vlinders. Amsterdam Vol. 1, p. 9. 

WeEstwoop, J. O. 1850. The genera of diurnal Lepidoptera. London. Vol. 2, p. 
SPAN 


SOME OBSERVATIONS ON DRYAS IULIA IULIA (HELICONIIDAE ) 


Dryas iulia iulia (Fabricius) is a common species in El Salvador and is found 
flying in wide open territory or under low vegetation from sea level to about 2000 m. 
Both sexes are assiduous visitors of flowers. The females lay eggs individually 
(pers. obs.) on tender terminals of various species of wild Passifloraceae vines. 

My sons and I have reared D. i. iulia many times from egg to adulthood, the 
process taking about one month. We have not found any case of parasitism yet. 

Males are bright orange dorsally, while the females are a dull orange dorsally. 
Both sexes have black margins on both wings and a black subapical band. When 
handled, males extrude a double gland under the genitalia, and females a semicircular 
one above the genitalia. Due to the interference of these glands, we have been 
unable to obtain hand pairing with this species. Both sexes produce a punget scent 
when disturbed. 

Many times we have witnessed nuptial flights, and always the male has been the 
active flyer, the female hanging limp. On 14 August 1971, shortly before noon, a 
pair consisting of a fresh male and an old and damaged female was observed in 
copula on a low shrub. When disturbed with the handle of a net, the male took 
flight with the female hanging motionless. They alighted some 20 m away in low 
vegetation. Three times we forced the pair to move, and everytime the same thing 
happened. The pair was then netted and brought home, still in copula, in a plastic 
bag. Next day, the female was put in the bag on a Passiflora sp. vine and was 
left there until the morning of the 17th, when it was killed and dissected. It had 
laid 37 eggs on the vine, and no eggs were found in the abdomen. 

On 5 October 1972, another pair was observed copulating. This time both the 
male and female were recently emerged. They were found at 0920 on the vine where 
they had been reared. While being observed from some distance, the male took 
flight, carrying the passive female, and alighted some 10 m away at 6 m above 
the ground on a white wall, where the pair was very conspicuous. They stayed 
motionless, male above, for one hour, until forced to fly into a wire cage. They then 
stayed on the side of the cage, male above, without further movement until copula- 
tion was ended at 1455. At that time both butterflies started flying in the cage 
trying to escape. Again, the female was put in a plastic bag on the vine for two 


VOLUME 27, NUMBER 4 303 


days. Then it was killed and dissected. One egg had been deposited on the vine, 
and four were found in the abdomen. 

The vertically oriented eggs are yellow, about 15 mm long by 1 mm wide, 
covered by a reddish coating, and with ribs and sculpturings somewhat resembling 
Danaidae eggs. They hatch in four days. First instar larvae, about 2 mm long, 
are yellowish, head and body. At this stage the head shows no markings or spines, 
but has some scarce fine setae. The body has no markings or spines either, but each 
segment has a transverse row of fine, dark setae. From the second instar, on the 
head shows white and reddish markings and two spines, one on each epicranium. 
The body is covered by rows of spines and shows white and reddish markings, such 
as described in great detail by Richard (1968, J. Lepid. Soc. 22: 75-76) for Dryas 
iulia delia ( Fabricius ). 

The larva of D. i. iulia, upon emerging from the egg, eats the egg-shell, at least 
partially, and then moves to the edge of the leaf where it starts feeding. It leaves 
small hanging sections of the leaf that soon wilt and dry, forming an excellent 
camouflage. The first, second and sometimes third instar larvae are easily located by 
examining the leaves that show these ragged edges. Larger larvae move about the 
vine and can cause a mild rash on the skin when touched with the back of the hand. 

It is interesting to note that in the two cases of copulation we are reporting, the 
young female laid only one egg and had four in the abdomen, whereas the old and 
damaged female laid 37 eggs and had none left in the abdomen. The females prob- 
ably copulate several times during their life span and lay considerably more than 30 
eggs, contrary to Labine’s suggestion (1968, in Young 1972, Acta Biol. Venez. 8: 
1-7) that most species of Heliconiinae probably lay less than 30 eggs during the 
average females lifetime. Another species that tends to contradict Labine’s sugges- 
tion is Dione iuno huascama Reakirt, whose female often lays groups of over 100 
eggs during a single sitting (pers. obs.). This is the only heliconiid with gregarious 
habits all through its developmental stages that we have found in EI] Salvador. 

As seen in the description above, in D. iulia iulia the male is the active flying 
partner. In most of the reports on butterfly copulation we have found, (Pronin 
1964, J. Lepid. Soc. 18: 35-41; Ferris 1969, J. Lepid. Soc. 23: 271-272; Carcason 
1970, J. Lepid. Soc. 24: 72; Jae 1972, J. Lepid. Soc. 26: 28; Priestaf 1972, J. 
Lepid Soc. 26: 104), the active partner usually has been the female. 

A final comment: the description by Richard (1968, op. cit.) of the early stages 
of Dryas i. delia matches exactly the early stages of Dryas i. iulia, except for the 
first instar and the fact that we have always found five larval stages instead of four. 
We suspect that the larvae he collected and thought were first instar were actually 
second instar larvae. 


ALBERTO MuysHonpT, 101 Avenida Norte #322, Lomas Verdes, San Salvador, 
El Salvador. 


NEW MEXICAN SPHINGIDAE RECORDS 


Since Hoffmann’s catalogue of Mexican sphingids (1942, An. Inst. Biol. Univ. Nal. 
Autoén. México 13: 213-256), few species have been added. Some of them like 
Phryxus caicus Cram. and Callionima nomius Wlk. were recorded in 1967 (Beutel- 
spacher & Vazquez 1967, An. Inst. Biol. Univ. Nal. Auton. México 38, Ser. Zool. (1): 
75-77). Hoffmann’s catalogue, however, requires numerous generic changes which 
we will be making soon, based on Hodges (in Dominick et al., 1971, Moths of America 
North of Mexico, fasc. 21, Sphingoidea). 

Thanks to Mr. Roberto de la Maza and his sons, we are leaming of more new 
species for Mexico, and at this time can add two new genera; one of Neotropical, and 
the other of Palearctic origin. 


304 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Amphimoea R. & J. A. walkeri Bdyv. (= staudingeri Drc., magnificus Rothsch. ) 


Amphimoea is a monotypic genus and the species A. walkeri, according to Druce 
(in Godman & Salvin, 1886, Biologia Centrali-Americana, Insecta, Lepidoptera 
Heterocera 2: 311; 3: pl. 65-4) and Draudt (in Seitz, Gross-Schmetterlinge der 
Erde, VI, Heter. Amer.: 847, 865), is distributed from Chontales, Nicaragua, Chiriqui 
Volcano, Panama to south of Brasil. These authors mention that the caterpillar 
feeds on a Jatropha species. 

One female specimen from Dos Amates, Catemaco, Veracruz, taken in August 
1965, is in the De la Maza collection. In Hoffmann’s catalogue, this genus must 
be situated before Manduca Hbn. 


Cressonia Grt. & Rob. C. juglandis (J. E. Smith) (Sphinx juglandis J. E. Smith) 


This genus of Palearctic origin, previously known only from the United States, is 
recorded now for the first time in Mexico. According to Hodges (op. cit.), “the 
larvae feed on various species of hickory (Carya species) and walnut or butternut 
(Juglans species), and perhaps beech (Fagus species ).” 

The following specimens are in the De la Maza collection: one male from 
Parque Funeral Guadalupe, Monterrey, Nuevo Leén, 17 June 1970; one male and 
one female from El Barrial, Monterrey, Nuevo Ledn, 10 September 1971. In 
Hoffmann’s catalogue, this genus must be situated after Paonias Hbn. 


Carios R. BEUTELSPACHER B., Instituto de Biologia, Apdo. Postal 70-233, México 
20, D.F. México, & ROBERTO DE LA MAzaA, Jr., Nicolas San Juan 1707, México 12, 
D.F., México. 


APODEMIA MORMO NEAR DIALEUCA (RIODINIDAE) FROM 
MONTANE SOUTHERN CALIFORNIA: NEW FOR U.S.A. 


Eleven male and 23 female unusual-appearing Apodemia mormo (Felder and 
Felder) were taken on 3 and 5 June 1966 by John Justice, Keith Hughes and the 
writer on Sugarloaf Ridge, north of Barton Flats, San Bernardino Mountains and 
County, California. They were closely associated with the prostrate buckwheat, 
Eriogonum wrightii subscaposum Wats. (Polygonaceae), growing on gravelly slopes 
above 2600 m elevation in open mixed deciduous-coniferous forest dominated by 
ponderosa pine, mountain mahogany and manzanita. The entire series was shown 
to Dr. Jerry Powell, who found them nearly identical to A. m. dialeuca Opler & 
Powell, described from similar habitats in the Sierra San Pedro Martir of Baja 
California, Mexico, nearly 500 km south of Barton Flats (Opler & Powell 1961, J. 
Lepid. Soc. 15: 145-171; Patterson & Powell 1959, J. Lepid. Soc. 13: 229-235). 
The wings dorsally have large white spots on a dark gray background with little if 
any red or orange suffusion, lending a distinctly tesselated appearance to the insect; 
the ventral surfaces are dusted generously with light gray scales, especially on the 
outer one-third of the primaries and outer two-thirds of the secondaries (Fig. 1). 

Subsequently other collectors have taken examples of this phenotype at the same 
and nearby localities, including some in September and early October suggesting 
multivoltinism (Opler, in litt). It is rather peculiar that this interesting population 
remained undiscovered until recent years. The fact that collectors like the Sperrys, 
Rindge, Comstock, Martin and others failed to turn it up in many years of intensive 
collecting during the last half-century causes one to wonder if it might be a very 
recent segregate from A. m. virgulti Behr populations to the south and west, or from 
A. m. near mormo and “blend-zone” populations to the east, north and northwest at 
lower elevations. The nearly exact similarity in facies, habitat and probable host- 
plant of this new population and topotypical dialeuca suggests either a common 
phenotypic response to similar environmental conditions, or previous ecologic and 


VoLUME 27, NuMBER 4 305 


RAL YT Oe 


Fig. 1. Male (upper left dorsal, and upper right ventral, surfaces of same speci- 
men) and female (lower left dorsal, and lower right ventral, surfaces of same 
specimen) Apodemia mormo near dialeuca Opler & Powell, Sugarloaf Ridge 8500’ 
(2600 m), San Bernardino Mountains, San Bernardino County, California, 5 June 
1958, leg R E Stanford. 


genetic continuity between the presently disjunct populations. Future studies wil! 
be necessary to clarify the taxonomic status of this new population, its geographical 
distribution and relationship to contiguous elements of the mormo complex. For 
the present, Apodemia mormo near dialeuca can be added to the butterfly fauna 
of California and the United States. (Thanks to Keith Hughes, Paul Opler and 
Jerry Powell for assistance in preparing this note, and to Louis Brunelle of Fort Dix, 
New Jersey for the illustration. ) 


Ray E. Stanrorp, MD, 720 Fairfax Street, Denver, Colorado 80220. 


BOLORIA SELENE (NYMPHALIDAE) AMBUSHED BY A TRUE BUG 
(HETEROPTERA ) 


Predator-prey relationships between butterflies and other insects have been in- 
frequently noted in the literature. An instance of a robber fly eating Papilio eurymedon 
Lucas is known to me (correspondence with T. Rogers) and dragonflies have been 
seen to capture and consume butterflies. But the following account is the first 
observation of which I am aware of a true bug preying upon a butterfly. 

On 20 August 1970, I visited the Moxee Bog Reserve in Yakima County, Wash- 
ington. This site, protected by The Nature Conservancy through the efforts of 


306 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Society member David McCorkle, nurtures one of the few known colonies of Boloria 
selene Denis and Schiffermueller in the state. The habitat here is an anomaly—a 
quaking bog in sagebrush desert. Both the bog and the insect seem to be glacially 
relict features. Various boggy pockets in the Columbia River Basin serve as refugia 
for formerly widespread species of plants and animals. These species were largely 
wiped out by regional flooding of the basaltic coulee country which happened 
following melting of the Pleistocene ice dams on Glacial Lake Missoula. 

I was aware of the historic enigma of Moxee and was visiting it as a general 
exercise in nature perception. My specific objective was to photograph selene and 
to observe its behavior for a work on Washington butterflies. I first encountered 
the insect along the ecotone where cattail marsh and sedge bog meet a drier, 
alkaline bench dominated by a native iris. Throughout the afternoon, selene was 
uncommon and perched infrequently after long, sustained and fairly strong flights. 
As sunset approached, the butterflies became suddenly very abundant (or rather, 
much more in evidence). Scores of fresh individuals of both sexes were now on 
the wing deeper into the wettest parts of the bog. Flights grew shorter and slower, 
perching occurred more frequently. Many of the Silver-bordered Fritillaries clung to 
grass blades, while others fed on five species of flowers: A Verbena, Rabbitbrush 
(Chrysothamnus nauseosus), two yellow daisies and a mint (Mentha). Coenonympha 
tullia was also abundant, but I observed no encounters between the two species of 
butterflies. Mating activity became intense during the last half hour of direct sun- 
light. Many pairs of selene were observed courting and copulating, the latter ac- 
complished in situ and never in flight. Immediate precopulatory behavior consisted 
of a gentle, slow fluttering of the wings as the two partners orbited one another 
by crawling around the grass blades to which they both clung. 

At about 1800 (PDT) I noticed an individual of selene in an unusual position. 
A fresh female, it was wedged upside down between two flower heads of Rabbit- 
brush, with its wings spread open as if mounted. Thinking at first it was simply a 
strange feeding posture, I set about exploiting the excellent photographic op- 
portunity thus afforded. Soon I noticed that the butterfly moved only feebly 
and indeed appeared to be immobilized. Looking closely, I was astonished to see 
several true bugs clustered on the same flower heads that bore the butterfly. Three 
of the heteropterans were visible, and two were in contact with the butterfly at the 
time. Their refined camouflage (yellow, mottled with black) accounted for my 
initial failure to see them against the Rabbitbrush florets. As I watched, the bugs 
clambered over the butterfly, presumably imbibing its fluids with their probosci, 
which often came into contact with the hapless selene. A small yellow crab spider 
occupied the same flower head, but did not appear to be implicated in the 
entomophagy. I collected the butterfly (which soon died) and one of the suspected 
assassins. When the selene became desiccated, its abdomen clearly showed a small 
hole in its side where it had been pierced. 

The bug turned out to be a member of the family Phymatidae, probably of the 
genus Phymata Latreille. These bugs are known, most aptly, as Ambush Bugs. 
W. S. Blatchley (The Heteroptera of Eastern North America, Indianapolis, 1926) 
wrote this account of Ambush Bugs and their tactics: “Only about 80 species of 
the family are known, 14 from North America. They are all predacious in habit, 
hiding themselves in the heads of flowers, especially Compositae, where they await 
the coming of bees and other nectar-seeking insects. When the prey is within reach 
the bug makes a quick stroke with its sabre-like fore tibiae, draws the victim within 
reach of its beak and then leisurely sucks it dry.’ The behavior of my predator 
obviously matched the family characteristics, and the means of actual capture, which 
had baffled me, is explained. Surely the Ambush Bug deserves its name if it can 
sieze and overpower an insect so relatively large, strong and fleet as a Silver- 
bordered Fritillary. However, one bite from a Phymata, quickly executed and well 
placed, might well disable an even larger prey than selene. I know. As I was 


VOLUME 27, NUMBER 4 307 


returning to my car after this hot, damp adventure in Moxee Bog, I felt an excruciating 
pain in my foot. Cursing, I tore off my boot and found the vector of my intense 
discomfiture: an Ambush Bug. I had shared selene’s fate, and came out only 
slightly better: my foot swelled and throbbed for hours afterward. 

Was there ethological significance to the prey-positioning? Do heteropterans 
usually prey communally when butterflies are the target? I would be most interested 
in reactions to these questions, and in reports of insect predation on butterflies in 
general. Thanks be to Dr. Dennis Paulson of the Department of Zoology, University 
of Washington, and to the library staff of the Royal Entomological Society of 
London, for aid. in identifying the Ambush Bug. 


Rosert MicHAEt Pyie, School of Forestry and Environmental Studies, Yale Uni- 
versity, New Haven, Connecticut 06520. 


BOOK REVIEW 


SoutH’s BRITISH BUTTERFLIES, by T. G. Howarth, illustrated by A. D. A. Russwurm 
and R. B. Davis. 1973. xiii + 210 p., 48 colour plates, 26 drawings, 57 maps. Cased. 
Frederick Warne & Co. Ltd., London and New York. Price £10.50 UK. 


PROVISIONAL ATLAS OF THE INSECTS OF THE BritisH Istes: Parr 1 LEPIDOPTERA 
RHOPALOCERA, BUTTERFLIES, edited by John Heath. 1970. iii p., 57 maps. Natural 
Environment Research Council. Agent: E. W. Classey Ltd., Middlesex, England. 
Price 50 p. UK. 


PROVISIONAL ATLAS OF THE INSECTS OF THE BRITISH ISLES: PART 2 LEPIDOPTERA 
(Morus—Part ONE). LASIOCAMPIDAE: SATURNIIDAE: ENDROMIDAE: DREPANIDAE: 
THYATIRIDAE: SPHINGIDAE: NOTODONTIDAE: LYMANTRIIDAE: ARCTIIDAE: NOLIDAE, 
edited by John Heath and Michael J. Skelton. 1973. iii+ 3 p., 102 maps. Natural 
Environment Research. Council. Agent: E. W. Classey Ltd., Middlesex, England. 
Price £1.00 UK. 


The butterflies of Britain must be the most intensely studied butterfly fauna in 
the world. The tradition of butterfly collecting is probably stronger there than any- 
where else, extending back to the parson-naturalists and other curious people in the 
eighteenth century, who have had to deal with a fauna of only 61 resident species 
(fewer than Long Island, and five of them now extinct), 3 regular migrants and a 
total of 41 assisted and unassisted strays and immigrants. This great band of bug- 
hunters has successfully recorded the distribution of the butterflies of most of England, 
about half of Scotland and scattered areas of Ireland; compiled by computer these 
records provide intensive distribution maps of all 56 residents, plotted on a 10 
kilometre grid covering the whole area. Intended as the basis for conservation of 
dwindling species (they clearly show the decline of many, and the apparent extinc- 
tion of one moth), these maps, now published provisionally for the butterflies and 
many of the more spectacular moths, are a triumphant combination of amateur 
natural history with professional data-processing; they will be of particular interest 
to the zoogeographer, and invaluable to the collector, observer or photographer, who 
with the aid of British Ordnance Survey maps or the AA Book of the Road, all 
bearing the same reference grid, can now pinpoint his quarry to within 100 square 
kilometres. Discreet enquiries with the Nature Conservancy can often produce the 
exact spot for the really localised species. 

The butterfly maps re-appear, considerably improved by new records for the com- 
mon or secretive species, and for the less scenic parts of the North and Scotland, 


308 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in the magnificient re-written version of Richard South’s The Butterflies of the 
British Isles. For more than half a century this knapsack-sized, charming but slightly 
rambling guide has remained the standard work, while many other books, bigger and 
smaller, came and went. The book now re-emerges as a properly arranged, large- 
format, popular monograph, the text for each species divided by sub-headings for 
easy reference, with a fine set of illustrations of the butterflies by A. D. A. Russwurm, 
and of the eggs, final instar larvae (including a highly magnified seventh segment ) 
and pupae, copied by R. B. Davis chiefly from original paintings by F. W. Frohawk; 
it is a pleasure to report that the printing of the beautiful Frohawk drawings is far 
better than in their first production in the two-volume limited edition of Frohawk’s 
Natural History of British Butterflies in 1924. 

There is no doubt that this will continue as the standard work on the subject, 
combining as it does the outstanding features of two earlier works with the latest 
information on the biology of the British species, and showing the influence of new 
attitudes to lepidopterology which were pointed by E. B. Ford’s seminal book 
Butterflies. Some of South’s original text from the more leisured world of 1906 is 
retained. To say of the Small Copper’s habit of flying at other butterflies, “Whether 
these seeming attacks are really due to pugnacity . . . or are merely of a sportive 
character, is not altogether clear; when the meeting is between two Small Coppers 
it usually results in a series of aerial evolutions by the pair, so it would seem that 
there is a good deal of playfulness in the business,” sounds strange today (though 
couched in the jargon of the behavioural sciences it would be little more informative ), 
but it makes pleasant reading. The short section on the history of each species, 
usually detailing the first discovery of the butterfly in Britain, and including a 
record of the Painted Lady from 1272, is also great fun. (I have a poser here for 
lepidopterists: what is the earliest, recognisable picture of a butterfly species? 
Any advance on the Meadow Brown and Small Tortoiseshell in The Garden of 
Delights by Bosch, ca 1503? ) 

British collectors, with their small fauna and accessible countryside, have lacked 
the lure of native rarities (although it is surprising what they missed in Scotland 
and Ireland), and have tended, like the not so wealthy stamp collector unable to 
afford the great prizes of philately, to go for “printing errors.” The space devoted 
to these, with their Latin names, is a thought overgenerous (though it will please 
a good many collectors), but it is good that considerable attention is now given 
to geographical variation as well. The amateur lepidopterist could well move with 
the times and become, in the real sense of the word, more conscious of ecology; 
there are signs that the distribution map scheme is providing a welcome stimulus 
by making the collection of data for those 3600 grid squares at least as challenging 
as hunting for mutants. I hope that by the time the next successful edition appears, 
it will be possible to compile more information on such things as habitat preferences 
and limits of distribution; there is for instance the marked coastal distribution of the 
Grayling, which few would have guessed at, and which goes unremarked in the text. 
I believe that the literature is extensively misleading on some species. Is it true 
that the Ringlet is a shade butterfly? And then every book I have seen states that 
the Green-veined White does not occur in cultivated land. Now while it is true that 
a Pieris in a wild place, particularly in Scotland where the other two species are 
comparatively rare, is overwhelmingly likely to be Green-veined, the reverse is not 
true: this species was common and co-existed with the others in the gardens and 
parks of suburban Liverpool when I was a boy. The impossibility of separating 
British Pieris on the wing may have allowed this error to go unchallenged. There 
is still much to be done on the distributions and voltinism of the two (or one-and-a- 
half) species of British Aricia. 

My only qualm about this lovely new book is for the plight of the beginner and 
the schoolboy, who need a little more help with identifications which are childsplay 
to Graham Howarth; how does one distinguish the various female Blues and Hair- 


VOLUME 27, NuMBER 4 309 


streaks at a glance, and for the rank beginner or general naturalist, what distinguishes 
the Grizzled and Dingy Skippers? Confusion of these last resulted in several in- 
correct records, now expunged, in the Provisional Atlas. The schoolboy (or school- 
girl, if she is liberated enough to enter this male preserve) might well forego this 
book till his pockets are larger, and find a second-hand copy of A Butterfly Book 
for the Pocket by Sandars, which despite its amateur illustrations and frankly sur- 
realist distribution maps, did come close to the Peterson identification system. For 
the visiting American collector, who will already be familiar with some of the 
species, the choice is between the Houghton-Mifflin Field Guide to the European 
butterflies, and this new work. Outside Britain, one must have the more extensive 
Guide, but “South” will give him far better service on a stay in Britain, for he will 
not be repeatedly tracking down his specimen to something found only in the Balkans 
(though in “South” beware the pictures of spectacular rare aberrations). And then 
there are those detailed maps, which could nearly double one’s number of species. 
Irish and Scottish Americans visiting their homelands should get a set of distribution 
report cards from the Nature Conservancy as well; the data are badly needed! 

And of course, if you have a library of standard works on the world’s butterflies, 
add this one. It is worth every decimalised penny. 


Joun R. G. Turner, Department of Ecology and Evolution, Division of Biological 
Sciences, State University of New York at Stony Brook, Stony Brook, New York 11790. 


AN INDEX TO THE DESCRIBED LIFE Histories, EARLY STAGES AND HOSTS OF THE 
MACROLEPIDOPTERA OF THE CONTINENTAL UNITED STATES AND CAnapA, by Harrison 
M. Tietz, 1972. v + 1041 p. in 2 volumes, cloth bound. Allyn Museum of Entomology, 
Sarasota, Florida. $25.00. Distributed by Entomological Reprint Specialists, P.O. 
Box 77971, Dockweiler Station, Los Angeles, California 90007. 


This large work of compilation by Harrison Morton Tietz (1895-1963) was in- 
tended to provide references to all published information on the early stages of 
macrolepidoptera for America north of Mexico, being the only index of this nature 
to appear since Henry Edwards’ “Bibliographic Catalogue of the Described Trans- 
formations of North American Lepidoptera” (1889, Bull. U.S. Natl. Mus. 35.). Its 
coverage appears to be reasonably thorough to about 1950, when the author left 
off his search of the literature to prepare the manuscript. Negotiation with the 
Smithsonian Institution to publish the work continued for a time thereafter, but 
final arrangements concerning changes in format, etc., were never completed. The 
text as now issued is just as it was left by Tietz, although retyped. With the exception 
of a 2-page introduction by William D. Field and J. F. Gates Clarke, which has 
the appearance of letterpress printing, the text was reproduced from typed copy 
by photo offset. The 2 volumes have good quality buckram bindings which in 
themselves are worth a large part of the purchase price. 

The work is divided into two major parts (which do not coincide with the bound 
volumes). Part 1 contains a list of entomological publications consulted (23 p.), 
a list of common names of Lepidoptera (33 p.), and the most important section, the 
list of references to published life history information, indexed alphabetically by 
species with hosts listed for each (636 p.). Part 2 contains an index of common and 
scientific names of food plants listed by common names (101 p.), and an index of 
hosts listed mainly by scientific names, each with a listing of all the species of 
macrolepidoptera reported to feed on it (221 p.). The work ends with a 12-page 
list of changes in nomenclature for plant names, giving the old name and the cor- 
rected equivalent used in the host index of part 2. 

The nomenclature follows rigidly that of the McDunnough check list of 1938, 
and no effort was made to correct names or revise the manuscript in any way. 
Admittedly, the up-dating of so large a work would have been a demanding and 
thankless task for anyone not credited with authorship and would have further 


310 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


delayed publication, perhaps indefinitely. The Lepidoptera species index includes 
synonyms and subspecies names, under each of which the reader is referred to the 
applicable species name. Inclusion of all such names seems unnecessary or even 
confusing, because life history information was only occasionally published under 
the synonymic name, and references cited may or may not apply to the subspecies 
listed. 

Sometimes the bibliographic references under a species name are missing; e.g., for 
Apaecasia atropunctata (p. 123), Eupithecia bradorata (p. 145), Septis castanea 
(p. 169), Elaphria chalcedonia (p. 181), and Catocala subnata (p. 620—no hosts 
or references). In a few instances the food plant list includes unlikely hosts not 
mentioned in any reference given; e.g., under Carsia paludata (p. 505) 7 hosts are 
listed, although in the one reference cited only Vaccinium is mentioned. The hosts 
of Apaecasia subaequaria (p. 617) are given as ferns and grasses, but neither of the 
references mentions grasses (Rupert, 1949, specified braken fern). 

The casual transliteration of plant names can be tricky, and I noticed one instance 
where this led to an obvious mistake. The eastern skunk cabbage, Symplocarpus 
foetidus (L.), is listed in error as one of the food plants of Arzama obliqua (p. 480). 
The host actually reported in the literature (Guppy, 1948) was Lysichiton (sic) 
kamtschatcense Schott, a different member of the Araceae that is known as skunk 
cabbage on the Pacific Coast. 

Throughout the work there are numerous typographic errors, especially in the 
spelling of scientific names, such as atlanta for atalanta (p. 119), Catabens for 
Catabena (p. 273), focosa for fucosa (p. 301), idalis for idalia (p. 342), uo for io 
(p. 363), lutea for luteata (p. 409), maculatz for maculata (p. 413), radicans for 
radians (p. 563), rockesi for ruckesi (p. 580), Agertum for Ageratum (p. 674), and 
curimacula for curvimacula (p. 809). I do not know whether these misspellings 
date from the final retyping or were reproduced verbatim from Tietz’s original copy. 

I noted the following errors of collation which owners of these volumes should 
carefully mark. They cannot be detected by the page numbering, which is un- 
interrupted. 

(1) Pages 147 and 148 are transposed (giving the false impression that poplar 
and willow are the food plants for Papilio brevicauda, instead of for Catocala briseis 
which begins at the bottom of p. 148). 

(2) Page 220 is misplaced and belongs between pages 210 and 211. 

(3) The left column on page 207 does not refer to Callophrys comstocki as it 
reads, but to Everes comyntas, the name of which does not appear at all. Pos- 
sibly a page is missing here but I have not found it. 

(4) Pages 264 and 265 are transposed. 

Although my criticism may seem lengthy, this is an important reference work 
of over 1,000 pages, promising a greatly simplified means of access to the literature 
of the first 150 years of research on the early stages of North American macro- 
lepidoptera. Despite its faults and lack of coverage for the last two decades, the 
work will undoubtedly be extremely useful. Few entomologists concerned with 
life history information on Lepidoptera can afford to ignore it. 


Doucias C. FERGcusON, Systematic Entomology Laboratory, USDA, c/o U.S. 
National Museum, Washington, D.C. 20560. 


BUTTERFLIES OF AUSTRALIA, by I. F. B. Common and D. F. Waterhouse. 1972. Angus 
and Robertson (Publishers) Pty. Ltd., Sydney, Australia, etc. 498 p., 41 plates (26 in 
color), 25 figs., 366 maps. Price: $37.50 (U.S.). Distributed exclusively in the 
United States by Entomological Reprint Specialists, P.O. Box 77224, Dockweiler 
Station, Los Angeles, California 90007. 


Australian lepidopterists are a fortunate breed: in the space of two years three 
fine books on their butterfly fauna have appeared. From a scientific viewpoint the 


VOLUME 27, NUMBER 4 ot 


present volume is the best of the lot, though from an esthetic standpoint the McCubbin 
book is more pleasing and the plates more readable, and the D’Abrera book covers 
more territory (see reviews of these books, J. Lepid. Soc. 26: 200-202). 

The introductory chapters on “Structure and Life History,” “Biology,” “Behaviour 
and Other Aspects of Physiology,’ “Geographical Distribution” and “Classification 
and Nomenclature” are well-written, concise and informative. The authors have 
wisely side-stepped the matter of continental drift in their explanation of the 
geographic distribution of Australian butterflies and the derivation of these insects 
from those in other regions. 

The general systematic accounts are excellent, but the arrangement is somewhat 
inconsistent. Many workers will have difficulty adjusting to the arrangement of 
families (Hesperiidae, Papilionidae, Pieridae, Nymphalidae, s. 1., Libytheidae and 
Lycaenidae), presumably from primitive to specialized. Within families different 
arrangements are followed. The Coeliadinae are generally considered more “ad- 
vanced” than are the Trapezitinae in the Hesperiidae, but following Evan’s 
Catalogue the Coeliadinae are placed first. Conversely, the satyrids are arranged 
from “primitive” to “derived” in accordance with my 1968 revision. These in- 
consistencies by no means detract from the usefulness of this book, they only serve 
to distress the taxonomist slightly. 

Each species description is accompanied by a distribution map, hopefully setting 
a pattern for similar works in other regions (Riley and Higgins also did this in 
their European book). Unfortunately, all the maps are uniform—more information 
on very localized species could have been derived from a map of a smaller area. The 
authors are nevertheless to be congratulated on the inclusion of maps. 

The figures are quite usable. Genitalic figures are given for only those species 
that cannot be distinguished superficially. The larval and pupal figures could have 
been stronger had they been accompanied by line representations of chaetotaxy, 
perhaps in the text. 

The section on “Collection and Study” is informative and applicable to any 
region, the “Glossary” defines technical terms and the “Food Plant List” summarizes 
the known butterfly-plant relationships. The currency of the book is attested to by 
inclusion of new information that came to the authors’ attention after completion 
of the manuscript. 

There are few typographical errors, one of the most glaring being “pleural” in- 
stead of “plural” on page 56. Such errors must be accepted; some are completely 
unavoidable. 

On balance, the authors have done an excellent job, and if one is interested in 
Australian butterflies, he must have this book. It can stand alone as a reference for 
the area and is a worthy successor to G. A. Waterhouse’s forty-year-old What 
Butterfly Is That? This work should be a standard forty years from now. 


Lee D. Mitxer, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, 
Florida 33580. 


NOTES AND NEWS 


FIRST KARL JORDAN MEDAL AWARDED TO HENRI STEMPFFER 


The Karl Jordan Medal (J. Lepid. Soc. 26: 207-209) was awarded for the first 
time at the annual banquet during the Society’s 24th Annual Meeting in Sarasota, 
Florida, on 23 June 1973. The Jordan Medal was presented to M. Henri Stempffer 
of Paris, France, by the sponsor of the medal, Mr. A. C. Allyn. 

M. Stempffer was awarded the medal for his various works on the Lycaenidae, 
especially those of Africa, culminating in his “The Genera of the African Lycaenidae 


312 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


y yyy ff) Yj” 
WY Y Yui Yj 
Ui, y 


Gy gY 


Presentation of the Karl Jordan Medal, Sarasota, Florida, 23 June 19 
left: Henri Stempffer, Mme. Stempffer, A. C. Allyn. 


(Lepidoptera: Rhopalocera)” (1967, Bull. British Mus. [Nat. Hist.], Suppl. 10: 
322 p., illustrated). These careful pieces of work have become standards for students 
of the African fauna in a relatively short period of time. 

It was gratifying to many of us who had long used M. Stempffer’s works to have 
the opportunity to meet him in person. Though he has never had a formal academic 
connection, M. Stempffer’s work has placed him in the forefront of workers on the 
African fauna, and he is truly representative of the type worker for whom the Karl 
Jordan Medal was intended. 


Lee D. Muter, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, 
Florida 33580. 


It is time again to thank a number of persons for their assistance to me during the 
past year. The members of the Editorial Committee of the Journal willingly reviewed 
most of the submitted manuscripts. In addition, the following individuals reviewed 
one or more manuscripts upon request: D. E. Berube, L. P. Brower, H. K. Clench, 
H. A. Freeman, D. F. Hardwick, C. G. Kellogg, C. D. MacNeill, E. G. Munroe, J. A. 
Powell, F. H. Rindge, and O. R. Taylor. The helpful assistance of all of these people 
is gratefully acknowledged. Two editorial assistants, Nancy Wells and Elaine Doyle, 
served with patience and skill, and I especially thank them for their efforts. The 
cover drawing (Sphinx vashti Strecker) was kindly provided by William H. Howe. 


THEODORE D. SARGENT 


VOLUME 27, NUMBER 4 ols 


INDEX TO VOLUME 27 


Author Index 


Becker, V. O., 160 
Beutelspacher B., C. R., 303 
Blanchard, A., 103, 141, 219, 278 
Borch, H.., 196 

Brown, ik, N., 136, 238 

Burris, D. on 84 

Clarke, C. A., 73 

Clarke, J. F. Gates, 99, 240, 268 
Collins, M. M., 225 

Covell, C. V., Jr., 144, 206 
Davis, D. R., 159 

Dimock, T. E., 274 

Dominick, R. B., 1 

Downes, J. A., 89 

Erickson, J. M., 16 . 

Ferguson, D. C., 288, 309 
Perms. D. 57, 112, 279 
Fisher, M. S., 112, 239 

Freeman, H. A., 40 

Godfrey, G. L., 119 
Goodpasture, C., 109 

Kean. P. j., 122 

Masters, J. H., 33, 78, 80, 86, 235 
Miller, L. D., 310, 311 

Munroe, E., 290 

Muyshondt, A., 15, 210, 294, 302 
Neck, R. W., 22 


Newcomer, E. J., 13 
Nicolay, S. S., 243 

Rankse Dy hes ky 

Perkins, E. M., Jr., 291 
Rite AS Pe 122, 

Pyle, R. M., 305 

Rawson, G. W., 206 

Rehr, S. S., 237 

Rindge, F. H., 140 
Sargent. Dao. E75. ol? 
Schmid, F., 196 

Scott, J. A., 283 

Selman, C. L., 87 
Sevastopulo, D. G., 143, 155, 157 
Shapiro, A. M., 79, 85, 157, 159, 238 
Sharp, M. A., 17 
Sheppard, P. M., 73 
Sibatani, A., 161 

Smith, A. G., 73 

Stanford, R. E., 304 
Straley, G. B., 144 
Turner, J. R. G., 130, 307 
Van Buskirk, M. D., 83 
Wagner, W. H., Jr., 192 
Welling M., E. C., 154 
Young, A. M., 258 


Name Index 


(New names in boldface) 


Acherontia atropos, 143 
Amblyscirtes aesculapius, 51, 146 
alternata, 54 
anubis, 55 
belli, 54 
carolina, 51, 146 
cassus, 47 
celia, 53 
elissa, 49 
eos, 52 
erna, 47 
exoteria, 46 
fimbriata, 56 
florus, 55 
fluonia, 49 
folia, 44 
immaculatus, 46 
linda, 48 
nereus, 52 
nisulae-pinorum, 46 


nysa, 52 

oslari, 48 

phylace, 55 

prenda, 50 

raphaeli, 44 

reversa, 51 

samoset, 49 

simius, 46 

tolteca, 50 

texanae, 50 

vialis, 53 
Amphimoea walkeri, 304 
Amplypterus donysa, 103 
Anacamptodes vellivolata, 94 
Anaea itys, 294 
Apodemia mormo, 304 
Appias drusilla, 209 
Arcyophora sylvatica, 97 
Argyria nivalis, 94 
Argyrotaenia quadrifasciana, 94 


314 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Argyrotoxa semipurpurana, 94 Eumarozia beckeri, 269 
Ascia monuste, 208 eleanthes, 271 
Aspilates donataria, 289 malachitana, 268 
Asterocampa celtis, 239 Eunica tatila tatilista, 208 
clyton, 239 Euphyes dion alabamae, 146 
Atalopedes campestris, 193, 238 dion dion, 146 
Atlides halesus, 284, 292 dukesi, 146 
Atrytonopsis hiana, 146 palatka, 146 
Battus philenor, 239 Euphyia centrostrigaria, 94 
polydamas, 208, 239 Eupithecia misturata frostiata, 94 
Blepharomastix ranalis, 93, 94 Euptychia gemma, 239 
Boloria selene, 305 hermes sosybia, 239 
Caberodes vavadaria, 288 Eurema lisa, 292 
Calephelis spp., 240, 241 Euthisanotia grata, 93, 94 
Callionima nomius, 303 Euxoa xasta, 103 
Callipsyche behrii, 13 Evergestis consimilis, 107 
Callithomia hezia perija, 80 Feniseca tarquinius, 89, 90, 91, 92 
skinneri, 78 Givira redtenbacheri, 109 
Callophrys augustinus, 284 Glenoides lenticuligera, 141 
johnsoni, 284 texanaria, 140, 143 
polios, 112 Glympis concors, 107 
polios obseurus, 115 Godyris caesiopicta, 261 
sheridanii, 279, 282 Gonodonta sinaldus, 105 
Calpe eustrigata, 97 Graphium marcellus, 239 
minuticornis, 97 Harkenclenus titus, 89, 91 
Catocala spp., 175 Heliconius aoede aoede, 131 
relicta, 9, 177 charitonius tuckeri, 208, 239 
Celastrina argiolus, 89, 91, 93 demeter sspp., 130, 133, 134 
Cercyonis pegala ariane, 292 egeria homogena, 131 
Ceratiscada canaria, 264 elevatus sspp., 130, 132 
Chlosyne lacinia, 22 erato sspp., 130, 131 
Choristoneura rosaceana, 94 eratoformis, 135 
Colias alexandra, 57 melpomene, 130 
alexandra columbiensis, 68 Hemiargus ammon bethunebakeri, 208 
eurytheme, 192 ceranus zachaeina, 292 
philodice, 89, 192 Herpetogramma pertextalis, 94 
Cosymbia pendulinaria, 11 Hesperia leonardus, 148, 193 
Cressonia juglandis, 304 metea, 148 
Cynthia annabella, 85, 274 pahaska, 239 
Damas immacula, 253 viridis, 239 
Danaus chrysippus, 73 Hyalophora cecropia, 226, 240 
gilippus berenice, 208, 239 colombia, 226, 240 
gilippus strigosus, 292 euryalus, 225 
plexippus, 92 gloveri, 225 
Desmia funeralis, 89, 94 “kasloensis,” 225 
Dircenna relata, 258 Hydria prunivorata, 93, 94 
Dryas iulia iulia, 302 Hylephila phylaeus, 238 
julia cillene, 208 Hymenitis nero, 262, 264 
Emariannia cucullidea, 105 Incisalia eryphon, 13 
E,piscepsis inornata, 103 fotis, 13 
Eriopyga mulina, 103 Iscadia daemonalis, 105 
Erora laeta quaderna, 285 Isoparce cupressi, 1, 83 
Erynnis lucilius, 89 Itame argillacearia-occiduvaria, 389 
zarucco, 239 exaospiciata, 94 


Euchlaena johnsonaria, 94 varadaria, 288 


VOLUME 27, NUMBER 4 


Kricogonia lyside castalia, 292 
Laspeyresia pomonella, 122 
Lerodea eufala, 238 
Lethe appalacia, 238 
Leuconycta diptheroides, 94 
Libytheana bachmanni larvata, 292 
Limenitis archippus, 89, 123, 239 
arthemis, 91, 123 . 
astyanax, 123 
lorquini, 127 | 
weidemeyerii, 127 
Lobocraspis griseifusa, 97 
Loxagrotis kyune, 119 
Lycaena dispar, 206 
helloides, 79 
phlaeas, 89, 90, 91 
Macaria inapta, 288 
Manduca, sexta, 122 
Marpesia petreus, 209 
Matigramma psegmapteryx, 105 
Mechanitis isthmia, 261, 265 
Mecyna submedialis, 94 
Megathymus yuccae, 144 
Meropleon titan, 105 
Metamorpha elissa, 235 
Mocis undata, 97 
Myscelia ethusa, 292 
Napeogenes tolosa amara, 262 
Nematocampa limbata, 94 
Neophanis respondens, 105 
Nymphalis antiopa, 193 
milberti, 193 
Ochlodes sylvanoides, 238 
Oeneis chryxus, 287 
uhleri, 287 
Ornithoptera priamus, 196 
Orthogramma prona, 105 
Ouleus dilla baru, 243 
Pamba boyaea, 249 
Panoquina ocola, 146 
panoquin, 145 
Pantographa limata, 94 
Papias trimacula, 249 
Papilio andraemon bonhotei, 136 
aristodemus ponceanus, 136, 206 
cresphontes, 207, 208, 239, 292 
glaucus, 239 
polyxenes, 16, 192, 237, 239 
Pentobesa valta, 107 
Phigalia titea, 10 
Phoebis agarithe maxima, 208 
Phyciodes frisia, 208 
phaon, 292 
tharos, 89, 91, 239 
Phyrxos caicus, 303 


Pieris rapae, 89 
Pingasa chlora crenaria, 96 
Platyptilia pallidactyla, 94 
Plebejus acmon, 109 
acmon lutzi, 282 
lupini sspp., 11, 109, 111 
neurona, 109 
saepiolus, 17 
Poanes yehl, 146 
Polites mystic, 89 
peckius, 89 
sabuleti, 238 
themistocles, 89 
Polygonia comma, 89, 91, 193 
interrogationis, 193, 239 
leo, 209 
Polygrammodes sanguinalis, 107 
Precis lavinia, 193 
Prepona omphale octavia, 210, 301 
Problema bulenta, 144 
Profilinota phillita, 101 
Pteronymia notilla, 262, 264 
Pyrausta orphrisalis, 89, 94 
Pyrgus oileus, 239 
Radara anartoides, 107 
Rhescipha servia, 107 
Satyrium calanus, 239 
caryaevorus, 144 
kingi, 144 
saepium, 283 
Scopula ancellata, 94 
enucleata, 94 
Scordylia atalanta, 107 
Siegela basipunctaria, 105 
Siproeta epaphus gadoui, 235 
epaphus trayja, 235 
stelenes sspp., 237 
superba sspp., 237 
Smyrna karwinskii, 15 
Sparganothis sp., 94 
Speyeria atlantis dodgei, 292 
cybele, 89, 292 
mormonia erinna, 292 
Sphinx juglandis, 304 
Staphylus mazans hayhurstii, 148 
Strymon saepium, 14 
Syntomeida melanthus, 103 
Tarachidia erastrioides, 93, 94 
Tephrosia texanaria, 140, 141 
Thespius inez, 255 
Thymelicus lineola, 144, 148 
Tosta sapasoa, 245 
Triozosneura dorsonotata, 222 
Urbanus proteus, 239 


316 


Vanessa atalanta, 239, 274 
virginiensis, 89 

Vettius chagres, 251 

Virga paraiba, 247 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Wallengrenia otho, 89, 148, 208, 239 
Xanthotype urticaria, 94 

Xystrota rubromarginaria, 94 

Zerene cessonia, 292 


Subject Index 


Aberrations, 79, 238 
Acraeidae, 169 
Agaristidae, 95 
Amathusiidae, 168 
Arctiidae, 95 
Behavior, 
communal resting, 15 
copulatory behavior, 291 
cryptic moths, 8, 175 
down valley flight, 283 
feeding at puddles, dung and carrion, 
90 
habitat selection, 17 
larval behavior, 261 
Book reviews, of, 
Commons, I. F. B. and Waterhouse, 
Dee olO 
Heath, J., 307 
Howarth, T. G., 307 
Owen, D. F., 86 
Tietz, Harrison M., 309 
Chromosome numbers, 109 
Collections, 
Dennis, 33 
Issiki, 159 
McAlpine, 240 
Newcomer, 241 
Vallin, 241 
Conservation, 137, 206 
Cossidae, 103 
Ctenuchidae, 103 
Danaidae, 36, 73, 167, 192 
Genetics, 73 
Genitalia (illustrations), 45, 82, 101, 142, 
221, 293, 246, 252, 254, 271, 272 
Geometridae, 10, 13, 95, 140, 141, 288 
Heliconiidae, 302 
Hesperiidae, 35, 40, 145, 167, 238, 239, 
343 
Hybridization, 231, 274 
Ithomiidae, 78, 80, 168, 258 
Karl Jordan Medal, 311 
Keys, 40, 82 
Larval foodplants, 
Anaea (Zaretis) itys, 296 
Brephidium exilis, 157 


Callophrys sheridanii neoperplexa, 282 
Callophrys sheridanii sheridanii, 279 
Chlosyne lacinia, 22 
Colias alexandra, 59 
Dircenna relata, 261 
Isoparce cupressi, 1, 83 
Loxagrotis kyune, 119 
Papilio polyxenes, 237 
Plebejus (Icaricia) acmon lutzi, 282 
Prepona omphale octavia, 210 

Libytheidae, 169 

Life histories, 
Anaea (Zaretis ) itys, 294 
Callipsyche behrii, 13 
Callophrys sheridanii sheridanii, 279 
Dircenna relata, 258 
Dryas iulia iulia, 302 
Incisalia eryphon, 13 
Incisalia fotis, 13 
Isoparce cupressi, 1, 83 
Ornithoptera priamus boisduvali, 196 
Ornithoptera priamus caelestis, 196 
Ornithoptera priamus demophanes, 196 
Prepona omphale octavia, 210 
Strymon saepium, 14 

List, state, 84, 144, 239 

Lycaenidae, 17, 39, 79, 109, 150, 157, 

159, 169, 279, 283 

Megathymidae, 145 

New genera, 49, 160, 219, 224 

New records (range extensions ) 
Aeschropteryx olivata, 107 
Amblyscirtes aesculapis, 146 
Amblyscirtes carolina, 146 
Amphimoea walkeri, 304 
Amphypterus donysa, 103 
Anartia jatrophae guantanamo, 85 
Apodemia mormo, 304 
Appias drusilla neumoegenii, 85 
Atlides halesus, 152 
Atrytonopsis hianna, 146 
Calephelis virginiensis, 148 
Callophrys augustinus croesiodes, 152 
Callophrys henria, 151 
Callophrys irus, 151 
Colias eurytheme, 148 


VoLUME 27, NUMBER 4 


Cressonia juglans, 304 
Emariannia cucullidea, 150 
Episcepsis inornata, 103 
Eriopyga mulina, 103 

Erora laeta, 152 

Euphyes dion dion, 146 
Euphyes dukesi, 146 

Euphyes palatka, 146 
Euptychia cymela viola, 85 
Euptychia gemma gemma, 85 
Euristrymon ontario, 152 
Euxoa xasta, 103 

Evergestis consimilis, 107 
Givira redtenbacheri, 109 
Glympis concors, 107 
Gonodonta sinaldus, 105 
Harkenclenus titus mopsus, 150 
Herminodes stigmaphiles, 105 
Hesperia leonardus, 148 
Hesperia metea, 148 

Hesperia pahaska martini, 239 
Iscadia daemonalis, 105 
Kricogonia lysidae, 85 

Lethe appalachia, 153, 238 
Lethe portlandia anathedon, 153 
Matigramma psegmapteryx, 105 
Megathymus yuccae, 145 
Meropleon titan, 105 
Neophanis respondens, 105 
Orthogramma prona, 105 
Panoquina ocola, 146 
Panoquina panoquin, 145 
Panthiades m-album, 152 
Phescipha servia, 105 

Phoebis statira floridensis, 85 
Phycoides frisia frisia, 85 
Poanes yehl, 146 

Polygonus leo, 85 
Polygrammodes sanguinalis, 107 
Problema bulenta, 146 
Problema byssus, 84 

Radara anartoides, 107 
Satyrium calanus falacer, 151 
Satyrium caryaevorus, 151 
Satyrium edwardsii, 151 
Satyrium kingi, 151 

Satyrium liparops strigosa, 151 
Scordylia atalanta, 107 

Sigela basipunctaria, 105 
Speyeria diana, 152 


Staphylus mazans hayhursti, 84, 148 


Syntomeida melanthus, 103 
Thymelicus lineola, 148 
Urbanus dorantes dorantes, 84 


317 


Urbanus proteus Linnaeus, 148 
Wallgrenia otho otho, 148 
New species, 44, 78, 102, 141, 222, 224, 
245, 247, 249, 251. 253, 255, 269 
New subspecies 
Callithomia (Callithomia) hezia perija, 
80 
Callophrys (Incisalia) polios obscurus, 
Ine 
Colias alexandra columbiensis, 68 
Ouleus dilla baru, 243 
Noctuidae, 9, 95, 103, 119, 175 
Notes and news, 86, 241, 278, 311 
Notodontidae, 95, 103 
Nymphalidae, 15, 22, 37, 85, 96, 122, 
130, 152, 159, 169, 210, 235, 274, 
291, 294, 305 
Obituary, 87 
Oecophoridae, 99 
Olethreutidae, 268 
Orchid attractant, 192 
Papilionidae, 16, 36, 95, 136, 155, 167, 
196, 206, 237 
Pieridae, 36, 57, 148, 154, 167 
Polymorphism, 
Anaea (Zaretis ) itys, 296 
Catocala spp., 176 
Danaus chrysippus, 73 
Population distributions, 
Callophrys (Incisalia) polios, 112 
Colias alexandra, 57 
Papilio andraemon bonhotei, 136 
Plebejus saepiolus, 17 
Predation 
by ants, 196 
by arachnids, 83 
by birds, 16, 175 
by midges, 143 
by heteroptera, 305 
on catocala, 175 
Pupal color, 155 
Pyralidae, 95, 103, 160, 219, 290 
Riodinidae, 149, 240, 304 
Saturniidae, 225 
Satyridae, 37, 95, 153, 168, 238, 291 
Sex ratios, 19 
Sphingidae, 1, 83, 103, 303 
Taxonomy and systematics 
Amblyscirtes, 40 
Colias alexandra, 68 
Eumarozia elaeanthes, 271 
Eumarozia malchitana, 268 
Glenoides texanaria, 140 
Heliconius, 130 


318 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Hyalophora columbia, 225 Techniques 

Itame varadaria, 288 background selection, 9 
Macaria inapta, 288 sexing of larvae and pupae, 122 
Metamorpha, 235 Thyridae, 95 

Ornithoptera priamus, 196 Uraniidae, 95 


Siproeta epaphus gadoui, 235 U.V. reflectance, 57 


EDITORIAL COMMITTEE OF THE JOURNAL 


Editor: THEODORE D. SARGENT, Department of Zoology, 
University of Massachusetts, Amherst, Massachusetts 01002 


K. S. Brown, J. M. Burns, R. H. Carcasson, J. P. DONAHUE, 
J. F. Gates Care, C. D. Ferris, R. O. KENDALL, J. H. MASTERS, 
L. D. Mrtter, A. P. Piatt, J. R. G. Turner 


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, 844 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, 1961la, 1961b) and all must be listed alphabetically 
under the heading LirErRATuRE 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 444 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 article'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 FIGURES, 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 approx- 
imate 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. 

Page Charges: Authors with grant or institutional funds are requested to pay a 
charge of $24.00 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 
case of authors who have no grant or institutional funding, as it is not intended that 
any author should pay this charge from personal funds. However, all authors will 
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. erika LAWRENCE, KANSAS 
usgn~ 


CONTENTS 


Descriptions oF NEw NrEotropicaL HesperupaE. S. S. Nicolay 


Tue Lire CycLe oF DIRCENNA RELATA (ITHOMIDAE) IN Costa RICA. 
Allen M. Young). 0 a 


THE Genus EuMAROZzIA HEINRICH (OLETHREUTIDAE). J. F. Gates 
Claghie ois NN dT EN 


Turee NATURAL Hysprips OF VANESSA ATALANTA RUBRIA X CYNTHIA 
ANNABELLA (NYMPHALIDAE). Thomas E. Dimock __ 


Lire History OF CALLOPHRYS S. SHERIDANII (LYCAENIDAE) AND 
Notes ON OTHER Species. Clifford D. Ferris 


Down-VALLEY FLIGHT oF ADULT THECLINI (LYCAENIDAE) IN SEARCH 
oF NouRISHMENT. James A. Scott. 


THe IDENTITY OF MACARIA INAPTATA WALKER AND ITAME VARADARIA 
(WALKER) (GEOMETRIDAE). Douglas C. Ferguson ___ 


TRANSFER OF CYMORIZA ABROTALIS WALKER, 1859, FROM NYMPHULA 
SCHRANK TO DISMILILA DyarR (PyYRALIDAE: NYMPHULINAE, 
MipiuinaE), Eugene’ Munroe) 


UNusuAL CopuULATORY BEHAVIOR IN THE NYMPHALIDAE AND 
SATYRIDAE. Edwin M. Perkins, Jr. 2 


NOTES ON THE LirE CycLE AND NatTuraL History oF BUTTERFLIES 
oF Ex Santvapor. II]. ANAEA (ZARETIS) ITYS (NYMPHALIDAE). 
Alberto Muyshondp) 28) OO 


GENERAL NOTES 


Some observations on Dryas iulia iulia (Heliconiidae). Alberto 
Muyshond$ i 0028) Mee LN IN A OOO 


New Mexican Sphingidae records. Carlos R. Beutelspacher B. __-- 


Apodemia. mormo near dialeuca (Riodinidae) from montaine southern 
Califo¢nia: /new for U.S.A.) Ray E. Stanford 


Boloria selene (Nymphalidae) ambushed by a true bug (Heteroptera). 
Robert Miglseel Pyle) 0 ee 


Book ‘REVIEWS oi Bee a EN See ioe es cate Use) ON 


NOTES AND! NEWS (00) UCONN AU) el A a 278, 


First Karl Jordan medal awarded to Henri Stempffer. Lee D. Miller _._ 


243 


279 


283 


288 


290 


291 


294 


311 


x ~ ; | 
| 
| 


Mi 


ws 


‘ Ka. 
eee ig: 
ET eae 
ELENA) 
Py 
in 
F 


/ 


eet 


— 


ee ee 


“USK Hey? AAD aan 4 7 Lh i: , i rift : psfrntieah 9§ oe 


9 
v 5) ‘ 
A AT ‘ WMS ee : aye A fs ‘ ‘ 7 


a 


it be Sc 
BETS CB ECR 0 ay warded \ 
‘ teh Hi i Di an Ope: ae 
ite tet 4 9 nile i Ni 


' We a) 
ee) 

oh, wee 
Wed he 


1 
Bigg? 
nae 


aii 
Un 
a) 
ep bit 
at Whe uth 
Digit eft 
Heid na he 


fe Neem 
PD ae 
Fi Ae 
fh ey beh ibibe 7 
hope 
i fe hay 
Wi Reed 


H Ay 
TGA eRe Dah 
: ae Mb iepmd & biped : : Be 
FuGan at Ch Wedel : th) Mormece H i du vacwete ht ie AM ti ieee 
A ( 1h the if a) " th, ( ihe y? f ce Wh 
} p, ‘ r ‘ Mere Ms, 


4 
Bap ay ee ON CU Hc ntibutitar any" 
f pig i j ‘ iis aH HO tite 


1 ca a ” 
u Woe Wet Wate sth 
Ue 
Wey HN Reg 


Bee yy H 
» “he % "WAM Eby 
SY 0 eee PS Hyde