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Full text of "Journal of the Lepidopterists' Society"

5^1 



bi 



9 



Volume 60 Number 3 
2 October 2006 
ISSN 0024-0966 



Journal of the 

Lepidopterists' Society 




Published quarterly by The Lepidopterists' Society 



THE LEPIDOPTERISTS' SOCIETY 

Executive Council 

William E. Conner, President Thomas J. Simonsen, Vice President 

Felix A.H. Sperling, Immediate Past President Ernest H. Williams, Secretary 

William E. Miller, Vice President Kelly M. Richers, Treasurer 
Yen Shen-Horn, Vice President 

Members at large: 

Robert M. Pyle Richard A. Anderson Michelle DaCosta 

John A. Shuey John V. Calhoun John H. Masters 

Andrew D. Warren Amanda Roe Michael G. Pogue 

Editorial Board 

John W. Brown (Chair, at large) 
Michael E. Toliver {journal) 

Brian Scholtens (Journal) 

Lawrence F. Gall (Memoirs) 

Dale Clark (Neivs) 

John A. Snyder (Website) 

Honorary Life Members of the Society 

Charles L. Remington (1966), E. G. Munhoe (1973), Ian F B. Common (1987), 
Lincoln P. Brower (1990), Frederick H. Rindge (1997), Ronald W Hodges (2004) 

The object of The Lepidopterists' Society, which was formed in May 1947 and formally constituted in December 1950, is "to pro- 
mote 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 mea- 
sures" 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. Prospective members should send to the Assistant Treasurer full dues for the current year, to- 
gether with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is is- 
sued, with addresses and special interests. 

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Life members — single sum $1,800.00 
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Send remittances, payable to The Lepidopterists' Societij, to: Kelly M. Richers, Treasurer, 9417 Carvalho Court, Bakersfield, CA 
93311; and address changes to: Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. 
For information about the Society, contact: Ernest H. Williams, Department of Biology, Hamilton College, Clinton, NY 13323. To 
order back issues of the Memoirs, write for availability and prices to Kenneth R. Bliss, 28 DuPont Avenue, Piscataway, NJ 08854. 



The additional cost for members outside the U.S. is to cover mailing costs. 



Journal of The Lepidopterists' Societij (ISSN 0024-0966) is published quarterly by The Lepidopterists' Society, % Los Angeles 
County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA 
and at additional mailing offices. POSTMASTER: Send address changes to The Lepidopterists' Society, % Natural History Museum, 
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Cover illustration: Photograph of Euribia lycisca (Westwood) Blue-winged Sheenmark. Photographed on March 9, 2006 at Bosque 
del Cabo Lodge on Cabo Matapalo, the Osa Peninsula, in Costa Rica. Photo Credit: Hank and Priscilla Brodkin, Hereford Arizona, 
email: hbrodkin@cox.net 



Volume 60, Number 3 



121 



Journal of the Lepidopterists' Society 
60(3), 2006, 121-137 



FIVE NEW SPECIES OF PAUCIVENA DAVIS, 1975 (LEPIDOPTERA: TINEOIDEA 

PSYCHIDAE) FROM CUBA 

Rayner Nunez Aguila 

Division de Colecciones Zoologicas y Sistematica, Instituto de Ecologia y Sistematica, Carretera deVarona 
Capdevila, Boyeros, C. de La Habana, Cuba. AP 8029. CP 10800, Cuba. 
E-mail: rayner@ecologia.cu 




ABSTRACT. Five new species of Paucivena Davis (Lepidotera: Tineoidea: Psychidae), P. ferruginea , P. pinarensis, P.fusca, P. cubana and P. 
orientalis are described from Cuba and compared with relatives. The females of P. ferruginea and P. orientalis are described being the first fe- 
males known widiin genus; their characteristics confirm the intermediate position of the genus among the American psychids. Notes on natural 
history of new species (e.g. hosts, habitat) are given as available. Keys for identification of known stages of all Paucivena species are provided. 

Additional key words: Tineoidea, Psychidae, bagworm, natural history, West Indies. 



The Neotropical region has the richest Lepicloptera 
diversity but one of the least studied psychid fauna widi 
only 61 known species (Heppner, 1998; Davis, 2000). 
This species number is extremely low compared with 
that of other faunal regions. The Paleartic, for example, 
has more than 300 described species (Heppner, 1998). 
Members of this family were last reviewed by Davis 
(1975) who described two genera and five species from 
die West Indies. However, none of these taxa was 
known from Cuba. Recent collections from Cuba, 
chiefly on the three main mountain chains, have 
resulted in the discovery of new records including 
several new species. In this work, five new species 
belonging to Paucivena Davis, 1975 are described. Keys 
for identification of known stages of all species and 
information on natural history are also provided. 

Materials and Methods 

Individuals of all species, except one whose 
representatives were taken flying at day, were collected 
as larvae and pupae in the field and reared in the 
laboratory. Lab-reared larvae were provided with field 
collected hosts until pupation. All type material is 
deposited at Instituto de Ecologia y Sistematica 
(CZACC). 

Diagnostic morphological characters employed 
follows Davis (1964, 1975) and Henderickx (1982). Setal 
maps of larvae follow Hinton (1946) and Stehr (1987). 
Measurements were taken with an ocular micrometer in 
a Carl Zeiss Stemi 2000 stereoscopic microscope. 
Interocular index of head was calculated as a ratio 
between the vertical diameter of the compound eye and 
interocular distance measured at a point across die frons 
midway between the base of antennal sockets and the 
anterior tentorial pits (Davis, 1975): 

Interocular index= vertical eye diameter/ interocular 
distance 

Characters of the two previously known species, 



Paucivena reticulata Davis, 1975 and Paucivena 
hispaniolae Davis, 1975, were taken from original 
descriptions and illustrations. Additionally, one P. 
reticulata specimen placed at CZACC, was examined. 

Abbreviations: x- mean, SD- standard de\iation, CV- 
coefficient of variation. 

Results 
Paucivena Davis, 1975 

This genus is known only from die western part of the 
Antilles. Davis (1975) described P. hispaniolae from 
Dominican Republic, Hispaniola, and P. reticulata from 
Puerto Rico and Jamaica. In die same work, tiiis author 
mentioned the possible presence of Paucivena on Cuba. 
This was confirmed recendy by Nunez (2004) based on 
unknown species from Topes de Collantes, in die 
Cuban central mountains, which are described here. 

Characters diat best define Paucivena males are labial 
palpi with a single segment not fused, origin of antennal 
rami at base of each antennal segment, two pairs of 
tibial spurs on mid and hindlegs, reduced wing venation 
and the abbreviated genitalia. Females mav be 
recognized by the possession of compound eves and 
functional legs, and the lack of antennae and wings. 
Larvae feed on several hosts including mosses and 
lichens growing on rocks and bark, and detritus. 

Paucivena ferruginea Nunez, new species 

(Figs. 1, 6, 11, 16, 17, 22, 24, 26. 2S-35. 41. 45-16, 52^ 

Diagnosis: Paucivena ferruginea male differs from 
all other Paucivena by its brown coloration with slight 
ferruginous iridescence. Other diagnostics characters 
are the acute and heavily sclerotized sacculus and die 
bifid saccus of its genitalia. 

Male (Figs. 1, 6, 11, 16, 17). Head: brown. Antennae with 23 
segments; lateral pectinations about 2-2.5 times length of supporting 
segment. Vertical diameter of eve 0.8 the interocular distance. Thorax 
(Figs. 6, 11): anterior half dark brown, posterior half brown with slight 
ferruginous iridescence; underside pale brown except inner surface of 



122 



Journal of the Lepidoptehists' Society 






* 




: 




Figs. 1— .5. Paucivena spp. adult males. 1 P. ferruginea, n. sp.; 2 P. pinarensis, n 
orientalis, n. sp. Scale= 3 mm. 



sp.; 3 P. fits 



sp.; 4 P. cubana, n. sp.; 5 P. 



coxa and femur of forelegs which are dark brown. Vestiture dense, 
scales hairlike. Wings brown widi slight ferruginous iridescence; basal 
two thirds of costa on FW dark brown. Tibial spurs approximately 0.35 
the length of basal tarsal segments (Fig. 6). Scales at discal cell 
variable in shape: oblanceolated, ovobated, with rounded or acute 
apices, or hairlike. FW with 9 veins, all veins separated (Fig. 11); 
accessory cell present; CuP not reaching inner margin; one anal vein. 
HW with 7 veins, all veins separated except M and CuA t which have 
a unique origin; cross vein between Sc and Rs absent; two anal veins. 
Wing expanse: 11 mm. Abdomen: brown with slight ferruginous 
iridescence; underside pale brown. Vestiture dense, scales hairlike. 
Genitalia (Figs. 16-17): tegumen broad, widi a pair of sparsely setose 
apical lobes. Valvae with pulvilli setose; apex of sacculus acute and 
heavily sclerotized, armed with three spines; cucullus apically rounded 
and sparsely setose. Saccus bifid, apices blunt; approximately 0.1 the 
length of main body. Aedeagus simple, cylindrical, 0.7 times the 
length of valvae. 

Female (Figs. 22, 24, 26). Lengdi: 7.5 mm. Vermiform. 
Stramineous, with two longitudinal bands of brown spots on dorsum. 
Head (Fig. 22): stramineous, eyes black. Slightly sclerotized. Shape 
near ovoid (ventral view); eyes compound, well developed, subventral. 
Labial palpi 1-segmented, 100% fused; antennae absent. Thorax (Fig. 
24): patterned as above; body wall slightly sclerotized. Legs functional, 
armed widi numerous tiny spines; tarsi 1-segmented widi a pair of 
claws at distal end (Fig. 24); wings absent. Abdomen: color pattern 
disappearing at A2-A3; membranous and naked except for a ring of 
dense brownish ochre hairlike scales around A7. External genitalia 
reduced (Fig. 26), largely membranous. Two pairs of apophyses 
present; anterior pair elongated, free except bifid base fused with 
tegument; posterior pair straight and free. 

Larva (Figs. 28-34). Length of largest larva 11 mm, maximum 
width of head capsule 1.2 mm. Head and diorax whitish with dark 



fuscous longitudinal bands continued on thorax forming a striated 
pattern; spiracle on TI as large as spiracle on AS, both larger than 
spiracles on A1-A7. Head (Figs. 28-32): patterned as above, lateral 
area with five elongated bands; labrum ochre; an elongated band on 
adfrontal sclerite and frons extending from CI to slightly beyond AF1; 
AF2 and P2 absent (Fig. 28). Six stemmata present; five arranged in 
an inverted semicircle, a sixdi more distant and ventrad, immediately 
anterior to S3 (Fig. 29). Labrum (Figs. 30, 31) with LAS isolated. 
Mandibles with four acute teeth and a fifth, blunt tooth (Fig. 32). 
Thorax (Fig. 33): patterned as above, three longitudinal bands 
between body axis and lateral margin of shield, interrupted on 
metathorax. TI with shield bearing D, SD, XD and L groups; XD- 
group and L2 in vertical line near anterior margin of shield, SD-group 
slightly posterior; XD1 about equal in length to SD1, about 2 times 
longer dian XD2 and D2; Dl dorsoposterior to XD1, about 1/4 its 
length; SD2 above SD1, slightly posterior and about 1/4 its length; L- 
group trisetose, LI about 3 times longer than L2 and L3, 
posteroventral to L2; L3 slightly longer than L2; spiracle 
dorsoposterior to L-group, diagonal; SV-group in horizontal line on 
elongated pinnaculum, SV2 about 3/5 length of SV1; MV2 on same 
pinnaculum, anterior to SV2; VI about equal in length to SV2, 
posteroventral to SV1. TII-TIII: D and SD groups in a vertical line on 
same pinnaculum; D2 about 2-21/2 times longer than Dl; SD1 about 
3 times longer than SD2; L2 separated from pinnaculum bearing LI 
and L3, about 1/2 length of L3; LI 3/5 length of L2, dorsoposterior to 
L3; SV group on same pinnaculum; SV1 about twice lengdi of SV2; VI 
slightly shorter dian SV1. Abdomen (Figs. 33, 34): integument dark- 
brown, pinnacula brownish ochre. Al: D-group on separated 
pinnacula, Dl dorsoposterior to D2 and about 3 1/2 dmes longer; SD1 
above and slightly anterior to spiracle, slightly shorter than D2; SD2 
minute, anterodorsal to spiracle; L-group trisetose, on separated 
pinnacula; LI posterior to L2 and about twice its length; L3 below, 



Volume 60, Number 3 



123 



midway between LI and L2, equal in lengtir to LI; SV-group bisetose 
and on same pinnaculum, SV2 anterodorsal to SV1 and about 1/3 its 
length; VI anteroventral to SV1 and about half its length (Fig. 33). A2 
(not shown) equal to Al except SV- group trisetose, SV3 on 
pinnaculum bearing VI, below SV2 and about equal its length. A3-A6 
with four pairs of prolegs, crochets (22—24) uniordinal, uniserial, 
arranged in a lateral penellipse; setae as above except SV-group on 
pinnaculum containing proleg. A7 (not shown) as above except SV- 
group bisetose. AS with setae as above except L-group arranged in a 
more or less vertical line, LI on same pinnaculum bearing SD-group 
and spiracle; SV-group unisetose. A9 with all setae arranged in a more 
or less vertical line; SD1 and D2 on same pinnaculum, SD1 about 5 
times longer than D2 and about equal in length to Dl. A10 (Fig. 34): 
anal plate with SD1 slightly longer dian Dl, about 2-2 1/2 times 
longer than D2; prolegs bearing 24 uniordinal crochets, uniserial, 
arranged in a lateral penellipse; anterior margin of shield irregular. 

Larval case (Fig. 41). Dimensions: length of main body: 6 12 
mm (x=12, SD=0, CV=0, n=2), total lengdi of projecting fragments: 
17-18 mm (X= 17.5, SD=0.71, CV=4%, n=2); 9 13 mm, total length 
of projecting fragments: 22 mm; maximum diameter: 6* 2.6 mm 
(x=2.6, SD=0, CV=0, n=2), 9 3.3 mm. Almost cylindrical in its entire 
length; soft. External cover formed by a basal layer of tiny vegetal 
fragments covered by large fragments of thin herbaceous stems, 
lengthwise arranged and parallel, various projecting backward from 
case. 

Male pupa (Figs. 45-46). Length 5.8 mm. Uniform yellowish 
ochre. Frontal ridge absent, frons rounded. Antennal sclerites 
extending slighdy beyond apex of prothoracic legs (Fig. 45). Wing 



sheaths extending midway along A3. Sclerites of metathoracic legs 
extending to anterior margin of A4. Cremaster reduced, consisting in 
a pair of small spines, ventrally curved, arising form a broad conical 
base; anal groove Y-shaped. Dorsum of A3— A7 with 2—3 irregular rows 
of spines directed caudad on anterior margin, both end of rows 
thickened (Fig. 46); AS with spines grouped in an elliptical patch: 
areas surrounding rows covered by hundreds of tiny spines arranged 
in 2-5 series or solitaire. Dorsum of A5-A7 with single posterior row 
of slender spines oriented caudad. Tabulation of spines shown in Table 
1. 

Female pupa (Figs. 50^51). Length 7.5 mm. Uniform yellowish 
ochre. Head with eyes and labial palpi distinct (Fig. 50). Thorax with 
leg sclerites distinct; wings absent. Cremaster vestigial, reduced to a 
coarse and rough area around anal groove; anal groove Y-shaped. 
Dorsum of A6-A7 with 2—i irregular rows of small spines directed 
caudad on anterior margin, AS with spines grouped in an elliptical 
patch. A4— A6 with a single posterior row of slender spines cephalad 
oriented (Fig. 51). Tabulation of spines shown in Table 2. 

Types. Holotype 6 (with associated larval case and pupal exuvium), 
reared from larva (emerged 22 June 2003), CUBA: Sancti Spiritus 
province, Topes de Collantes, Pico Potrerillo, 973 m, 18 Mav2003 (R. 
Niinez), slides RNA 014, 038, 046, 077, 078. Paratypes, 9 (with 
associated larval case and pupal exuvium), reared from pupa (emerged 
20 May 2003), CUBA: Sancti Spiritus province. Topes de Collantes, 
Parque Codina, 800 m, (R. Nunez), slide RNA 042; 2 larvae (with 
associated larval case), same data as holotype; 3 larval cases, same data 
as holotype; 1 larval case, same data as holotype except 17 June 2004. 




FIGS. 6—10. Paucivena spp. male legs. 6 P. ferruginea , n. sp.; 7 P. pinarensxs, n. sp.; 8 P. fusea, n. sp.; 9 P. cubana, n. sp.; 10 P. ori- 
entals, n. sp. Top- prothoracic leg, middle- mesothoracic leg, bottom-metathoracic leg. Scale= 2 mm. 



124 



Journal of the Lepidopterists' Society 



Natural history observations. Larvae were found 
feeding on mosses, Orthostichidium guyanense (Mont.) 
V.F. Brotherus (Pterobryaceae) and another 
unidentified species, on bark of an unidentified bush. 

Two adults were reared from larvae: a male emerged 
from the pupa after a month; a female emerged and was 
observed hanging from distal end of the case. 

The species was found in two localities with veiy 
different vegetation and climatic conditions. Parque 
Codina is characterized by a secondary and very humid 
evergreen forest with the lower strata protected by a 
dense canopy. A diy scrub, included in the mogote 
vegetational complex, grows on top of Pico Potrerillo, an 
enviroment very exposed to wind and solar radiation. 

Distribution (Fig. 52). Known only from two 
localities at Topes de Collantes region, Trinidad 
Mountains, central Cuba. 

Etymology. The species name is derived from the 
slight ferruginous iridiscence of male wings. 

Remarks. The female may be separated from that of 
P. orientalis, n. sp., by its greater size (length 7.5 versus 
5 mm), its more elongated genitalia, the presence of 



only two longitudinal spot bands on tegument (six in the 
other species) and its unswollen tibiae. Larvae may be 
recognized by the loss of AF2 on head and the isolation 
of LA3 on lab rum. The larval case is also diagnostic for 
this species within the genus. Davis (1964, 1975) 
reported similar cases from Haiti and Trinidad. Davis 
noted the similarity of these cases with diose 
constructed by species of Epichnopteryx Hiibner and 
Psyche Schrank, two Old World genera, considering 
them as a probable introduction from the Old World. 
Larval cases of P. ferniginea prove that this construction 
pattern is not exclusive to Old World species. 

Paucivena pinarensis Nunez, new species 

(Figs. 2, 7, 12, 18, 42, 47, 52) 

Diagnosis: Paucivena pinarensis males possess a 
color pattern similar to that of P. cubana, sp. n., P. 
orientalis, sp. n., and P. reticulata. However, the 
reticulated pattern is weaker in P. pinarensis due to its 
more obscure background color. Compared to other 
Paucivena within this group, P. pinarensis males exhibit 
several diagnostic characters: absence of saccus in its 





12 





FIGS. 11—15. Paucivena spp. wing venation. 11 P. ferniginea, n. sp.; 12 P. pinarensis, n. sp.; 13 P.fusca, n. sp.; 14 P. cubana, n. sp.; 
15 P. orientalis, n. sp. Scale= 3 mm. 



Volume 60, Number 3 



125 



Table 1. Rows and spines numbers per rows on dorsum of abdominal segments of Cuban Paucivena male pupae. 





Rows 








Abdominal 


segments 










I 


II 


HI 


IV 


V 


VI 


VII 


VIII 


Paucivena ferruginea 


anterior 








269 


135 


140 


176 


122 


48 


n=l 






















posterior 














42 


44 


48 





Paucivena pinarensis 


anterior 








57 


63 


64 


60 


49 


49 


n=l 






















posterior 








1 


7 


38 


38 


14 





Paucivena cubana 


anterior 











52-70 


47-58 


42-55 


29^5 


22-24 


n=3 






















posterior 








31 


35-44 


37-^16 


36-42 


32-36 





Paucivena orientalis 


anterior 











102-136 


96-125 


95-111 


90-96 


25-32 


n=3 






















posterior 











27-38 


23-39 


24-32 


23-50 






Table 2. Rows and spines numbers per rows on dorsum of abdominal segments of Cuban Paucivena female pupae. 





Rows 








Abdominal 


segments 










I 


II 


III 


IV 


V 


VI 


MI 


Mil 


Paucivena ferruginea 


anterior 

















61 


90 


21 


n=l 






















posterior 











7 


58 


53 








Paucivena cubana 


anterior 











3 


14 


20 


21 





n=l 






















posterior 








6 


56 


78 


71 








Paucivena orientalis 


anterior 











8-10 


9-10 


29-30 


92-96 


1S-22 


n=3 






















posterior 











16-17 


23-27 


21-22 









126 



Journal of the Lepidoptemsts' Societv 



tegumen 



valva 1 




aedeagus 




17 







Figs. 16—21. Paucivena spp. male genitalia, ventral view. 16 P. ferruginea, n. sp., main body; 17 P. ferruginea, n. sp., aedeagus; 18 
P. pinarensis, n. sp.; 19 P.fusca, n. sp.; 20 P. cubana, n. sp.; 21 P. orientalis, n. sp. Scale= 0.5 mm. 





Figs. 22-23. Paucivena spp. female head, ventral view. 22 P ferruginea, n. sp.; 23 P orientalis, n. sp. Scale= 1 mm. 



Volume 60, Number 3 



127 




24 




25 



Figs. 24-25. Paucivena spp. female legs. 24 P. ferruginea , n. sp., scale= 0.5mm; 25- P. orientalis, n. sp., scale= 0.25 mm. Top- 
prothoracie leg, middle- mesodioracic leg, bottom-metathoracic leg. 



posterior 
apophyses 





27 



Figs. 26-27. Paucivena spp. female genitalia, ventral view. 26 P. ferruginea, n. sp.; 27 P. orientalis, n. sp. Scale= 0.5 mm. 



128 



Journal of the Lepidopterists" Society 





Figs. 28-34. Chaetotaxy ofPaucivenaferruginea, n. sp., largest larva (last instar?). 28 Head, dorsal view; 29 Stemmata of left side, 
lateral view; 30 Labmm, dorsal view; 31 Labrum, ventral view; 32 Right mandible, ventral view; 33 Prothorax, mesothorax and ab- 
dominal segments 1, 3-6, 8 and 9, lateral view; 34 Anal shield, dorsal view. Scale, 28-29= 0.5 mm; 30-32= 0.2 mm. 



Volume 60, Number 3 



129 








40 



Figs. 35^0. Chaetotaxy of Paucivena cubana, n. sp., largest larva (last instar?). 35 Head, dorsal view; 36 Stemmata of left side, 
lateral view (scale= 0.5 mm); 37 Labrum, dorsal view; 38 Labrum, ventral view; 39 Right mandible, ventral view (scale= 0.2 mm); 
40 Prothorax and mesothorax, lateral view (abdomen damaged). Scale. 35-36= 0.5 mm: 37-39= 0.2 mm. 



genitalia; the relative length of its tibial spurs, 0.3 versus 
0.15 (P. cubana, P. orientalis) and 0.5 (P. reticulata); and 
interocular index 0.7 versus 1.1 (P. reticulata) and 1.5 (P. 
cubana). 

Male (Fig. 2, 7, 12, 18).: Head: pale brown, labial palpi dark- 
brown. Antennal tips broken; lateral pectinations 3 about times length 
of supporting segment. Vertical diameter of eye 0.7 the interocular 
distance. Thorax (Figs. 7, 12): dark brown; underside pale brown, 
inner surface of fore and midlegs dark brown, joints pale brown. 
Vestiture dense, scales hairlike. Tibial spurs approximately 0.3 the 
length of basal tarsal segments (Fig. 7). FW with basal two thirds of 
costa dark brown; ground brown streaked with dark brown forming a 
faint reticulated pattern; fringe with various tones of brown. Scales at 
discal cell variable in shape: oblanceolated and ovobated, with 
rounded or acute apices, with scattered hairlike scales. Venation (Fig. 
12) as in P. ferruginea. HW uniform brown; fringe with various tones 
of brown. Venation as in P ferruginea except M 2 . and CuA, which 
arise separate from cell, M, <3 equidistant from Cu/ 1 ! and M only one 
anal vein present. Wing expanse: 9.2 mm. Abdomen: pale brown. 
Vestiture dense, scales hairlike. Genitalia (Fig. 18): tegumen broad, 
with a pair of sparsely setose apical lobes. Valvae with pulvilli setose; 
apex of sacculus armed with five strong spines; cucullus apically 
rounded and sparsely setose. Saccus absent. Aedeagus simple, 
cylindrical, 0.6 times length of valvae. 



Female. Unknown. 

Larva. Unknown. 

Larval case (Fig. 42). Dimensions: length: e 9.8 mm; maximum 
diameter: 6 3.4 mm. Fusiform in outiine; soft. Exterior heavilv 
covered with vegetal fragments, leaves and short stems, and mosses 
various shaped and oriented lengthwise. Cases were found hanging 
from silk filaments (3.5 mm in lengdi) attached to rocks and tree 
trunks. 

Male pupa (Fig. 47). Length 5.1 mm. Ochre, wing sheaths reddish 
brown. Frontal ridge absent, frons rounded. Antennal sclerites 
extending slightly beyond apex of protiroracic legs. Wing sheaths 
extending to posterior margin of A3. Sclerites of metathoracic legs 
extending to posterior margin of A4. Cremaster consisting in a pah - of 
strong spines ventrally curved; anal groove Y-shaped. Dorsum of 
A3-A8 with an irregular row of spines on anterior margin; areas 
surrounding rows covered by hundreds of tiny, solitaire spines. 
Dorsum of A3-A7 widi single posterior row of slender spines. 
Tabulation of spines shown in Table 1. 

Female pupa Unknown. 

Types. Holoti/pe, 6 (with associated larval case and pupal 
exuvium), reared from larvae (emerged 28 February 2004\ CUBA: 
Pinar del Rio province, Sierra del Rosario, Trco Taco River shore 1 km 
northeast from Jardm de Aspiro, 200 m, 28 November 2003 (R. 
Nunez), slides RNA 054, 079, 080. Paratypes: 4 larval cases. CUBA. 
Pinar del Rio province, Sierra del Rosario, Jardm de Aspiro, 150 m. 



130 Journal of the Lepidopterists' Society 



2S-.29 November 2003, (R. Nunez). localities at Sierra del Rosario, Pinar del Rio province. 

Natural history observations. The single larva was Etymology. The species name is derived from the 

found on limestone rock near die Taco Taco River shore name of the Cuban province where the type locality, 

and its food source can not be accurately identified. In Pinar del Rio, is located. 

the lab, the larva was fed with several crustose lichens Remarks. The larval case is identical to that of P. 

and mosses collected on its substrate. Other larval cases cubana. The male pupa may be easily identified by the 

were found on rocks and tree trunks in an old unique arrangement of spine rows on the dorsum of 

abandoned Botanical Garden (Jardin de Aspiro). The abdominal segments (Table 1). 
unique adult emerged after a month; emergence took 
place between 0900 and 1 130h. Paucivena fusca Nunez, new species 

At the Taco Taco River shore secondary remnants of Paucivena sp. n. 1: Nunez, 2004: 155 
gallery forest are present whereas at Jardin de Aspiro '" s ' ' ' ' ' 
several introduced and native trees grow forming groves Diagnosis. Males of P. fusca may be recognized by- 
separated by cleared areas occupied by camping their uniform dark brown coloration. Within the genus 
installations. only P. hispaniolae exhibits a similar coloration but it has 

Distribution (Fig. 52). Known only from two close 

Key to the adult males of Paucivena 

1. Wings dark brown with a ferruginous shine; genitalia with an acute and heavily sclerotized sacculus 
and a bifid saccus (Fig. 16) P. ferniginea 

- Wings with a different color pattern; genitalia with sacculus not acute and weakly sclerotized, saccus 
not bifid 2 

2. Dorsum of wings and body entirely dark brown, almost black 3 

- Dorsum of wings and body with light color pattern 4 

3. Body whitish grey ventrally; eyes of medium size (vertical diameter of eye 1.1 the interocular 
distance); genitalia with the margins of apex of sacculus and the apical lobes of tegumen smooth 

P. hispaniolae 

- Body dark brown ventrally; eyes small (vertical diameter of eye 0.8 the interocular distance); 
genitalia with the margins of apex of sacculus and the apical lobes of tegumen serrulated (Fig. 19) 

P. fusca 

4. Tibial spurs much reduced, approximately 0.15 the length of basal tarsal segment 5 

- Tibial spurs less reduced, approximately 0.3 the length of basal tarsal segment or longer 6 

5. Eyes very large (vertical diameter of eye 1.5 the interocular distance); wing expanse: 12 mm 

P. cubana 

- Eyes very small (vertical diameter of eye 0.7 the interocular distance); wing expanse: 8.2-9.1 mm 

P. orientalis 

6. Eyes of medium size (vertical diameter of eye 1.1 the interocular distance); tibial spurs large, 
approximately 0.5 the length of basal tarsal segment; FW reticulated pattern distinct P. reticulata 

- Eyes veiy small (vetical diameter of eye 0.7 the interocular distance); tibial spurs reduced, 
approximately 0.3 the length of basal tarsal segment; FW reticulated pattern weak, indistinct 

P. pinarensis 

Key to the known larvae of Paucivena (excludes P. reticulata, P. pinarensis, P. fusca and P. orientalis, 

which are unknown) 

1. Head and thorax whitish to light tan with irregular patches of dark fuscous; meso and metathorax 
with an extra seta (SDla?) P hispaniolae 

- Head and diorax whitish with longitudinal dark fuscous bands arranged in a striated pattern; meso 
and metathorax without an extra seta 2 

2. Head with AF2 absent (Fig. 28); LA3 on labrum isolated from the rest (Fig. 30); abdominal 
integument dark brown P ferniginea 

- Head with AF2 present (Fig. 35); LA3 on labrum not isolated (Fig. 37); abdominal integument dirty 
white P. cubana 



Volume 60, Number 3 131 



Kev to the known male pupae of Paucivena (excludes P. reticulata and P. fusca, which are unknown) 

1. Anterior margin of A3 without spines 2 

- Anterior margin of A3 with at least one row of spines 3 

2. Spines absent from posterior margin of A3; rows at anterior margin of A4-A7 with 90 or more spines 
(Table 1); length 3.8-4.8 mm P. orientalis 

- Spines present on posterior margin of A3; rows at anterior margin of A4-A7 with 70 or fewer spines 
(Table 1); length 5.0-5.4 mm P. cuhana 

3. Rows on anterior margin of A3-A7 with more than 100 spines (Table 1); length 5.8 mm 

P. ferruginea 

- Rows on anterior margin of A3-A7 with less than 100 spines (Table 1); length 5.0-5.1 mm 4 

4. Spines present, although reduced, on posterior margin of A3-A4; cremaster consisting in a pair of 
strong spines ventrally curved P. pinarensis 

- Spines completely absent from posterior margin of A3-A4; cremaster consisting in a pair of small 
spines ventrally curved P. hispaniolae 

Key to the female pupae of Paucivena (excludes P. reticulata, P. pinarensis and P. fusca, which are 

unknown) 

1 . Anterior rows of spines absent from dorsum of A3-A4 2 

- Anterior rows of spines present on dorsum of A3-A4 3 

2. Posterior row of spines reduced but present on dorsum of A4; cremaster vestigial, reduced to a 
coarse and rough area around anal groove; length 7.5 mm P. ferruginea 

- Posterior row of spines absent from dorsum of A4; cremaster relatively well developed, consisting in 
pair of short acute spines; length 10-11 mm P. hispaniolae 

3. Anterior margin of A8 with a row of spines surrounded by hundreds of tiny spines; length 4.9-5.1 
mm P. orientalis 

- Anterior margin of A8 covered only by hundreds of tiny spines; length 8.0 mm P. cuhana 

the underside of body whitish grey. Other useful a single specimen seen flying between rocky walls at 

characters are FW shape (more rounded in the Cuban peak access. This species shares its habitat, drv scrub on 

species), eye size (interocular index 0.8 in P. fusca and the top of Pico Potrerillo, with P. ferruginea and P. 

1.1 in P. hispaniolae) and the serrated margins at apex of cuhana. 

sacculus and apical lobes of tegumen in the genitalia of Distribution (Fig. 52). Known only from Pico 

P. fusca, both smooth in P. hispaniolae. Potrerillo at Trinidad Mountains, central Cuba. 

Male (Figs. 3 8, 13,19)- Head: dark brown. Antennae with 21-22 Etymology. The species name is derived from its 

segments; lateral pectinations 2 times length of segment. Vertical .,. , P , , 

diameter of eye 0.8 the interocular distance. Thorax (Figs. 8, 13): uniform dark brown color. 
uniform dark brown. Vestiture dense, scales hairlike. Tibial spurs 
approximately 0.2,5 the length of basal tarsal segments (Fig. 8). Scales 

at discal cell of FW oblanceolated and ovobated with rounded or Paucivena Cllbana Nunez, new species 

acute apices. Venation (Fig. 13) as in P. ferruginea. HW venation as in Paucivena SD n 2- Nunez 2004- 155 

P. pinarensis, except the origin of M, 3 which is closer to CuA, than to '• 

M r Wing expanse: 10-11 mm (x=10.2, SD=0.23, CV=4%, n=9). Figs. 4, 9, 14, 20, 35^0, 43, 48, 52 

Abdomen: dark brown. Vestiture dense, scales hairlike. Genitalia (Fig. . , 

19): tegumen broad with apical cleft, lobes minutely serrated and Diagnosis. Males of P. cuhana may be Separated 

sparsely setose. Valvae with pulvilli setose; apex of sacculus strongly from other Paucivena with reticulated wins; pattern bv 

serrated; cucullus rounded, apex sparsely setose Saccus reduced, ^ f H ow ing characters: 12 mm of wing expanse (the 
approximately 0.2 the length ol main body Aedeagus simple. ° . . . . ii 

cylindrical, 0.6 times the length of valvae. largest within the genus), elongated legs witii tiny tibial 

Female. Unknown. spurs (approximately 0.15 the length of basal tarsal 

Inmature stages. Unknown. segment) and large eyes (interocular index 1.5. the 

Larval case, unknown. ° \ 

Types. Holotype, 6 CUBA: Sancti Spiritus province, Topes de largest within the genus). 

Collantes, Pico Potrerillo, 973 m, 6 May 2002 (R. Nunez), slides RNA Male < Fi g s - 4 . 9 - 14 . 20 >- Head - P^e yellowish ochre. Antennae 

011, 015, 039. Paratopes, 3 6, same data as holotype, slides RNA 010, ™ t!l 18 segments; lateral pectinations 1.5-2 times the length of 

022. 5 6, same data as holotype except 17 June 2004, slides RNA 026, segment. Eyes large, vertical diameter of eye 1.5 the interocular 

029, 041. distance. Thorax (Fig. 9, 14): pale yellowish ochre with scattered dark 

xt a- lL-«. u M-i AH1--J1 brown scales. Vestiture dense, scales hairlike. Tibial spurs 

Natural history observations. A II individuals were m nici i j. a i, i .. n?- o\ \*f 

J approximately 0.15 the length of basal tarsal segments (Fig. 91. \\ ings 

found flying at noon on the top of Pico Potrerillo except thinly scaled. FW (faded) yellowish ochre with scattered dark brown 



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Journal of the Lepidopterists' Society 





42 



43 




44 



Figs. 41-44. Paucivena spp. larval cases. 41 P. ferruginea, n. sp.; 42 P. pinarensis, n. sp.; 43 P. cubana, n. sp.; 44 P orientalis, n. 
sp. Scale= 3 mm. 



scales; dark brown scales concentrated at basal half of anterior and 
posterior margins, forming a faint reticulated pattern on basal two 
thirds; fringe ochre. Scales at discal cell variable in shape: 
oblanceolated and ovobated, with rounded or acute apices, or hairlike. 
Venation as in P. ferruginea, except accessory cell which is wider, 
closing below origin of R (Fig. 14). HW pale yellowish ochre with 
scattered dark brown scales, paler than FW; fringe pale yellowish 
ochre. Venation as in P.fusca. Wing expanse: 12 mm. Abdomen: pale 
yellowish ochre with scattered dark brown scales. Vestiture dense, 
scales hairlike. Genitalia (Fig. 20): tegumen broad, apex damaged. 
Valvae with pulvilli sparsely setose; apex of sacculus armed with three 
spines; cucullus rounded with apex sparsely setose. Saccus reduced, 
approximately 0.2 the lengdi of main body. Aedeagus simple, 
cylindrical, 0.7 times the lengdi of valvae. 



Female. Unknown. 

Larva (Figs. 35^0). Length of longest larva 6.9 mm, maximum 
width of head capsule 1.0 mm. Head and thorax whitish widi dark 
fuscous longitudinal bands continued on thorax forming a striated 
pattern. Head (Figs. 35-39): as in P. ferruginea except, AF2 present; 
adfrontal sclerite with elongated spot on upper diird covering origin of 
AF2; frons with spot covering origin of Fl and C2 (Fig. 35); AFa 
closer to AF2 than to AF1. Sixth stemma immediately anterior to S2 
and S3, midway between them (Fig. 36). Labrum (Figs. 37, 38) with 
setae approximately mesad except, LAI and M2 distinctly closer to 
border. Mandibles with four acute teeth and a fifth, blunt tooth (Fig. 
39). Thorax (Fig. 40): as in P. ferruginea with the following exceptions. 
TI with XD1 about equal in length to XD2 and SD1, and about 1 1/2 
times longer than D2; MV2 separated from pinnaculum bearing SV- 



Volume 60, Number 3 



133 



group. TII-TIII: D2 about 3 times longer than Dl; SD1 about 4 times 
longer than SD2. Abdomen damaged, integument dirty white. 

Larval case (Fig. 43). Dimensions, length: 6 10-11 mm (x=10.5, 
SD=0.T1. CV=7%, n=2), 9 18 mm; maximum diameter: 6 4.0 mm 
(x=4.0, SD=0, CV=0, n=2), 9 6.5 mm. Fusiform in outline, soft. 
Exterior densely covered by fragments of leaves, small herbaceous 
stems and mosses of different shape and lengthwise oriented. Cases 
were found attached to rocky walls at Pico Potrerillo and banana trees, 
Musa paradisxaca L. (Musaceae), at Mogote Mi Retiro, hanging from 
silk threads (length: o 3.5 mm, 9 5 mm). 

Male pupa"(Fig. 48). Length 5.0-5.4 mm (x=5.2, SD=0.23, 
CV=4%, n=2). Uniform brownish ochre. Frontal ridge absent, frons 
rounded. Antennal sclerites extending slightly beyond apex of 
profhoracie legs. Wing sheaths extending to anterior margin of A4. 
Sclerites of metathoracie legs extending midway along A5 or its 
posterior margin. Cremaster consisting in a pair of strong and very 
close spines, ventrally curved and abruptly tapered at apex; anal 
groove Y-shaped. Dorsum of A4-A8 with 1-2 irregular rows of spines 
on anterior margin; anterior margin A3 and areas surrounding spine 
rows on A4-A8 covered by hundreds of tiny, solitaire spines. Dorsum 
of A3-A7 with single posterior row of slender spines. Tabulation of 
spines shown in Table 1 . 

Female pupa. Lengdi 8 mm. Uniform ochre. Head with eyes and 
labial palpi distinct. Thorax with leg sclerites distinct; wings absent. 



Cremaster vestigial, reduced to a small pair of blunt, widely separated 
spines; anal groove Y-shaped, with a pair of small rounded tubercles 
on either side. Dorsum of A4 with single anterior row of reduced, 
widely spaced spines: A5-A7 with 2-3 irregular rows of spines on 
anterior margin; anterior margin of A3, areas surrounding rows on 
A4-A8 and anterior margin of A8 covered by hundreds of tiny spines, 
solitaire or in 2—4 series. A3-A6 with single posterior row of slender 
spines. Tabulation of spines shown in Table 2. 

Types. Holott/pe, 6 (with associated larval case and pupal 
exuvium), reared from larva (emerged June 2002), CUBA: Sancti 
Spiritus province, Topes de Collantes, Cabumi River depression, 500 
m, 30 April 2002 (R. Nunez), slides RNA 013, 016, 023. 040, 045. 
Paratypes, 1 larva, same data as holotype, slides RNA 065, 066, 070: 6 
larval cases (some with associated pupal exuvium), CUBA: Sancti 
Spiritus province, Topes de Collantes, mogote Mi Retiro northern 
base, S00 m, 17 May 2003 (R. Nunez); 4 larval cases (some with 
associated pupal exuvium), CUBA: Sancti Spiritus province, Topes de 
Collantes, soudiern side of rocky outcrop at Pico Potrerillo access, 850 
m, 18 May 2003 (R. Nunez), slides RNA 075, 076: 3 larval cases (one 
with associated pupal exuvium), same data as preceding except 17 
June 2004; 2 larval cases, CUBA: Sancti Spiritus province. Topes de 
Collantes, Parque Codina, 800 m, 20 May 2003 (R. Nunez). 

Natural history observations. Larvae fed on 

Plagiochila sp. (Plagiochilaceae), a hepatic growing on 




45 46 

Figs. 45^16. Paucivena spp. male pupa. 45 P. ferruginea , n. sp., ventral view; 46 P.ferruginea, n. sp.. dorsal view 



134 



Journal of the Lepidopterists' Society 






Figs. 47^9. Paucivena spp. male pupa. 47 P. pinarensis, n. sp., ventral view; 48 P. cubana, n. sp., ventral view; 49 P. orientalis, 
n. sp. Scale= 0.25 mm. 





50 \y 51 

Figs. 50-51. Paucivena ferruginea, n. sp., female pupa. 50 Ventral view; 51 Dorsal view. Scale= 0.25 mm. 



Volume 60, Number 3 



135 



rocks on the Caburni River shore. However, larvae 
probably use other hosts since cases were found on 
different substrates at other localities. At die Pico 
Potrerillo access cases were found attached to rocky 
walls covered by crustose lichens, at mogote Mi Retire 
base they were located on banana plants whereas at 
Parque Codina they live on trunks of native trees. The 
single adult emerged after a month as pupa. 

Paucivena cubana inhabits localities with very 
different vegetation and climatic conditions. At Parque 
Codina it was found in the lower strata of secondary 
evergreen forest, a very humid habitat, and at the 
mogote Mi Retire base in cultivated land, banana and 
coffee (Coffea arabica L., Rubiaceae), close to the 
mogote rockv wall. On the other hand, Caburni River 
shores are covered by gallery forest remnants, today 
dominated by an introduced tree (Syzigium jambos L., 
Myrtaceae). whereas at the Pico Potrerillo access cases 
were attached to rockv walls surrounded by evergreen 
forest. 

Distribution (Fig. 52). Known from four localities at 
Topes de Collantes, Trinidad Mountains, central Cuba. 

Etymology. The species name is derived from die 
name of the Cuban island. 

Remarks. The larval color pattern is identical to that 
of P. ferruginea; however, differences in chaetotaxy and 
abdominal coloration are present. Male and female 
pupae are easily distinguished by the arrangement of 
abdominal dorsal spines (Tables 1, 2). 

Paucivena orientalis Nunez, new species 

Figs. 5, 10, 15, 21, 23, 25, 27, 44, 49, 52 

Diagnosis. Adult males of P. orientalis possess a 
color pattern similar to that of P. pinarensis, P. cubana, 
and P. reticulata. P. orientalis may be separated from P. 
pinarensis by its smaller tibial spurs (0.15 versus 0.3 the 
length of basal tarsal segment), the presence of a saccus 
in its genitalia and its more distinct FW reticulated 
pattern. From P. reticulata, it may be distinguish by its 
smaller eyes (interocular index 0.7 versus 1.1) and tibial 
spurs (0.15 versus 0.5 the length of basal tarsal 
segment). From P. cubana it differs by its smaller size 
(8.2—9.1 versus 12 mm of wing expanse) and eye size 
(interocular index 0.7 versus 1.5). 

Male (Figs. 5, 10, 15, 21). Head: pale greyish brown. Anten- 
nae with 16 segments; lateral pectinations 2—2.5 times the length 
of segment. Eyes small, vertical diameter of eye 0.7 the interoc- 
ular distance. Thorax (Figs. 10, 15): dorsum dark brown; under- 
side pale greyish brown, inner surface of legs dark brown. Vesti- 
ture dense, scales hairlike. Tibial spurs approximately 0.15 the 
length of basal tarsal segments (Fig. 10). Wings thinly scaled. 
FW variously rounded; pale grayish brown streaked with dark 
brown scales to form a reticulated pattern; fringe with various 
tones of brown. Scales at discal cell mostly oblanceolated and 
ovobated, with rounded, or rarely acute apices. Venation (Fig. 
15) as in P. ferruginea . HW uniform pale greyish brown; fringe 



with various tones of brown; venation as in P. fusca. Wing ex- 
panse: 8.2-9.1 mm (x=8.8, SD=0.49 CV=6%, n=3). Abdomen: 
dorsum dark brown at bodi ends, remainder pale greyish brown. 
Vestiture dense, scales hairlike. Genitalia (Fig. 21): tegumen 
broad, with a pair of sparsely setose apical lobes. Valvae with 
pufvilli sparsely setose; apex of sacculus armed with three to four 
spines; cucullus with apex rounded and sparsely setose. Saccus 
reduced, approximately 0.1 the length of main body. Aedeagus 
simple, cylindrical, 0.6 times die length of valvae. 

Female (Figs. 23, 25, 27). Length 5.0 mm. Vermiform. 
Stramineous with six longitudinal bands of brown spots on dorsum 
and sides of body. Head (Fig. 23): stramineous, eyes black. Slightly 
sclerotized. Shape near ovoid (ventral view); eyes compound, well 
developed, subventral. Labial palpi 1-segmented, almost 100% fused; 
antennae absent. Thorax (Fig. 25): patterned as above; body wall 
slightly sclerotized. Legs functional, armed with numerous tinv spines; 
tibiae swollen; tarsi 1 -segmented with a pair of claws at distal end (Fig. 
25); wings absent. Abdomen: color pattern disappearing at A2-A3; 
membranous and naked except for a ring of dense brownish ochre 
hairlike scales around A7. External genitalia reduced (Fig. 27), largely 
membranous. Two pairs of apophyses present; anterior pair elongated, 
free except, bifid base fused with tegument; posterior pair straight and 
free. 

Larval case (Fig. 44). Dimensions, length: a" 6.5-7.1 mm 
(x=6.7, SD=0.32, CV=5%, n=3), 9 9-12 mm 
(-s=10.8, SD=1.10, CV=10%, n=5); maximum diameter: 6 1.5-1.7 mm 
(x=1.6, SD=0.12, CV=8%, n=3), 9 1.8-2.7 mm (x=2.2, SD=0.34, 
CV=15%, n=5). Fusiform in outline, soft. Exterior densely covered by 
elongated and divergent fragments of leaves and stems of brvophvtes 
and herbaceous plants, occasionally hair fragments are added: 
material is arranged lengthwise. 

Male pupa. Length 3.8-18 mm (*=4.3, SD=0.71, CV=17%, 
n=2). Uniform brownish ochre. Frontal ridge absent, frons rounded. 
Antennal sclerites usually extending beyond apex of prothoracic legs 
(Fig. 49). Wing sheaths extending midway along A3. Sclerites of 
metathoracic legs usually extending to A4 posterior. Cremaster 
consisting in a pair of strong and widely separated, ventrallv curved 
spines; anal groove Y-shaped. Dorsum of A4-A8 with 2-3 irregular 
rows of spines on anterior margin; anterior margin of A3 and areas 
surrounding rows at A4— A5 covered by hundreds of tinv spines, 
solitaire or in 2—5 series. Dorsum of A4— A7 widi single posterior row 
of slender spines. Tabulation of spines shown in Table 1. 

Female pupa. Length 4.9-5.1 mm (x=5.0, SD=0.1, CV=2%, 
n=3). Uniform ochre. Head with eves and labial palpi distinct. Thorax 
with legs sclerites present; wings absent. Cremaster vestigial, reduced 
to a coarse and rough area around anal groove; anal groove Y-shaped. 
Dorsum of A4-A6 and A8 with single row of spines on anterior 
margin, A7 with two rows; areas surrounding rows at A4— A6 and 
anterior margin of AS covered by hundreds of tinv spines, solitaire or 
in 2-5 series. A4-A6 with single posterior row of slender spines. 
Tabulation of spines shown in Table 2. 

Types. Holotype, i (widi associated larval case and pupal 
exuvium), reared from pupa (emerged 28 April 2004). CUBA: 
Santiago de Cuba province, La Gran Piedra. Estacion Meteorologica 
La Gran Piedra, 1100 m, 23 April 2004 (R. Nunez), slides RNA 024, 
031, 035, 036. Paratijpes, 3 larval cases (some with associated pupal 
exuvium), CUBA: Granma province, La Bayamesa, abandoned coffe 
plantation at Nuevo Mundo stream shore, 1600 m, 21 April 2004 (R. 
Nunez); & (with associated larval case and pupal exuvium), reared 
from pupa (emerged 2 May 2004), same data as holotvpe: ■:' (widi 
associated larval case and pupal exuvium), reared from pupa ( emerged 
9 May 2004), same data as holotvpe; 2 (widi associated larval case and 
pupal exuvium), reared from pupa (emerged 24 May 2004), same data 
as holotype, slides RNA 030. 034; 1 larva widi its larval case, same data 
as holotype; 9 larval cases (some with associated pupal exuvium), same 
data as holotype. 

Natural historv observations. Larvae were 

collected on substrates covered bv several lichens and 

mosses and on external walls of edifications; thus, this 



136 



Journal of the Lepidopterists' Society 



species probably also feeds on detritus. The adult reared 
from a larva, a female, emerged after three weeks. One 
case was collected widi an egg cluster and 53 larvae 
hatched 1 May 2004. 

At Nuevo Mundo larval cases were found on trunks of 
old isolated Mangifera indica L. (Anacardiaceae) trees, 
in an abandoned coffee plantation. At Gran Piedra all 
individuals were located on edification walls. 

Distribution. Known only from two widely 
separated localities at Sierra Maestra, southeastern 
Cuba (Fig. 52). 

Etymology. The species name is referred to its 
distribution range, restricted to the oriental Cuban 
region. 

Remarks. The larval case is diagnostic for this 
species. Although constructed with small dry vegetal 
fragments like those of P. pinarensis and P. cubana, 
material is always cut in slender pieces and arranged 
lengthwise but in a divergent way. Pupae of both sexes 
are easily distinguished by the arrangement of 
abdominal dorsal spines (Tables 1, 2). 

Discussion 
All new species described here fit the genus 



description. However, interespecific variation in the 
interocular index was noted with values between 0.7 and 
1.5. This is the only character that shows significant 
deviation. Measurement given by Davis (1975) was 1.1. 
Deviation in Cuban representatives may be due to 
differences in daily activity between species. Powell 
(1973) also used an "eye index" in his study on New 
World Ethmia Hiibner (Oecophoridae: Ethmiinae). 
Index values were between 0.9 and 1.2 in moths known 
or presumed to be nocturnal whereas diurnal species 
exhibited ratios from 0.7 to 0.8. All adult males of 
Paucivena species described here were lab-reared, 
except those oiP.fusca that always were collected flying 
during the day. Data on daily activity of species 
described by Davis (1975) are unavailable so diis matter 
will only be clarified with future work. 

The female, described here for the first time, shows 
characters that confirm Paucivena intermediate position 
among the American psychids proposed by Davis (1975) 
based on male characters. Female primitive features are 
the presence of well developed compound eyes, 
functional legs and the behaviour of leaving the larval 
case and climbing on it, as occurs in some primitive Old 
Word forms (Davis, 1964). Specialization evidences are 




O Paucivena ferruginea 
u Paucivena pinarensis 
♦ Paucivena fusca 
-k Paucivena cubana 
O Paucivena orientalis 



Fig. 52. Distribution of Cuban Paucivena spp. Scale bar in kilometers. 



Volume 60, Number 3 



137 



tlie reduced genitalia, the complete loss of antennae and 
wings and the slightly sclerotized bodv wall. 

In the larvae, the color pattern of the head and thorax 
apparently lack diagnostic value, at least in Cuban 
species. In P. ferruginea, P. cubana, P. pinarensis and P. 
orientalis (the last two not described but observed 
during rearing), the pattern is white to greyish white 
with dark fuscous longitudinal bands. However, the 
abdominal integument is differently colored in P. 
ferruginea (dark brown) and P. hispaniolae and P. 
cubana (dirty white). Differences in chaetotaxy may be 
also used for species recognition. 

Davis (1975) mentions that P. hispaniolae larvae feed 
on crustose lichens growing on the bark of an 
unidentified tree. Larvae of Cuban Paucivena feed on a 
wide variety of hosts including mosses, hepatics, lichens 
and detritus. All diese food preferences have been 
observed before in the Psychidae (Davis, 1964; 
Hattenschwiler, 1985; Davis & Robinson, 1998) and 
perhaps explain in part genus diversification together 
with geographic isolation. 

Paucivena appears to be well expanded on Cuba 
compared to other Antillean islands such as Hispaniola, 
Jamaica or Puerto Rico. However, this may due to lack 
of sampling on these islands. In Cuba, more collect 
effort on Psychidae is also needed. The Nipe-Sagua- 
Baracoa Mountains, in the northeast part of the island, 
and other habitats like coastal forests, ultramafic scrub, 
and white sand savannahs are yet unexplored. 

Acknowledgements 

I wish to thank Adriana Lozada, Arturo Avila, Carlos Sanchez 
and Ledis Regalado for their collaboration in the expeditions 
and to executive and administrative personnel of Facultad 
Agropecuaria de Montana del Escambray (FAME), at Topes de 



Collantes, for support during surveys. Also, I wish to thank 
Ricardo Herrera and Rigel Fernandez for the photographs and 
Luis Manuel Diaz for his help with the Corel Draw computer 
program. Luis F. De Armas, Emily V. Saarinen and Alejandro 
Barro read die manuscript suggesting valuable changes and help 
with literature. 

Literature Cited 

Davis, D. R. 1964. Bagworm Moths of the Western Hemisphere. Bull. 
Unit. St.. Nat. Mus. 244:1-385. 

. 1975. A review of the West Indian moths of the family Psychi- 
dae with Descriptions of New Species. Smithson. Contr. Zool. 
188:1-66. 

. 2000. Brachygyna incae, a new genus and species of Psychidae 

from Peru with atypical larval biology (Lepidoptera: Tineoidea). 
Trap. Lepid. 10(2): 51-58. 

Davis, D.R. & G.S. Robinson. 1998. The Tineoidea and Gracillari- 
oidea, pp 91-117. In N.P. Kristensen (ed.), Lepidoptera, Moths 
and Butterflies. Handbuch derZoologie IV (Arthropoda): Insects 
35. Walter de Gruyter and Co., Berlin and New York. 

HAttenschwiler, P. 1985. Psychidae, pp 128-151. In J. Heath (ed.), 
The Moths and Butterflies of Great Britain and Ireland. Vol. 2 
Cossidae-Heliodinidae. Black-well Scientific Ltd. Oxford, and The 
Curwen Press, London. 

HENDERICKS, H. 1982. Possible determination characters in psvchid 
females. SHILAP Rev. Lepid. 10(39): 174. 

Heppner, J.B. 1998. Classification of Lepidoptera. Part 1 Introduc- 
tion. Hoi. Lepid. 5 (Suppl. 1): 1-148. 

Hinton, H.E. 1946. On the homology and nomenclature of the 
setae of lepidopterous larvae, with some notes on the phvlogenv 
of the Lepidoptera. Trans. Roy. Entomol. Soc. Lond. 97: 1-37. 

Nunez, R. 2004. Lepidoptera (Insecta) de Topes de Collantes. Sancn' 
Spiritus, Cuba. Bol. S.E.A. 34: 151-159. 

Powell, J. A. 1973. A Systematic Monograph of New World Ethmiid 
Moths (Lepidoptera: Geleehioidea). Smiths. Contrib. Zool. 
120:1-301. 

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(ed.), Immature Insects. Kendall/Hunt Publishing Co., Dubuque. 

Received for publication 3 May 2005; revised and accepted 30 May 
2006 



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journal of the Lepidopterists' Society 
60(3), 2006, 138-142 

A REMARKABLE NEW RIODINID SPECIES, STALACHTIS HALLOWEENI (RIODINIDAE: 
STALACHTINI), FROM MOUNT AYANGANNA, GUYANA 

JASON P. W. HALL 

Department of Entomology, National Museum of Natural History.Smithsonian Institution.Washington, DC 20560-0127, USA 

Email: hallja@si.edu 



ABSTRACT. A new riodinid species, Stalachtis halloweeni Hall n. sp. (Stalachtini), is described from Mount Ayanganna, a tepui in western 
Guyana. A preliminary hypothesis of phylogenetic inter-relationships within the small genus Stalachtis Hiibner is suggested, based on an 
informal study of external morphology and male genitalia. Three species groups are proposed, the phlegia, calliope and euterpe groups, and 
S. halloweeni is hypothesized to be sister to the remaining members of the euterpe group. 

Additional key words: endemism, montane forest, morphology, South America 
Situated in the Pakaraima Mountain Range of Guyanan 



Range 

western Guyana, Mount Ayanganna (2042m) is one of 
the easternmost tepuis in the Guiana Shield. Like the 
other tepuis in western Guyana and eastern Venezuela, 
Mount Ayanganna consists of eroded sandstone 
remnants of the Roraima Formation (MacCulloch & 
Lathrop 2001), and its isolated high-elevation habitats 
harbor significant numbers of endemic plants and 
animals. During the last five years alone, several new 
endemic species of frogs, snakes and lizards have been 
described from Mount Ayanganna (MacCulloch & 
Lathrop 2001, 2002, 2004). 

During the last ten years, there have been numerous 
Lepidoptera expeditions to the upland regions of 
western and southern Guyana (Fratello 1996, 2001, 
2003, 2005). In 1999, accompanied by several US and 




Figs. 1—2. Stalachtis halloweeni adults (dorsal surface on 
left, ventral surface on right). 1. Holotype male. Mount Ayan- 
ganna, Guyana (USNM). 2. Paratype female, Mount Ayan- 
ganna, Guyana (USNM). 



colleagues, S. Fratello led the first 
Lepidoptera collecting expedition to the upper slopes of 
Mount Ayanganna. Among the many new taxa of 
butterflies collected there were several new species of 
Riodinidae, including a new species of Stalachtis 
Hiibner, 1818 (Stalachtini), although most of these 
species were represented by only a small number of 
female specimens. Fortunately, a second expedition to 
Mount Ayanganna in 2002, by a different team that 
included one of the 1999 Guyanan expedition members 
(R. Williams), produced additional Stalachtis 
specimens, including males. 

This new Stalachtis species is remarkable in several 
respects. It is the first new species to be described in 
this small, essentially South American genus of 
aposematic species for over 150 years, since Westwood 
(1851) described S. magdalena; it is the first known 
Stalachtis species to apparently occur exclusively in 
montane habitats; and its wing pattern differs 
substantially from that of its congeners. I herein 
describe this new Stalachtis species and attempt to 
establish its phylogenetic position within the genus by 
informally constructing a preliminary hypothesis of 
phylogenetic relationships for Stalachtis and proposing 
a new species-group classification. 

Stalachtis halloweeni Hall, new species 

(Figs. 1-2; 3; 4) 
Description: MALE: Forewing length 29.5 mm. Forewing 
elongate, costal and distal margins approximately straight, four 
forewing radial veins, discal cell elongate; hindwing rounded and 
slightly elongate, hindwing veins Rs and Mj stalked. Dorsal surface: 
Forewing ground color black, discal cell orange with some black 
scaling medially, broad orange rays in cells 2A and Cu„ extending from 
(Cu.,) or near (2A) wing base to join submarginal band, orange 
rectangle at base of cell Cu r small orange spot at base of cell M 3 , 
orange streaks in cells M, to IU. and R„ immediately distal to discal 
cell end, decreasing in size from cell M. to cell R„ broad submarginal 
orange band extending from costa to tornus, with an enlarged 
rectangular orange patch nearly encircling a black spot in cells M 3 and 
Cu 2 , distal fringe black; hindwing ground color black, pale orange spot 
at wing base, a broad orange streak through discal cell and cells Cu, to 
M and along anal margin, those in cells Cu, and M 3 shorter than 



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139 



others and that in cell Cu, joining submarginal band, small distal 
orange spot in cell Sc+R r broad and uneven submarginal orange band 
extending from apex to tornus, distal fringe black. Ventral surface: 
Forewing differs from dorsal surface bv having a small orange spot at 
costal wing base, only distal portion of orange streak present in cell 
2A, orange streak in cell M„ joining submarginal band, orange spot in 
cell R, replaced bv a sparse scattering ol whitish scales; hindwing 
differs by having a verv narrow line of orange scaling at middle of 
costal margin, with a sparse scattering of whitish scales distally, a 
larger orange spot in cell Sc+R 1; a much larger and darker orange spot 
at wing base, and a sparse scattering of whitish scales along anal 
margin. 

Head: First segment of labial palpi a mixture of black and white 
scaling, segment two black with a broad white lateral band and some 
white scaling ventrally. and third segment black; eyes bare and black, 
with a mixture of black and white scaling at margins; frons black, with 
white scaling laterally; antennae 60% of forewing length, segments 
black with white scaling at ventral base and narrow nudum region 
along inner ventral margin, clubs black. 

Body: Dorsal surface of diorax black, tegula black with dark orange 
scaling at base, ventral surface of thorax black with a white band 
between legs and a large orange patch near base of forewing; 
abdomen black dorsally, with a broad orange band laterally, and white 
ventrally, with narrow black lines on either side (patterning virtually 
indiscernible in Fig. 1 due to a covering of mold), all legs black. 

Genitalia (Fig. 3): Uncus in lateral view rectangular and vertically 
elongate, lateral "window" anterior to uncus very narrow, tegumen 
very narrow, with a triangular ventral margin; falces extremely long 
and ventrally directed, with a weakly bent "elbow"; vinculum narrow, 
anteriorly bowed, broadest medially and slightly posteriorly indented 
near ventral tip, with a posteriorly directed triangular section of 
sclerotized tissue at ventral margin and no anterior saccus; aedeagus 
narrow, convex and ol medium length, gradually tapering to a slightly 
upturned and finely pointed tip, vesica exits along ventral margin of 
posterior third of aedeagus, comutal patch consists of a short 
sclerotized rod with about six prominently curved and anteriorly 
directed spines densely positioned along all but its anterior tip, 
comutal patch positioned about one third distance from posterior to 
anterior tip of aedeagus on uneverted vesica; pedicel in lateral view 
broad basally, becoming narrower in angular posterior section, pedicel 
joins aedeagus about one third distance from anterior to posterior tip; 
valvae in lateral view consist of a large rectangular basal section, a 
narrow, posteriorly elongate and round-tipped lower process, a slightly 
broader, more rectangular and posteriorly elongate upper process, 
with a posteriorly and upwardly curved, finely pointed terminal 
projection, a very broad and rounded inner process, slightly shorter 
than lower process, and a pointed transtilla of medium length 
extending posteriorly between pair of inner processes and across top 
of aedeagus; narrow tuft of long, posteriorly directed, pale brown 
setae around outer margin of genital capsule; eighth abdominal tergite 
and sternite rectangular. 

FEMALE: Differs from male in following ways: Forewing length 
28 mm. Distal forewing margin convex. Wings: Orange on both wings 
very slighdy paler, medial black scaling in forewing discal cell more 
prominent, several forewing postdiscal orange spots do not extend as 
far proximally to cell bases, orange ray in ventral forewing cell M„ does 
not distally join submarginal band. 

Head: Second palpal segment slightly more elongate, third 
segment about twice as long; nudum region on antennal segments 
slighdy larger. 

Genitalia (Fig. 4): Corpus bursae somewhat narrow and elongate, 
with a pair of small, sclerotized, invaginated spine-like signa; ductus 
bursae consists of a large, creased, hardened swelling immediately 
posterior to corpus bursae containing about four pieces of rectangular 
sclerotization, a short membranous section posteriorly, dren a long and 
twisted lightly sclerotized section, followed by a short, concave ventral 
section of sclerotization; membranous ductus seminalis exits ductus 
bursae dorsally immediately anterior to ventral section of 
sclerotization; ostium bursae in dorsal view consists of a small, round, 
sclerotized entrance hole, with a broad and prominently convex band 




Fig. 3. Male genitalia of Stalachtis halloweeni holotype in lat- 
eral view. Scale bar = 1 mm. 

of sclerotization dorsally curving anteriorly into an invaginated pouch 
below papillae anales that is membranous except for a broad, 
triangular, horizontal band of sclerotization along dorsal "roof" of 
pouch (perhaps a protected resting place for extremely long male 
genital falces during copulation); papillae anales proportionately small 
and round; very broad, semicircular tuft of long, posteriorly directed, 
pale brown setae around posterior margin of eighth tergite. 

Types: Holotype d, GUYANA: Cuyuni-Mazaruni, Mount 
Ayanganna, 1120"m, 5°22.22'N 59°57.34'W 12-16 Oct 2002 ^R. 
Williams) (National Museum of Natural History, Smidisonian 
Institution, Washington, DC, USA [USNM]). 

Paratypes: lc5, 19, same data as holotype. 39, GUYANA: Cuyuni- 
Mazaruni, Mount Ayanganna, 4500-5500 ft, 5°24.1'N 59°57.4'W, 
13-18 Apr 1999 (S. Fratello et al.) (USNM). 

No additional specimens have been located in the major museums 
of Europe and North America (as listed in Hall 1999, 2005). 

Etymology: The name is derived from the middle 
English word halloween, in reference to the fact that 
the wing pattern is composed of the traditional orange 
and black colors of Halloween, and is reminiscent of a 
carved pumpkin. 

Systematic placement and diagnosis: Stalaclitis ( = 
Nerias Boisduval, 1836) is the sole genus currently 
treated in the tribe Stalachtini. The familv-group name 
was proposed by Bates (1861), as a subfamilv, for an 
undefined number of genera whose species possessed a 
"pupa not flattened beneath, secured rigidly bv the tail 
in an inclined position, without girdle", a set of 
characters quickly discovered bv Bates (1868: 368) 
himself not to be phvlogeneticallv informative in die 
context of the higher classification of the Riodinidae. 
Stichel (1910-11) used the Stalachtini as a tribal name 
for the first time (as the Stalachtidi). and followed Bates 
(1868) in including only Stalachtis within it. Most 
recently, Harvey (1987) defined the monotvpic 
Stalachtini by the presence of a tuft of long setae around 



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Journal of the Lepidopterists' Society 





Fig. 4. Female genitalia of Stalachtis halloweeni paratype in 
dorsal view. Scale bar = 1 mm 



the posterior margin of abdominal segment eight in 
males and, to a greater extent, females, and mentioned 
that its species are unusual in the Riodinidae in having 
hindvving veins Rs and M { stalked rather than arising 
separately from the discal cell end (Bates 1S68; Stichel 
1910-11). Additional unique characters that are 
universal within the Stalachtini include extremely long 
and weakly bent (i.e. ventrally directed) falces, a 
complex arrangement of inner valve processes, and a 
long straight comb of prominently curved, anteriorly 
directed spines on the aedeagal vesica in the male 
genitalia (Hall unpubl. data). The new Stalachtis 
species, S. halloweeni, possesses all of the above 
characters. 

Traditionally, Stalachtis has been treated as 
containing eight species (e.g. Stichel 1910-11, 
1930-31), although some authors have listed as many as 
ten species (D'Abrera 1994; Bridges 1994). However, 
the genus is now justifiably recognized to contain only 
six species (Callaghan & Lamas 2004). Hemming (1964, 
1967) selected the type species of Hiibner's (1818) 
Stalachtis to be S. phaedusa (Hiibner, [1813]). It is 
worth mentioning that the name Stalachtis funereus 
alb ul us Lathy, 1958, which occasionally appears in the 
riodinid literature (e.g. Rebillard 1958; D'Abrera 1994; 
Bridges 1994), actually refers to a pericopine moth 
(Arctiidae) (Hall unpubl. data). Stichel (1910-11, 
1930-31) divided Stalachtis into two sections, the 
"Adiorati" for S. calliope (Linnaeus, 1758) and S. 
magdalena Westwood, 1851, in one subgroup and S. 
phlegia (Cramer, 1779) (+ S. susanna (Fabricius, 1787)) 
and S. euterpe (Linnaeus, 1758) in another subgroup, 
and the "Diaphanes" for S. phaedusa (+ S. zephyritis 
(Dalman, 1823)) and S. lineata (Guerin-Meneville, 
[1844]). Based on a study of wing pattern and male 
genitalia characters in all six Stalachtis species, an 
alternative species-group classification and preliminary 
hypothesis of phylogenetic relationships for the genus is 
proposed here. 

As S. calliope, S. magdalena, S. euterpe, S. phaedusa 
and S. lineata all share a pair of large, rounded, inner 
valve processes, with an elongate, pointed transtilla 
extending posteriorly between them and across the top 
of the aedeagus, S. phlegia is hypothesized to be sister 
to the remaining species in the genus. Stalachtis phlegia 
has a much smaller pair of inner valve processes, 
without the intervening transtilla, and has the least 
derived wing pattern, with a full complement of white 
basal, postdiscal and submarginal spots. Stalachtis 
calliope and S. magdalena, like S. phlegia, but unlike 
any odier Stalachtis species, have a complete row of 
submarginal white spots on the forewing, an orange 
patch at the base of the dorsal forewing and hindwing, 



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141 



and an entirely checkered black and white hindwing 
fringe. Their similar wing patterns, with mottled orange 
markings at the base of the forewing and parallel orange 
bands on the hindwing, shared possession of an upper 
valve process that is broadest medially (instead of 
basallv in S. phlegia and distallv in the remaining three 
species), and parapatric geographic ranges strongly 
suggest that they are sister species. Stalachtis euterpe, S. 
phaedusa and S. lineata appear to form a monophyietic 
group, as all three species share similarly positioned 
white wing markings that are consistently formed into 
rays instead of spots, the absence of an orange patch at 
the base of the dorsal forewing and hindwing, the 
absence of a complete row of submarginal white spots 
on the forewing, largely black wing fringes, and an 
upper valve process that is broadest distallv. As 
Stalachtis phaedusa and S. lineata both have elongate 
hvaline rays on both wings they are probably sister 
species. 

Stalachtis halloweeni appears to exhibit external 
pattern characters that are somewhat intermediate 
between those of members in the calliope and euterpe 
groups. It shares with the two calliope group species the 
presence of orange markings at the base of the dorsal 
forewing, and with S. magdalena the absence of a lateral 
white line above as well as below the lateral orange band 
on the abdomen. It shares with the three euterpe group 
species a similar pattern of rays at the base of the 
forewing and particularly the hindwing, even if these are 
orange instead of white, the absence of a complete row 
of submarginal white spots on the forewing, and black 
wing fringes. Based on the above characters, and the 
fact that S. halloweeni has the full complement of inner 
valve processes and an upper valve process that is 
broadest distally, this new species is tentatively 
suggested to be the most basal member of the euterpe 
group. The male genitalia of Stalachtis species are 
rather homogeneous, with the most significant 
interspecific variation exhibited by the upper and, to a 
lesser extent, lower valve processes. Although S. 
halloweeni appears to be most closely related to S. 
euterpe, its male genital valvae are probably most 
similar to those of S. phaedusa, but its upper valve 
process does not have such a prominent ventral 
protrusion at the posterior tip, and the posterior margin 
extends at about a 45° angle instead of vertically. 
Elsewhere in the euterpe group, S. euterpe can be 
characterized by its dorsal as well as ventral swelling to 
the distal tip of the upper valve process and atypically 
small and straight terminal projection, and S. lineata can 
be characterized by its broadly triangular instead of 
narrower rod-shaped lower valve process. 

In conclusion, Stalachtis seems to be best divided 



into three species groups, the phlegia group for S. 
phlegia, the calliope group for S. calliope and S. 
magdalena, and the euterpe group for S. halloweeni, S. 
euterpe, S. phaedusa and S. lineata. Thus, only two of 
Stichel's (1910-11, 1930-31) three proposed species 
groups for Stalachtis appear to be monophyietic. 

Biology: This new species appears to be restricted to 
lower montane forest habitats, where it is currently 
known from between about 1100 and 1700 m. 
Specifically, the type series was collected in wet, low 
(canopy approximately 10-15 m), evergreen high-tepui 
forest, a vegetation type that was described and 
illustrated by Maguire (1970), Huber et al. (1995) and 
MacCulloch & Lathrop (2001). 

Steve Fratello (pers. comm.) observed approximately 
ten individuals of S. halloweeni on Mount Ayanganna, 
most of which were probably females, judging by the 
fact that all individuals captured were of that sex. Within 
the forest, these individuals consistently flew at about 5 
to 7 m above the ground, although two individuals were 
observed flying only 2 to 3 in above the ground over a 
patch of low tepui scrub at 1700 m. Individuals were 
seen flying over a wide area from mid-morning to mid- 
afternoon, with a rather slow, steady flight, and were not 
observed alighting or resting. No other Stalachtis 
species were seen flying in the same habitats as S. 
halloweeni, but S. phaedusa, S. calliope and S. euterpe 
have been commonly collected in neighboring lowland 
areas of Guyana (Fratello pers. comm.). 

Stalachtis is one of the most well known groups of 
aposematic riodinids (Seitz 1916-20; DAbrera 1994). 
Given that the known caterpillars are gregarious and 
aposematic (Callaghan 1986), and members of at least 
some of the known foodplant families (e.g. 
Simaroubaceae) contain toxic phytochemicals (e.g. 
Moretti et al. 1982; Polonsky et al. 1984), it seems likely 
that some or all of the Stalachtis life stages are to some 
extent distasteful to predators, and adults may thus be 
predominantly Mullerian rather than Batesian mimics. 
However, I am not aware of any sympatric butterflies or 
moths that specifically closely resemble S. halloweeni. 

Distribution: Stalachtis halloweeni is eurrentiy 
known only from the middle slopes of Mount 
Ayanganna, in the uplands of western Guvana. 
However, the geographic range of diis species probably 
extends to neighboring highland areas in Guvana and 
extreme eastern Venezuela. There continues to be 
debate about whether most Guiana highland endemics 
are relicts of a widespread pantepui fauna or 
descendants of lowland ancestors (e.g. Mvers & 
Donnelly 1996; MacCulloch & Ladirop 2001). Given 
that all six described Stalachtis species have entirely 
lowland distributions, the ancestor of S. halloweeni 



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Journal of the Lepidopterists" Society 



presumably colonized Mount Ayanganna from the 
surrounding lowlands. 

Acknowledgements 

I thank The National Science Foundation (DEB # 0103746) 
for financial support; G. Lamas and S. Fratello for carefully re- 
viewing the manuscript; S. Fratello and R. Williams for collect- 
ing and donating Stalachtis specimens to the USNM; and S. 
Fratello for generously providing notes on the ecology of Sta- 
lachtis halloweeni from his arduous and highly productive 1999 
Mount Ayanganna expedition, which was funded in part by the 
Smithsonian Biological Diversity of the Guianas Program. 

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Received for publication 9 August 2005; revised and accepted 2 June 
2006 



Volume 60, Number 3 



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Journal of the Lepidopterists' Society 
60(3), 2006, 143-148 

A NEW SPECIES OF AURATONOTA (LEPIDOPTERA: TORTRICIDAE: CHLIDANOTINAE) 
FORMERLY CONFUSED WITH A. HYDROGRAMMA (MEYRICK) 

John W Brown 

Systematic Entomology Laboratory, P.S.I., Agricultural Research Service, U.S. Department of Agriculture 

National Museum of Natural History, P.O. Box 37012, Washington, DC 20013-7012 

Email: jbrown@sel.barc.usda.gov 

ABSTRACT. Auratonota pharata, new species, is described and illustrated from Costa Rica, Panama, Venezuela, and French Guiana. The 
species is most similar to A. hydrogramma (Meyrick), with which it formerly was contused. It can be distinguished superficially from the latter 
by the absence of the narrow pale curved band beyond the distill end of die discal cell of the forewing. The male genitalia of the new species 
differ by a slighdv expanded, dorsally convex, and ventrallv flattened distal portion of the uncus. The female genitalia possess numerous short 
curved bands of 5-6 microtrichia around a larger single seta set in a shallow pit on the surface of the middle of the papillae anales compared 
widi die more semicircular bands of microtrichia nearly surrounding the seta in A. hydrogramma. A survey of wing coupling in numerous gen- 
era of Chlidanotini and Hilarographini revealed diat the female frenulum consists of two bristles usually separated throughout dieir length in 
all representatives examined in diese two tribes: three brisdes are present in females of most other Tortricidae. This character state represents 
an additional putative svnapomorphy uniting those two tribes. 

RESUMEN. Auratonota pharata, especie nueva, es descrita e ilustrada de Costa Rica, Panama, Venezuela y Guyana Francesa. Este especie 
es mas similar a A. hydrogramma (Meyrick), con la cual ha sido con tundida. Puede ser identificada de una manera superficial de esta ultima 
por la ausencia de una banda curva fina clara detras de la portedistal de la celda discal de las alas anteriores. La genitalia del macho de la nueva 
especie puede ser distinguida por la presencia en la partediseal del uncus de una area ligeramente expandida, convexa dorsalmente v plana ven- 
tralmente. La genitalia de la hembra puede ser distinguida por la presencia de numerosas bandas curvas cortas de 5-6 espinas pequenas arriba 
de una sola seta en media superficie de los papillae anales en comparaeion con bandas mas semicirculares de espinas pequenas casi rodeando la 
seta en A. hydrogramma . 

Additional key words: Systematica, genitalia, morphology, Costa Rica, Central America, Venezuela, French Guiana, inventory. Chlidan- 
otini, frenulum 



Auratonota Razowski is the largest and most diverse 
genus in Chlidanotini (Tortricidae: Chlidanotinae) with 
30 described species (Razowski & Becker 2000, Brown 
2005) and numerous undescribed species present in 
collections. The genus is restricted to the New World 
tropics, ranging from Mexico and the Caribbean (Cuba, 
Dominica) south through Brazil. 

A previously undescribed species of Auratonota has 
been concealed in entomological collections for many 
years under the name A. hydrogramma (Meyrick). The 
similarity of the new species to A. hydrogramma in size, 
forewing pattern, and genitalia, along with their 
geographic sympatry, have combined to inhibit their 
recognition. The two species can be separated by a 
subtle feature of the forewing pattern, but recently 
discovered features of the male and female genitalia 
provide convincing evidence that they are indeed 
distinct and diagnosable. The purpose of this paper is to 
name, describe, and illustrate the new species. This 
work was stimulated, in part, by the desire to associate 
scientific names with morpho-species collected during 
the NSF-funded ALAS (Arthropods of La Selva, Costa 
Rica) project in order to more easily discuss differences 
and similarities among the tortricid faunas of transect 
sites (at different elevations) sampled over the course of 
the project (1993-2005). 

Dissection methodology followed that presented in 
Brown and Powell (1991). Images of adults and genitalia 



were captured using a Microptics digital camera system 
and enhanced using Adobe Photoshop and Illustrator 
software. Terminology for genital structures follows 
Horak (1984). The following institutional abbreviations 
are used for the deposition of specimens examined: 
AMNH = American Museum of Natural History, New 
York, New York, U.S.A.; BMNH = The Natural History 
Museum, London, United Kingdom: INBio = Insrituto 
Nacional de Biodiversidad, Santo Domingo de Heredia. 
Costa Rica; UCB = Essig Museum of Entomology, 
University of California, Berkeley, USA; and USNM = 
National Museum of Natural History, Washington, 
D.C.. U.S.A. 

Auratonota pharata Brown, new species 

(Figs. 2, 4, 5) 
Diagnosis. Among described species of Auratonota. 
A. pharata is most similar to A. hydrogramma in 
forewing pattern, size, and genitalia. However, die latter 
is superficially distinguishable from all described 
congeners, including A. pharata, by the presence of a 
slender, pale, arched fascia in die subterminal region of 
the forewing that roughly parallels the apical half oi die 
termen, intersecting the costa subapicallv (Fig. 1); the 
fascia is lacking in A. pharata (Fig. 2). The male 
genitalia of A. pharata can be distinguished from those 
of A. hydrogramma by the shape of the distal one- 
fourth of the uncus: attenuate and apically pointed in A. 



144 



Journal of the Lepidopterists' Society 



hydro gramma (Fig. 3); slightly broadened, convex 
dorsally, and flattened ventrally in A. pharata (Fig. 4). 
The female genitalia are extremely similar in both but 
can be distinguished by a subtle feature of the papillae 
anales. In A. pharata microtrichia on the internal, 
middle portion of the papillae anales are arranged in 
short, slightly curved bands of 5-6 immediately dorso- 
posterad of a small, pale-colored pit from which arises a 
single larger seta (Fig. 5). In A. hydro gramma 
microtrichia are arranged in circular or semicircular 
bands of 8-9 nearly surrounding a more rounded pit 
(Fig. 6). Auratonota pharata also is similar superficially 
to A. aporema (Dognin), described from Colombia, but 
has a considerably smaller forewing length (mean 11.9 
mm for pharata vs. 15.9 for aporema) and lacks the 
yellow-gold scaling of the forewing pattern elements of 
A. aporema; the latter feature is more characteristic of 
members of Pseudocomotis Brown (Chlidanotini), with 
which A. aporema is more similar superficially. 

Description. Head: Vertex rough sealed, mostly pale cream with a 
few pale cream-brown scales; frons smooth scaled, pale cream white; 
labial paplus relatively slender, short, length (all segments combined) 
ca. 1.2 times horizontal diameter of compound eye, brown externally, 
pale cream on inner surface; antenna thickened, with setae extremely 
short, inconspicuous (typical of Chlidanotini). Thorax: Dorsum 
clothed in reddish-brown scales, anterior and posterior regions with 
considerable scattered white and pale brown scales; tegula pale 
brown, with variably expanded patch of long scales posteriorly, 
frequently expressed as an erect scale patch in both sexes. Legs 
unmodified; no hairpencil or secondary scale patches in male. 
Forewing length 11.5-12.9 mm (x = 11.9; n = 10) in male, 12.2-14.1 
mm (x = 13.1; n = 2) in female; costa nearly straight, apex obtuse, 
termen slightly convex, rather oblique; ground color ferruginous, widi 
light silvery-gray reticulations formed by irregular interrupted streaks 
on veins and a series of indistinct transverse fascia crossing them, 
scattered widi iridescent green scales in interspaces (the green scales 
inconspicuous on flight worn specimens); basal portion from near base 
to ca. 0.66 distance to apex largely suffused with blackish brown 
between the reticulations; a small, ill-defined, irregularly-shaped, 
ferruginous spot near distal end of discal cell bordered basally by a 
narrow bluish silvery-gray line; termen uniform red-brown; fringe 
olive-ferruginous [lacking in most specimens examined]. Hindwing 
uniform dark brown, fringe concolorous; female frenulum with two 
bristles separated throughout their length. Abdomen: Dark brown. 
Male genitalia (Fig. 4; image of JWB slide 806, Costa Rica; n = 6) with 
uncus strong, long, mostly rod-shaped, slightly broader at base, curved 
near middle, slightly expanded in distal 0.25 with dorsum convex and 
venter flattened or weakly concave; socius moderately short, broad, 
subrectangular, clothed in long, fine scales; hami long, ca. 0.75 times 
length of uncus, weakly attenuate from base to tip, distal 0.1 bent 
dorsally; gnathos extremely reduced, lateral arms membranous, mesal 
portion inconspicuous; valva large, broad, expanding distallv. distal 
0.75 covered widi fine, long setae, costa sclerotized; transtilla a simple, 
narrow band; juxta a broad, shield-like plate; saccus well developed, 
narrow, attenuate distally; aedeagus short, stout, mostly straight, 
slightly curved at phallobase, a tiny scobinate patch of small setae on 
each side near distal end, vesica with small linear patch of 
sclerotization. Female genitalia (Fig. 5; image of USNM slide 95264, 
Panama; n = 3) with papillae anales large, bearing numerous tiny, 
weakly curved lateral bands of spines in middle portion, which are 
situated immediately dorso-posterad of a small pale-colored pit from 
which a single seta arises; perimeter and basal portions of papillae 
anales with much larger setae arising from elongate, warty bases; 
sterigma simple, weakly sclerotized. ventral posterior edge of ostium 



with narrow row of 5-6 long setae on each side; ductus bursae 
broadest at ostium, narrowed at about 0.2 distance from ostium to 
corpus bursae, then nearly uniform in width to junction with corpus 
bursae; corpus bursae, large, pear-shaped sac, junction with ductus 
bursae slightly anterior ot posteriormost end, signum a patch of 30-35 
slender, slightly curved spines originating from sclerotized plate near 
middle ol corpus; ductus seminalis from posteriormost end of corpus 
bursae near junction of coqius bursae and ductus bursae; a frail 
accessory bursae from a long, slender ductus originating just anterad 
of signum. 

Holotype. Male: Costa Rica, Estacion Biologica La Selva, Puerto 
Viejo de Sarapiqui, 50-150 m, 10°26'N, 84°01W, 7 Feb 2002, Wagner, 
Rota & Kawahara (INBio). 

Paratypes (24c?, 39). BRITISH GUIANA: Potaro River, 
Anundubaru, 2000 1 , Jan 1928 (Id") (AMNH). COSTA RICA: Heredia 
Province: Estacion Biologica La Selva, 10°26'N, 84 o 01'W, 50-150 m, 
Jan 1998 (19), INBio-OET, J. Powell (UCB), 8-25 Mar 1999 (Id*), 
22-31 Mar 2001 (19), Wagner & Rota, 7 Feb 2002 (let), Wagner, Rota 
& Kawaliara (INBio); Ciebo, 11 km ESE La Virgen, 250-350 m, 
10°21'N, 84°0rW, 18 Mar 2004 (Id*) (INBio); 10 km SE La Virgen, 
450-550 m, 10 20N, 84 05W 17 Mar 2003 (2c!), 19 Mar 2003 (Id"), 22 
Mar 2003 (1<J), INBio-OET-ALAS transect (INBio). Lim6n Province: 
Cerro Tortuguero, P.N. Tortuguero, 0-120 m, Oct 1989 (let), J. Solano 
(INBio), Jul 1991 (Id*), J. Solano (INBio), Jul 1993 (Id'), R. Delgado 
(INBio); Sector Cerro Cocori, Finca de E. Rojas, 150 m, Aug 1991 
(2c*), E. Rojas (INBio). Unknown Province: Carchi [possibly Sarchi, 
Alajuela Province], [no date] (Id"), Wm. Schaus (USNM). FRENCH 
GUIANA: St. Jean, Maroni, [no date] (2c!), Wm. Schaus (USNM); Rio 
Maroni, [no date] (Id*), Le M[oult] (USNM). PANAMA: Canal Zone, 
Barro Colorado Island, 17 Sep 1941 (19), J. Zetek (USNM). 
VENEZUELA: Aragua: Rancho Grande, 1100 m, 16-23 Oct 1966 
(3d"), 24-31 Oct 1966 (Id"), 1-5 Nov 1966 (Id"), S. S. & W. D. 
Duckworth (USNM), 15-21 Jun 1967 (Id"), 22-31 Aug 1967 (Id*), R. 
W. Poole (USNM). 

Distribution and Biology. Auratonota pharata is 
recorded from Costa Bica, Panama, Venezuela, British 
Guiana, and French Guiana. Although it appears to be a 
species of the lowlands (i.e., below 500 m), it has been 
recorded on several occasions at Bancho Grande, 
Venezuela, at 1100 m and once in British Guiana from 
660 m. During a multi-year survey in Costa Bica along 
an elevation transect, specimens were collected at 
50-150 m (n = 4), 250-350 m (n = 1), and 450-550 m (n 
= 4), with none recorded from 1000, 1500, or 2000 m. 
Adults have been recorded in January through March, 
and June through November. 

A female collected by J. Powell in January 1998 at La 
Selva, Costa Bica was confined in a plastic vial where it 
laid several huge, bulky eggs, 3.3 x 2.8 mm, ca. 10 times 
the size of comparable-sized tortricine females (J. 
Powell, pers. coram.). First instars likewise were large, 
2.8-3.0 mm in length. Various leaves, syndietic diet, and 
raw carrot were offered to the larva; the last has been 
used successfully with other "borers" such as 
Hepialidae. The only feeding was by one larva on the 
carrot, and it ceased to feed before reaching second 
instar. 

Etymology. The specific epithet is from the word 
"pharate," meaning cloaked or hidden. 

Auratonota hydrogramma (Meyrick, 1912) 



Volume 60. Number 3 



145 



(Figs. 1, 3, 6) 
Cnephasia hydrogramma Meyrick, 1912: 683. 
Eulia hydrogramma: Clarke, 1958: 128. 
Auratonota hydrogramma: Razowski, 1987: 62; Brown, 
1990: 156; Powell et al. 1995: 151; Razowski & 
Becker, 2000: 1151; Brown, 2005: 144. 

Auratonota hydrogramma was described from a 
single specimen from Dutch Guiana (= Surinam) 
(BMNH). The adult and male genitalia of the holotype 
are figured by Clarke (1958); Razowski & Becker (2000) 
provided a color illustration of an adult and line 
drawings of the male and female genitalia. Based on 
material in several museum collections (AMNH, 
BMNH, INBio, USNM), it has been recorded from 
Costa Rica, Panama, Colombia, French Guiana, British 
Guiana, and Ecuador and at many of the same localities 
as A. pharata (e.g., Costa Rica, Estacion Biologica La 
Selva; Panama, Barro Colorado Island; French Guiana, 
St. Jean, Rio Maroni). Razowski & Becker (2000) 
reported it from Brazil. As is the case in A. pharata, A. 
hydrogramma appears to be a species of the lowlands, 
rarely collected above 600 m. During the multiple-year 
transect surveys of the ALAS project in Costa Rica 
(1993-2005), A. hydrogramma was collected only at the 
50-150 m elevation site. 

Auratonota hydrogramma can be distinguished 
superficially from all other congeners by the presence of 
a narrow, curved, pale fascia in the apical region of the 
forewing. The male genitalia (Fig. 3, image of USNM 
slide 84889, Panama; n = 6) are slightly smaller and 
have a slightlv broader base of the saccus than those of 
A. pharata; but the most conspicuous feature that 
distinguishes the two species is the pointed tip of the 
uncus of A. hydrogramma. The papillae anales (Fig. 6, 
image of JWB slide 745, Costa Rica; n = 2) of A. 
hydrogramma also have the tiny curved bands of 
microtrichia described above for A. pharata, but in A. 
hydrogramma the bands are more semicircular and 
nearly surround a rounded, pale-colored pit. Even 
though only two females of each species were examined, 
these differences appear to be consistent. 

Holotype 6, Dutch Guiana [Surinam], Paramaribo, Aug 1892 
BMNH). 

Additional specimens examined: BRITISH GUIANA: Bartica 
District, Kartabo, 2 Apr 1926 (Id") (AMNH). COLOMBIA: Dept. 
Valle, Anchicaya, 600 m, 76° 53'W, 3° 33'N, 20-24 Jan 1992 (16), J. B. 
Sullivan (USNM). COSTA RICA: Cartago Province: Turrialba, 13-17 
Mar 1965 (19), S. S. & W. D. Duckworth (USNM). Heredia Province: 
Finca La Selva, Puerto Viejo de Sarapiqui, 50 m, 6-9 Mar 1985 (16), 
D. Janzen & W. Hallwachs (USNM); La Selva Biological Station, 
10"26'N, 84°0rw, Jan 1998, INBio-OET J. Powell (UCB),18 Feb 
2003 (19), 27 Feb 2003 (lc5), 28 Feb 2003 (16), D. Wagner (INBio); 
Estacion Magassay, P.N. Braulio Carrillo, 200 m, Feb 1991 (19), M. 
Barrelier. Limon Province: Rio Sardinas. R.N.F.S. Barra del Colorado, 
18-29 Feb 1993 (16), F. Araya; 30 km N Cariari, Sector Cocori, 100 



m, Dec 1993 (16), Nov 1993 (Id"), E. Rojas; Finca de E. Rojas, Sector 
Cerro Cocori, 150 m, Sep 1993 (16), Apr 1991 (16), Aug 1991 (16), E. 
Rojas; Cerro Tortuguero, P.N. Tortuguero, 100 m, Apr 1989 (lei*), R. 
Aguilar & J. Solano. Jan 1993 (Id*), R. Delgado. Puntarenas Province: 
Estacion Esquinas. Peninsula de Oso, 0-200 m, Feb 1993 (46), Sep 
1993 (26), Oct 1993 (Id*), Aug 1993 (16, 19), May 1993 (16), J. 
Quesada, Feb 1993 (26), Mar 1994 (16), 1 Apr 1993 (1<J) M. Segura, 
Jan 1993 (16), G. Fonseca; Est. Esquinas, Peninsula de Osa, 200 m, 
Aug 1993 (16), J. Quesada (INBio); Bosque Esquinas, Peninsula de 
Oso, 200 m, Jan 1993 (Id"), Apr 1993 (26), J. Quesada, Mar 1994 (16), 
M. Segura; Albergue Cerro de Oro, 150 m, 30 Aug 1995 (16), L. 
Angulo; Estacion Sirena, P.N. Corcovado, 0-100 m, Jan 1993 (16*), 
Nov 1989 (16), Jul 1991 (Id), Jun 1990 (Id*), Jun 1993 (Id*), G. 
Fonseca, 1-19 Aug 1980 (Id*), 10-12 Aug 1980 (2d"), 5-11 Jan 1981 
(Id*), D. Janzen & W. Hallwachs, Aug 1991 (2d"), J. C. Saborio; Golfito, 
R.V.S. Golfito, Sector El Tajo, 15 May 2002 (Id*), M. Moraga; Rancho 
Quemado, Peninsula de Oso, 200 m, Oct 1991 (Id*), Oct 1991 (Id*), 
Nov 1990 (Id*), F. Quesada; Cerro de Oro, 200 m. 26-30 May 1995 
(Id*), E. Phillips (all INBio). Unknown Province: Sixola River, [no 
date] (Id*) (USNM). ECUADOR: Pichincha, Tinalandia, 16 km E 
Santo Domingo de los Colorados, 600 m, 5-11 May 1990 (Id*), R. 
Leuschner (USNM). FRENCH GUIANA: St. Jean, Maroni, [no date] 
(4d*), Wm. Schaus (USNM), [no mondi] 1926 (d, paralectotype), 
LeM[oult] (USNM); Piste de la Montagne des Singes, km 10, 5°05'N, 
52°42'W, 8 Jan 1985 (Id*), J.-F. Landry (USNM); Godebert-Maroni, 
[no date] (Id*), Collection Le Moult (USNM); Saint-Jean du Maroni, 
[no date] (19), Janvier (USNM). PANAMA: Canal Zone: Barro 
Colorado Island, 1-9 May 1964 (56), 25-28 Mar 1965 (2d*), W. D. & S. 
S. Duckworth (USNM), 11 Mar 1941 (Id"), J. Zetek. [no date] (d*), J. 
Zetek (USNM), 21 Mar 1933 (Id*) (AMNH), 19-22 Jul 1951 (Id"), R. 
M. Laughlin (AMNH), 14 Feb 1936 (Id") (AMNH); Navy Res. nr. 
Gamboa, 29 Mar 1965 (Id*), S. S. & W. D. Duckworth (USNM). 

Discussion 

In most tortricids, the setae of the papillae anales 
arise from variably sized, papillose protuberances. In A. 
lu/drogramma and A. pharata these protuberances are 
present only around the perimeter of the papillae 
anales, with most of the papillae anales bearing short, 
thin setae from weakly depressed pits bordered by a 
straight or curved row of microtrichia. This unusual 
arrangement of setae also is present in A. petalocrossa 
and is suspected to occur in A. aporema — diese four 
species are all similar in size, forewing markings, and 
structures of the male genitalia. These features are 
easily observed using a dissecting microscope because 
of the large size of the moths. In Auratonota dominica 
Brown there is a similar configuration of setae but at a 
much smaller scale, requiring observation using a 
compound scojae. Similar arrangements of setae appear 
to be lacking in other Chlidanotini genera examined, 
leading to the possibility' that it is a feature unique to 
Auratonota. 

The structure of the frenulum in the female of A. 
pharata, with two distinct bristles clearlv separated at 
their base, is somewhat unusual in Tortricidae where 
the female frenulum typically consists of three brisdes, 
usually coalesced basally. A two-bristled frenulum was 
hypothesized by Komai (1999) to represent a 
syanpomorphv for Stropliedra Herrich-Schaffer and 



146 



Journal of the Lepidopterists' Society 





2.0 mm 




FIGS. 1-2. Adults of Auratonota. 1, A. hydrogramina, 2, A. pharata. 

Figs. 3-4. Male genitalia of Auratonota, with valve spread and aedeagus remove (inset of uncus). 3, A. hydrogramma, 4, A. pharata. 



Volume 60, Number 3 



147 



mS^ 



2.0 mm 







i 



Fics. 5-6. Female genitalia of Auratonota, with inset of details of papillae anales. 5, A. pharata, 6. .A. hydrogran 



148 



Journal of the Lepidopterists' Society 



Anclrioplecta Obraztsov (Grapholitini), and more 
recently Brown and Baixeras (2006) discussed its 
distribution among species of several genera of 
Grapholitini. However, to my knowledge it previously 
has not been reported in Chlidanotinae. A survey of 
various (but not all) genera within that subfamily 
revealed that all Hilarographini and Chlidanotini 
examined have a female frenulum that consists of two 
bristles, potentially representing an additional 
synapomorphy for that putative sister-group pair. Its 
distribution is less consistent within Polyorthini where 
females of Polyortha Dognin, Ardeutica Meyrick, 
Lopharcha Diakonoif, and Cnephasitis Razowski have 
two bristles and those of Isotrias Meyrick, Olindia 
Guenee, Chlorortha Razowski, Ebodina Diakonoff, and 
Lypothora Razowski have three; it is variable among 
females of Histura Razowski. 

Acknowledgements 

I thank the following for allowing me to examine material in 
their care: Eugenie Phillips (formerly INBio), Jerry Powell 
(UCB), Kevin Tuck (BMNH), and Randall Schuh (AMNH). I 
thank Marie Metz, USDA, Systematic Entomology Laboratory, 
Washington, D.C., for preparing the illustrations and plates. The 
following provided helpful reviews of the manuscript: Ronald 
Ochoa, USDA, Systematic Entomology Laboratory, Beltsville, 
Laryland; Thomas Henry, USDA, Systematic Entomology Lab- 
orator)', National Museum of Natural History, Washington, 
D.C., USA; Jozef Razowski, Polish Academy of Sciences, Insti- 
tute of Systematic Zoology, Krakow, Poland; Richard Brown, 
Mississippi State University, Mississippi State, USA; and Jerry 
Powell, University of California, Berkeley, California, USA. 
Field work in Costa Rica was supported by NSF grant 
LTLSI/ALAS IV, a long-term, large-scale inventory of rainforest 
arthropods, through John Longino, Evergreen State University, 
Olympia, Washington. 



Literature Cited 

Brown, J. W. 1990. New species and first U.S. record of Auratonota 
(Lepidoptera: Tortricidae). Florida Entomologist 73: 153-157. 

Brown, J. W. 2005. World Catalogue of Insects 5. Lepidoptera: Tortri- 
cidae. Apollo Books. 741 pp. 

Brown, J. W. & J. Baixeras. 2006. Macrocydia divergens, a new genus 
and species of Grapholitini (Lepidoptera: Tortricidae: Olethreuti- 
nae) from Central America. Zootaxa 1200: 1-6. 

Brown, J. W. & J. A. Powell. 1991. Systematics of the Chrysoxena 
group of genera (Lepidoptera: Tortricidae: Euliini). University of 
California Publication Entomology 111. 87 pp. + figs. 

Clarke, J. F. G. 1958. Catalogue of the Type Specimens of Microlepi- 
doptera in the British Museum (Natural History) Described by 
Edward Meyrick, Volume 3. Trustees of the British Museum, 
London. 600 pp. 

Horak, M. 19S4. Assessment of taxonomically significant structures 
in Tortricinae (Lep., Tortricidae). Mitteilungen der Schweiz- 
erischen Entomologischen Gesellschaft 57: 3 64. 

Komai, F. 1999. A taxonomic review of the genus Grapholita and 
allied genera (Lepidoptera: Tortricidae) in the Palaearctic region. 
Entomologica Scandinavica Supplement 55: 1-226. 

Meyrick, E. 1912. Descriptions of South American Microlepidoptera. 
Transactions of the Entomological Society of London 1911: 
673-718. 

Powell, J. A., J. Razowski. and J. W. Brown. 1995. Tortricidae: Tortric- 
inae, Chlidanotinae, pp. 138-151. In: Heppner, J. B. (ed.), Atlas 
of Neotropical Lepidoptera, Checklist Part II: Hyblaeoidea - 
Pyraloidea - Tortricoidea. Association for Tropical Lepidoptera, 
Scientific Publishers. Gainesville, Florida. 

Razowski, J. 1987. Neotropical Chlidanotini (Lepidoptera: Tortrici- 
dae). Bulletin of die Polish Academy of Sciences, Biological Sci- 
ences 35: 61-71. 

Razowski, J. and V. O. Becker. 2000. A review of the New World 
Chlidanotini (Lepidoptera: Tortricidae). Revista Brasileira de Zo- 
ologial6(1999): 1149-1182. 

Received for publication 8 September 2005; revised and accepted 13 
April 2006 



Volume 60, Number 3 



149 



Journal of the Lepidopterists' Society 
60(3), 2006, 149-155 

A REVISION OF PSOLOPTERA BUTLER, INCLUDING A REDESCRIPTION OF ITS KNOWN SPECIES 

(ARCTIIDAE: ARCTIINAE: EUCHROMIINI) 

Rebecca B. Simmons 

Dept. of Biology, University of North Dakota, P. O. Box 9019, Grand Forks, ND 58202, rebecca.simmons@und.nodak.edu 



ABSTRACT. Psoloptera Butler, a genus of three species within Euchromiini, was previously described based on wing venation and overall 
appearance, resulting in a polvphvletie assemblage. Here, species of Psoloptera are redescribed and illustrated. One species, P. aurifera (Her- 
rich-Schaffer), is transferred to Calonotos Hiibner as a new combination. Comparisons of male and female genitalia of Psoloptera with other 
historically associated genera refute previous hypodieses on euchromune relationships. 

Additional key words: Neotropical fauna, taxonomy, mimicry 



The tribe Euchromiini (Lepidoptera: Arctiidae: 
Arctiinae) consists of highly specialized mimics of 
Hvmenoptera, Diptera, and Coleoptera. This 
specialized mimicry has confounded efforts to classify 
these animals; distantly related species may mimic the 
same model, resulting in a convergence in appearance. 
Historically, classifications were formed based on overall 
wing venation and similar coloration. This practice often 
results in polvphvletie genera. One such genus is 
Psoloptera Buder (Figs. 1-3). 

Buder (1876) erected Psoloptera, and originally 
placed two species in the genus, P. thoracia (Walker) 
and P. leucosticta (Hiibner). Later, Schaus (1894) 
described P. ba.sifulva, citing that it was allied to P. 
thoracia (Walker), the type of Psoloptera. Hampson 
(1898) placed P. aurifera (Herrich-Schaffer) in 
Psoloptera because of similar overall appearance and 
wing venation. 

Butler (1876) placed Psoloptera in the subfamily 
Euchromiinae (= Euchromiini of Jacobson & Weller 
2002), and stated that it was closely allied to Calonotos 
Hiibner and Amijcles Herrich-Schaffer (= Pompiliodes 
Hampson 1898; = Sphecosoma Simmons & Weller in 
press), but distinct because of its plumose antennae and 
different hindwing venation. Hampson (1898), in his 
phylogeny of the Syntominae, placed Psoloptera as 
ancestral to Metaloba Hampson, and sister to Calonotos, 
Chrysocale Walker, Microgyria Butler, Mystrocneme 
Herrich-Schaffer, Orcynia Walker, Paramya Druce ( = 
Methysia Butler; Simmons & Weller in press), Saurita 
Herrich-Schaffer, and Scena Walker. Forbes (1939a) 
allied Psoloptera to Saurita based on wing venation, and 
noted that the genera differed in the branching of the 
radial veins. Though a consensus of these views 
indicates a close relationship with Calonotos, this 
assertion has never been examined using any characters 
other than external features. 

This paper redescribes Psoloptera and revises its 
composition by transferring one species to Calonotos. 
The individual species are diagnosed and figured. The 



genitalia ol both sexes of all Psoloptera species are 
figured for the first time here as well. Relationships of 
Psoloptera and other euchromiines are discussed. 




Fig. 1. Male P. basifulva Schaus, tvpe specimen (USNM). 

Fig. 2. Adult P. leucosticta (Hiibner), lectotxpe specimen 

(USNM). Fig. 3. Male P. thoracia (Walker), tvpe specimen 
(BMNH). 

Materials and Methods 

Standard genitalic and whole-bodv dissections were 
performed (Winter 2000). Bodies were softened in 
warm 10% KOH for 5-15 minutes and dien cleaned 
(scales and viscera removed) in several rinses of 30-40% 



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Journal of the Lepidopterists' Society 



ethanol. Structures were stained with chlorazole black E 
(Sigma, St. Louis, MO) dissolved in distilled water 
(saturated). Specimens were viewed in 30^0% ethanol. 
Wings were bleached and then neutralized with dilute 
acetic acid. They were then rinsed in distilled water, and 
stained overnight with Eosin Y (1% in distilled water; 
Fisher Scientific, Pittsburgh, PA). Permanent slide 
mounts of wings, abdominal pelts, appendages, genitalia 
and thoraces were made with Euparol (Bioquip, Rancho 
Dominguez, CA). 

Male-female pairs of each species of Psoloptera 
(sensu Zerny 1912) were dissected to describe genitalia. 
These species included P. thoracia (Walker), P. aurifera 
(Herrich-Schaffer), P. basifulva Schaus, and P. 
leucosticta Hiibner. Camera lucida drawings were made 
from these preparations. Forewing measurements were 
made from specimens representing each species of 
Psoloptera. Type specimens for P. aurifera (BMNH), P. 
basifulva (USNM), and P. thoracia (BMNH) were 
examined to verify species identifications. 

Genera previously associated with Psoloptera were 
examined to identify putative sister taxa. Male and 
female genitalia were examined for Calonotos phlegmon 
(Cramer), Chnjsocale principalis (Walker), Metaloba 
argante (Druce), Methysia notabilis (Walker), Orcynia 
calcarata Walker, Saurita cassandra (L.), Scena styx 
(Walker), and Spliccosoma aliena (Walker). 

Terminology for abdominal and genital morphology 
follows Klots (1970) and Forbes (1939b). Collections 
consulted include BMNH, the Natural Histoiy 
Museum, London (M. Scoble), and NMNH, National 
Museum of Natural Histoiy, Smithsonian Institution, 
Washington, D.C. (D. Harvey). 

Results and Discussion 

Species composition. Psoloptera basifulva, P. 
leucosticta, and P. thoracia are medium-sized black 
moths with lateral white spots on the first abdominal 
segment, which may mimic a wasp waist (Weller et al. 
2000). Upon examination of Psoloptera species sensu 
Hampson, I found that P basifulva, P. leucosticta, and P. 
thoracia can be distinguished from P aurifera by both 
genitalic and nongenitalic characters. Psoloptera 
aurifera is colored metallic green and is larger than die 
other species of Psoloptera. Further, P. aurifera males 
have an enlarged juxta bearing large spines, bilobed 
valves, and a trilobed uncus (Fig. 4). These characters 
are not shared with the other members of Psoloptera 
(see following species descriptions), but are diagnostic 
for Calonotos males. Based on these observations, I 
propose the following change: 

P. aurifera (Herrich-Schaffer), 1854 = Calonotos 
aurifera (Herrich-Schaffer), new combination 



Phylogenetie placement. In Euchromiini, male 
genitalia are useful for assigning species to genera, 
grouping genera, and for verification of tribal affiliation. 
Psoloptera shares genital features with some other 
euchromiines: projections on the tegumen, slightly 
bilobed valves, and enlarged juxta. Calonotos phlegmon, 
Chn/socale principalis, and Metaloba argante share an 
enlarged juxta with spines, as in Fig. 4. This character is 
also present in Macrocneme Hiibner and certainly 
represents a synapomorphy for this group of genera 
(Deitz 1994). Methysia notabilis appears to have highly 
specialized genitalia with trilobed valves and a spirelike 
juxta (figured in Simmons & Weller in press). Males of 
Saurita cassandra, Scena styx, and Sphecosoma aliena 
do not have genitalia similar to Psoloptera males (not 
figured). The valves of Orcynia are bilobed, while diose 
of Psoloptera are unilobed; however, Orcynia calcarata 
has projections on the tegumen similar to those of 
Psoloptera and its juxta is slightly butterflv-shaped. 




Fig. 4. Calonotos aurifera (Herrich-Schaffer). male genitalia. 
1 = juxta, U = uncus, V = valve. 



As in other Lepidoptera, the female genitalia of 
Euchromiini tend to be more conserved than those of 
the males. Psoloptera females have genitalia with two 
signa placed opposite each other on the corpus bursae 
and an accessoiy bursa originating from the ductus 
bursae (Figs. 8, 11, 14). These traits are shared with 
other euchromiines examined here including Calonotos 
phlegmon, Chnjsocale principalis, Metaloba argante, 
and Saurita cassandra. Based on tirese observations, 
Butler's (1876) placement of Psoloptera in Euchromiini 



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151 



is justified. Metaloba seems to be related to Calonotos, 
Chrysocale, and Macrocneme because of the presence 
of a knob-shaped projection on the antevagellar plate, 
which is not found in Psoloptera. The sister taxon of 
Psoloptera is probably not Saurita, as proposed by 
Forbes (1939a) or Calonotos (Buder 1876, Hampson 
1898). Female Orcynia possess a ridged bursae, like 
that of P. basijulva (Fig. 8), but the signa differ. 
Although the male and female genitalia are not identical 
to diose of Psoloptera, Orcynia seems to be the most 
likely candidate for the sister taxon. This relationship 
remains open to further exploration with other data, 
such as molecular or larval characters. 

Taxonomy. Here, I redescribe Psoloptera and 
provide illustrations of the habitus, wing venation, and 
male and female genitalia of its three species. 

Psoloptera Butler, 1876 

Psoloptera Buder, 1876: 369. 

Type: Euchromia thoracia Walker 1854: 243, by 
original designation. 



Ductus bursae (DB) short, membranous. Corpus bursae (CB) with 
two patches of spinose signa (S) located opposite one another. 




Diagnosis. This genus superficially resembles many 
other euchromiine and ctenuchine genera (Figs. 1-3). 
The male genitalia of Psoloptera, especially the 
ornamented, V- or butterfly-shaped juxta is distinctive 
widiin Ctenuchini and Euchromiini. The highly 
sclerotized, curved uncus is also unique to Psoloptera. 

Adult habitus (Figs. 1-3). Wings entirely Mack, or with scarlet at 
the base. Male forewing length 12—17 mm (average = 14.5 mm, SD = 
1.2 mm, n = 20); female forewing length 12-17 mm (average = 14.9 
mm, SD = 1.5 mm, n = 11). Antennae black. Ground color of head 
and thorax black or red; abdomen black with paired lateral spots on 
first abdominal segment. 

Head and thorax. Antennae biserrate and ciliate in males: filiform 
and ciliate in females; ocelli present with melanized outer ring. 
Proboscis longer than head. Epiphyses and tibial spurs short and 
smooth. Tarsal claws simple. 

Wings. Forewing venation (Fig. 5A): Sc sinuous, extending 3/4 
of the costa. R, and R, branched, arising from discal cell. R stalked 
with R 5 arising closer to cell than R . Mj arising from the cell; M., 
arising from die cell medially between M } and M„. M, arising from the 
base of the cell. CuA, and CuA„ widely separated; A, present. 
Hindwing venation (Fig. 5B): Sc + R, absent. Rs and M, connate. 
Discal cell cross vein asymmetrical, V-shaped. M, 3 fused. CuAj and 
CuA, stalked, branching close to wing margin. A, and A, present. 

Abdomen. In both sexes, second stemite with short, stubby 
apodemes. Structural modifications for wasp waist absent (Weller et 
at. 2000). Male lacking androconia. 

Genitalia. Males (Figs. 6, 7, 9, 10, 12, 13): Tegumen bearing 
curved projections (TP) (Figs. 6, 9, 12), one on either side of uncus; 
uncus strongly sclerotized, hooked, setose. Valve (V) setose, 
spindlelike or slightly bilobed. Juxta (J) enlarged, V- or butterfly- 
shaped. Aedagus with sclerotized vesicular region (SP) (Fig. 7), 
cornuti (C) or both (Fig. 10), depending on the species. 

Females (Figs. 8, 11, 14): Papillae anales (PA) laterally flattened; 
membrane surrounding ovipore folded; posterior apophyses (PP) long 
and narrow; anterior apophyses (AP) long and narrow or reduced and 
thickened; dorsal pheromone glands reduced. Seventh sternite (VII) 
unmodified; antevagellar plate unmodified or bearing crescent-like, 
heavily sclerotized lip; ostium bursae symmetrical or assymetrical. 



Fig. 5. Wing venation of Psoloptera Butler. A. Forewing, B. 
Hindwing. A = anal vein, CuA = cubital vein, M = medial vein, 
R = radial vein, Rs = radial sector, Sc = subcosta, 1-5 = vein 
number. 

Appendix bursa (AB) from ductus bursae, irregular in shape, lacking 
ornamentation. Ductus seminalis (DS) from appendix bursa. 

Discussion. Species of Psoloptera are easily 
distinguished from each other by habitus and male and 
female genitalia. The larval host plants and stages are 
not known. Tymbal organs are present, indicating that 
ultrasound is utilized lor either defense or intraspecific 
communication; they do not appear to be sexually 
dimorphic. 

Psoloptera basifulva Schaus. 1894 

(Figs. 1, 6, 7, 8) 

Psoloptera basifulva Schaus, 1894. Proc. Zool. Soc. 
Lond. 1894: 225. 

Psoloptera thoracia Druce, 1884. Biol. Centr.-Am.. 
Het. 1: 46, preoccupied by Psoloptera thoracia Walker 
(1854). 

Diagnosis. Though P. basijulva resembles P. 
thoracia, P. basifulva has orange-red patches at the base 
of the fore- and hindwings (Fig. 1). These patches are 
lacking in P. thoracia (Fig. 3). Psoloptera basifulva also 
has an orange-red diorax and head: the head and thorax 

are black in P. leucosticta (Fig. 2). 

Description. Medium-sized, black modis that have red heads and 
thoraces (Fig. 1). 

Male. Head Red; antenna black, biserrate; frons and labial palpus 
black. 

Thorax. Patagium, meso-, metathorax orange-red; fore, mid, and 



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Journal of the Lepidopterists' Society 



hindlegs black with white spots on coxae and femur; tarsi black. 

Wings. Forewing. Length = 14-16 mm (average = 15 mm, SD = 1 
mm, n = 3). Black with basal scarlet area. Hindwing. Black, costal area 
slightly hyaline, basal scarlet area present. 

Abdomen. Black with white spots on the first abdominal segment. 

Genitalia (Figs. 6 & 7). Tegumen heavily sclerotized, bearing 
spirelike projections (TP), one on each side of uncus; uncus curved, 
setose; saccus square-shaped; valve unilobed, spatulate, setose; juxta 
enlarged, butterfly-shaped; base of phallus rounded; vesica with 
sclerotized area. 

Female. As in male, except antennae filiform and ciliate; forewing 
length = 12-17 mm (average = 14.4 mm, SD = 1.7, n = 7). 

Genitalia (Fig. 8). Papillae anales, posterior apophyses unmodified; 
anterior apophyses greatly reduced, nublike; S7 unmodified; 
antevagellar plate U-shaped, highly sclerotized; asymmetrical; ductus 
bursae sclerotized tube; corpus bursae membranous, slightly ridged, 
bearing two patches of signa near accessory bursa; ductus seminalis 
from middle of accessory bursa. 

Type material. Psoloptera basifulva Schaus. The holotype male 
(USNM) is labeled: Peru; 520; not [? text unreadable]; Collection Wm 
Schaus. Type locality: Peru (Fig. 1). 

Psoloptera thoracia Druce. The holotype male (BMNH) is labeled: 
Panama: Chiriqui; Godman-Salvin Coll. Tvpe locality; Panama. 

Specimens examined. BRAZIL: Amazones: Villa Franca 
(BMNH: 2d*, 29); BRITISH HONDURAS: Punta Gorda (BMNH: 
16, 59) ; No Data (BMNH: 19); COSTA RICA: Guapiles (USNM: 
19); San Mateo (BMNH: 19), Tuis (USNM: 16); GUATEMALA: 
Cayuga (USNM: 86, 69; BMNH: 26, 29); HONDURAS: Cambre 



(BMNH: 36, 19); MEXICO: Tabasco (BMNH: 16, 29): Teapa 
(USNM: 19); PANAMA: Bugaba (BMNH: 26); Chiriqui (BMNH: 
16); NICARAGUA: No Data: Mobile 752 on a ship (USNM: 19); 
PERU: No Data (USNM: 19); VENEZUELA: Palma Sola 

(BMNH: 16); NO DATA: (USNM: 26). 

Psoloptera leucosticta (Hiibner, 1827) 
(Figs. 2, 9, 10, 11) 
Glaucopis leucosticta Hiibner, 1827. Samml. Exot. 
Schmett. 1: t. 162. 

Psoloptera leucosticta (Hiibner) Hampson 1898. Cat. 
Lep. Phal. 1: 285. 

Diagnosis. Psoloptera leucosticta, unlike its 
congeners, does not have any red markings on its head 
or thorax (Fig. 2). 

Description. Medium-sized, black to purplish-black moths (Fig. 
2). 

Male. Head. Black; antenna black, biserrate and ciliate; palpus 
black; vertex and frons with white spots. 

Thorax. Patagium black with lateral white spot; mesothorax black; 
metathorax black; fore, mid, and hind legs black with white spots on 
tire coxa. 

Wings. Forewing. Length = 13-16 mm (average = 14.4 mm, SD = 
1.1 mm, n = 8). Black, with two white spots at base. Hindwing. 




Fig. 6. Psoloptera basifulva Schaus, male genitalia, ventral view. J = juxta, S = saccus, TP = tegumenal process, U = uncus, V = 
valve. Fig. 7. Psoloptera basifulva Schaus, aedagus, lateral view. PB = phallic base, SP = sclerotized plate, V = vesica. Fig. 8. 
Psoloptera basifulva Schaus, female genitalia, ventral view. VII = seventh sternite, AB = accessory bursa, AP = anterior apophysis, 
AW = antevagellar plate, CB = coipus bursae, DB = ductus bursae, DS = ductus seminalis, O = ostium, PA = papillae anales, PP 
= posterior apophysis, S = signa. 



Volume 60, Number 3 



153 



Ground color black; lighter shade of black in costal area. 

Abdomen. Black. Dorsal lateral white spots on first abdominal 
segment. Medial white, faint patch on venter of first three abdominal 
segments. 

Genitalia (Figs. 9 & 10). Tegumen heavily sclerotized, bearing 
spirelike projections, one on each side of uncus; uncus curved, setose; 
saccus rounded; valve terminating in blunt projection, setose; juxta 
elongate and narrow, V-shaped; phallus blunt at base; vesica bearing 
single patch of large comuti apicallv. 

Female. As in male, except antennae filiform and ciliate; forewing 
length = 15-16 mm (average = 15.5 mm, SD = 0.7 mm, n = 2). 

Genitalia (Fig. 11). Papillae anales, posterior and anterior 
apophyses unmodified; SS unmodified; antevagellar plate 
asymmetrical, without ornamentation; ductus bursae membranous; 
corpus bursae membranous, bearing two patches of signa; accessory 
bursa membranous, from ductus bursae; ductus seminalis from 
accessory bursa. 

Type material. The type of Glaucopis leucosticta Hiibner is 
apparendv lost. Tvpe locality: Venezuela. The lectotype male (USNM, 
here designated) is labeled: 60 m. up Maroni River, Psoloptera 
leucosticta Hbn. from BM. Collection Wm Schaus. The lectotype is 
designated to ensure nomenclatural stability in this genus. 

Specimens examined. BRAZIL: Cayenne (USNM: 19); 
BRITISH GUIANA: Rio Potaro. Tumatumari (USNM: 4d); 
MEXICO: No Data (USNM: 2d, 1?). VENEZEULA: Maroni 
River (USNM: 16", 19). NO DATA (USNM: 2d', 69). 



Psoloptera thoracica (Walker, 1854) 

(Figs. 3, 12, 13, 14) 

Euchromia thoracica Walker, 1854. List Lep. Ins. Br. 
Mus. 1: 243. 

Psoloptera thoracica Butler, 1876. J. Linn. Soc. Lond. 
Zool 12: 369. 

Diagnosis. Bright scarlet coloration is restricted to 
the head and thorax in P. thoracia (Fig. 3); in P. 
basifulva, the base of the fore and hind wings are 
orange-red, as well as the thorax and portions of the 
head (Fig. 1). Psoloptera leucosticta lacks red coloration 
altogether (Fig. 2). 

Description. Medium-sized, black moths with red heads and 

thoraces (Fig. 3). 

Male. Head. Vertex black with pair of white spots; galae scarlet; 
antenna black, biserrate and ciliate; palpus black. 

Thorax. Patagium scarlet; mesothorax scarlet with scattered, long 
black scales; metathorax scarlet; fore, mid and hind legs black with 
white spots on coxae. 

Wings. Forewing. Length = 12-17 mm (average = 14.6 mm, SD = 
1.3 mm, n = 9). Black. Hindwing. Costal region gray, grading to black 
at CuAj and CuA,. 

Abdomen. Pair of white lateral spots present on first abdominal 
segment; remainder of abdomen black dorsally; medial white spot 




Fig. 9. Psoloptera leucosticta (Hiibner), male genitalia, ventral view. J = juxta. TP = tegumenal process, U = uncus, V = valve. 
Fig. 10. Psoloptera leucosticta (Hiibner), aedagus, lateral view. C = cornuti, PB = phallic base, V = vesica. Fig. 11. Psoloptera leu- 
costicta (Hiibner), female genitalia, ventral view. VII = seventh sternite, AB = accessory bursa, AP = anterior apophysis, A\~P = an- 
tevagellar plate, CB = corpus bursae, DB = ductus bursae, DS = ductus seminalis, PA = papillae anales, PP = posterior apophysis, 
S = signa. 



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Journal of the Lepidofterists' Society 




Fig. 12. Psoloptera thoracia (Walker), male genitalia, ventral view. J = juxta, TP = tegumenal process, U = uncus, V = valve. Fig. 
13. Psoloptera thoracia (Walker), aedagus, lateral view. C = cornuti, PB = phallic base, SP = sclerotized plate, V = vesica. Fig. 14. 
Psoloptera thoracia (Walker), female genitalia, ventral view. VII = seventh sternite, AB = accessory bursa, AP = anterior apophysis, 
AVP = antevagellar plate, CB = corpus bursae, DB = ductus bursae, DS = ductus seminalis, PA = papillae anales. PP = posterior 
apophysis, S = signa. 



present on venter of first three abdominal segments; remainder black 
ventrally. 

Genitalia (Figs. 12 & 13). Tegumen rounded with elongate 
spindlelike projection on either side of uncus; uncus strongly hooked, 
setose; saccus rounded; valve apically narrowed, ending in rounded 
projections, setose; juxta with butterfly-shaped structure, strongly 
sclerotized; vesica with sclerotized plate and medial patch ol medium- 
sized cornuti. 

Female. As in male, except antennae filiform and ciliate; forewing 
length = 16 mm (average = 16 mm, SD = 0, n = 2). 

Genitalia (Fig. 14). Papillae anales, posterior and anterior 
apophyses unmodified; sternite 7 unmodified; antevagellar plate with 
heavily sclerotized lip, symmetrical; ductus bursae membranous; 
coipus bursae membranous, bearing two patches of signa; accessory 
bursa large, irregular in shape, from ductus bursae; ductus seminalis 
from base of accessory bursa. 

Type material. The holotype male of Euchromia thoracia Walker 
(BMNH) is labeled: Ega, Bates, 51-43. Type locality: Amazones 
[Brazil]. 

Specimens examined. BRAZIL: Amazones: Fonte Boa 
(BMNH: 5d, 39), Pegas (BMNH: Id), Rio Ucayla (BMNH: 2c!), S. de 
Villa Franca (BMNH: 12c!), S. Paulo (BMNH: 19); Cundinmarca: 
Cananche (BMNH: Id'); Ega: (BMNH: 4o*); Humayta (BMNH: 5c?, 
29); Lower Amazon and R. Madeira (BMNH: Id); Rio Maderia: 
Allianca below S. Antonio (BMNH: Id); San Juan: Solhnoens 
(BMNH: Id); S. Paulo de Olivenca (BMNH: 4c!, 59; USNM: 4c!); 
Sao Paulo de Amazones (USNM: 2(5, 19); Teffe (BMNH: 6c! ; 
USNM: 19). COLOMBIA: Caqueta: Rio Orteguaza nr. Rio Peneya 
(USNd: lc!, 19); Chiriguana District: Lake Sapatoza Region 
(BMNH: Id); Llanos of Rio Meta: S. Martin (BMNH: Id); 



Magdalena Valley (BMNH: 19); Ort.: Medina (USNM: 19); R. 
Cantinere Muzo (BMNH: 39); Rio Negro (BMNH: Id); Villaricua 
(USNM: 19). COSTA RICA: Guanacaste: Santa Rosa National Park 
(BMNH: Id, 19); Turrialba (USNM: 2d. 19). ECUADOR: Napo 
Prow: Yasuni Research Station, Rios Tivacuno & Tiputini, 76° 36' W, 
0° 38' S, 250 m (USNM: 29); Sarayacu (BMNH: Id, 19). 
GUATEMALA: No Data (BMNH: 19); PANAMA: Alhajuelo 
(USNM: 19); Barro Colorado Island (USNM: 59); Canal Zone 
(USNM: Id): Corozal (USNM: 2d); Canno Saddle (USNM: 29); La 
Chorrera (BMNH: 29); Matachin (BMNH: 1?); PortoBello 
(USNM: Id, 29); No Data (BMNH: Id; USNM: Id). PERU: 
Amazones: Cavallo-Cocho (BMNH: 29); Pebas: Loreto (BMNH: 
19); Rio Udayali: Contamama (BMNH: 19); Tarapoto (BMNH: Id); 
No Data (BMNH: Id). VENEZUELA: Aroa (USNM: 6d, 59); La 
Cruees Colon. (BMNH: 29); Las Quigas: San Eseban Valley (8d; 
39); Las Quigas nr. San Esteban (BMNH: 3d, 29); Palma Sol 
(BMNH: Id); San Esteban (BMNH: lid, 39); Valencia (BMNH: 
id); Valera (USNM: Id); No Data (BMNH: 2d, 19). NO DATA: 
(BMNH: Id; USNM: Id, 19). 

Acknowledgements 

Revisionary work is impossible without cooperation from col- 
lections and associated curators. I would like to thank Dr. Don- 
ald Harvey (USNM), Mr. Martin Honey (BMNH) and Dr. Mal- 
colm Scoble (BMNH) for access and loans of specimens, as well 
as hospitality during museum visits. I would like to thank Drs. 
William Miller and Susan Weller and Ms. Michelle DaCosta for 
helpful discussion and suggestions during the formation of diis 
manuscript. Ms. DaCosta also provided editorial aid. Funding 



Volume 60, Number 3 



155 



for associated travel was provided by ND-EPSCor and Univer- 
sity of North Dakota, Dept. of Biology. 

Literature Cited 

Butler, A. G. 1876. Notes on the Lepidoptera of the family Zy- 
gaenidae, with descriptions of new genera and species. J. Linn. 
Soc. Lond. Zool. 12: 342-407. 

Dietz. R. E. 1994. Systematics and biology of the genus Macrocneme 
Hiibner (Lepidoptera: Ctenuchidae). University of California 
Press, Berkelev. 

Forbes, W. T. M. 1939a. The Lepidoptera of Barro Colorado Island, 
Panama. Bulletin of die Museum of Comparative Zoology at Har- 
vard College. 85: vii + 974322, 8 plates. 

Forbes, W. T. M. 1939b. The muscles of die lepidopterous male gen- 
italia. Arm. Entomol Soc. Amer. 32: 1-10. 

Hampson, G. F. 1898. Syntomidae. Catalogue Lepid. Phalaenae in the 
British Museum. Vol. I. Trustees of the British Museum, London. 

JACOBSON, N. J. & S. J. Weller. 2002. A cladistic study of the tiger 
modi family Arctiidae (Noctuoidea) based on larval and adult 
morphology. Thomas Say Monograph Series, Entomol. Soc. 
America. 



Klots, A. B. 1970. Lepidoptera, pp. 115-130. In Tuxen, S. L. (ed.), 
Taxonomist's glossary of genitalia in insects, 2nd ed. Munksgaard, 
Copenhagen. 

Schaus, W. 1894. On new species of Heterocera from Tropical 
America. Proc. Zool. Soc. Lond.: 225-243. 

Simmons, R. B. & S. J. Weller. 2006. Review of the Sphecosoma 
genus group using adult morphology (Lepidoptera: Arctiidae). 
Thomas Sav Monograph Series. Entom. Soc. America, 108 pp.. 
108 figs. 

Walker, F. 1854. Lepidoptera Heterocera, Part 1. List of the speci- 
mens of lepidopterous insects in the collection of the British Mu- 
seum. 278 pp. 

Weller, S. J., R. B. Simmons, R. Boada, & W. E. Conner. 2000. Ab- 
dominal modifications occurring in wasp mimics of the Ctenu- 
chine-Euchromiine clade (Lepidoptera: Arctiidae). Ann. Ento- 
mol. Soc. Am. 93: 920-928. 

Winter, W. D. 2000. Basic techniques for observing and studying 
moths and butterflies, Lepidopterists' Society. 

Received for publication 25 October 2005; revised and accepted 7 July 
2006 



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Journal of the Lepidopterists' Society 



Journal of the Lepidopterists' Society 
60(3), 2006, 156-160 

FOREST TENT CATERPILLAR: MATING, OVIPOSITION, AND ADULT CONGREGATION AT TOWN 
LIGHTS DURING A NORTHERN MINNESOTA OUTBREAK 

William E. Miller 

Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108 email: mille014@umn.edu 



ABSTRACT. Mating and opposition of the forest tent caterpillar, Malacosoma disstria (Hiibner) (Lasioeampidae), were studied in two sites 
=2 ha each at levels 1- 5 m above the ground, and adult congregation at lights was studied in brightly lit town districts. The observations were 
made near or at Ely, St. Louis Co., Minnesota, in the latter years of an outbreak peaking during 1999-2003. Descent of the nomadic larvae from 
the forest canopy to less defoliated vegetative strata for feeding and cocooning later enabled mating to be observed conveniently from the 
ground. Adult activity began or intensified at =5:30 pm CDT, when males vigorously sought mates in foliage harboring cocoons. Mating pairs 
were captured and caged in jars containing a host branchlet for oviposition; mean duration of copulation was 202 min. Captive mated females 
always oviposited the morning after mating; mean duration of oviposition was 88 min. Dispersal behavior always preceded oviposition. Five new 
oviposition hosts were recorded. Naturally deposited egg rings were found most often on branchlets oriented nearer horizontal than vertical. 
Females congregating in brightly lit districts were usually gravid, but seldom oviposited. 



Additional key words: Malacosoma disstria, Lasioeampidae, defoliation. 



In northern Minnesota and elsewhere, the forest tent 
caterpillar, Malacosoma disstria (Hiibner) 
(Lasioeampidae), builds up periodically to outbreak 
densities (Witter 1979). During Minnesota outbreaks, 
quaking aspen, Populus tremuloides Michx. 
(Salicaceae), and most other broad-leaved trees and 
shrubs except red maple, Acer rubrum L. (Aceraceae), 
are severely defoliated. Red maple contains an 
antifeedant (Nicol et al. 1997), but the nomadic larvae 
readily spin cocoons among its leaves. The forest tent 
caterpillar is univoltine, and the larvae hatch in spring at 
host bud-break (Batzer and Morris 1971). 

Many thousands of hectares of forest, primarily 
quaking aspen, were defoliated in the outbreak that 
peaked during 1999-2003 in northern Minnesota when 
this study was done (Minnesota Dept. of Nat. Res. For. 
Ins. Disease Newsletter 1999-2004). Although 
defoliation seldom kills trees, it reduces their wood- 
volume growth, and the larvae are considered a 
nuisance by landowners and outdoor recreationists 
(Duncan et al. 1956). The eastern tent cateipillar, 
Malacosoma americanum (Fabricius), also occurs in 
Minnesota (Stehr and Cook 1968), but mostly in the 
eastern part of the State, and it was not seen in the 
outbreak area. Fitzgerald (1995) provides an exhaustive 
compendium of information for all Malacosoma. 

At low, nonoutbreak densities, forest tent cateipillar 
adults mate high above the ground in the forest canopy; 
during outbreaks they mate on lower, understory 
vegetation. Mating nearer the ground occurs because 
larvae descend to less defoliated strata to find food and 
cocooning sites (Batzer et al. 1995), and males 
concentrate their search for mates where female 
cocoons occur. The larvae require angled surfaces for 



cocooning, and they commonly create cocoon structures 
by drawing and spinning leaves together (illustrated by 
Batzer and Morris 1971 and Fitzgerald 1995, p. 60). In 
this study, many cocoons were spun on herbaceous 
plants of the ground stratum as well as shrubs. The 
downward shifting later enabled mating to be 
conveniently observed from the ground. 

Previous reports touch on topics treated here. 
Bieman and Witter (1983) described field mating 
behavior at low and high levels of mate competition, 
reflecting low and high population densities. Stehr and 
Cook (1968) noted that forest tent cateipillar 
oviposition proceeds helically and in a layer one egg 
deep. Shepherd (1979) reported daily rhythms of male 
and female activity. Hodson (1941) noted adult 
congregation at town lights. As elaborated in the 
discussion section, the present study supplies more or 
different details about these and related topics. 

Materials and Methods 

This study was conducted near and in the town of Ely, 
St. Louis Co., Minnesota. Mating and oviposition were 
observed in two multilayered stands =40 yr old. 
Overstories in these stands were dominated by quaking 
aspen, but scattered examples of other tree species were 
present, in descending order of abundance, balsam 
poplar, Populus balsamifera L. (Salicaceae); red pine, 
Pinus resinosa Ait.; jack pine, P. banksiana Lamb.; 
eastern white pine, P. strobus L. (Pinaceae); white birch, 
Betula papijrifera Marsh. (Betulaceae); and balsam fir, 
Abies balsamea (L.) Mill. (Pinaceae). The understories 
were dominated by willow, Salix sp. (Salicaceae), but 
contained scattered examples of other species, in 
descending order of abundance, red-osier dogwood, 



Volume 60, Number 3 



157 



Cornus stolonifera Michx. (Cornaceae); speckled alder, 
Alnus nigosa (Du Roi) Spreng (Betulaceae); pin cherry, 
Primus pennsijlvanica L. (Rosaceae); beaked hazel, 
Corylus cornuta (Marsh.) (Betulaceae); and American 
cranberry-bush, Viburnum trilobum Marsh. 
(Caprifoliaceae). The vertical range of observation was 
1-5 m above ground, and the sites were each =2 ha. 

Copulating pairs captured in late afternoon were 
caged in wide-mouthed, 1-liter glass jars with gauze- 
covered tops containing an aspen or willow branchlet 
3—6 mm in diameter. Cages were moved in the evening 
to a darkened basement with temperatures constantly at 
17-19°C, which approximated cool, outdoor evening 
temperatures. Cages were returned in the morning to 
shaded outdoor spaces where temperatures ranged 
19-26°C. Observations of copulation and oviposition 
durations were made at =s30-min intervals. 

Gender identification of adults was based on antennal 
rami, which are >2x longer in males than in females. 
Males also are noticeably smaller than females (sample 
measurements given later). Early in the study, active 
males were netted and released after their gender was 
confirmed (n> 20). Females were dissected for 
verification of their gravid or spent condition. Egg 
fertilitv was determined bv the presence of embryos. 

Egg numbers/ring were estimated as described by 
Witter and Kulman (1969), and spumaline — the frothy 
substance that covers egg rings — was removed as these 
authors recommended by brushing with a discarded 
toothbrush. Forewings were removed for accurate 
length measurement. Angles of egg-bearing branchlets 
in the field were referenced to horizontal and were 
estimated with a large protractor. All clock hours refer 
to Central Daylight Time. Statistics were computed 
with SYSTAT (1992) software, except for G ad ., which was 
computed according to Sokal and Rohlf (1981). The 
abbreviation SD refers to standard deviation. 

Mean daily temperatures during the late larval stage 
were computed from daily maxima and minima 
recorded at the Winton meteorological station <5 km 
from Ely (Minnesota State Climatology Office 2005). 

Results 

Mating. The first adults of the flight season were 
males, as the species is protandrous. Throughout daily 
adult activity, starting or intensifying typically =5:30 pm, 
males visible by the hundreds in any directional view 
flew vigorously and constantly in vegetative strata 
containing cocoons. They flew mainly in a zone within 
<0.5 m of tree and shrub canopies, circling, zigzagging, 
alighting and crawling along branchlets before taking 
flight again if they did not find an opportunity to 
copulate. The number of active, searching males around 



foliage that contained cocoons was ~4x that around 
foliage that contained no cocoons. On windy evenings, 
males confined their activity closer to canopies. During 
two cool, rainy, late afternoons widi ambient 
temperatures < 15° C, only a small fraction of the males 
known to be present were active. 

Many female pupae appeared to emit their calling 
pheromone (Struble 1970) before they completed 
eclosion, as up to 6 males often concentrated activity 
around a single cocoon until the female eclosed. Males 
copulated with such females within 1-2 sec after the tip 
of the female abdomen cleared the cocoon. The wings 
of such females inflated during copulation. Most mating 
pairs were found within 10 cm of a cocoon structure 
presumed to be that of the female, and meconium was 
often seen on such structures. Some copulating females 
walked several cm as they were being observed before 
capture, and because of their larger size pulled the 
smaller males along. None of the females observed in 
this study attempted to fly during copulation. Females 
did not always begin calling before completing eclosion. 
The locations of two such females found resting quietlv 
on cocoon structures were marked for continuing 
observation during the evening. At first, captured males 
released within centimeters of these females flew away. 
However, by onset of darkness both females were in 
copula, apparently having eventuallv emitted their 
calling pheromone. The sexes were always positioned 
end to end during copulation. 

A total of 69 pairs were seen in copula during this 
study, of which 48 were successfully caged for further 
observation. Some copulating pairs separated on 
capture, presumably because copulation had just begun 
or was nearly completed. Copulating pairs were seen in 
the field as soon as daily male activity commenced and 
pairs known to be newly copulating were captured as 
late as 9:30 pm. Neither active males nor mating pairs 
were seen during early dawn hours, presumablv 
because of too cool temperatures. Caged pairs remained 
in copula following capture for 150-255 min, averaging 
202 min (n = 10), but it should be noted that beginning 
and ending of copulation was not always preciselv 
timed, and that some durations are subject to an error of 
±30 min. Copulating pairs occurred on all die trees and 
shrubs named earlier, including die conifers. None of 
the conifers had been fed on, but thev provided 
cocooning sites under conditions of extensive foodplant 
defoliation. Based on forewing lengths — a proxy for 
body size — there was a statistically - significant relation 
between gender body sizes of naturallv mating pairs: 
forewing lengdi averaged 16.8 ± SD 1.42 mm for 
females, and 12.4 ± SD 1.05 mm for dieir mates 
(Pearson's r = 0.49, df = 23, P = 0.013). 



158 



Journal of the Lepidopterists' Society 



Oviposition. As in other species of Malacosoma, the 
forest tent caterpillar deposits its eggs in one batch 
during one oviposition event unless disturbed during 
the process. Of the 48 captive mated pairs, 88% of the 
females began ovipositing the day after copulation, most 
starting in early morning. The earliest an oviposition was 
completed was 7:05 am. Other females began 
ovipositing later, the latest at noon. If not ovipositing the 
day after copulation, captive females failed to do so 
entirely. Duration of individual oviposition in captivity 
monitored during daily hours of observation, which 
began at 7:00 am, ranged 65-134 min, averaging 88 min 
(n = 11). Shorter and longer oviposition durations were 
associated with smaller and larger egg rings, which 
imply smaller and larger females and cooler and warmer 
ambient temperatures. 

Just before beginning to oviposit, captive females 
always exhibited dispersal behavior. For 5-15 min they 
flew upward and around inside their cages. Further, 
naturally deposited egg rings were never observed near 
cocoon structures (n = 74). After attempting to disperse, 
females became quiet, positioning themselves more or 
less diagonally on the caged branchlet head up with 
wings partly spread, and extended abdomenal tips 
around the branchlet nearly 180° before starting to 
oviposit. As they oviposited, they gradually moved 
around and down the branchlet, coating eggs and the 
egg ring surface with spumaline. The second turn of the 
egg ring was aligned with the first so that the top margin 
of rings seldom had gaps (Fig. Id). By the completion of 
oviposition, several turns of the egg ring had been 
deposited and the females were standing on the egg ring 
surface. Directions of female progression around 
branchlets during oviposition were clockwise in 12 cases 
(60%) and counterclockwise in 8 (40%), with only 3 
reversals during the process, 2 from clockwise to 



counterclockwise, and 1 the opposite. Within minutes of 
completing oviposition females flew vigorously again as 
before oviposition. 

Whether termed 'mass' or 'ring', eggs were deposited 
as a continuous helical band each turn of which was 
tightly fitted with the previous turn, the width of the 
band generally diminishing downward (Fig. 1). When 
egg ring deposition was observed closely in captivity, 
seams in the spumaline covering between turns of the 
band were usually faintly evident as slight depressions, 
sometimes as ridges. Depending on female size, 
number of eggs, and branchlet diameter, rings consisted 
of up to five turns (Fig. la, c). After spumaline removal, 
margins between turns were not evident. After two days 
of rainy, windy weather, an increase in irregularly 
deposited eggs was evident in field egg rings 
presumably because oviposition had been interrupted 
by such weather (Fig. le). Spumaline was more or less 
clear when dispensed, but darkened after =24 h. 

For egg rings deposited in captivity, number of 
eggs/ring averaged 285 ± SD 112 and for diose 
deposited naturally, 246 ± SD 66, and the difference 
was not significant (Student's t = 1.20, df = 13.8, 
separate variances, P = 0.25). Based on 5-39 eggs/ring 
dissected in late July and early August, 94% of rings 
contained fertile eggs, with an average of 94% of 
dissected eggs/ring fertile (n = 14). Corresponding 
values for eggs collected in the field were 100 and 98% 
(n = 39). These differences between captive and field 
eggs were close and inferred to be nonsignificant 
statistically. 

Species on which naturally deposited egg rings were 
found included quaking aspen (76%), willow (12%), 
with the remainder (12%) on red osier dogwood, balsam 
poplar, beaked hazel, speckled alder, pin cherry, and 
American cranberrybush (n = 72). 







FlG. 1. Delineated margins of the band or bands of eggs in egg rings of the forest tent caterpillar on aspen branchlets. Delineation 
is based on seams in corresponding spumaline deposition. In the absence of female disturbance, oviposition proceeds in a down- 
ward helical manner. 



Volume 60, Number 3 



159 



Branchlets on which egg rings were found in the field 
ranged 2-6 mm in diam. Orientation of egg-bearing 
branchlets in the field tended to be nearer horizontal 
than vertical. All 5 samples of 28-74 branchlets 
containing naturally deposited egg rings in late July and 
early August were distributed similarly in each sample 
among three equally progressing angle classes. When 
pooled, 50% were on branchlets angled 0-30° relative to 
horizontal, 18% on those 30-60°, and 32% on those 
60-90° (n = 291), with departure from a no-preference 
distribution highly significant (G. i(1 . = 42.9, df = 2, P < 
0.001). The distribution through 0-90° thus was 
backward-J-shaped. 

Adult congregation at lights. Like many moths, forest 
tent caterpillar adults are photopositive. During flight 
periods, thousands of adults congregated near 
streetlights an d on brightlv fit exterior walls of buildings 

o - o J o 

in towns in the outbreak area. In Ely, a few congregated 
males had frayed wings, indicating much flight history, 
but the females showed little or no wing wear. The 
source was almost certainly infested quaking aspen that 
were <1 km distant. Once congregated, the adults 
remained notably inactive, and seemed to be present 
night and day until dying or becoming prey of birds and 
bats. Both genders seemed equally represented, but 
copulating pairs were seldom seen. Eggs were 
sometimes seen on building walls, but were scarce 
relative to the numbers of females present. 

Congregated females collected at various times of day 
for close examination came from the 400-600 and 900 
blocks of Sheridan Street in downtown Elv. Their yearly 
forewing lengdis during 2000-2003 averaged 20.0, 18.9, 
19.1, and 18.9 mm, respectively, and appeared to 
indicate temporally decreasing body size (F = 13.5, df = 
3, 166; P < 0.001). Corresponding percentages that 
were fully gravid were 96, 90, 97 and 58. 

Discussion 

Mating and oviposition. The present study indicates 
strongly that the vegetative stratum in which most 
mating occurs is the one containing the most cocoons. 
The high frequency with which males were observed to 
copulate with females at the moment of female eclosion 
is more likely to occur at high than at low population 
densities, and was observed earlier in Minnesota 
(Hodson 1941, Bieman and Witter 1983). Female 
calling pheromone doubtless stimulated such intense 
male activity. Bieman and Witter (1983) also reported 
that some males fingered near empty or parasitized 
cocoons, as well as brown objects, which suggests they 
also use visual cues. 

Typically, females eclosed and copulated in late 
afternoon and evening, and oviposited early the next 



day. The absence of early morning flight activity in this 
study is not surprising for northern forest tent 
caterpillar populations. Shepherd (1979) also reported 
little or no male activity near dawn in Alberta and 
British Columbia, and found that males ceased flying at 
<11°C. Such temperatures were common near dawn in 
the present study. Daily patterns of male activity, female 
eclosion, and evening female calling closely match those 
reported by Shepherd. 

Shepherd (1979) and Bieman and Witter (1983) also 
reported attempted dispersal by newly mated captive 
females. Lack of proximity between naturallv deposited 
egg rings and cocoon structures is further evidence of 
preoviposition dispersal. 

Positive and significant correlation between gender 
body sizes of mating pairs indicates diat the sexes do not 
mate randomly with respect to size. Reasons for diis are 
unknown, but one might speculate that larger males 
out-compete smaller ones to mate with larger females. 
The mean duration of copulation observed in captivity, 
202 min, is near the mean duration of 197 rnin extracted 
from Table 1 in Bieman and Witter ( 1983) for dieir high 
population density. 

Highly variable numbers of eggs/ring averaging 285 
in captivity did not seem to differ significantly from the 
similarly variable mean of 246 observed in the field, nor 
did the apparent fertility means of 94% in captivity 
differ significantly from the corresponding 100 and 98% 
observed in the field. Both higher and lower fertilities 
were recorded earlier in Minnesota (Witter and Kulman 
1972). After close observation confirmed die helical 
pattern of oviposition reported bv Stehr and Cook 
(1968) and revealed corresponding seams in spumaline 
deposition, it became possible to reconstruct the course 
of oviposition in detail (Fig. 1). 

Of naturally deposited egg rings found at studv sites, 
88% were on quaking aspen and willow. The remaining 
12% were on 6 lesser utilized oviposition species, of 
which 5 — red-osier dogwood, speckled alder, balsam 
poplar, beaked hazel, and American eranberrybush — 
were absent from Fitzgerald's (1995) foodplant 
tabulation for the forest tent caterpillar and odier 
Malacosoma. (Flowering dogwood, Cornus florida L.. 
an occasional foodplant of southern populations [Gover 
et al. 1987], was presumably omitted accidentally from 
the tabulation.) The new oviposition host records 
reported here involved no more than 2 egg rings/host, 
and they are probably explainable bv the presence of 
these plants in areas of high forest tent caterpillar 
population density. 

The backward-J-shaped distribution of angles of 
branchlets bearing naturallv deposited egg rings 1-5 m 
above the ground is reported here for the first time. Its 



160 



Journal of the Lepidopterists' Society 



significance, if any, is unclear. One possibility is that 
branchlets nearer horizontal than vertical — where most 
egg rings were deposited — may be more abundant in 
midcrowns especially of the overstory and thus provide 
larvae easier access to the larger foliage volumes at 
midcrown. 

Adult congregation at lights. Hodson (1941) earlier 
reported adult congregation at town lights where the 
nearest source infestations were several kilometers 
distant. In the present study, sources were si km 
distant. Such short-range attraction to lights at high 
population densities does not seem surprising. Female 
movement to lit areas probably occurred during the 
evening after late-aftemoon eclosion and mating. Most 
congregated females were gravid, and while their status 
with respect to mating is unknown, they seem likely to 
have already mated because of the generally assiduous 
male pursuit. Congregation results in little reproduction 
because females do not appear to return to foodplants. 
The paucity of eggs at lights may result from lack of 
suitable oviposition sites (branchlets). The decreasing 
yearly body sizes of congregated females during 
2000-2003 may be explainable by either or both of two 
effects: decreasing diet quality with repeated foodplant 
defoliation, and the generally inverse relation between 
body size and direction of temperature difference 
during late larval development (Miller 2005). During 
late larval development, 10-19 June, in 2000, when 
female forewing length was greatest, daily mean 
temperature averaged a cool 12.4°C, whereas in 
2001-2003, when female forewing length decreased, 
daily mean temperature was higher by an average of 
4.4°C. Less common than adult congregation at lights is 
convective transport of adults in the turbulent air of 
cold fronts for hundreds of kilometers in a few hours as 
Brown (1965) reported in Alberta and elsewhere. The 
population dynamics significance of convective 
transport is unclear as the gravid and mated statuses of 
transported females have not been reported. 

Literature Cited 

Batzer, H. O. and R. C. Morris. 1971. Forest tent caterpillar. U. S. 
Dept. Agric. For. Serv. For. Pest Leaflet 9, 5 pp. 

Batzer, H. O., M. P. Martin, W. J. Mattson and W. E. Miller. 
1995. The forest tent caterpillar in aspen stands: distribution and 
density estimation of four life stages in four vegetation strata. For. 
Sci. 41: 99-121. 

Bieman, D. N. and J. A. Witter. 1983. Mating behavior of Malaco- 
soma disstria at two levels of mate competition. Fla. Entomol. 66: 



272-279. 

Brown, C. E. 1965. Mass transport of forest tent caterpillar moths, 
Malacosoma disstria Hiibner, by a cold front. Can. Entomol. 97: 
1073-1075. 

Duncan, D. P., A. C. Hodson, A. E. Schneider, H. Batzer, R. 
Froelich, D. Meyer and C-J. Shiue. 1956. Influence of the for- 
est tent caterpillar (Malacosoma disstria Hbn.) upon the aspen 
forests of Minnesota. Office of Iron Range Resources and Reha- 
bilitation, St. Paul, Minnesota, 45 pp. 

Fitzgerald, T. D. 1995. The tent caterpillars. Comstock, Idiaca, New 
York. 303 pp. 

Goyer, R. A., G. J. Lenhard, J. D. Smith and R. A. May. 1987. Esti- 
mating the number of eggs per egg mass of the forest tent cater- 
pillar, Malacosoma disstria, on three tree species in the southern 
U. S. J. Entomol. Sci. 22: 188-191. 

Hodson, A. C. 1941. An ecological study of the forest tent caterpillar, 
Malacosoma disstria Hbn., in northern Minnesota. Univ. Minn. 
Agric. Expt. Stn. Tech. Bull. 148, 55 pp. 

Miller, W. E. 2005. Extrinsic effects on fecundity-maternal weight 
relations in capital-breeding Lepidoptera. J. Lepid. Soc. 59: 
143-160. 

Minnesota Department of Natural Resources. Forest Insect and 
Disease Newsletter. 1999-2004. Available online at 
http://www.dnr.state.mn.us/fid/. 

Minnesota State Climatology Office. 2005. Closest station 
climate data retrieval. Available online at 

http://climate.umn.edu/HIDradius/radius.asp. 

Nicol, R. W., J. T. Arnason, B. Helson and M. M. Abou-Zaid. 1997. 
Effect of host and nonhost trees on the growth and development 
of the forest tent caterpillar, Malacosoma disstria (Lepidoptera: 
Lasiocampidae). Can. Entomol. 129: 991-999. 

Shepherd, R. F. 1979. Comparison of the daily cycle of adult behav- 
ior of five forest Lepidoptera from western Canada, and their re- 
sponse to pheromone traps. Mitt. Schweiz. Entomol. Ges. 52: 
157-168. 

Sokal, R. R and F. J. Rohlf. 1981. Biometry, ed. 2. Freeman, New 
York. 859 pp. 

Stehr, F. W. and E. F. Cook. 1968. A revision of the genus Malaco- 
soma Hiibner in North America (Lepidoptera: Lasiocampidae): 
systematics, biology, immatures, and parasites. Smidis. Inst. Bull. 
276, 321 pp. 

Struble, D. L. 1970. A sex pheromone in the forest tent caterpillar. J. 
Econ. Entomol. 63: 295-296. 

SYSTAT: statistics, version 5.2 ed. 1992. SYSTAT Inc., Evanston, Illi- 
nois. 

Witter, J. A. 1979. The forest tent caterpillar (Lepidoptera: Lasio- 
campidae) in Minnesota: a case history review. Great Lakes En- 
tomol. 12: 191-197. 

Witter, J. A. and H. M. Kulman. 1969. Estimating the number of 
eggs per egg mass of the forest tent caterpillar, Malacosoma diss- 
tria (Lepidoptera: Lasiocampidae). Mich. Entomol. 2: 63-71. 

Witter, J. A. and H. M. Kulman. 1972. Mortality factors affecting 
eggs of the forest tent caterpillar, Malacosoma disstria (Lepi- 
doptera: Lasiocampidae). Can. Entomol. 104: 705-710. 



Received for publication 20 May 2005; 
2006 



■lised and accepted 18 July 



Volume 60, Number 3 



161 



Journal of the Lepidopterists' Society 
60(3), 2006, 161-164 

A NEW SPECIES OF EUCOSMA HUBNER (TORTRICIDAE: OLETHREUTINAE) FROM THE TALL 
GRASS PRAIRIE REGION OF MIDWESTERN NORTH AMERICA 

Donald J. Wright 

3349 Morrison Ave., Cincinnati, Ohio 45220-1430, USA email: wrightdj@fuse.net 



ABSTRACT. Eucosma haydenae, new species, is described from Iowa and Illinois. This species, which seems to be a tall grass prairie 
obligate, is similar in appearance to E. rusticana (Kearfott) but is much smaller. A review of rusticana is included, and illustrations are provided 
for the adults and genitalia of both species. 

Additional key words: Eucosmini. 



The once pervasive tall grass prairie of the North 
American Midwest is now reduced to a scattered 
assortment of small disjunct patches, but those 
remnants still harbor insects that are rarely encountered 
elsewhere. This paper proposes a name for one such 
insect, a small brown moth described below as Eucosma 
haijdenae, new species. It was discovered during fauna! 
surveys in five prairie preserves, one in northeast Iowa, 
four in the vicinity of Chicago, Illinois. In general 
appearance it is most similar to Eucosma rusticana 
Kearfott, but the two species are separated easily by 
their marked difference in size. 

Eucosma rusticana is distributed widely in eastern 
North America but is rather poorly represented in 
collections. Kearfott (1905) described the species based 
on six specimens. Klots (1942) reported two svntypes in 
the American Museum of Natural Histoiy (AMNH), 
including one labeled LECTOTYPE, a designation he 
attributed to Heinrich (1923). I examined the lectotype. 
The second syntype in die AMNH, reported by Kearfott 
(1905) from Algonquin, Illinois, was not found. I also 
examined four syntypes at the United States National 
Museum (USNM). 

Materials and Methods 

This study is based on an examination of 97 adult 
specimens and 10 genitalia preparations from the 
following collections: AMNH, Loran D. Gibson, Todd 
Gilligan (TG), Mississippi Entomological Museum 
(MEM), USNM, Ron Panzer, and Donald J. Wright 
(DJW). Forewing length (FWL), defined as distance 
from base to apex (including fringe), was measured to 
the nearest one tenth of a millimeter. A rough indication 
of forewing geometry is provided by the aspect ratio 
(AR), defined as FWL divided by medial forewing 
width, the later measurement taken perpendicular to 
the dorsal margin. Costal fold ratio (CFR) is defined as 
costal fold length divided by FWL. Reported values of 
AR and CFR are averages, rounded to two decimal 



places, ol the corresponding values calculated for a 
small sample of specimens. The number of 
measurements or observations supporting a particular 
statement is indicated by n. Forewing pattern 
terminology follows Brown & Powell (1991) and 
Baixeras (2002). 

Species Accounts 

Eucosma rusticana Kearfott 
(Figs. 2, 4, 7, 8) 
Eucosma rusticana Kearfott 1905:358; Barnes and 
McDunnough 1917:170; Heinrich 1923:125. Fig. 162; 
McDunnough 1939:47; Powell 1983:35. 

Types. Lectotype designated by Heinrich (1923): 6", Kerrville, Tex., 
AMNH. Paralectotypes. NORTH CAROLINA: Trvon, Fiske, 13 May 
1903 (1 d, USNM), 28 May 1904 (1 d, USNM), 1 August 1904 (1 o, 
USNM, genitalia slide 70461), no date (1 9, USNM). 

Remarks. Eucosma rusticana is identified bv the 
following forewing characteristics: dorsal surface (Fig. 
2) divided longitudinally into a blackish-brown anterior 
region and a brownish-tan dorsoterminal region, the 
line of separation running roughly along the cubital vein 
from base to ocellus and from there obliquelv outward 
to costa just short of apex; region between said line and 
dorsal margin crossed longitudinallv bv three or four 
brown streaks; ocellus with pale-tan central field. 
bordered basally and distally by indistinct, transverse, 
silvery-gray bars and crossed longitudinally bv two, 
variably expressed, dark-brown dashes; nindi costal 
strigula white and conspicuous, odier costal strigulae 
gray and obscure. In melanic specimens die anterior 
region of the forewing is nearly all black, with black- 
suffusion extending to dorsum, but the streaked 
appearance of the dorsal region is still apparent. 
Forewing statistics: 6 FWL 9-12 mm (mean = 10.3. ri = 
25), AR = 2,57, CFR = 0.47,2 FWL 9.9-12.2 mm (mean 
= 11.2, n = 5), AR = 2.43. 

Male genitalia (Fig. 4) (n = 2): Uncus convex and moderatelv 
developed, socii long and setose, dorsolateral shoulders of tegumen 
well developed, gnathos a narrow band; vesica with ca. 22 deciduous 
comuti, valva with dorsal margin strongly concave, apex rounded. 



162 



Journal of the Lepidopterists' Society 




Figs 1^: Adults and male genitalia. 1, haydenae, holotype. 2, rusticana, 
slide DJW 556. 4, rusticana, Wyandot Co., Ohio, slide DJW 167. Scale bar: 



Rowan Co., Kentucky. 3, haydenae, Howard Co., Iowa, 
= 0.5 mm 



ventral two thirds of distal margin convex, dorsal one third mildly 
inset, producing narrowing of apical one third of cucullus, anal angle 
rounded, neck with scooped out invagination of ventrolateral margin 
(indicated by dashed line in Fig. 4), corner of sacculus rounded and 
nearly right-angled, margin of basal opening with weakly developed 
medial projection supporting a small patch of spines. Female 
genitalia (Fig. 8) (n = 2): papillae anales small, facing laterally and 
sparsely setose; sterigma (Fig. 7) semirectangular, length ca. 1.5x 
width, witii shallow trough from center of posterior margin to ostium, 
lamella antevaginalis ringlike and very weakly sclerotized, lamella 
postvaginalis with lateral and posterior surfaces densely 
microtriehiate; posterior margin of sternum VII invaginated to three 
lourths length of sterigma and closely approximate diereto; ductus 
bursae short, of nearly uniform width, with variably sclerotized ring 
posterior to juncture with ductus seminalis; corpus bursae with two, 
large, fin-shaped signa, inner surface of membrane microtriehiate. 

Biology and distribution. I examined 52 specimens 
(46 6, 69) from Arkansas, Illinois, Indiana, Kentucky, 
Missouri, Mississippi, North Carolina, Ohio, Tennessee, 
Texas and Wisconsin. The flight period extends from 
mid April to mid August, the earliest records coming 
from Mississippi. Midwest records are predominantly 
from June and July. No larval host has been reported, 
but other members of the genus are known to be stem 
and root borers of Asteraceae. 



Eucosma haydenae new species 
(Figs. 1, 3, 5, 6) 
Diagnosis. Size and forewing maculation distinguish 
haydenae from other eastern North American species of 



Eucosma. Mean FWL of superficially similar rusticana 
is ca. 3 mm longer than that of haydenae. Distinctive 
male genitalic characters include the scooped out 
invagination of the medioventral surface of the valval 
neck and the rounded anteroventral projections of the 
anellus. The sclerotized twist in die female ductus 
bursae is prominent but not unique to this species. 

Description. Head: Lower frons creamy white, upper frons 
creamy white to light brown, vertex brown, scales adjacent to eye 
lighter; labial palpus with basal segment white, second segment with 
medial surface and dorsal margin white, lateral surface gray brown 
with white medial mark, third segment brown, often with blackish- 
brown apex; antenna with dorsal surface brown, posterior surface 
white. Thorax: Dorsal surface orange brown, scales at posterior 
extremity of tegula shading to tan, ventral surface creamy white, fore 
and midlegs with anterior surfaces pale gray brown, posterior surfaces 
white, hindlegs white to tan, midtibia with white, oblique, medial 
mark on anterior surface, fore and mid tarsi with pale white 
annulations. Forewing (Fig. 1): 6 FWL 6-8.2 mm (mean = 7.3, n = 
29), AR = 2.82, CFR = 0.48,9 FWL 7.5-8.2 mm (mean = 7.9, n = 2), 
AR = 2.71; costa weakly convex, apex nearly right-angled, termen 
weakly convex; dorsal surface with blackish-brown region bounded 
anteriorly by basal one half of costa, posteriorly by cubital vein and 
distally by end of discal cell, region between cubital vein and dorsum 
white to tan, with orange-brown suffusion near base and along basal 
margin of ocellus, a narrow, sometimes interrupted, brown streak 
along dorsal margin; ocellus bordered basally and distally by silvery- 
gray transverse bars, central field light tan, crossed longitudinally by 
up to four, variably expressed, thin, black dashes; orange-brown 
scaling along distal one half of costa, crossed by five, paired, white to 
gray strigulae and their associated silvery-gray stria; ninth costal 
strigula white; fringe gray brown anteriorly, becoming paler toward 



Volume 60, Number 3 



163 



tomus. Male genitalia (Fig. 3) (n = 4): Uncus a rounded, dorsally 
setose lobe, divided medially by shallow indentation; socii long and 
densely setose; gnathos a narrow band, aedeagus tapered distallv, 
vesica with ca. 22 deciduous comuti; anellus with small rounded 
projections at anteroventral extremities; valva with costal margin 
strongly concave, apex evenly rounded, distal margin convex, ventral 
angle narrowly rounded, neck widi strongly scooped out invagination 
of medioventral margin, cucullus with medial surface densely setose 
and with ca. 10 stout setae evenly distributed along distal margin, 
sacculus moderately setose, margin of basal opening with narrow 
raised pulvinus, die latter connected to neck by weakly developed 
ridge. Female genitalia (Fig. 5) (n = 2): papillae anales small, facing 
ventrallv and moderately setose; sterigma (Fig. 6) long and 
semirectangular, lengdi more than 2x width, lamella antevaginalis 
ringlike and weakly sclerotized, lamella postvaginalis with lateral 
ridges bordering shallow medial trough, surface microtrichiate; 
sternum VII widi posterior margin deeply and narrowly invaginated to 
lengdi of sterigma, closely approximate to sterigma; ductus bursae 
with sclerotized twist posterior to juncture with ductus seminalis; 
corpus bursae with two fin shaped signa posterior to mid bursa, inner 
surface microtrichiate. 

Holotype. 6, IOWA, Howard Co., Hayden Prairie, 23 June 1997, 
D. J. Wright, deposited in USNM. Type locality at 43°26' 35" N, 
92°22' 58" W 



Paratypes. ILLINOIS: Dupage Co., W. Chicago Prairie, R. 
Panzer, 23 May 2004 (5 6, 29, 6 genitalia slide DJW129L2 genitalia 
slides DJW1290 & 1296), 6 June 2004 {16); Lee Co., Green River E., 
6 July 2002 (1 6). IOWA: Same locality as holotoype, D. J. Wright, 21 
June 2000 (1 6), 23 June 1997 (8 A, genitalia slide DJW 556), 28 June 
1995 (2 £, genitalia slides DJW 131 & 206), T. Gilligan (2 6). Paratvpe 
depositories: AMNH, TG, MEM, USNM, DJW 

Etymology. Dr. Ada Hayden (1884-1950) was a 
botanist at Iowa State College (now Iowa State 
University) who devoted much of her professional life to 
the study of the native Iowa prairie (Isely, 1989). Her 
advocacy in die 1940's for conservation of prairie habitat 
was largely responsible for the preservation of the 240 
acre tract in northeast Iowa that now bears her name 
and serves as the type locality for the moth described 
here. It's a joleasure to name this insect after Dr. 
Hayden. 

Distribution and biology. I examined 45 specimens 
(43 6", 29), documenting a flight period from late May to 




Figs. 5-8: Female genitalia. 5-6, haydenae, Dupage Co., Illinois, slide DJW 1290. 7-8, rusticana, Cook Co., Illinois, slide DJW 
1289. Scale bars = 0.5 mm. 



164 



Journal of the Lepidopterists' Society 



the beginning of July. All were collected in remnant tall 
grass prairie habitat in Howard County, Iowa, and in 
Cook, Dupage, and Lee Counties in Illinois. The larval 
host is not known but, as with msticana, is probably a 
species of Asteraceae . 

Remark. The haydenae specimens from the Chicago 
area have a generally blacker appearance than those 
from Iowa. 

Acknowledgements 

I thank J. W. Brown, R. L. Brown, and R. T. Schuh for the 
loan of specimens under their care. Karl Gnaedinger generously 
supplied me with specimens from the Chicago area, and D. 
Howell, Iowa Department of Natural Resources, helped with 
permits to sample the Hayden Prairie fauna and with biograph- 
ical information on Ada Hayden. Two anonymous reviewers of- 
fered helpful comments on the manuscript. 

Literature Cited 

Baixeras, J. 2002. An overview of genus level taxonomic problems 
surrounding Areyroploce Hiibner (Lepidoptera: Tortricidae), 
with description of a new species. Ann. Entomol. Soc. Am. 
95(4);422-431. 

Barnes, W. & J. McDunnouch. 1917. Checklist of the Lepidoptera 
of Boreal America. Herald Press, Decatur, Illinois. 392 pp. 



Brown, R. L. & J. A. Powell. 1991. Description of a new species of 
Epiblema (Lepidoptera: Tortricidae: Olethreutinae) from coastal 
redwood forests in California with an analysis of the forewing 
pattern. Pan-Pacific Entomol. 67:107-114. 

HEINRICH, C. 1923. Revision of the North American moths of the 
subfamily Eucosminae of the family Olethreutidae. U.S. Nat. 
MusBull. 123:1-298. 

Isely, D. 1989. Ada Hayden: A Tribute. Jour. Iowa Acad. Sci. 96(1):1- 
5. 

Kearfott, W. D. 1905. Descriptions of new species of Tortricid 
moths from North Carolina, with notes. Proc. U.S. Nat. Mus., 
28:349-364. 

Klots, A. B. 1942. Type material of North American microlepi- 
doptera other than Aegeriidae in the American Museum of Nat- 
ural History. Bull. Amer. Mus. of Nat. Hist. 79:391-424. 

McDunnough, J. 1939. Check List of die Lepidoptera of Canada and 
the United States of America. Part II Microlepidoptera. Mem. 
South. Calif. Acad. Sci. 2:3-171. 

Powell, J. A. 198.3. Tortricidae, pp. 31-41. In Hodges, R. W. et al. 
(eds.), Check list of the Lepidoptera of America north of Mexico. 
E. W. Classey & Wedge Entomol. Res. Foundation. London, 
England. 

Received for publication 6 January 2006; revised and accepted 19 June 
2006 



Volume 60. Number 3 



165 



Journal of the Lepidoptensts' Society 
60(3), 2006, 165-170 

MONARCH (DANAUS PLEXIPPUS L, NYMPHALIDAE) MIGRATION, NECTAR RESOURCES AND 
FIRE REGIMES IN THE OUACHITA MOUNTAINS OF ARKANSAS 

D. Craig Rudolph, Charles A. Ely, Richard R. Schaefer, J. Howard Williamson, and Ronald E. Thill 

Wildlife Habitat and Silviculture Laboratory (maintained in cooperation with the Arthur Temple College of Forestry, 

Stephen F. Austin State University), Southern Research Station, USDA Forest Service, 

506 Hayter Street, Nacogdoches, Texas 75965 USA Email: crudolph01@fs.fed.us 



ABSTRACT. Monarchs (Danaus plexippus) pass through the Ouachita Mountains in large numbers in September and October on their 
annual migration to overwintering sites in die Transvolcanic Belt oi central Mexico. Monarchs are dependent on nectar resources to fuel their 
migratory movements. In die Ouachita Mountains of west-central Arkansas migrating monarchs obtain nectar from a variety of plant species, 
especially Bidens aristosa and other composites. Fire suppression has greatly altered the structure of forest communities with major implica- 
tions for ecological relationships. Sites diat are undergoing restoration to a shortleaf pine-bluestem grass community following thinning and fre- 
quent prescribed fire, and diought to closely resemble pre-European conditions, support increased abundances of nectar resources and 
migrating monarchs compared to untreated controls. These results suggest that widespread fire-suppression since the early 1900s has substan- 
tially reduced nectar production for migrating monarchs in the Ouachita Mountains Physiographic Region. 

Additional key words: Interior Highlands, surveys, restoration 

The eastern North American population of the 
monarch butterfly, Danaus plexippus L., undertakes one 
of the most remarkable migrations of any lepidopteran 
(Urquhart 1976, Brower & Malcolm 1991). During the 
fall most individuals of this population migrate to 
extremely restricted sites in the Transvolcanic Belt of 
central Mexico (Urquhart 1976, Calvert & Brower 
1986). Concern has been expressed about the 
continued health of this population and the persistence 
of the massive migration phenomenon (Wells et al. 
1983, Brower & Malcolm 1991). Changes in abundance 
and quality of larval hosts (Zalucki & Brower 1992), loss 
of critical overwintering sites due to logging and fire 
(Brower 1996, Brower et al. 2002), vehicle mortality 
(McKenna et al. 2001), pesticides (Oberhauser 2004), 
introduced species (Calvert 2004), and transgenic Bt 
modified crops (Losey et al. 1999, Jesse and Obrycki 
2004) have been identified as actual or potential threats. 
Less attention has been directed to landscape-level 
changes in nectar availability which ultimately fuels the 
extended fall migration to central Mexico (Garcia & 
Equihau-Zamora 1997, Brower & Pyle 2004). 

Land use changes and management protocols in 
more natural habitats have drastically altered essentially 
all the land base that constitutes the breeding range and 
migration corridors of D. plexippus in eastern North 
America. The Ouachita Mountains Physiographic 
Region of west-central Arkansas and southeastern 
Oklahoma, encompassing 3,237,600 ha, remains 
primarily forested (Bukenhofer and Hedrick 1997). 
However, logging, fire suppression, and silvicultural 
management have altered vegetation structure and 
composition throughout the region (Foti and Glenn 
1991, Masters et al. 1995). Fire-maintained shortleaf 
pine (Pinus echinata) forests were widespread in the 



Ouachita Mountains until the earlv 20th century (Foti & 
Glenn 1991). Since the original harvest of these pine 
forests, most forested sites have been altered using 
intensive short-rotation pine production or remain as 
more natural forests, but have been subjected to fire 
suppression for several decades (Bukenhofer and 
Hedrick 1997). In either case, the abundance and 
quality of nectar resources available to Lepidoptera has 
been drastically altered (Thill et al. 2004). 

The managers of the Ouachita National Forest have 
initiated a landscape scale restoration of tire fire- 
maintained shortleaf pine-bluestem (Schizachrium spp., 
Andropogon spp.) ecosystem on 48,706 ha (U. S. Forest 
Service 1996). Restoration involves thinning tire 
overstory, reduction of midstory vegetation, and 
prescribed burning on a three-year return interval. This 
restoration was undertaken to restore habitat for the 
endangered red-cockaded woodpecker (Picoides 
borealis) and to restore what is thought to be the pre- 
European structure and composition of the vegetation 
(Foti and Glenn 1991, Bukenhofer and Hedrick 1997). 
A number of authors have examined the eflect of these 
restoration efforts on a diversity of taxa (see Thill et al. 
2004). 

As part of ongoing studies of the effects of restoration 
of fire-maintained shortleaf pine-bluestem habitat on 
lepidopteran communities, butterfly and nectar 
resource surveys were conducted in restored and 
untreated control plots. This paper reports results for D. 
plexippus in relation to the fire regime and suggests 
implications for fall migration and over-winter survival. 

Materials and Methods 

This study was conducted on the Poteau Ranger 
District (34°45'N, 34°15'W) of the Ouachita National 



166 



Journal of the Lei>idopterists' Society 



Forest in west-central Arkansas. Topography in this 
region consists of east-west trending ridges and valleys 
with an elevational range of 150-820 m. Mixed 
hardwood forests dominate north-facing slopes and pine 
and mixed pine-hardwood forests dominate south- 
facing slopes. Prior to the initiation of fire-suppression 
activities, much of the landscape, especially the more 
xeric pine communities on south- and west-facing 
slopes, burned on a regular basis (Foti and Glenn 1991, 
Masters etal. 1995). These were primarily low intensity 
ground fires ignited bv lightning. Native Americans, and 
more recently by European colonists. The resulting 
forest structure was characterized by pine-dominated 
overstories, sparse midstories, and diverse herbaceous 
understories (du Pratz 1975, Featherstonhaugh 1844, 
Nuttall 1980, Foti and Glenn 1991). 

The Ouachita Mountains are still predominately 
forested; however, logging and fire-suppression have 
dramatically altered vegetation structure (Bukenhofer 
and Hedrick 1997). Compared to pre-European 
conditions, existing forests are typically characterized by 
a younger and denser canopy, a dense woody midstory, 
and a very suppressed herbaceous understory (Fenwood 
etal. 1984, Masters 1991, Sparks 1996). These changes 
have profoundly altered the original biodiversity of the 
Ouachita Mountains (Neal and Montague 1991, Smith 
and Neal 1991, Wilson et al. 1995, Sparks et al. 1998, 
Thill et al. 2004). 

Landscape scale restoration was initiated in 1979, 
formally incorporated into the Ouachita National Forest 
Plan in 1996, and currently projects the restoration of 
48,706 ha (7.3% of the Forest) to a shortleaf pine- 
bluestem condition (U. S. Forest Sendee 1996, 
Bukenhofer and Hedrick 1997). 

Bestoration consists of thinning of the overstory, 
removal of most midstory vegetation, and prescribed 
burning on approximately a 3-year rotation (U.S. Forest 
Service 1996). Ultimately, regeneration of canopy trees 
will be accomplished primarily through the 
implementation of irregular shelterwood and seed-tree 
harvests with a portion of the overstory retained 
indefinitely (U. S. Forest Service 1996). In addition, 
rotation age will be lengthened, primarily to provide 
sufficient older trees to support red-cockaded 
woodpecker recovery (Rudolph and Conner 1991). At 
the initiation of our studies, approximately 9,071 ha had 
been restored in a 42,148 ha landscape on the Poteau 
Ranger District (W. G. Montague pers. com.). We use 
"restored" in a relative sense and recognize that stands 
are on a trajectory toward an ecological state that 
resembles pre-European conditions. 



Materials and Methods 

We censused butterflies and nectar resources 
annually in nine restored (treatment) sites and three 
un-restored (control) sites. Sites varied from 10.5 to 
42.1 ha. All treated sites had received a minimum of 
four prescribed burns prior to the initiation of the study. 
Restored sites were included as portions of larger areas 
(range 65 to 2226 ha) burned on the same day. A total 
of three restored sites were prescribe-burned each 
spring. Consequently, in any given year, first, second, 
and third growing seasons post-fire were each 
represented by three sites. Additional details 
concerning treatment and control sites can be found in 
Thill et al. (2004). 

Adult butterflies (only D. plexippus data reported 
here) were censused using a time-constrained walking 
census along 500- m triangular transects centrally 
located within each site (Pollard 1977, Pollard and Yates 
1993). Individual transects were censused by slowly 
walking the length of transects for approximately 20 
min. Time involved in counting butterfly aggregations, 
netting individuals for identification, or waiting for sun 
or wind conditions to conform to set variables was not 
included in the 20-min period. During 2000-2003, 
census counts were conducted four times each year 
(first week of April, June, August, October). Only 
October data for D. plexippus are reported here. 
Census counts were replicated three times each month, 
each replicate conducted by a different observer on 
different days during the survey week. Individual 
censuses were conducted between 0900 and 1330 hrs 
CST when temperatures were between 18-36° C and 
wind velocity beneath the canopy was not too high to 
suppress butterfly flight (Beaufort Scale <4). Censusing 
was further restricted to periods when sunlight was 
sufficient to cast discernable shadows. The response of 
butterflies to light, wind, temperature and cloud cover 
varies seasonally and daily in complex ways. 
Consequently, observer judgement further constrained 
censusing to those periods when butterfly activity 
appeared to be substantial. Additional details 
concerning treatment and control sites can be found in 
Thill et al. (2004). 

In addition to census counts of butterflies, we 
recorded all observations of nectaring and other feeding 
activities by butterflies observed during this and other 
studies in the Ouachita Mountains between 1999 and 
2004. Voucher specimens were deposited in the 
herbarium of Stephen F Austin State University. Plant 
nomenclature uses the nomenclature found in Smith 
(1994). 

Nectar resources were quantified by counting flowers 



Volume 60, Number 3 



167 



in three lxlOO-m plots at each study site. Plots were 
located parallel to each side of the triangular 500-m 
butterfly census transect. Within these plots all 
potential nectar resources were counted each week that 
butterfly censuses were conducted. For most plant 
species, individual flowers or composite heads (capitula) 
were enumerated. Inflorescences, or portions thereof, 
were counted for a few species with small and/or dense 
aggregations of flowers (e.g. Ceanothus americanus, 
Allium spp.. Solidago spp., Apiaceae). These 
enumeration decisions were based on estimating the 
structure that most closely approximated a separate 
landing site for a typical butterfly. 

Table 1. List of nectar plants, inclusive nectaring dates by 
plant species, and number of nectaring observations for 
monarchs (Danaus plexippus) in the Ouachita Mountains, 
Arkansas during 1999-2005. 



Plant Species 


Dates 


# Observations 


Bidens aristosa 


9/7-10/5 


1890 


Eupatorium serotinum 


8/20-10/5 


164 


Solidago spp. 


8/1-10/5 


101 


Vernonia baldwinii 


6/30-10/2 


93 


Cunila origanoides 


9/9-10/2 


69 


Solidago petiolaris 


9/27-10/2 


60 


Liatris elegans 


7/2-10/4 


44 


Helenium amarum 


7/2-10/2 


40 


Aster spp. 


9/8-10/5 


39 


Aster ericoides 


9/27-10/4 


2S 


Aster anomalus 


9/27-10/2 


22 


Senecio obovatus 


4/5-4/23 


21 


Solidago rugosa 


10/1-10/5 


19 


Asclepias tuberosa 


5/17-8/2 


14 


Polygonum pensylvanicum 


10/2-10/5 


13 


Eupatorium spp. 


8/20-10/5 


10 


35 other species 




74 



Results 

In the Ouachita Mountains D. plexippus adults were 
rarely observed in spring and summer. However, during 
the fall migration in September and October, D. 
plexippus were common to abundant. We obtained a 
total of 2701 D. plexippus nectaring observations from 
1999 to 2004. All but 101 of these observations were 
made during September and October (Table 1). 
Danaus plexippus nectared most frequently on Bidens 
aristosa (1890, 70.0%), Solidago spp. (180, 7.0%), 
Eupatorium serotinum (164, 6.1%), Aster spp. (98, 



3.6%), Vernonia baldwinii (93, 3.4%), Cunila 
origanoides (69, 2.6%), Liatris spp. (59, 2.2%), and 
occasionally on an additional 31 species. The 1890 
nectaring observations on Bidens aristosa were during 
the fall migration, and were concentrated primarily on 
the verges of the extensive road system within the 
region. 

Due to low regional abundance we counted only 15 
D. plexippus during April, June, and August censuses. 
During October, generally the peak of migration, a total 
of 1019 D. plexippus were detected during transect 
surveys. These observations occurred most often on 
restored treatments, especially during the first growing 
season post-fire, rather than on controls (Table 2). 
Significant differences were detected across treatments 
(X 2 = 1637.6, df= 3, P< 0.001). 

Nectar resources were also more abundant on 
restored treatments than on controls (Table 3). Within 
restored treatments, nectar resources were more 
abundant on first year post-burn treatments and least 
abundant in drird year post-burn treatments during the 
October surveys. Significant differences were 
frequently detected across these four treatments (Table 
3). 

Table 2. Number of Danaus plexippus, summed across plots 
and observers, detected during October surveys on restored 
sites and control sites on the Ouachita National Forest during 
2000-2002. 



Year 



B-l" 



B-2 B-3 Control 



2000 

2001 
2002 
Total 



5.5 

746 

8 

809 



29 

101 

3 

133 



53 

13 

1 
67 





1 

10 



a B-l, B-2, and B-3 correspond to 1 st , 2 nd , and 3 rd growing 
seasons post-bum. 

Discussion 

Restored sites in the Ouachita Mountains consisting 
of a fire-maintained shortleaf pine-bluestem community 
supported a higher abundance of D. plexippus during 
migration than un-restored, fire-suppressed controls. 
This pattern was most noticeable during the first 
October post-fire. Roth aspects of this pattern were 
similar to the abundance of nectar sources. Numbers 
of D. plexippus detected varied considerably across 
years. This may have been due to timing of tire 
migration relative to our survey times, or changes in D. 
plexippus numbers in the eastern North American 



168 



Journal of the Lepidopterists' Society 



Table 3. Response of nectar resources to pine-bluestem restoration on the Ouachita National Forest during October 2000-2002. 
Mean number of nectar resources per 300 m 2 plots in restored and control stands. Means in the same row sharing the same letter 
were not significantly different in a I -way ANOVA with REGYVQ at P < 0.1 (SAS Inst. Inc. 198S:598). Because abundance values 

varied greatly within treatments, data were rank traiislonnrd prior to analysis. 



B-l" 



B-2 



B-3 



Control 



Year 



SE 



SE 



SE 



SE 



2000 


746.3A 


66.4 


84.3B 


45.9 


59.3B 


21.9 


7.3C 


5.0 


0.0010 


2001 


2745.0A 


604.9 


3219.0A 


1605.5 


515.3B 


106.2 


183.7B 


130.4 


0.0047 


2002 


4587.3A 


2987.2 


1746.7A 


689.1 


719.3AB 


355.0 


35.0B 


8.3 


0.0181 



1 B-l, B-2, and B-3 correspond to 1", 2 nd , and 3 rJ growing seasons post-bum 



population. The very low numbers in 2002 followed 
catastrophic winter storm mortality at the Mexican over- 
wintering sites the previous January (Brower et al. 
2004). 

In the absence of frequent fire, nectar resources in 
the forested portions of the Ouachita Mountains 
Physiographic Region are generally low. Improved 
pastures and intensively managed pine plantations 2-3 
years after planting, the only other significant land uses 
in the region, also typically support a low abundance of 
nectar resources. Limited areas of increased nectar 
abundance occur in disturbed sites, i.e. road verges, 
utility rights-of-way, fence rows, and areas of recent 
timber harvest. Thus, at the regional level, in this 
primarily forested area, nectar resources may be 
limiting for lepidopteran species that require them 
(Thill et al. 2004, Rudolph et al. In prep.). Similar 
results were found for fire-maintained pine 
communities in eastern Texas (Rudolph and Ely 2000). 

Due to the very patchy distribution of nectar sources 
in relation to roads, rights-of-way, and forest 
management we were unable to assess the relative 
abundance of nectar sources across the entire 
landscape. The relative numbers of nectaring 
observations in Table 1 are probably biased toward 
those occurring along road verges. However, 
butterflies, including D. plexippus, were frequently 
observed nectaring along the transect surveys, especially 
in the restored treatments. 

Data presented here indicate that fire suppression 
has resulted in a landscape that is currently depauperate 
in nectar availability. We suggest that reduced nectar 
availability, compared to probable pre-European 
conditions, limits use of the widespread fire-suppressed 
habitats by monarchs and other lepidopteran species 
(Thill et al. 2004). The large numbers of monarchs 
passing through the Ouachita Mountains each fall 
obtain nectar resources primarily from concentrations 



of flowers located in disturbed sites, mainly the 
abundant Bidens aristosa growing on road verges. How 
this current availability of nectar resources compares 
with the pattern of nectar resources dispersed across the 
forest landscape in the pre-European fire-maintained 
shortleal pine forests is unknown, but this pattern 
suggests that fire suppression has greatly limited the 
availability of nectar resources across most of the 
forested landscape. 

Lipid-loading by D. plexippus during the fall 
migration, both to fuel the migration and sustain winter 
survival, is crucial (Brower 1985, Alonso-Mejia et al. 
1997). Brower and Pyle (2004) suggest that lack of 
nectar resources might be a limiting factor for migrating 
D. plexippus. In addition, a significant proportion of the 
monarchs inhabiting the wintering sites in the 
Transvolcanic Belt lack sufficient lipids to survive until 
spring (Brower & Pyle 2004). A detailed understanding 
of the current quality and availability of nectar resources 
along the migration corridor, and effects of historical 
changes, would allow an improved understanding of the 
remarkable migration of D. plexippus. Our results 
suggest that historical changes have been substantial, 
even in a landscape still dominated by "natural" plant 
communities. These changes may have potential 
consequences for migrating monarchs. Habitat and 
critical ecological processes, fire in this instance, may 
both be important in maintaining a major biological 
phenomenon on a continental scale. 

Acknowledgments 

We thank R. Buford, A Carter, C. Collins, and R. Perry for as- 
sistance in various aspects of fieldwork. N. Koerth provided as- 
sistance with data analysis and T. Trees maintained databases. 
Partial funding for this work was provided by the Poteau District 
of the Ouachita National Forest and the Ouachita Mountains 
Ecosystem Management Research Project. We thank R. M. 
Pyle, P. A. Opler, W. G. Montague, L D. Hedrick and two anony- 
mous reviewers for comments on an earlier draft of this manu- 
script. 



Volume 60. Number 3 



169 



Literature Cited 

Alonso-Mejia, A., E. Rendon-Salinas, E. Montesinos-Patino, and 
L. P. BftOWER. 1997. Use of lipid reserves by monarch butterflies 
overwintering in Mexico: Implications for conservation. Ecol. 
App. 7:934-947. 

Brower. L. P. 19S5. New perspectives on the migration biology of 
the monarch butterfly. Danaus plexippus L. Pp. 748-7S5 in M. 
Am. Rankin (ed.), Migration: Mechanisms and adaptative signifi- 
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Brower, L. P. 1996. Forest dunning increases monarch butterflv 
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Brower. L. P., G. Castelleja, A. Peralta, J. Lopez-Garcia, L. Bo- 
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Brower, L. P., D. R. Kust, E. Rendon-Salinas, E. G. Serrano, K. 
R. Kust, J. Miller, C. Fernandez del Rey, & K. Pape. 2004. 
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Brower, L. P.. and R. M. Pile. 2004. The interchange of migratory 
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Bukenhofer, G. A. & L. D. Hedrick. 1997. Shortleaf pine/bluestem 
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Fenwood, J. D., D. F. Urbston & R. F. Harlow. 1984. Determin- 
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170 



Journal of the Lepidopterists' Society 



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Received for publication 16 December 2005; revised and accepted 28 
June 2006 



General Notes 

Journal of the Lepidopterists' Society 
60(3), 2006, 171-174 

OBSERVATIONS OF KRICOGONIA LYSIDE (PIERIDAE) IN THE FLORIDA KEYS 

Additional kev words: West Indies, migration, seasonal dispersal, Florida Keys 



Kricogonia hjside (Godart), (Fig. 1) a butterfly native 
to the West Indies, the southwestern United States, and 
Central and South America, has historically occurred on 
an irregular basis on several of the middle and upper 
Florida Kevs, as well as on the southeastern Florida 
peninsula (Young 1938, Minno and Emmel 1993, Smith 
et al. 1994, Glassberg et al. 2000, Minno et al. 2005). A 
highly migratorv species, K hjside often travels en masse 
within and between the islands of the Caribbean 




FlG 1. Kricogonia hjside on Vaca Key, 23 July 2005 (Photo 
Credit: H. L. Salvato). 



(Wolcott 1927, Williams 1930, Smith et al. 1994, Miyata 
2000). The species also demonstrates similar mass 
movements in die southwestern United States with 
large influxes entering southern Texas from Mexico 
(Clench 1965, Gilbert 19S5). Dispersal of K. lyside into 
and within southern Texas appears to be triggered by 
reductions in new hostplant growth (Gilbert 1985). 
However, whether or not similar ecological cues 
encourage K. lyside migrations, both within the 
Caribbean and to southern Florida, remains unknown. 
Luis R. Hernandez (pers. comm.) suggests tropical 
storm activity in the Caribbean may play a large role in 
the dispersal of K. lyside within the West Indies, and 
perhaps to southern Florida. Smith et. al. (1994) 
witnessed several K lyside making landfall on Upper 
Matecumbe Key via strong, although not storm-related, 



easterly winds, indicating the species is capable of 
dispersal into the Keys during typical seasonal 
conditions. Young (1938), based on examination of fresh 
specimens collected near Biscayne Bay, suggested that 
the species reproduces within southern Florida and the 
Kevs. However, neither oviposition nor larval activity 
has ever been observed for K. lyside in the region. A 
known hostplant of this species, lignum vitae, Guaiacum 
sanctum L., (Zygophvllaceae) occurs commonly within 
hardwood hammocks throughout the Keys and is also 
widely used in the region as an ornamental species in 
landscaping. 

Following an apparent decade-long absence in 
Florida K. hjside was observed locallv from June 
through September 2002 on Plantation Key (Salvato 
and Salvato 2002) and Key West (MHS unpublished 
data) in the Florida Keys. After these initial 
observations of K. lyside re-occurrence in die Keys 
MHS and HLS continued to survey for this species as 
part of a larger ongoing long-term study to determine 
the status and distribution of butterflies throughout die 
Keys. Surveys were conducted on warm, clear davs 
under conditions that were considered sufficient for 
butterflies to be flying. Each sampling date included 
approximately 8-9 hours of field time (between 
0800-1700 h) in which two surveyors (MHS and HLS) 
walked a standard route at survey sites widiin die study 
area to visually record K. hjside activity. On each 
sampling date a selected span of the Keys was 
monitored in either the Upper (Key Largo to Upper 
Matecumbe Key), Middle (Lower Matecumbe Kev to 
Vaca Key) or Lower Keys (Baliia Honda Kev to Kev 
West). Survey sites on each island were \isited monthly 
during May 2002 to December 2005, with die exception 
of Lisnumritae Kev, which due to its inaccessibility, was 
only surveyed during June of each survey year. The 
amount of time spent surve\ing for K lyside on each 
Key varied based on island size and number of survey 
sites. Overall, a total of 25 survey sites, of varring sizes, 
were monitored monthly during this study. These 
locations included State and County' parks, National 
Wildlife Refuges and roadsides. Guaiacum sanctum 
was observed either directly within or adjacent to die 
majority of our study areas. 

Although there were unconfirmed reports of K lyside 
during 2003, we did not observe die species again in die 



172 



Journal ok tin-: Lltidoptlrists' Society 



Keys until 13 June 2004, after which K. lyside was 
frequently encountered throughout the Florida Keys, as 
well as southern Miami-Dade County, marking the first 
known occurrences for the species on mainland Florida 
in several years. Kricogonia lyside was absent from the 
majority of the islands we surveyed by mid-August 2004; 
however, local occurrences remained on Bahia Honda 
within the lower Keys into November of that year. 

In 2005, we began to re-encounter K. lyside during 
the early summer months with our first observations 
occurring on Lignumvitae Key on 11 June. Despite an 
abundance of G. sanctum on Lignumvitae Key, K. lyside 
had never been reported from this island (Minno and 
Emmel 1993, Smith et al. 1994). To our knowledge 
these observations of K. lyside on Lignumvitae Key 
represent the first reports from this island. From June 
through August of 2005 K. lyside was observed on every 
island we surveyed within the Keys (Fig. 2). 
Throughout this time frame the species was witnessed 
actively dispersing towards the east on both the 
northern and southern coastlines of the Keys, as well as 
on outer islands such as Lignumvitae Key, indicating a 
wider migratory swath than was noted earlier in either 
the 2002 or 2004 observations. As was observed in 



2004, K lyside remained on Bahia Honda until later in 
the season (at least through 24 September 2005) than 
elsewhere in the Keys. Table 1 provides an overview of 
K. lyside observations made on various islands within 
the Florida Keys during this survey. As with 
Lignumvitae Key, several of the K. lyside observations 
listed in Table 1 appear to be the first documented 
occurrences of the species on select islands, particularly 
within the Lower Keys (Minno and Emmel 1993). 

We observed an estimated 10, 74 and 238 individuals 
of K lyside during monthly surveys of the Keys in 2002, 
2004 and 2005, respectively. In all years the majority of 
individuals were found on Key West, Bahia Honda, Vaca 
Key and Plantation Key with varying numbers of 
individuals occurring on 15 other islands in the Keys. 
Although the sexes are similar, males from western 
Cuba and the Bahamas are distinctive from other 
known K. lyside populations in that these individuals 
generally lack a black bar near the apex of the upper 
hindwing (Biley 1975, Minno and Emmel 1993, 
Hernandez 2004, J. Y. Miller, pers. comm.). 
Examination of photographs (n = 25), collected 
specimens (n = 2) and field observations (n = 322) 
indicate that the K. lyside males encountered during 



Gulf of Mexico 




FIG 2. Islands in the Florida Keys on which Kricogonia lyside was observed during die survey period. 



Volume 60, Number 3 



173 



Table 1. Kricogonia lyside observations from the Florida 
Kevs during June through September of 2002, 2004 and 2005. 



Island 




2002 


2004 


2005 


Kev Largo 








3 


Plantation Key 




7 


12 


10 


Upper Matecumbe Key 




2 


1 


Windley Key 






1 


2 


Lignumvitae Key 








2 


Lower Matecumbe 


■ Key 




2 


1 


Grassv Key 








1 


Long Key 






7 


3 


Vaca Key 






3 


40 


Knights Key 








12 


West Summerland 


Key 






3 


Bahia Honda Key 






32° 


94 


Big Pine Kev 








28 


No Name Key 








2 


Cudjoe Key 








4 


Sugarloaf Key 








2 


Saddlebunch Key 






1 


6 


Stock Island 






2 


4 


Key West 




3 


12 


20 


° Indicates observations 


continued ur 


itil November 





these studies were morphologically similar to the 
western Cuban and Bahamian populations, suggesting 
that the seasonal occurrences of K, lyside in the Keys 
and southern Florida observed in this study were likely 
of West Indian origin. 

During June to mid-August of 2004 and 2005 
Kiicogonia lyside was consistently observed in active 
dispersal, traveling in either an easterly or northeasterly 
direction between islands in search of, and in frequent 
interaction with, G. sanctum. Conversely, K lyside 
observed later in the season during 2004 and 2005, 
specifically those remaining on Bahia Honda, occurred 
only locally and took nectar from any available sources, 
not just those in the immediate proximity of the 
hostplant. 

Despite extensive searches throughout the study 
period for signs that K. lyside had reproduced or 
underwent diapause in southern Florida it was not until 
9 July 2006 that MHS, HLS and Dennis J. Olle 
observed the species mating and ovipositing on the fruit 
and fresh growth of G. sanctum on Stock Island and Key 
West. Additionally on Key West, K. lyside was observed 
actively ovipositing on the leaves of Maracaibo lignum 
vitae, Bulnesia arborea (Jacq.)Engl., (Zygophvllaceae) a 



non-native species that occurs as an ornamental in the 
Keys and was not common in our study areas. To our 
knowledge these observations of oviposition provided 
the first accounts of K lyside reproduction in southern 
Florida. However, K. lyside larval activity was not 
observed during these studies. Larvae of K lyside feed 
nocturnally (Biley 1975, Hernandez 2004) and earlv 
instars are similar in coloration to that of the new G. 
sanctum growth on which they feed (Allen et al. 2005). 
The mature larvae are darker green with white and 
brown stripes (Minno et al. 2005). The cryptic 
coloration and nocturnal feeding habits of K lyside 
larvae makes the species difficult to find on the 
hostplant (Luis B. Hernandez, pers. comm.) and may 
explain our inability to locate them. Additional studies 
are needed to better determine the occurrence and 
natural history of immature K. lyside in southern 
Florida. Furthermore, there were no signs of adult K 
lyside activity across die Keys by mid-fall of 2002, 2004 
and 2005, suggesting the species may disappear from 
Florida and then sporadically re-colonize, probably 
from Cuba. The tropical storm seasons in 2004 and, for 
the Keys particularly 2005, were extremely active. 
However, the influxes and northeastern movements of 
K. lyside across the Keys and into southeastern Florida 
preceded any significant storms occurring during these 
study years. 

Our observations indicate that K lyside periodicallv 
disperses to a greater array of islands in the Florida Keys 
than previously noted. We have documented that 
during some years large numbers of K. lyside adults 
occur in the Keys. Additionally, K. lyside appears to 
breed in southern Florida, but to what extent the 
species colonizes widiin die state requires further 
investigation. 

The authors thank Luis R. Hernandez for sharing Iris field ob- 
servations and insights on die biology of K. lyside in die West In- 
dies. Jacqueline Y. Miller examined and presented die authors 
with her observations of K. lyside specimens within die collec- 
tion of the McGuire Center for Lepidoptera and Biodiversity. 
Florida Museum of Natural History Gaines\ille, Florida. The 
authors also extend thanks to Tim Adams, Don Stillwaugh Jr. 
Lyn and Brooks Atherton, Byrum and Linda Cooper. Dennis J. 
Olle and David L. Lysinger for sharing dieir field observations of 
K. lyside during the 2004-05 migrations. We also thank an 
anonymous reviewer for comments that helped improve die 
manuscript. Finally, die authors thank Barn Wood for creating 
and editing Figure 2. 

Literature Cited 

Allen, J. T, P. Brock & J. J. Glvssberg. 2005. Caterpillars in die 
Field and Garden; A Field Guide to die Butterfly Caterpillars of 



North America. Oxford University Press, New York. 232 



IT- 



Clench, H. K. 1965. A migration of Libytheana and Ki-icogonia in 

southern Texas. J. Lepid. Soc. 19:223-224. 
Gilbert, L. E. 19S5. Ecological factors which influence migratory 

behavior in two butterflies of the semi-arid shrublands of soudi 



174 



Journal of the Lepidoptehists' Society 



Texas. Contrib. Mar. Sci. 27: no. suppl. 
Glassberc, J., M.C. Minno & J. V. Calhoun. 2000. Butterflies 

through binoculars. Oxford University Press, New York. 242 pp. 
Hernandez, L. R. 2004. Field guide of Cuban-West Indies Butter- 
flies. Ediluz, Maracaibo. 269 pp. 
Kimball, C. P. 1965. The Lepidoptera of Florida — an annotated 

checklist. State of Florida Department of Agriculture, Division 

of Plant Industry. Gainesville. Florida. 363 pp. 
MlNNO„M. C. &T. C. Emmel. 1993. Butterflies of the Florida Keys. 

Scientific Publishers, Gainesville, Florida. 16S pp. 
Minno, M. C, J. F. Butler, & D. W. Hall. 2005. Florida Butterfly 

Caterpillars and Their Host Plants. University Press of Florida, 

Gainesville. 341 pp. 
Miyata, A. 2000. Mass migration of Kricogonia hjside (Lepidoptera, 

Pieridae) in Santa Domingo, Dominican Republic, in 1995. 

Trans. Lepid. Soc. Japan. 51: 281-286. 
Riley, N. D. 1975. A field guide to the butterflies of the West Indies. 

Collins. London, England. 224 pp. 
Salvato, M. H. & H. L. Salvato,. 2002. A new host record for 

Automeris io and the re-occurrence of Kricogonia hjside in 

Florida. News Lepid. Soc. 44:128. 

Journal oftlie Lepidopterists' Society 
60(3), 2006, 174-176 



Smith, D. S., L. D. Miller & J. Y. Miller. 1994. The Butterflies of 

the West Indies and South Florida. Oxford University Press, New 

York. 264 pp. 32 pi 
Wolcott, G. N. 1927. Notes on the pierid butterfly, Kricogonia 

castalia Fab. (Lepid.). Entomol. News. 38:97-100'. 
Williams, C. B. 1930. The Migration of Butterflies. Oliver and Boyd, 

London, p. 133. 
Young, F. N. 1938. Some interesting butterfly records for south 

Florida. Entomol. News. 49:115. 

Mark H. Salvato, 1765 17th Avenue SW, Vera 
Beach. Florida, 32962, USA: Email: 

anaea_99@yalioo.com, John V. Calhoun, 977 Wicks 
Drive, Palm Harbor, Florida, 34684, USA: Email: 
bretcall@verizon.net and Holly L. Salvato, 1765 17th 
Avenue SW, Vero Beach, Florida, 32962, USA 

Received for publication 29 December 2005, revised and accepted 21 
June 2006. 



A PRECAUTIONARY TALE ABOUT RARITY 7 : ON THE LARVA AND LIFE HISTORY 
OF LITHOPHANE JOANNIS (LEPIDOPTERA: NOCTUIDAE) 

Additional key words: shelter-forming. Aesculusflava, Lithophane innominata, Lithophane patefacta 



This note is about rarity, and how species that are 
regarded as scarce may be anything but, once aspects of 
their life history are better understood. Lithophane 
joannis Cove 11 and Metzler was not described until 
1992. Prior to the authors' distribution of paratypes 
there were no specimens of L. joannis in any major 
eastern institution, i.e., the Smithsonian, American 
Museum, and Carnegie Museum. Not William Forbes; 
nor Jack Franclemont, Doug Ferguson, Michael Pogue, 
Eric Quinter, or Tim McCabe has collected the moth. 
Dale Schweitzer wrote his dissertation on the tribe — he 
has yet to see the moth alive. Despite year-round 
surveys in Great Smoky Mountains National Park 
(GSMNP) — and especially over the last five years 
during which time die Park has been die focus of 
intensive surveys as part of its "All Taxon Biodiversity 
Inventory" — the moth escaped detection. Yet 
Lithophane joannis is among the Park's most common 
lepidopterans in middle elevation cove forests. 

On 19 May 2001 I collected two Lithophane larvae 
crawling up the trunk of a small yellow buckeye tree 
(Aesculus flava Ait.) (Hippocastanaceae) while 
collecting modis at a sheet and mercury light (with 
Doug Ferguson), above the Chimneys Campground 
(1000m) in Great Smoky Mountains National Park, 
Sevier County, Tennessee. The caterpillars looked 
similar to diose of the innominata group (e.g., L. 
hemina Grote, L. patefacta (Walker), L. petulca Grote 
and L. innominata (Small), and others), but different 



enough to raise doubt. Based on the host association 
and phenotype, Dale Schweitzer guessed diat the larvae 
were those of Lithophane joannis. Return trips to the 
same pullout along Newfound Gap Road in 2002, 2003, 
and 2004, yielded additional examples of the 
Lithophane. Typically, only one or two caterpillars were 
collected each year. Unfortunately, I failed repeatedly 
to rear examples through to the adult stage- 
inappropriate foliage was offered or larvae were lost 
during the obligator) - , four-month prepupal diapause 
common to Lithophane and other xylenines. In 2005, 
while light trapping at the same site above die 
Chimneys Picnic area, I thoroughly searched the same 
4m yellow buckeye tree that had Yielded caterpillars in 
every year previous. Nine Lithophane caterpillars were 
found in 20 minutes of searching (by flashlight). The 
larvae were feeding, perched on die underside of leaves, 
or observed walking along the trunk, with the exception 
of two larvae that were recovered from within leaf 
shelters. Both of these latter individuals were in the 
process of molting. 

Returning to the same area two days later (20 May, 
2005), I happened upon a buckeye tree widi numerous 
leaf shelters. Upon opening the first, I found a last instar 
Lithophane. Searching this same tree I counted more 
than 20 additional Lithophane joannis caterpillars in less 
than 10 minutes by opening other leaf shelters. Nearly 
every shelter had a caterpillar and some two (few if any 
of these Yvere in the process of a molt). No additional 



Volume 60. Number 3 



175 



caterpillars were obtained bv beating limbs of the same 
tree over a large queen-sized bed sheet. In late 
September, a series of Lithophane joannis issued from 
diis collection (Fig. 1). 

The larva of Lithophane joannis is strongly mottled 
(Fig. 2). There is often a straw to yellow tint where 
adjacent segments overlap and/or a tan to straw flush to 
the middorsal and lateral stripes. The white dorsal 
pinacula (Dl and D2 setae) are edged with black; both 
dorsal pinacula are often embedded in a diffuse dark 
patch diat is best developed over the eighth abdominal 
segment. The well-differentiated prothoracic shield is 
heavily blackened above die subdorsal stripe. Below the 
lateral stripe the subventer and venter are pale and 
largely unmarked. The head bears a dark coronal bar, a 
black spot within the frons (frontal triangle), and a black- 
bar above each antenna. Fully mature last instars are 
about 4 cm in length. The middle and penultimate 
instars are lime green, translucent, with a strong, 
somewhat creamy spiracular stripe and a broken, white 
middorsal stripe; die body bears numerous minute 
white spots over the trunk (Fig. 3). In appearance the 
last instars resemble those of L. hemina and L. 
innominata and may not be separable from them, 
although most individuals will be recognizable by their 
pale ground color, especially those individuals that have 
a pale green, yellowish, or steely blue cast. In most 
instances, larvae of L. joannis will be identifiable by 
their host association (with buckeye). 

No odier eastern Lithophane is known to consistently 
take up residence in leaf shelters. Other members of 
the innominata complex typically rest in bark crevices 
by day (Wagner 2005) Trunks of Aesculus flava — 
particularly on the understory trees where one can 
expect to find larvae in numbers — are often smooth and 
without fissures in which larvae could conceal 
themselves. It is not clear if L. joannis ever spins its 
own shelters or only uses those of other leps. In May 
2006 a collection of 13 additional larvae was made from 
the Chimneys area of the Park — all came from leaf 
shelters of microlepidopterans. Eight were collected 
from abandoned and pupal shelters made by 
Choristoneura fractivittana (Clemens) (Tortricidae) and 
the remainder from prepupal and pupal shelters spun 
by Yponomeuta multipunctella Clemens' 

(Yponomeutidae). 

Several Lithophane are known to be both predatory 
and cannibalistic, including the apparently closely- 
related L. patefacta (Schweitzer 1979; Wagner, 2005). 
While there would seem to be clear advantages to 

'While Yponomeuta multipunctella larvae normally feed on 
Euomjmus, since 2004 I have increasingly noted larvae on other 
hosts in the Smokies. 




Figs 1-3. Lithophane joannis: all from Chimneys area of 
Great Smoky Mountains National Park, Sevier County, Ten- 
nessee. 1, Reared adult. 2. Lithophane joannis last instar. 3. 
Penultimate instar. 

having the ability to take over previously spun shelters, 
cannibalism was not observed in five pint rearing 
containers that housed 3-5 middle and late instars. And 
as noted above, I occasionally found shelters with two 
larvae. Similarly, 10 of 13 larvae collected in 2006 came 
from occupied microlepidopteran shelters (see 
above) — no evidence of predation was noted in these 
(or any of the other shelters opened on the dav of die 
initial collection in the Park). 

In the middle elevation cove forests of GSMNP 
where caterpillars of Lithophane joannis were 
discovered, the insect is among the most abundant 
noctuid caterpillars — on 20 May, 2005, L. joannis was 
arguably the most common noctuid caterpillar present 
in the Chimneys area. At die tvpe localitv in Ohio. L. 
joannis outnumbered all odier members of die genus 
Lithophane at bait (Covell and Metzler 1992). 
Interestingly, adults ignored the light traps diat were 
run at the same location (Eric Metzler pers. comm.). L. 
joannis provides a noteworthy case of apparent rarity — 



176 



Journal of the Lepidopterists' Society 



if one were to depend on standard light trapping 
methods one would conclude that the moth is among 
the rarest lepidopterans in eastern North America. 
However if one employs bait or searches for caterpillars 
one could conclude just the opposite, that L.joannis is 
among the most common noctuids in Appalachian 
forests where its foodplant, Aesculus flava, grows in 
abundance. 

Identification of the adults was confirmed by Eric 
Metzler. Vouchers of both larvae and adults have been 
deposited at the University of Connecticut; adults have 
also been deposited at the United States National 
Museum. 

James Adams, Dale Schweitzer, and Bo Sullivan offered 
suggestions on an earlier draft of the paper and Rene Twarkins 
assisted with the larval images. 



Literature Cited 

Covell, Jr., C. V. and E. H. Metzler. 1992. Two new species of 
moths (Noctuidae: Acronictinae, Cuculliinae) from midland 
United States. J. Lepid. Soc. 46: 220-232. 

Schweitzer, D.F., 1979. Predatory behavior in Lithophane quer- 
quera and other spring caterpillars. J. Lepid. Soc. 33: 129-134. 

Wagner, D. L. 2005. Caterpillars of Eastern North America. Prince- 
ton University Press. 

David L. Wagner Department of Ecology and 
Evolutionary Biology, University of Connecticut, 
Storrs, Connecticut 06269, USA, E-mail: 
davicl.wagner@uconn.edu 

Received for publication 7 November 2005; revised and accepted 9 
June 2006 



Journal of the Lepidopterists' Society 
60(3), 2006, 176-178 

DIURNAL HERBIVORY DOCUMENTED FOR SPEYERIA IDALIA (NYMPHALIDAE) LARVAE ON 
VIOLA SAGITTATA (VIOLACEAE) IN PENNSYLVANIA 



Additional key words: regal fritillary, violet, foraging 

Only two extant populations of the regal fritillary, 

Spei/eria idalia Drury (Nymphalidae), are documented 
east of Indiana (Barton 1996 for Pennsylvania, Hobson 
1999 and Chazal 2002 for Virginia). The larger of the 
two populations occurs inside National Guard Training 
Center-Fort Indiantown Gap (NGTC-FIG), an 
approximately 6,925-ha military base located in south- 
central Pennsylvania. Comprehensive descriptions of 
the old-field successional habitats occupied by S. idalia 
at NGTC-FIG are presented in Barton (1996) and TNC 
(2001). Morphologic and genetic evidence indicates that 
eastern populations may deserve specific or subspecific 
status and designation as an evolutionary significant unit 
(Williams 2001a, 2001b, 2002). In light of the 
conservation status of S. idalia, research is warranted on 
its life history. 

Nocturnal foraging on Viola species has been 
reported or referenced for S. idalia larvae by Holland 
(1898), Ferris & Brown (1981), Opler & Krizek (1984), 
Schull (1987), Royer (1988), Iftner et al. (1992), Royer 
& Marrone (1992), and West (1998). However, Kopper 
et al. (2001) documented diurnal feeding on V. 
pedatifida G. Don (Violaceae) in three out of 12 S. 
idalia larvae observed in Kansas. At NGTC-FIG, Barton 
(1995) reported diurnal movements of S. idalia larvae 
and noted the predominance of V. sagittata Aiton 
relative to the presence of other Viola species but did 
not describe larval foraging behavior. 

Because S. idalia larvae have been challenging to 



locate in the field across the species' range (Scudder 
1889 for New England, TNC 2001 for Pennsylvania, 
Kopper et al. 2001 for Kansas, Debinski pers. com. for 
Iowa), behavioral observations of larvae have been 
difficult to obtain (Kopper et al. 2001). A combination of 
factors such as low population density (Barton 1995), 
small body size, solitary distribution, cryptic coloration 
and behavior (Stamp & Wilkens 1993), high mortality 
rates (Mattoon et al. 1971, Wagner et al. 1997), and 
concealing vegetation may partially explain the modest 
numbers of field-documented larvae. Previous surveys 
conducted at NGTC-FIG to detect larvae have resulted 
in very small sample sizes (n = 9; Barton 1995) or failure 
(n = 0; TNC 2000, 2001). 

On May 14, 2001, one S. idalia larva was 
unintentionally discovered at the Pennsylvania site 
during a vegetation study. Shortly thereafter, a 
qualitative survey of selected grasslands, known to be 
inhabited by S. idalia adults during previous years, was 
performed in an attempt to detect more larvae. 
Typically conducted between 0900 and 1600 hrs, the 
survey followed a generalized protocol: searching for 
individuals and groups of V. sagittata (including arrow- 
and ovate-leaved varieties), inspecting violets for 
evidence of strip-feeding herbivory (typical of S. idalia), 
and visually scanning violets and the surrounding area 
for larvae. Images of S. idalia larvae in Allen (1997) and 
Richard & Heitzman (1987) assisted with positive 
species identification. 



Volume 60, Number 3 



Table 1. Spatio-temporal and behavioral data for S. idalia larvae observed at NGTC-FIG, Pennsylvania, May 2001. Spatial 
locations have been normalized bv subtracting the coordinate values of the first sighting. 



Individual 
larvae 



Date 



Time 



Spatial location 
(UT.M meters) 



Behavior at time of discovery 



Northing Easting 



Diurnal feeding on 
leaves of V. sagittate 



5/14 1200 0.0 0.0 motionless; on bare ground" observed in field 

5/1S 0945 2617.3 5049.6 motionless; < 30 cm from Viola observed in captivity 



5/1 8 


1102 


2618.4 


5047,5 


motionless; on Viola 


not observed 


5/22 


1200 


361.8 


72.5 


motionless; on Viola 


observed in captivitv 


5/24 


1110 


3557,5 


6057.0 


feeding on Viola 


observed in field 


5/25 


1440 


2705.6 


5041.1 


feeding on Viola 


observed in field 



"distance to nearest Viola was unrecorded for larva 1 

Five additional S. idalia larvae were opportunistically 
discovered after more dmn 30 observer-hours of search 
effort (Table 1). Digital photographs were taken of each 
larva encountered and of the habitat in the immediate 
vicinitv of each sighting. Spatial coordinates of larvae 
were determined through Global Positioning System 
(GPS) technology (hardware: Pro XR Trimble receiver 
unit and TSC1 Asset Surveyor Trimble datalogger; 
software: Pathfinder Office version 2.80) and expressed 
in Universal Transverse Mercator (UTM) meters. 
Larvae were not marked so as to avoid potential 
handling effects. Because newly and previously 
identified larvae were indistinguishable, duplication was 
possible but unlikely, due to the relatively long distances 
between sightings on preceding days [i.e., -80 m 
(minimum), -7000 m (maximum)]. The two larvae 
observed simultaneously on May 18 were separated by 



approximately two meters. 

Diurnal foraging on V. sagittate was documented in 
five of the six larvae observed (Table 1 and Figure 1). a 
strong trend despite the extremely small sample size. 
Similar to those reported in Kopper et al. (2001), 
feeding bouts were short-lived. S. idalia larvae were 
observed to consume only leaves of V. sagittata, eidier 
partially or completely. In addition to foliar herbivory, 
Beattie & Lyons (1975) and Kopper et al. (200±) 
reported floral consumption of Viola spp. by the larvae 
of Argynnis spp. and S. idalia, respectively. Such 
behaviors, as well as nocturnal foraging bouts, may have 
been exhibited by larvae at the NGTC-FIG site but 
were not observed. Nonetheless, given the observations 
presented here for Pennsylvania, in conjunction with 
those for Kansas by Kopper et al. (2001), diurnal 
foraging by S. idalia larvae may be more prevalent than 




FIG. 1. Images of two S. idalia larvae diumally foraging on V sagittata in the field (a) and in controlled conditions (b) at NGTC-FIG. 
Pennsylvania, May 2001. 



Journal oftiii: Lli'iooptlrists' Society 



previously described in the scientific literature. 

Drs. Brian Kopper and Barn- Williams (University of Wis- 
consin), Dr. Roger Latham (Continental Conservation), and 
Joseph Hovis (Pennsylvania Department of Military and Veter- 
ans Affairs or DMVA) offered insightful comments on a preced- 
ing draft. John Emmett, NGTC-FIG GIS Analyst, provided the 
Trimble GPS receiver and datalogger for field use and assisted 
with geospatial data management. Special thanks go to the 
Pennsylvania Army National Guard (PAARNG) for granting ac- 
cess to field sites and vehicles for on-base transportation. This 
project was sponsored by the PAARNG (Cooperative Agree- 
ment # DAHA36-01-2-9001), and funding was provided by the 
Pennsylvania DMVA. The content of the information presented 
does not reflect the position or policy of the U.S. Government, 
and no official endorsement should be inferred. 

Literature Cited 

Allen, T 1997. The butterflies of West Virginia and their caterpillars. 
University of Pittsburgh Press, Pittsburgh. 388 pp. 

Barton, B. 1995. Report on the life history of the regal fritillary 
(Speijeria idalia) and interspecific competition with other 
Speyeria species. Unpublished report to die U.S. Department of 
Defense. 34 pp. 

. 1996. Final report on the regal fritillary, 1992-1995, Fort 

Indiantown, Annville, Pennsylvania. Unpublished report to the 
U.S. Department of Defense. 

Beattie, A., & N. Lyons. 1975. Seed dispersal in Viola (Violaceae): 
adaptations and strategies. Amer. Jour. Bot. 62(7): 714-722. 

Chazal, A. 2002. Status survey of the regal fritillary (Speijeria idalia) 
in 2002 on the Radford Army Ammunition Plant. Natural Her- 
itage Technical Report 02-20. Virginia Department of Conserva- 
tion and Recreation, Division of Natural Heritage, Richmond. 20 
pp. 

Ferris, C, & F. Brown. 1981. Butterflies of the Rock)- Mountain 
States. University of Oklahoma Press, Norman. 442 pp. 

HOBSON, C. 1999. Conservation status assessment for the regal 
fritillary (Speijeria idalia) in Virginia. Natural Heritage Technical 
Report 99-25. Virginia Department of Conservation and 
Recreation, Division of Natural Heritage, Richmond. 23 pp. 

Holland, W. 1898. The butterfly book. 1st edition. Doubleday and 
McClure. New York. 382 pp. 

Iftner, D., Shuey, J., & J. Calhoun. 1992. Butterflies and skippers of 
Ohio. Bulletin of the Ohio Biological Survey 9(1). The Ohio lepi- 
dopterists research report No. 3. Ohio State University, Colum- 
bus. 212 pp. 

Kopper, B., Margolies, D., & R. Charlton. 2001. Notes on the 
behavior of Speijeria idalia (Drury) (Nymphalidae) larvae with 
implications that they are diurnal foragers. J. Lep. Soc. 54(3): 
96-97. 

Mattoon, S., Davis, R., & O. Spencer. 1971. Rearing techniques for 
species of Speyeria (Nymphalidae). J. Lep. Soc. 25(4): 247-256. 

Opler, P., & G. Krizek. 1984. Butterflies east of the Great Plains: an 
illustrated natural history. Johns Hopkins University Press, Balti- 
more. 294 pp. 

Richard, J., & J. Heitzman. 1987. Butterflies and moths of Missouri. 
Missouri Department of Conservation, Jefferson City. 385 pp. 

Royer, R. 1988. Butterflies of North Dakota: an atlas and guide. 
Minot State University Science Monograph No. 1. 192 pp. 

Royer, R., & G. Marrone. 1992. Conservation status of the regal frit- 
illary (Speyeria idalia) in North and South Dakota, a report to 
United States Department of the Interior, Fish and Wildlife Serv- 
ice, Denver. 51 pp. 

Schull, E. 1987. The butterflies of Indiana. Indiana Academy of Sci- 
ence, Indiana University Press, Bloomington/Indianapolis. 179 
pp. 

SCUDDER, S. 1889. Butterflies of die eastern United States and 
Canada with special reference to New England. Published by the 
autiior. Cambridge. Vol. 1, pp. 1-776; vol. 2, pp. 777-1774; vol. 3, 
pp. 1775-1958. 



Stamp, N., & R. Wilkens. 1993. On the cryptic side of life: being un- 
apparent to enemies and the consequences for foraging and 
growth of caterpillars, pgs. 283-330. In Stamp, N., & T. Casey 
(Eds.), Caterpillars: ecological and evolutionary constraints on 
foraging. Chapman 6c Hall, New York. 

[TNC] The Nature Conservancy. 2000. Population monitoring and 
life history studies of the regal fritillary (Speyeria idalia) at Fort 
Indiantown Gap Military Reservation, Annville, Pennsvlvania: ac- 
tivity summary and report of findings (January - December 
1999). Prepared by The Nature Conservancy. Unpublished re- 
port to the Pennsvlvania Department of Military and Veterans Af- 
fairs. 

. 2001. Population monitoring and life history studies ot the regal 

fritillary (Speyeria idalia) at Fort Indiantown Gap National 
Guard Training Center, Annville, Pennsylvania: activity summary 
and report of findings (January - December 2000). Prepared bv 
The Nature Conservancy Unpublished report to the Pennsylva- 
nia Department of Military and Veterans Affairs. 

Wagner, D., Wallace, M., Boettner, J., & J. Elkinton. 1997. Sta- 
tus update and life history studies on the regal fritillary (Lepi- 
doptera: Nymphalidae), pgs. 261-275. In Vickery, P., Dunwiddie, 
P., & C. Griffin (Eds.), The ecology and conservation of grass- 
lands and heathlands in northeastern North America. Massachu- 
setts Audubon, Lincoln. 

West, P. 1998. Establishing long-term monitoring of die regal fritillary 
(Speyeria idalia Drury) in Wisconsin. RJ/KOSE Report, WIFO: 
regal fritillary' monitoring. 12 pp. 

Williams, B. 2001a. Patterns of morphological variation in Speyeria 
idalia (Lepidoptera: Nymphalidae) with implications for taxon- 
omy and conservation. Ann. Ent. Soc. Amer. 94(2): 239-243. 

. 2001b. Recognition of western populations of Speyeria idalia 

(Nymphalidae) as a new subspecies. J. Lep. Soc. 55(4): 144—149. 

. 2002. Conservation genetics, extinction, and taxonomic status: a 

case history of the regal fritillary. Cons. Biol. 16(1): 148-157. 

P. MOORESIDE 1 , D. ZERCHER 2 , AND P. McELHENNY. 

The Nature Conservancy, Fort Indiantown Gap Office, 
Department of Military and Veterans Affairs, 
Environmental Unit, Budding 11-19, Annville, PA 
17003. 



Received for publication 28 March 2003; revised and accepted 26 
June 2006 

'Current address: American Association for the Advancement of 
Science, 1200 New York Avenue, NW, Washington, DC 20005. E- 
niail: pmooresi@aaas.org 

-Current address: The Nature Conservancy, Peoria Office, 301 SW 
Adams Street, Suite 1007, Peoria, IL 61602. 



Book Review 



Journal of the Lepidopterists' Society 
60(2), 2006, 179-180 



THE GEOMETRID MOTHS OF EUROPE (A. 
Hausmann, ed.), Volume 2. Sterrhinae. Axel Hausmann. 
2004. Apollo Books, Stenstrup, Denmark. 600pp. 
Hai-dback ISBN 87-88757-37-4. Priced at 140 Euros on 
http://www.pensoft.net/notes/12090.stm. 

This volume represents another significant 
contribution to a series on European Geometridae that 
will be seen in retrospect as perhaps the most significant 
milestone in our understanding of that fauna. Not onlv 
does it pro\ide a comprehensive review of our 
knowledge to date, coupled to an immense amount of 
fresh research based on exhaustive examination of 
copious material, but it probably comes at a turning 
point in the way taxonomic information is presented, 
with the emphasis moving more towards the electronic 
media, away from print on paper. The editorial preface 
indicates the way the volume is embedded within the 
context of existing or imminent initiatives in electronic 
informatics for Geometridae. But, as someone who 
sometimes wonders whether the rush to embrace the 
electronic age may be more headlong than judicious, 
given our lack of experience of the security/vulnerability 
of the internet to all sorts of factors, I am reassured to 
know that there are hard-copy products such as this to 
fall back on under any worst case scenario. 

The Sterrhinae, after the Larentiinae, are probablv 
the most successful subfamily of the Geometridae in 
their proportional representation at temperate latitudes, 
though the Ennominae may have higher numerical 
representation. They also contain, after Eupithecia in 
the Larentiinae (covered in Vol 4 of this series), the two 
next most species-rich geometrid genera: Scopula and 
Idaea. All three genera pose significant difficulties in 
identification of their species, so publication of these 
two volumes has removed this impediment for the 
European fauna. 

The preface notes the strong (particularly western) 
Mediterranean focus of the subfamily in a European 
context, relative to the more even distribution of the 
Larentiinae, but this aspect is not explored to any great 
extent in the main text, though all the data are there to 
enable interested readers to analyse it for themselves. 
For example, there is endemism at a generic level, 
albeit from genera that are monotypic such as 
Anthometra (eastern Mediterranean) and Emmiltis 
(central Mediterranean), or Oar, if regarded as distinct 
from Scopula, with just two species in the 



Mediterranean. The five species of Cleta are also 
essentially Mediterranean, and also the genus 
Glossostrophia. Brachyglossina includes major X. 
African and Levantine species groups, with one species 
in Spain considered to be an outlier of the latter group. 

Apart from these smaller genera, the Mediterranean 
focus is seen to an extreme in Idaea, where over two- 
thirds of the species are Mediterranean, 73% of these 
restricted to the western part. In Scopula, as in the 
larentiine Eupithecia, only one diird are Mediterranean, 
and those are more evenly distributed across that 
region. 

The book reviews extensively the recent taxonomic 
advances made for the subfamily, but diere is still not a 
full consensus on the generic classification, as indicated 
by Sihvonen (2005, Nota lepid., 28: 70-71) in his review. 
There is also discussion of relationships of the 
Sterrhinae within the family as a whole, where there is 
growing consensus that the Sterrhinae and Larentiinae 
are the most basal groups, though relationships between 
the two lineages of Sterrhinae and the Larentiinae still 
need further study. 

All species are copiously illustrated by color plates 
showing the range of variation, often accompanied bv 
diagnostic half-tone figures in the body of the text. 
There are clear line-drawings of male and female 
genitalia, and some stereoscans of critical features of 
antennae and legs. As with other volumes, the 
description of the adult is headed as a diagnosis, and the 
true diagnosis is found as an account of differences of 
similar species. The text for each species includes 
extensive reviews of available biological and ecological 
information, drawing on the breadth of literature and 
current expertise on the fauna. Most species have larvae 
that are herbaceous feeders, those of Idaea tending to 
prefer dried or withered foliage. However, die larvae of 
the genus Cyclophora are arboreal defoliators. 

The systematic checklist at the end of the book also 
lists, with asterisks, species from neighboring areas, a 
particularly useful addition given the Mediterranean 
nature of the group, enabling the reader to appreciate 
how the diversity extends to N. Africa, Turkey or die 
Levant. Species from Madeira and the Canaries are also 
listed. The distributions of all species are illustrated bv 
maps with shaded areas indicating die general range 
within which are placed black circles or odier symbols 
indicating more precise localities from which material 



180 Journal of the Lepidopterists' S< « 1 1:1 1 



has been examined. Jeremy Holloway Entomology Department, Natural 

The book brings together an immense and diverse History Musem, Cromwell Road, London, SW7 5BD. 

amount of information in a clear and cohesive manner UK Email: j.holloway@nhm.ac.uk 
that will retain its value for many years to come. It is 
therefore an investment well worth making. 



COVER PHOTOS SOLICITED 

The Journal solicits high-quality color photographs for consideration 

as cover illustrations. The photographs may illustrate any aspect of 

Lepidopteran biology. 

Digital image files saved to a CD are preferred. Submit digital files 

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at a resolution of at least 120 pixels/cm in °.tif format. If you wish to 

submit printed photographs, please ensure that they are vertical in 

orientation (approximately 5.5 inches wide X 6.75 inches long 

or 14 cm wide x 17 cm long;). 



s 



Date of Issue (Vol. 60, No. 3): 2 October 2006 



EDITORIAL STAFF OF THE JOURNAL 

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NOTICE TO CONTRIBUTORS 

Contributions to die Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, 
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3 9088 01278 3155 
CONTENTS 

Five New Species of Paucivena Davis, 1975 (Lepidoptera: Tineoidea: Psychidae) From Cuba 
Rayner Nunez Aguiki 121 

A Remarkable New Riodinid Species, Stalachtis halloweeni (Riodinidae: Stalachtini), From 

Mount Ayanganna, Guyana Jason P.W. Hall 138 

A New Species of Auratonota (Lepidoptera: Tortricidae: Chlidanotinae) Formerly confused 
with A. Hydrogramma (Meyrick) John W. Brown 143 

A Revision of Psoloptera Butler, Including a Redescription of Its Known Species (Arctiidae: 

Arctiinae: Euchromiini) Rebecca B. Simmons 149 

Forest Tent Caterpillar: Mating, Oviposition, and Adult Congregation at Town Lights 

during a Northern Minnesota Outbreak William E. Miller 156 

A New Species of Eucosma Hubner (Tortricidae: Olethreutinae) From the Tall Grass Prairie 
Region of Midwestern North America. Donald J. Wright 161 

Monarch (Danaus Plexippus L. Nymphalidae) Migration, Nectar Resources and Fire Regimes in 
the Ouachita Mountains of Arkansas. D. Craig Rudolph, Charles A. Ely, Richard R. 
Schaefer, J. Howard Williamson, and Ronald E. Thill 165 

General Notes 

Observations of Kricogonia Lyside (Pieridae) in the Florida Keys Mark H. Salvato, John 

V. Calhoun, and Holly L. Salvato 171 

A Precautionary Tale about Rarity: On the Larva and Life History of Lithophane Joannis 
(Lepidoptera: Noctuidae) David L. Wagner 174 

Diurnal Herbivory Documented For Speyeria idalia (Nymphalidae) Larvae on Viola 
sagittata (Violaceae) in Pennsylvania P. Mooreside, D. Zercher, and P. McElhenny 176 

Book Review 

The Geometrid Moths of Europe Jeremy Holloway 179 



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