Journal of the Lepidopterists' Society 5,3 (4), 1999, 142- 152

EARLY STAGES OF CALICO ILLIONEUS AND C. lDOMENEUS (, BRASSOLINAE) FROM PANAMA, WITH REMARKS ON LARVAL FOOD FOR THE SUBFAMILY.

CARLA M. PENZ Department of Invertebrate Zoology, Milwaukee Public Museum, 800 West Wells Street, Milwaukee, Wisconsin 53233, USA , and Curso de P6s-Gradua9ao em Biocicncias, Pontiffcia Universidade Cat61ica do Rio Grande do SuI, Av. Ipiranga 6681, FOlto Alegre, RS 90619-900,

ANNETTE AIELLO Smithsonian Tropical Research Institute, Apdo. 2072, Balboa, Ancon, HEPUBLIC OF PANAMA

AND

ROBERT B. SRYGLEY Smithsonian Tropical Research Institute, Apdo. 2072, Balboa, Ancon, REPUBLIC OF PANAMA, and Department of Zoology, University of Oxford, South Parks Road, Oxford, OX13PS, ENGLAND

ABSTRACT, Here we describe the complete life cycle of Galigo illioneus oberon Butler and the mature larva and pupa of C. idomeneus (L.). The mature larva and pupa of each species are illustrated. We also provide a compilation of host records for members of the Brassolinae and briefly address the interaction between these and their larval food plants,

Additional key words: Central America, host records, monocotyledonous plants, larval food plants.

The nymphalid subfamily Brassolinae includes METHODS Neotropical species of large body size and crepuscular habits, both as caterpillars and adults (Harrison 1963, Between 25 May and .31 December, 1994 we Casagrande 1979, DeVries 1987, Slygley 1994). Larvae searched for ovipositing female butterflies along generally consume large quantities of material to Pipeline Road, Soberania National Park, Panama, mo­ reach maturity, a behavior that may be related as much tivated by a study on Caligo mating behavior (Srygley to the low nutrient content of their larval food plants & Penz 1999). The study area was a mosaic of old sec­ (Auerbach & Strong 1981) as to their large body size ondary and primary forests and pasture grasses with ap­ (e.g., 15 g for a living, mature larva of Caligo memnon proximately 2.2 m annual precipitation (Ridgely 1976) (Felder), DeVries 1983). Several species of brassolines and a wet season extending from late April to mid De­ lay eggs in clusters, and their larval feeding activity may cember. Our ohservations showed that C. illioneus produce remarkable damage to their food plants. For ex­ oberon oviposit mostly at dusk (approx. 1700-1900 h) ample, two Caligo memnon females were repOlted to lay, and only occasionally at dawn (0530- 0630 h), and we jOintly, 165 eggs on phmts in approximately three therefore concentrated our observations in the twilight weeks in an outdoor enclosure (Young & Muyshondt hours. Field collected early stages were reared in plastic 1985 ). Caligo caterpillars were reported to cause severe containers at ambient temperature (25-29°C). damage to banana plantations (Malo & Willis 1961, Har­ Wild female Caligo illioneus oberon were captured rison 1962, 1963, 1964), and larvae of isthmia in two traps at the edge of the forest along Pipeline Bates are known to defoliate palms (Dunn Road. To induce oviposition, captured females were re­ 1917; R. B. Srygley, C. M. Penz pers. obs.). leased into an outdoor insectary (3 x 3 x 3 m) inside of Apart from studies of population control (Malo & which grew sapientum L. and latis­ Willis 1961, Harrison 1962, 1964), few investigations patha Benth.(), latifolia (Link) K. have focused on the early stage biology of Caligo (see (), and three species of unidentified palms Young & Muyshondt 1985, and references therein). (). Females were supplied also with fresh cut Here we describe the complete life cycle of C. il­ of Saccharum spontaneum L. () and lioneus oberon Butler, describe a mature larva of C. Cyrtostachys sp. (Arecaceae), both exotics on which we idomeneus (L.), and review larval food plant records had observed oviposition by female Caligo illioneus and for 39 brassoline species. sp. respectively (R. B. Srygley pers. obs.). VOLUME 53, NUMBER 4 143

Preserved larvae and pupal skins of C. illioneus minating at upper end of front; frontal suture whitish. oberon are in the collection of the Milwaukee Public Body: predominantly green; brown middorsal stripe Museum. The head capsule and pupal skin of C. divided by discontinuous white midline stripe; thoracic idomeneus are currently in the Smithsonian Tropical and abdominal legs grayish white; ventral side grayish Research Institute, to be relocated to the National white; caudae held separated, pink with black tips and Museum of Natural History in the future. numerous short white setae. The larvae molted syn­ chronously. RESULTS Third instar (duration = 5-6 days, n = 4). Head: as in second instar. Body: as in second ins tar, except for a Caligo illioneus oberon Butler broad, red lateral line divided by a thin, white spiracu­ Oviposition behavior and food plants. In the lar stripe; Single dark brown triangular middorsal pro­ field, females laid clusters of9- 13 eggs (n = 4 clusters) jection at posterior end of abdominal segment A3. in a row along the midvein on the underside of Shed caudae were not eaten after molt to fourth instar. medium-aged to old blades of Saccharum sponta­ Larvae molted asynchronously. neum, an introduced Asian grass that invaded natural Fourth instar (duration = 6-7 days, n = 4). Head: grasslands of Panama during the 1970's. FollOwing its patterned in creamy-white and dark brown; dorsal introduction, S. spontaneum gradually replaced the scoli light brown anteriorly and reddish-brown poste­ pasture grasses Hyparrhenia rufa (Nees) Stapf and riorly at the base (approximately same height as the Panicum maximum Jacq. (both introduced from head); subdorsal scoli creamy-white (approximately Africa) on Pipeline Road (N. Smith pers. comm.). The two-thirds the length of dorsal scali); lateral scoli native larval food plant for C. illioneus is not known in creamy-white (approximately one-half the length of Panama, and our captive females did not oviposit on the subdorsal scoli); one pair of creamy-white tuber­ any of the plants available in the insectary. cles below lateral scoli; epicranial suture darkened; Egg (developmental time = 6 days, n = 13). White, front creamy-white with vertical, medial brown stripe; spherical, approximately 1.5 mm wide, adorned with two brown stripes arise from the base of dorsal scoli vertical ribs; description refers to a cluster of 13 eggs converge toward and flank epicranial and frontal su­ laid 27 August 1994. tures, terminating halfway down the length of the First instar (duration = 8 days, n = 4) . Head: front; adfrontal region dark brown above stemmatal brown with simple black setae; two dark brown verti­ region; post-genae reddish-brown; base of head red­ cal stripes flank epicranial and frontal sutures from dish-brown from occiput to mandibles; mandibles apex of head, terminating at approximately halfWay the creamy-white, darkened at the cutting edge. Body: length of the front. Body: translucent green; broad, color varied from light mustard to greenish; thoracic lemon-yellow middorsal stripe bordered by an irregu­ segments T1 and T2 with middorsal white midline lar reddish-brown stripe that is prominent on the tho­ stripe, flanked by reddish stripes; remaining segments rax and divided by a thin white discontinuous midline with thin grayish middorsal stripe; large reddish­ stripe that is more prominent on the thorax than on brown, triangular middorsal projection located at poste­ the abdomen; two thin, lateral, lemon-yellow longitu­ rior end of abdominal segment A3; very small middorsal dinal stripes; thoracic and abdominal legs grayish­ projection at posterior end of A5; supra-spiracular white white; ventral side grayish-white; caudae held sepa­ stripe along the entire length of the body; white sub­ rated, reddish to dark brown, each with a black spiracu lar stripe on a continuous longitudinal swelling; sub-terminal seta arising at one-third to one-half the thoracic and abdominal legs reddish; ventral side red; length of the caudae, and a terminal seta which is caudae pale pinkish-brown patterned with reddish­ white at base and black at tip. The larvae molted syn­ brown dorso-Iaterally, where pattern develops into chronously. thin broken lines. The larvae molted asynchronously Second ins tar (duration = 5 days, n = 4). Head: and three out of four aggregated at rest. dark brown anteriorly, lateral and post-genal regions Fifth instar (duration = 6-7 days, n = 4, Fig. 1a). translucent white; three pairs of scoli: dorsal scoli light Head: as in fourth ins tar, except for a brown stripe brown (approximately half the height of the head), arising from dorsal scoli that flanks the epicranial and subdorsal scoli whitish (two-thirds the height of the frontal sutures, and terminates above antennal socket; dorsal scoli), lateral scoli whitish (approXimately one­ head densely covered with short creamy-white setae. third the height of the subdorsal scoli ); front dark Body: predominantly beige; dark brown middorsal brown; two thin whitish lines arise from base of dorsal stripe runs along entire length of body; on segments scoli, converge toward and flank epicranial suture, ter- T1, T2, and anterior end of T3, the dark brown mid- 144 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

Flc. 1 Caligu illiuneus oberon, pbotograpbs by R. Srygley; (a) fiftb instar (October, 1994); (b) pupa (24 October, 1994). FTC. 2. Caligu idomeneus (Aiello Lot 81-77), pbotograpbs by A. Aiello; (a) final ins tar (2 December, 1981); (b) final instar, bead (2 December, 1981); (e) final instar, body detail (2 December, 1981); (d) pupa (23 December, 1981). dorsal stripe is divided by a thin creamy-white stripe; mately two-thirds the length of the head. Larvae triangular middorsal projections same as in fourth in­ molted asynchronously and did not aggregate at rest. star; body patterned with brown from dorsal midline Sixth ins tar (duration = 14- 16 days, n = 3), Head: towards the sides, where pattern develops into thin, as in fifth instar, except for four pairs of scoli; addi­ broken, longitudinal brown lines; dark brown supra­ tional stripe from dorsal scoli terminating above stem­ spiracular line bordered by creamy-white lines; creamy­ matal region; dorsal scoli dark brown with white tips white subspiracular stripe on a continuous longitudinal and basal creamy-white spots posteriorly; subdorsal swelling; ventral side pink; caudae held separated, light and lateral scoli posteriorly brown at base; sub-lateral brown at base, becoming dark brown at tip, approxi- scoli dark brown posteriorly (one-half of the length of VOLUME 53, NUMBER 4 145 the lateral scoli). Body: as in fifth instar, except for length of the dorsal scali, clothed in long setae, gently small dark brown triangular middorsal projection at curved upwards to pointed apex; lateral scoli smallest, posterior end of abdominal segment A2; large triangu­ about one half the length of the subdorsals, conical; lar middorsal projection at posterior end of A3; and front with dark vertical dash; adfrontal area dark small triangular middorsal projections at posterior brown; upper section of epicranial suture dark brown; ends of A4 and A5; ventral side brown; caudae held broad stripe lateral to adfrontal area, terminating at separated, slightly longer than head height. stemmatal level with a darker vertical dash; broad Larval development and behavior. Egg develop­ stripe from base of dorsal scoli, terminating on stem­ ment took 6 days (n = 13) and larval development mata with a darker vertical dash; dark brown stripe (from hatching to pupation) took 44-49 days (n = 3). from mid point of inside of each dorsal scolus, termi­ First through fourth instar larvae fed gregariously and nating at epicranial suture; dark brown stripe along generally rested together on the leaf blade. Fifth and curve from mid point of outside of each dorsal scolus, sixth instal' larvae rested away from each other on the to sides of head just in front of subdorsal scolus; base stem of the plant. Late ins tar larvae found in the field of head dark brown from occiput to mandibles. Body: were solitary (R. B. Srygley pers. obs.). brown, except paler dorsally on abdominal segments Pupa (duration = 13-15 days, n = 3, Fig. Ib). Beige A2-A6; broad subspiracular white stripe on AI-A7 with fine, dark brown cryptic markings, giving the gen­ with oblique brown intrusions from above, just poste­ eral appearance of a dried, curled and sun-bleached rior to each spiracle on A2-A 7; four soft laterally flat­ leaf; head with transverse keel at apex; long, black se­ tened triangular middorsal projections, one on each of tae located immediately above and on the surface of A3-A6; large oval middorsal spot, lying between the the eyes; antennae with a black longitudinal stripe, and projections of A3 and A4, dark brown with a beige pos­ a black transverse line marking each antennal seg­ terior-pointing arrow; caudae held separated, brown, ment; thoracic segment T2 with a prominent keel broad. along dorsal midline, more developed in female than Pupa (17 days, Fig. 2d). About 4.5 cm long, and 2 in male pupae; posterior edge of wing pad forming a cm wide at widest point; beige with fine brown cryptic crest; prominent hump at the base of wing pad; wing marldngs, giving the general appearance of a dried, surface with two small silver spots located near base of curled and sun-bleached leaf; head ridged from center wing; abdomen with conspiCUOUS long black setae of eye to vertex; eye area adorned with stout dark along dorsal midline; abdominal segments A5-10 with brown upright setae; antennae with a median black dark brown lateral line, A4-10 with brown ventral line; stripe for their entire length, and with cross lines set A6 humped; brown middorsal stripe arising at head approXimately 0.5 mm apart; mesothorax mid-dorsally and terminating at cremaster; abdominal segments humped and keeled; a lateral keel begins near the tho­ with transverse oblique markings that resemble leaf racic spiracle, passes along the forewing, parallel to the venation. Pupal mass: 2.3 g (n = 1, male). inner margin, and, at a level with abdominal segment A!, smoothes to become a raised area following the Caligo idomeneus (L.) forewing inner margin to the tornus; each forewing A wandering final instal' larva of C. idomeneus was bears two white enameled triangles toward the base of found off the food plant by R. Kimsey at Fort Clayton the inner margin and just ventral to the wing keel; (Canal Area, Panama) on 1 December, 1981 and clear patches and a small dark triangle are found at the reared to adult (Aiello Lot 81-77). The larva had about midpOint of each mesothoracic leg; on the abdomen a 1.5 white fly eggs cemented to the underside of the dark brown midventralline terminates at the tip of the thorax and head. The eggs were removed with forceps cremaster, as does an oblique dark brown line that be­ and preserved in 80% ethanol. The oviposition behav­ gins at the spiracle on abdominal segment A6; dorsum ior and larval food plants of C. idomeneus are un­ adorned with dark brown, upright setae from the known, but the captive mature larva readily accepted mesothoracic hump through abdominal segment A8, Benth. (Musaceae) and Calathea on which the setae are somewhat appressed; spiracles latifolia (Link.) K. (Marantaceae) which it ate for 20 narrowly elliptical, and that of A8 is obscure. days prior to pupation. Diagnostic characters of early stage morphol­ Final instar (n = 1, Figs. 2a-c). Head: beige with ogy. The mature larvae of Central American species of brown stripcs; three pairs of beige scali: largest scoli Caligo can be easily diagnosed by head and body color, dorsal, clothed in long setae, enlarged towards the and by the number and morphology of the head scoli. pointed apex and abruptly curved outward toward the The mature larva of sulanus Fruh­ sides of the head; subdorsal scoli about two thirds the storfer has a dark tan head adorned with four pairs of 146 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

TABLE 1. Larval food plants of the subfamily Brassolinae (Nymphalidae). Abbreviations: ARE = , BRO = Bromeliales, eyC = Cyclanthaceae, GEN = Gentianales, gym = gymnosperm, POA = Poales, ZIN = , ovp = oviposition record. References: Aiello, unphll; Aiello & Silherglied, 19782; Bm'cant, 19703; Biezanko et aL , 19744; Burmeister, 18735; Casagrande, 19796; Condie, 19767; Cubero, 19858; d'Almeida, 1922"; d'Aralljo e Silva et aL, 196810 ; DeVries, 1985", 198712; Fontaine, 191313; Harrison, 196314 ; Hayward, 196915; Moss, unphl (in Ackery, 1988)16; Miiller, 1886' 7; Rothschild, 191616; Small, unpbjl9; Slygley, lInpbI 20; Srygley & Penz, unpbJ21; Stauffer et al., 199322 ; Urich & Boos, 198FJ; Urich & Emmel, 199]24; Yepez et aL, 1985"; Young, 197726, 198627; Young & Muyshondt, 197528, 19852".

Butterfly speCIe, Pl ant ordel Keft'fcnCf'

L Blepolenis (as Opsiphanes) batea (Hubner) ARE Areeaceae "palm" 18 POA Poaceae Panicum lanatum Sw. (as capim amargosa) 10 (no 2388) Blepolenis batea (Hubner) ARE Arecaceae Arecastrum (as Syagrus) romanzoJfianum 4 (Cham.) Bece. ARE Arecaceae Butia (as Syagrus) capitata (Marl.) Becc. 4 2. Bmssolis astyra Godart ARE Arecaceae "diversas especies cle Palmae" 15 ARE Arecaceae "palrnell" 17 Bmssolis astyra astym Godart ARE Arecaceae Arecastrum (as Cocos) romanzoJfianum lO (no. 2.362) (Cham.) Bece. (as gerivii) ARE Arecaceae ayri Mart. (as brejauva) 10 (no 2362) ARE Arecaceae sp. lO (no 2.3(2) ARE Arecaceae Butia (as Cocos) eriospatha (c. Mart. ex Drucle) 10 (no. 2.362 ) Bece. (as butiazeiro) ARE Arecaeeae Cocos nucifem L. (as baba de boil 10 (no. 2.3(2) ARE Arccaceae Cocos nucifera L. (as coqlleiro da Bahia) 10 (no. 2362) ARE Arecaceae Cocos nucifera L. (as eoqueiro anita) lO (no. 2.362) ARE Areeaceae Copernicia cerifera Mart. (as earnaubeira) 10 (no 2362) ARE Arecaceae chinensis (Jaeq.) R. Br. 10 (no. 2.362 ) (as pent-sao cla China) ARE Arecaceae Livistona rotundifolia (Lamurck) Mart. 10 (no 23(2) ARE Arecaceae Phoenix da ctylifera L. (as tamareira) lO (no. 2.362) ARE Arecaceae Roystonea (as Oreodoxa ) oleracea (J acg.) lO (no. 2.362) O.F. Cook (as palmeira imperial) ARE Arecaecae Hoystonea (as Or-eodoxa) regia (Kunth) 10 (no 2.362 ) O.F. Cook (as palmeira real) POA Poaceae Saccharum. oJficinarum L. (as cana de a91lCar) 10 (no 2362) ZIN Musaceae Musa sapientum L. (as bananeira) 10 (no. 2.362) 3. Brassolis isthmia Bates ARE Arecaceae Chaemodora sp. 27 ARE Arecaceae Cocos !tucifem L. 12 4. (L) ARE Arecaceae (Jacq.) Locld. ex Mart. 22 ARE Arceaeeae Archontophoenix alexandrae (MueH.) 22 H. Wend!. & Drucle ARE Arecaceae Arecastrum rorrumzoffianum (Cham.) Bece. 22 ARE Arecaceae Astrocaryum spp. lO (no 2.365) ARE Arecaceae sp. 10 (no. 2365) ARE Arecaceac Jaeq. 22 ARE Arecaceae Bactris 'pp. 10 (no. 2.365) ARE Arccaceae Butia (as Cocos ) eriospatha (C. Mart. lO (no 23(5) ex Drude) 13ecc. (as butiazeiro) ARE Arecaceae mitis Lour. (as C. plumosa horticola) 22 ARE Areeaceac L. 22 ARE Arecaceae Chrysalidoearplls lutescens H. Wend!. 22 ARE Arecaceae Chrysalidocarpus lutescens H. Wend!. 10 (no. 23(4) (as areca bambll) ARE Areeaceae Cocos nucifem L. 3,15,22 ARE Arceaceae Cocos nueifem L. (as coqueiro cla Bahia) 10 (nos 2.36.3- 2.365 ) ARE Arecaceac Cocos nucifera L. (as coquciro anao) 10 (no. 23(5) ARE Arecaceae Copernicia cerifera Mart. (as carnallba) 10 (no. 2365) ARE Arecaeeae Desm.oncus spp. 10 (no. 2365) ARE Arecaceae Euterpe spp. 10 (no. 2365) ARE Arecaceae (LH. Bailey) H.E. Moore 22 ARE Arecaceae Livistona chinensis (Jaeq.) R. Br. (as pent- 10 (nos 2.364, sao cla China) 2.365 ) ARE Arecaceae Livistona sp. 22 ARE Arecaceae Mauritia jlexuosa L. f. 22 ARE Arecaceae Neodypsis decaryi Jumelle 22 ARE Arecaceae Orbignya spp. 10 (no. 23(5) ARE Arecaceae "palms" 18 ARE Arecaceae hart. ex Chabaucl 22 ARE Arecaceae Phoenix dactylifera L 22 ARE Arecaeeae Phoenix dactylifem L. (as tamareira) lO (no. 2.365) VOLUME 53, NUMRER 4 147

TABLE 1. Continued.

Butterfly ~pecle~ Plant unJer farm! ), specIes Reference

ARE Arecaceae Jacq. 22 ARE Amcaceae pacifica Seemann and Wendland 22 ARE Arecaceae Ptychospemw macarthurii (A.A. Wendl.) 22 C. Nicholson ARE Arecaceae Roystonea (as Oreodoxa) oleracea (J acq. ) 10 (nos 2364. O.F. Cook (as palmeira imperial) 2365) ARE Arecaceae Ruystonea (as Oreodoxa ) regia (Kunth) 10 (nos 2364, O.F. Cook (as palmeira real) 2365) ARE Arecaceae (Jacq. ) O .F. Cook 25 ARE Arecaceae (Kunth) O.F. Cook 22 ARE Arecaceae Roystonea venezuelana L.H. Bailey 22 ARE Arecaceae mauritiiformis (H. Karst.) Griseb. 22 and H. Wend!. ARE Arecaceae Sabal umbraculiferus Mart. 22 ARE Arecaceae Scheelea macrocarpa Karsten 22 AHE Arecaceae (Linden ex Andre) H. Wenell. 22 POA Poaceae L. (as cana de a<;ucar) 10 (no. 2365) ZIN Musaeeae Mllsa sapientum L. (as bananeira) 10 (no. 2365) ZIN Musaeeae madagascariensis Sonn. 22 ZIN Musaeeae Streiitzia nicolai Hegel & Korn. 22 5. Caligo {[1isbe [-Wbner ZIN Marantaceae Caetc 10 (no. 2369) 6. Caligo atreus Kollar ARE Areeaeeae Asterogyne martiana H. Wend!. (H. Wend!.) 11 ex Hems!. ZIN Marantaeeae Calathea sp. 7 ZIN Musaeeae Heliconia spp. 11,29 Galigo atreus dionysos Fruhstorfer CYC Cyclanthaceae Cyclanths 12 ZIN Musaceae Heiiconia sp. 12 ZIN Musaceae Mllsa sp. 12 7. Caligo beltrao (Illigcr) ZIN Cannaceae L. 6 ZIN Marantaceae Caete 10 (no. 2367) ZIN Marantaceae Calathea zebrirlfl (Sims) Lind!. 6 ZIN Musaceae Musa sapientum L. (as bananeira) 10 (no. 2367) ZIN Musaceae "plusieurs Musaceae" 9 ZIN J. Konig (as Iirio do brejo) 10 (no 2367) 8. Caligo eurilochus (C ramer) ZIN M arantaceae Calathea latifalia (Willd. ex Link.) Klotzsch 20 ZIN Musaceae T-ledychillrn sp. 17 ZIN Musaceae T-l eliconia latispatha Benth. 1 (lot 80-26) ZIN Musaceae Heliconia latispatha Benth. 20 (ovp) ZIN Musaceae Musa sapientllm L. 20 (ovp) ZIN Musaceae "plusieurs Masaceas (sic)" 9 Caligo curilochus brasiliensis (Felder) ARE Arecaceae Euterpe edulis Mart. (as palmito) 10 (no. 2368) ZIN Musaceae banana 18 ZIN Musaceae Musa sapientum L. (as bananeira) 10 (no 2368) ZIN Zingiberaceae Hedyehium eorona'dum J. Konig (as lirio do brejo) 10 (no. 2368) Caligo eurilochus sulanus Fruhstorfer ZIN Marantaceae Calathea sp. 12 ZIN Musaceae Helieonia sp. 12 ZIN Musaceae Musa sp. 12 9. Caligo idomeneus (L.) ZIN Marantaceae unidentified 15 ZIN Musaceae Helieonia latispatha Benth. 1 (lot 81-77) ZIN Musaeeae Musa sp. 16 10. Caligo illioneus (Cramer) ZIN Marantaceae T-ledyehium coronarium J. Konig (as lirio do brejo) 10 (no. 2371) ZIN Musaeeae Helieonia spp. 11 ZIN Musaceae Musa sapientum L. (as bananeira) 10 (no. 2371) Caligo illioneus oberon Butler POA Poaceae Saccharum spontaneum L. 21 ZIN Musaceae Heliconia sp. 12 ZIN Musaceae Mllsa sp. 12 Caligo il/ioneus pampeiro Fruhstorfer ZII'i Musaceae banana 18 l1. Caligo nwrtia (Godart) POA Poaceae Echinochloa crus-galli (L.) P. Reauv. 10 (no. 2372) (as capim canivao) POA Poaceae Pennisetum purpWTl1m Schumach. 10 (no. 2372) (as capim elefante) 12. Caligo memnon (Felder & Felder) ZIN Cannaceae Canna indica L. 1 (lot 85-57) ZIN Cannaceae Canna sp. 7 ZIN Maran taeeae Calathea latifolia (Willd. ex Link.) Klotzsch 20 (ovp) ZIN Musaeeae Helieonia latispatha Bent\'. 20,29 ZIN Musaceac Heliconia spp. 7, 11 , 19 ZIN Musaceae Musa sapientum L. 20 (ovp) ZIN Musaceae Musa sp. 14,29 148 JOURNAL OF THE LEPIDOPTERISTS' SOCIE TY

TABLE 1. Continued.

Butterfly spec!!O' s Plant-order family specIes Reference

Caligo memnon memnon (Felder & Felder) ZIN Musaceae Heliconia sp. 12 ZIN Musaceae Musa sp. 12 Caligo memnon telamonius (Felder & Felder) GEN Rubiaceae Coffea sp. (as eafeeiro) [dubious record] 10 (no. 2373) 13. Caligo oberthurii oberthurii (Deyrolle) ARE Arecaceae "low, evergreen palm" 18 14. Caligo oileus (Felder & Felder) ZIN Musaceae Musa sp. 16 Caligo oileus scamander (Boisduval) ZIN Musaceae Heliconia sp. 12 15. Caligo placidianus Staudinger ZIN Musaceae Musa sp. 16 16. Caligo praxsiodus Fruhstorfer POA Poaceae Saccharum officinanJm L. (as cana de a<;;ucar) 10 (no. 2.374) 17. CrLligo prometheus epimetheus ZIN Musaceae banana 18 (Felder & Felder) 18. Caligo teucer (L.) ZIN Musaceae Heliconia sp. 16 ZIN Musaceae Musa sp. 3 Caligo sp. ARE Arecaceac Cyrtostachys sp. 20 ZIN Musaceae Musa sapientum L. (as bananeira) 10 (no. 2366) ZIN Zingiberaceae Hedychium coronarium J. Konig (as lirio do brejo) 10 (no. 2366) 19. Catohlepia amphirhoe (Hubner) ARE Arecaceae Arecastrum (as Cocos) rornanzoffianwn (Cham.) 10 (no. 2375) Becc. (as geriva) ARE Arecaceae "palmeras" 15 20. Catoblepia orgetorix championi Bristow ARE Arecaceae palms 12 21. Da~yophthalma rusina (as geraensis) (Godart) ARE Arecaceae Bactris tomentosa Mart. (as uricana) 10 (no. 2376) Dasyophthalma r'Usina (Godart) ARE Arecaceae Euterpe edulis Mart. (as palmito) 10 (no. 2377) POA Poaceae Bamhusa sp. (as bambu) 10 (no. 2378) 22. Dynastor darius (F.) BRO Aechmea fasciata (Lind!.) Baker 10 (no. 2381 ) BRO Bromeliaceae Aechmea nudicaulis (L.) Griseb. 24 BRO Bromeliaceae Anan~ comosus (as sativus)(L.) MelT. (as abacaxi) 10 (nos 2379- 2381) BRO Bromeliaceae Ananas sp. (as ananas selvagem) 10 (no. 2379) BRO Bromeliaceae Ananas sp. (as ananas) 10 (no. 2,379) BRO Bromeliaceae Billhergia nutans H. Wend!. ex Regel 10 (no. 2381) BRO Bromeliaceae Billbergia speciosa Thunb. 10 (no. 2381) BRO Bromeliaceae Billbergia spp. 10 (no. 2380) BRO Bromeliaceae Bromelia fastuosa Lind!. (as bananilha do mato) 10 (no 2381) BRO Bromeliaceae Bromelia fastuosa Lind!. (as caraguata) 10 (nos 2379, 2380) BRO Bromeliaceae Bromelia fastuosa Lind!. (as banana do mato) 10 (no. 2379) BRO Bromeliaceae Ortgiesia (as Aechmea) gamosepala (Wittm.) 10 (no. 2381) L.B. Sm. &WJ. Kress BRO Bromeliaceae Tillandsia zehrina hort. ex Baker 10 (no. 2379) BRO Bromeliaceae unidentifed 17 Dyn~tor darius mardonius Fruhstorfer BRO Bromeliaceae unidentified 15 Dynastor darius stygianus Butler BRO Bromeliaceae Aechmea magdalenae (Andre) Andre ex Baker 11 BRO Bromeliaceae Aechmea sp. 12 BRO Bromeliaceae Agallostachys pingl1in (J ,.) Beer (as Bromelia 11 pinguin L.) BRO Bromeliaceae Ananas comosus (L.) Merr. [accepted by larvae 2, 1 (lot 7A-84) in captivi ty1 BRO Bromeliaceae Anan~ sp. 12 BRO Bromeliaeeae Bramelia plumieri (E. Morren) L.B. Sm. 1 BRO Bromeliaceae Bromelia sp. 12 BRO Brorneliaceae "pineapple and other bromeliads, gravata·· 18 23. Dynastor macrosiris (Doubleday) BRO Bromeliaceae Aechrrwa nudicaulis (L.) Griseb. 23,24 24. Dyn~t01 ' napoleon (Doubleday) BRO Bromeliaeeae Aechmea nudicaulis (L.) Griseb. 24 BRO Bromeliaceae Aechmea sp. 10 (no. 2382) BRO Bromeliaceae Anan~ comosus (as sativus) (L.) Merr. (as abacaxi) 10 (no. 2382) BRO Bromeliaceae gravata 18 25. aesacus bubocula (Butler) ARE Areeaceae palms 12 POA Poaceae Bambusa vulgariS Schrad. ex J.e. Wend!. 8 POA Poaceae Chusquea scabra Soderstr. & C.E. Calderon 8 POA Poaceae Olyra caudata Trin. 8 26. (C ramer) POA Poaceae Bambusa sp. 3 27. Eryphanis polyxena lycomedon POA Poaceae bamboo 12 (Felder & Felder) POA Poaceae Barnbusa arundinacea (Retz.) Willd. 1 (lot 82-9) POA Poaceae Saccharum spontaneum L. 1,21 28. Eryphanis reevesii (Doubleday) POA Poaceae Barnhusa (as Guadua) sp. 10 (no. 2383) POA Poaeeae Bambusa vulgariS Schrad. ex rC. Wend!. 10 (no. 2383) (as bambu comum) VOLUME 53, NUMBER 4 149

TABLE L Continued.

Butterfly ~pec l es Plant" order family specJes Referen c.: t'

POA Poaceae Bamhusa vulgaris Schrad. ex J.C. Wend!. 10 (no 2383) (as bambu comum) POA Poaceae Olyra latifolia L. (as taquarinha) 10 (no. 2383) POA Poaeeae Pennisetum purpureum Schnmach. 10 (no 2383) (as capim elefante) Eryphanis reevesii (Doubleday) POA Poaceae Bambusa sp. 17 (as Caligo rivesii ) POA Poaceae Olyra latifolia L. 17 29. Narope cyllastros Donbleday POA Poaceae Bambusa spp. 1.5,17 Narope cyUastros cyllastros Doubleday POA Poaceae Bambusa (as Guadua) sp. 10 (no. 2384) POA Poaceae Bambusa sp. (as bambu) 10 (no. 2384) Narope cyllastros testacea Godman & Salvin POA Poaceae Bambusa sp. 12 30. aorsa (Godart) POA Poaceae Bambusa vulgaris Schrad. ex J.e. Wendl. 10 (no. 2387) (as Opsiphanes aorosa) (as bambi, comum) 31. Opoptera staudingeri (Godman & Salvin) POA Poaceae Chusquea longifolia Swallen 8 POA Poaceae Chusquea sp. 12 Opoptera (as Opsiphanes ) staudingeri POA Poaceae Chusquea sp. 11 (Godman & Salvin) 32. Opoptera (as Opsiphanes ) syme (Hubner) POA Poaeeae Bambusa (as Guadua) sp. 10 (no. 239.5) 33. Opsiphanes bogotanus Distant ARE Arecaceae palms 11, 12 ZIN Marantaceae Calathea inocephala (Kuntze) H.A. Kenn. 1 (lot 81 -41) & Nicolson ZIN Marantaceae Calathea latifolia (Willd. ex Link.) Klotzsch 20 Opsiphanes bogotanus bogotanus Distant ZIN Musaeeae banana 18 34. Opsiphanes cassiae (L.) ARE Areeaceae unidentified 13 ZIN Musaeeae Heliconia sp. 16 ZIN Musaeeae Helieonia sp. and "differentes plantes musacees" 5 ZIN Musaeeae Musa sapientum L. 15 Opsiphanes cassiae cassiculus Stichel ZIN Musaceae Musa sp. 3 Opsiphanes cassiae lucttllus Fruhstorfer ZIN Musaeeae banana 18 ZIN Musaeeae Musa sapientum L. (as bananeira) 10 (no 2389) 35. aiellae Bristow ARE Areeaceae Cocos nUcifera L. 1 (lots 77-76, 82-1,83-17, 87-3) ARE Arecaeeae Livistona sp. 1 (lot 91-25) ARE Arecaeeae palm 1 (lot 95-8) Opsiphanes cassino fabricii (Boisduval) ARE Arecaeeae Acrocomia vinifera Oerst. 11,12 ARE Arecaceae Bactris guineensis (L. ) H.E. Moore (as 28 Bactris minor) ARE Arecaceae Bactris sp. 11,12 ARE Arecaeeae Cocos nucifera L. 11,12,28 ARE Arecaceae Erythea salvadorensis (H.Wend!. ex Bece.) 28 H.E.Moore (as Brahea saldorensis) ARE Arecaeeae Roystonea regia (Kunth) O.F. Cook 28 36. Opsiphanes invirae (Hubner) ARE Arecaceae Arecastrum (as Cocos) romanzoffianum (Cham.) 10 (nos 2390, Beee. (as geriva) 2392) ARE Areeaceae Arecastrum (as Syagrus ) romanzoJfianum 4 (Cham.) Beec. ARE Arecaceae Butia (as Cocos) eriospatha (e. Mart. ex 10 (nos 2390, Drude) Bece. (as buthlzeiro) 2392) ARE Arecaceae Butia (as Syagrus) capitata (Mart.) Beee. 4 ARE Arecaceae Cocos nucifera L. (as coqueiro da Bahia) 10 (no. 2390) ARE Arecaceae Copemicia cerifera Mart. (as carnauba) JO (no. 2390) ARE Arecaceae Livistona australis (R. Br. ) C. Mart. 4 ARE Arecaceae Livistona australis (R. Br. ) C. Mart. 10 (nos 2390, (as pent-sao austral) 2392) ARE Areeaceae Livistona chinensis (.facq.) R. Er. 10 (no. 2392) (as pent-sao chines) ARE Arecaceae Livistona chinensis (Jaeq.) R. Br. 4 ARE Arecaceae Livistona rotundifolia (Lamarck) Mart. 10 (no. 2390) ARE Areeaceae Palmeira de leque 10 (no. 2392) ARE Arecaceae Phoenix canariensis hort. ex Chabaud 4 ARE Arecaceae Prestoea sp. 8 ARE Arecaceae Raphia sp. (as palmeira ornamental) 10 (no. 2391) ARE Arecaceae Roystonea (as Oreodoxa) oleracea (Jac'l.) 10 (no. 2390) O.F. Cook (as palmeira imperial) 1.50 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

TABLE 1. Continued.

Butterflr spc(' Jt~~ Plant· order fHlnd)' ~ppcl es Referenc..

gym Cycadaceae Cycas eireinalis L. (as palmcira de jardim) 10 (no 2390) [dubious record] ZIN Musaceae Musa sapientum L. (as bananeira) 10 (nos 2390, 2392) Opsiphanes invirae amplificatu8 Stiehel ARE Areeaeeae "giriva and palms" 18 (as 0. i. remolia/us) Opsiphanes invirac amplificatus (Stiehel ) ARE Arecaceae Phoenix sp. 1.5 Opsiphanes invirae amplificatus Stichel ZIN Musaceae Musa sapientum L. (as bananeira) 10 (no. 2386) (as amplipladta ) Opsiphanes invirae cuspidatus Stichel ARE Arecaceae Bactris major Jacg. 1 (lot 82-44) ARE Arecaceae palms 12 37. Opsiphanesmerianae Stiche.1 ARE Areeaeeae ornamental palm 3 38. Opsiphanes quiteria (Cramer) ARE Arecaccae Arecastrum ramanzojfianum (Chan!.) Eecc. 15 Opsiphanes quiteria badius Stichel ARE Areeaceae Baetris sp. 1 (lot 81-74) Opsiphanes quiteria meridionalis Staudinger ARE Arecaceae Chrysalidocarpus (as Areca) /utescens H. Wendl. 10 (no . 2393) Opsiphanes quiteria merielionalis (as philon) ARE Arecaceae Astroearyum ayri Mart. (as brejauva) 10 (no 2394) Staudinger ARE Arecaceae Euterpe eelulis Mart. (as palmito) 10 (no 2394) Opsiphanes quiteriLI quirinus Godman & Salvin ARE Arecaceae Chrysalieloearpl1s luteseens H. Wencll. 8 ARE Arecaceae Cocos nucifera L. 26 ARE Arecaceae Geonoma sp. 8 ARE Arecaceae palms 12 ARE Arecaceae Pres toea allenii H. E. Moore 8 39. Opsiphanes tamarindi Felder & Felder ZE\ Mllsaceae Musa sapientl1m L. (as bananeira) 10 (no. 2396) ZI N Mu saceae Musa sp. 14,17 Opsiphanes tamarindi sikyon Fruhstorfer ZIN Musaceac H eli-conia col/insiana Griggs 28 ZIN Musaceae Helieonia latispatha Bcnth. 28 ZIN Musaceae Musa sp. 28 Opsiphanes tarnarindi tamarineli ZIN Cannaceae Canna indica L. 1 (lot 84-9) Felder & Felder ZIN Musaceae Helieonia latispatha Henth. 1 (lots 80-40, 80-43) ZIN Musaeeae H eliconia sp. 12 ZIN Musaceae Musa sp. 12 Opsiphancs sp. ARE Arecaeeae Cocos nucifera L. (as coqueiro anao) 10 (no. 2385) ARE Arecaceae Cyrtostachys sp. 20 ZIN Musaceae He/ieonia latispatha Benth. 1 (lot 82-18) ZI N Mllsaceae Musa sapientum L. (as bananeira) 10 (no 2385) scoli , dark brown body with six middorsal projections with the dorsal pair enlarged at tip and curved outward (see figs. 6 and 7 in Malo & Willis 1961 p. 532). That of (see fig. 32, E1 in DeVries 1987 p. 248), a tan colored C. atreus dionysos Fruhstorfer has a tan colored head body with many fine striations on dorsum, and five with fine vertical striations and three pairs of scoli, middorsal projections (DeVries 1987). The head cap-

TABLE 2. Summary oflarval food plant records for brassoline butterflies. Numbers represent species as listed in Table 1.

Phnt urdCI ARE POA ZIN eye BRO fam ily Arecacen!? Poaceae fvlusaceae Marantaceae Canaceae Cycl"nthaceae Bromeh-

Butterfly genera Blepolenis 1 Brassolis 2,:3,4 2,4 2,4 Caligo 6,9,14 11, 12, 17 6,7,8,9,10 5,6,7,8,9, 7,8 7, 12 6 12,14,15,17,18 10, 12 Catoblepia 19,20 Dasyophthalma 21 21 Dynastor 22,23,24 Eryphanis 25 25,26,27,28 Narope 29 0pol'tera 30,31,32 Opsiphanes 33, 34, 35, 36, 33,34,36,39 3:3 39 37,38 VOLUME 5:3, NUMBEH 4 151 sule of the mature larva of C. memnon is banded with with Arecaceae and Musaceae, except for O. bo­ tan and dark brown with four pairs of scoli (see fig. 5 in gotanus and O. tarnarincli, whose food plant range also Young & Muyshondt 1985 p.162; fig ..32, E2 in De­ includes Marantaceae and Canaceae respectively. Vries 1987 p. 248), with the dorsal pair enlarged at tip. Available information suggests that other brassolines The body is light brown with a dark brown middorsal are restricted to a Single plant family (Blepolenis and stripe, dark brown striations, and six middorsal projec­ Catoblepia on Arecaceae, Narope and Opoptera on tions (note that fig. 5 in Young & Muyshondt 1985 Poaceae, Dynastor on Bromeliaceae). Noteworthy is p.162, and fig. :31 F in DeVries 1987 p. 248, do not that the food plant range of the putative basal portray the same body color pattern). The mature Brassolis includes Arecaeae, Poaceae and Musaceae; larva of C. illioneus has a head patterned in brown and the plant families upon which most brassolines feed as creamy white adorned with three pairs of scoli plus a immatures. Although patterns of host association can lateral tubercle, and the body is beige with a dark he recognized at the generic level, their examination in brown middorsal stripe and four middorsal projections an evolutionary context awaits a well supported phy­ (Fig. la). That of C. iclomeneus has a beige head pat­ logeny for this group of butterflies. terned with brown, three pairs of scoli with the dorsal We hope that the summary presented here encour­ pair enlarged at tip and curved outward (Fig. 2b). The ages research aimed at furthering our understanding body is brown with a lighter colored area dorsally and of the patterns of food plant utilization and evolution a large oval middorsal spot between A:3 and A4, a in brassoline butterRies. broad white subspiracular stripe, and four middorsal projections (Fig. 2a). The pupae of all species are very ACKNOWLEDGMENTS similar, and those of C. illioneus and C. iclomeneus We are grateful to: R. Kimsey, who brought in the C. iJomeneus seem to differ only in the size of the white triangular larva; C. Galdames, for identifYing plants collected by R.B. Srygley and C.M. Penz; W. Hahn for guidance on palm nomenclature; PJ. marking at the base of the wing (more prominent in C. DeVries, G. Lamas and A. Neild for their help and suggestions; and iclomeneus, Fig. 2d). Early stages of C. oileus have P. AckelY, P. J. DeVries and D. Jenkins for comments on the manu­ never been formally described. script. We also acknowledge InRe:'olaRe (now A.N.A.M.) for grant­ ing permission to conduct research and collect butterflies in Larval food plants of the Brassolinae. It is well Panama, and the Smithsonian Tropical Research Institute (STRl) known that brassoline immatures are restricted to for proViding a vehicle to conduct research. Financial support was monocotyledonous plants (Ehrlich & Raven 1965, provided by STRI (to C. Penz), National Geographic Society (to R.B. Srygley), and National Science Foundation DEB98-06779 (to Ackery 1988, Table 1) , but little correlation has been C.M. Penz) . found between plant use and brassolinc classification (Ackery 1988). We found that larval food plants in­ LITERATURE CITED clude four of the eight monocot superorders (Table 1), ACKERY, P. R. 1988. Hostplants and classification: a review of a distribution suggesting that brassolines are generalist nymphalid butterflies. BioI. J. Linn. Soc. 33:9.5- 203. AIELLO, A. & R. E. SILBERGLIED. 1978. Life history of Dynastor monocot feeders. However, all food plant records to­ darius (: Nymphalidae: Brassolinae) in Panama. gether indicate that the majority of species feed on Psyche 8.5:331-34.5. plants in the families Arecaceae, Musaceae, and AUERBACH, M. J. & D. n. STRONC. 1981. Nutritional ecology of He­ liconia herbivores: plant fertilization and alternate hosts. Ecol. Poaceae (Tables 1 and 2). Therefore, the apparent lack Monogr. .51:63-83. of correspondence between plant use and brassoline BARCANT. M. 1970. The butterflies of Trinidad and Tobago. Lon­ classification should be reexamined. don. Collins. BIEZANKO , C. M. , A. RUFFINELLl & D. LINK. 1974. Plantas y otras Available records are sufficient to show that brasso­ sustancias alimenticias de las ofllgas de los lepidopteros line genera vary both in diet breadth and their associa­ uruguayos. Rev. Centro de Ciencias Rurais 4: 107-147. tion with monocot families (Table 2; see Stichel1909 BRI STOW, C. R. 1981. A revision of the brassoline genus Catoblepia (Lepidoptera: Rhopalocera). Zool. J. Linn. Soc. 72:117- 163. and Bristow 1981, 1982, 1991 for taxonomic classifica­ BRISTOW, C. R. 1982. A revision of the brassoline genus Seleno­ tion of the butterRics). For instance, although individ­ phanes (Lepidoptera: Rhopalocera). Zool. J. Linn. Soc. ual species of Caligo have been reported to feed on 76:27.3-291. BRISTOW, C. R. 1991. A revision of the brassoline genus Opsiphanes 1-4 plant genera in 1-4 families, collectively Caligo (Lepidoptera: Rhopaloeera). Zool. J. Linn. Soc. 101:203- 293. has a larval food plant range that includes 11 genera in BURMEISTER, H. 1873. Description de Morphonides Bresiliens. 7 families (Table 2, see also Ackery 1988) suggesting Rev. el Mag. Zoo!. 3:17-47. CASAGRAN DE . M. M. 1979. Sobre Caligo beltroo (liliger). 1. Tax­ multiple events of host colonization during its evolu­ onomia, biologia, morfologia das fases imaturas e distribui<;:oes tionary history. Similar patterns occur in Eryphanes espacial e temporal (Lepidoptera, Satyridae, Brassolinae). Rev. and Opsiphanes : species of Eryphanes typically feed Brasil. BioI. 39: 17:3-193. CONDIE, S. 1976. Some notes on the biology and behavior of three on Poaceae, except for E. aesacus which has also becn specip,s of Lepidoptera (Satyridae: Brassolinae) on non-eco­ found on Arecaeae; and Opsiphanes tend to associate nomic plants in Costa Rica. Tebiwa 3:1-28. 152 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

CUBERO, R. 1985. Notes on the life cycle and natural histol)' of Op­ SRYGLEY, R. B. 1994. Shivering and its cost during reproductive be­ siphanes quiteria quirinus Godman and Eryphanis aesacus haviour in Neotropical butterfies Caligo and Opsiphanes buhoculus Butler (Brassolidae). J. Lep. Soc. 39:33-43. (Nymphalidae : Brassolinae). Anim. Behav. 47:23-.32. D'ALMElDA, R. F. 1922 . .\IIelanges Lepidopterologiques. Etudes sur SRYGLEY, R. B & c. M. PENZ. 1999. Lekking in Neotropical owl les Lepidopteres du Bresil. Berlin. butterflies, Caligo illioneus and C. oileus (Lepidoptera: Bras­ D'ARAUJO E SILVA, A. G. , C. R GO NGALVES, D. M. GALVAo , A. J. L. solinae). J. Behav. 12:81- 103. GONGALVES, J. GOMES, M. DO NASCIM EN TO E SILVA & L. DE SI­ STAUFFEI1., F. , A. J. CLAVIJO & M. BEVILACQUA. 1994. Ataque de MONI. 1968. Quarto Catalogo dos insetos que vivem nas plan­ Brassolis sophorae (L., 1758) (Lepidoptera: Nymphalidae: tas do Brasil. Rio dc Janeiro: Ministerio de Agricultura. Brassolinae) a las palmas (Palmae) del parque del este "Romulo DEVRIES, P. J. 1983. Checklist of butterflies, pp. 654- 678, 703- Betancourt", Caracas, . Bol. Ent. Venez. N S. 704, 722-723, 729- 732, 741- 742, 751- 752, 754-755. In, D.H. 8(1):95-103. Janzen (ed.) Costa Rican natural histOly. University of Chicago SncHEL, H. 1909. Brassolidae. Das Tierreich 25:1-244. Press, Chicago. URICH, F. C. & J. O. Boos. 1981. Metamorphosis of Dynastor DEVRIES, P. J 1985. Hostplant records and natural histol)' notes on macrosiris Westw. (Lepidoptera Brassolidae). Living World, Costa Rican butterflies (Papilionidae, Pieridae & Nymphali­ 1981:34. dae). J. Res. Lep. 24:290- 333. URICH , F. C. & T. C. EMMEL. 1991. Life histories of Neotropical DEVRIES, P. J. 1987. The Butterflies of Costa Rica and their natural butterflies from Trinidad: 5. Dynastor darius darius (Lepidoptera: histol)'. Princeton University Press, Princeton, New Jersey. "Iymphalidae :Brassolinae). Trop. Lepidoptera 2(2):145--149. D UNN, L. H. 1917. The coconut- caterpillar (Brassolis isthmia) YEPEZ, G., F. FERNADEZ Y. & J. CLAVIJO. 1985. Presencia de Brasso­ of Panama. J. Econ. Entomol. 10:473-488. lis sophorae (L.) (Lepidoptera: Brassolidae) causando danos en EHRLICH, P. R. & P. H. RAVEN. 1965. Butterflies and plants: a study palm as de chaguaramos, Roysotea oleracceae (Jacq.), en el estado in coevolution. Evolution 18:586--608. Carabobo, Venezuela. Bol. Entomol. Venez. N. S. 4(3):23--24. FOUNTAINE, M. E. 1913. Five months butterfly collecting in Costa YOUNG, A. M. 1977. Notes on the defoliation of coconut palm (Co­ Rica in the Summer of 1911. Entomologist 46:189-194,214-219. cos nUcifera) by the butterAy Opsiphanes quiteria quirinus in HARRISO N, J. O. 1962. The natural enemies of some banana pests north-eastern Costa Rica. Deutsche Entomologische Zeit­ in Costa Rica. J. Econ. Entomol. 56:282- 285. schrift, 24:353-365. HARRISO N, J. O. 1963. On the biology of three banana pests in YOUNG, A. M. 1986. Natural histol)' notes on Brassolis isthmia Costa Rica (Lepidoptera: Limacodidae, Nymphalidae). Ann. Bates (Lepidoptera: Nyrnphalidae: Brassolinae) in northeastern Entomol. Soc. Arner. 56:87- 94. Costa Rica. J. Res. Lep. 24:385< 392. HARRISON, J. O. 1964. Factors affecting the abundance of Lepi­ YOUNG, A. M. & A. MUYSHONDT. 1975. Studies on the natural his­ doptera in banana plantations. Ecology 45:508-519. tol)' of Central America butterflies in the family cluster Satyri­ HAYWARD, K. F. 1969. Datos para el cstudio de la ontogenia de dae-Brassolidae-Morphidae (Lepidoptera: Nymphaloidea). Ill. Lepidopteros Argentinos. Instituto Miguel Lillo, Tucuman. Opsiphanes tanwrindi and Opsiphanes cassina in Costa Rica MALO, F. & E. R. WILLIS . 1961. Life histol)' and biological control and EI Salvador. Studies on the Neotropical F.auna 10:19--56. of Caligo eurilochus, a pest of . J. Econ. Entomol. Yout\G, A. M. & A. MUYSHONDT. 1985. Notes on Caligo m.emnon 54:530-,536. Felder and Caligo atreus Kollar (Lepidoptera: Nymphalidae: MULLER, W. 1886. Siidamerikanische Nymphalidenraupen: ver­ Brassolinae) in Costa Rica and EI Salvador. J Res. Lepid. such eines natiirlichen systems der nymphaliden. Zoologische 24:154-175. Jahrbiicher 1:417-678. RIDGELY, R. S. 1976. Birds of Panama. Princeton University Press, Received for publication 23 April 1999; revised and accepted Princeton, New Jersey. 10 October 1999 ROTHSCHILD, W. 1916. Notes on Amathusiidae, Brassolidae, Mor­ phidae, etc, with descriptions of new forms. Novitates Zoologi­ cae 23:299- 318.