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Journal of the Lepidopterists' Society 53(4), 1999, 142-152

EARLY STAGES OF CALIGO ILLIONEUS AND C. IDOMENEUS

(NYMPHALIDAE, BRASSOLINAE) FROM PANAMA, WITH REMARKS ON LARVAL

FOOD PLANTS FOR THE SUBFAMILY.

Carla M. Penz

Department of Invertebrate Zoology, Milwaukee Public Museum, 800 West Wells Street, Milwaukee, Wisconsin 53233, USA, and

Curso de Pos-Graduacao em Biociencias, Pontificia Universidade Catolica do Rio Grande do Sul,

Av. Ipiranga 6681, Porto Alegre, RS 90619-900, BRAZIL

Annette Aiello

Smithsonian Tropical Research Institute, Apdo. 2072, Balboa, Ancon, REPUBLIC 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, OX1 3PS, ENGLAND

ABSTRACT. Here we describe the complete life cycle of Caligo illioneus oberon Butler and the mature larva and pupa of C. idorneneus (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 butterflies and their larval food plants.

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

The nymphalid subfamily Brassolinae includes Neotropical species of large body size and crepuscular habits, both as caterpillars and adults (Harrison 1963, Casagrande 1979, DeVries 1987, Srygley 1994). Larvae generally consume large quantities of plant material to reach maturity, a behavior that may be related as much to the low nutrient content of their larval food plants (Auerbach & Strong 1981) as to their large body size (e.g., 15 g for a living, mature larva of Caligo memnon (Felder), DeVries 1983). Several species of brassolines lay eggs in clusters, and their larval feeding activity may produce remarkable damage to their food plants. For example, two Caligo memnon females were reported to lay, jointly, 165 eggs on banana plants in approximately three weeks in an outdoor enclosure (Young & Muyshondt 1985). Caligo caterpillars were reported to cause severe damage to banana plantations (Malo & Willis 1961, Harrison 1962, 1963, 1964), and larvae of Brassolis isthmia Bates are known to defoliate coconut palms (Dunn 1917; R. B. Srygley, C. M. Penzpers. obs.).

Apart from studies of population control (Malo & Willis 1961, Harrison 1962, 1964), few investigations have focused on the early stage biology of Caligo (see Young & Muyshondt 1985, and references therein). Here we describe the complete life cycle of C. illioneus oberon Butler, describe a mature larva of C. idorneneus (L.), and review larval food plant records for 39 brassoline species.

Methods

Between 25 May and 31 December, 1994 we searched for ovipositing female butterflies along Pipeline Road, Soberania National Park, Panama, motivated by a study on Caligo mating behavior (Srygley & Penz 1999). The study area was a mosaic of old secondary and primary forests and pasture grasses with approximately 2.2 m annual precipitation (Ridgely 1976) and a wet season extending from late April to mid December. Our observations showed that C. illioneus oberon oviposit mostly at dusk (approx. 1700-1900 h) and only occasionally at dawn (0530-0630 h), and we therefore concentrated our observations in the twilight hours. Field collected early stages were reared in plastic containers at ambient temperature (25-29°C).

Wild female Caligo illioneus oberon were captured in two traps at the edge of the forest along Pipeline Road. To induce oviposition, captured females were released into an outdoor insectary (3 x 3 x 3 m) inside of which grew Musa sapientum L. and Heliconia latis-patha Benth.(Musaceae), Calathea latifolia (Link) K. (Marantaceae), and three species of unidentified palms (Arecaceae). Females were supplied also with fresh cut leaves of Saccharum spontaneum L. (Poaceae) and Cyrtostachys sp. (Arecaceae), both exotics on which we had observed oviposition by female Caligo illioneus and Opsiphanes sp. respectively (R. B. Srygley pers. obs.).

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Preserved larvae and pupal skins of C. illioneus oberon are in the collection of the Milwaukee Public Museum. The head capsule and pupal skin of C. idomeneus are currently in the Smithsonian Tropical Research Institute, to be relocated to the National Museum of Natural History in the future.

Results

Caligo illioneus oberon Butler

Oviposition behavior and food plants. In the

field, females laid clusters of 9-13 eggs (n = 4 clusters) in a row along the midvein on the underside of medium-aged to old leaf blades of Saccharum sponta-neum, an introduced Asian grass that invaded natural grasslands of Panama during the 1970s. Following its introduction, S. spontaneum gradually replaced the pasture grasses Hyparrhenia rufa (Nees) Stapf and Panicum maximum J acq. (both introduced from Africa) on Pipeline Road (N. Smith pers. comm.). The native larval food plant for C. illioneus is not known in Panama, and our captive females did not oviposit on any of the plants available in the insectary.

Egg (developmental time = 6 days, n = 13). White, spherical, approximately 1.5 mm wide, adorned with vertical ribs; description refers to a cluster of 13 eggs laid 27 August 1994.

First instar (duration = 8 days, n = 4). Head: brown with simple black setae; two dark brown vertical stripes flank epicranial and frontal sutures from apex of head, terminating at approximately halfway the length of the front. Body: translucent green; broad, lemon-yellow middorsal stripe bordered by an irregular reddish-brown stripe that is prominent on the thorax and divided by a thin white discontinuous midline stripe that is more prominent on the thorax than on the abdomen; two thin, lateral, lemon-yellow longitudinal stripes; thoracic and abdominal legs grayish-white; ventral side grayish-white; caudae held separated, reddish to dark brown, each with a black sub-terminal seta arising at one-third to one-half the length of the caudae, and a terminal seta which is white at base and black at tip. The larvae molted synchronously.

Second instar (duration = 5 days, n = 4). Head: dark brown anteriorly, lateral and post-genal regions translucent white; three pairs of scoli: dorsal scoli light brown (approximately half the height of the head), subdorsal scoli whitish (two-thirds the height of the dorsal scoli), lateral scoli whitish (approximately one-third the height of the subdorsal scoli); front dark brown; two thin whitish lines arise from base of dorsal scoli, converge toward and flank epicranial suture, ter-

minating at upper end of front; frontal suture whitish. Body: predominantly green; brown middorsal stripe divided by discontinuous white midline stripe; thoracic and abdominal legs grayish white; ventral side grayish white; caudae held separated, pink with black tips and numerous short white setae. The larvae molted synchronously

Third instar (duration = 5-6 days, n = 4). Head: as in second instar. Body: as in second instar, except for a broad, red lateral line divided by a thin, white spiracu-lar stripe; single dark brown triangular middorsal projection at posterior end of abdominal segment A3. Shed caudae were not eaten after molt to fourth instar. Larvae molted asynchronously.

Fourth instar (duration = 6-7 days, n = 4). Head: patterned in creamy-white and dark brown; dorsal scoli light brown anteriorly and reddish-brown posteriorly at the base (approximately same height as the head); subdorsal scoli creamy-white (approximately two-thirds the length of dorsal scoli); lateral scoli creamy-white (approximately one-half the length of the subdorsal scoli); one pair of creamy-white tubercles below lateral scoli; epicranial suture darkened; front creamy-white with vertical, medial brown stripe; two brown stripes arise from the base of dorsal scoli converge toward and flank epicranial and frontal sutures, terminating halfway down the length of the front; adfrontal region dark brown above stemmatal region; post-genae reddish-brown; base of head reddish-brown from occiput to mandibles; mandibles creamy-white, darkened at the cutting edge. Body: color varied from light mustard to greenish; thoracic segments Tl and T2 with middorsal white midline stripe, flanked by reddish stripes; remaining segments with thin grayish middorsal stripe; large reddish-brown, triangular middorsal projection located at posterior end of abdominal segment A3; very small middorsal projection at posterior end of A5; supra-spiracular white stripe along the entire length of the body; white sub-spiracular stripe on a continuous longitudinal swelling; thoracic and abdominal legs reddish; ventral side red; caudae pale pinkish-brown patterned with reddish-brown dorso-laterally, where pattern develops into thin broken lines. The larvae molted asynchronously and three out of four aggregated at rest.

Fifth instar (duration = 6-7 days, n = 4, Fig. la). Head: as in fourth instar, except for a brown stripe arising from dorsal scoli that flanks the epicranial and frontal sutures, and terminates above antennal socket; head densely covered with short creamy-white setae. Body: predominantly beige; dark brown middorsal stripe runs along entire length of body; on segments Tl, T2, and anterior end of T3, the dark brown mid-

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Fig. 1 Caligo ilUoneus oberon, photographs by R. Srygley; (a) fifth instar (October, 1994); (b) pupa (24 October, 1994). Fig. 2. Caligo idomeneus (Aiello Lot 81-77), photographs by A. Aiello; (a) final instar (2 December, 1981); (b) final instar, head (2 December, 1981); (c) final instar, body detail (2 December, 1981); (d) pupa (23 December, 1981).

dorsal stripe is divided by a thin creamy-white stripe; triangular middorsal projections same as in fourth in-star; body patterned with brown from dorsal midline towards the sides, where pattern develops into thin, broken, longitudinal brown lines; dark brown supra-spiracular line bordered by creamy-white lines; creamy-white subspiracular stripe on a continuous longitudinal swelling; ventral side pink; caudae held separated, light brown at base, becoming dark brown at tip, approxi-

mately two-thirds the length of the head. Larvae molted asynchronously and did not aggregate at rest.

Sixth instar (duration = 14-16 days, n = 3). Head: as in fifth instar, except for four pairs of scoli; additional stripe from dorsal scoli terminating above stem-matal region; dorsal scoli dark brown with white tips and basal creamy-white spots posteriorly; subdorsal and lateral scoli posteriorly brown at base; sub-lateral scoli dark brown posteriorly (one-half of the length of

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the lateral scoli). Body: as in fifth instar, except for small dark brown triangular middorsal projection at posterior end of abdominal segment A2; large triangular middorsal projection at posterior end of A3; and small triangular middorsal projections at posterior ends of A4 and A5; ventral side brown; caudae held separated, slightly longer than head height.

Larval development and behavior. Egg development took 6 days (n = 13) and larval development (from hatching to pupation) took 44-49 days (n = 3). First through fourth instar larvae fed gregariously and generally rested together on the leaf blade. Fifth and sixth instar larvae rested away from each other on the stem of the plant. Late instar larvae found in the field were solitary (R. B. Srygley pers. obs.).

Pupa (duration = 13-15 days, n = 3, Fig. lb). Beige with fine, dark brown cryptic markings, giving the general appearance of a dried, curled and sun-bleached leaf; head with transverse keel at apex; long, black setae located immediately above and on the surface of the eyes; antennae with a black longitudinal stripe, and a black transverse line marking each antennal segment; thoracic segment T2 with a prominent keel along dorsal midline, more developed in female than in male pupae; posterior edge of wing pad forming a crest; prominent hump at the base of wing pad; wing surface with two small silver spots located near base of wing; abdomen with conspicuous long black setae along dorsal midline; abdominal segments A5-10 with dark brown lateral line, A4-10 with brown ventral line; A6 humped; brown middorsal stripe arising at head and terminating at cremaster; abdominal segments with transverse oblique markings that resemble leaf venation. Pupal mass: 2.3 g (n = 1, male).

Caligo idomeneus (L.)

A wandering final instar larva of C. idomeneus was found off the food plant by R. Kimsey at Fort Clayton (Canal Area, Panama) on 1 December, 1981 and reared to adult (Aiello Lot 81-77). The larva had about 15 white fly eggs cemented to the underside of the thorax and head. The eggs were removed with forceps and preserved in 80% ethanol. The oviposition behavior and larval food plants of C. idomeneus are unknown, but the captive mature larva readily accepted Heliconia latispatha Benth. (Musaceae) and Calathea latifolia (Link.) K. (Marantaceae) which it ate for 20 days prior to pupation.

Final instar (n = 1, Figs. 2a-c). Head: beige with brown stripes; three pairs of beige scoli: largest scoli dorsal, clothed in long setae, enlarged towards the pointed apex and abruptly curved outward toward the sides of the head; subdorsal scoli about two thirds the

length of the dorsal scoli, clothed in long setae, gently curved upwards to pointed apex; lateral scoli smallest, about one half the length of the subdorsals, conical; front with dark vertical dash; adfrontal area dark brown; upper section of epicranial suture dark brown; broad stripe lateral to adfrontal area, terminating at stemmatal level with a darker vertical dash; broad stripe from base of dorsal scoli, terminating on stem-mata with a darker vertical dash; dark brown stripe from mid point of inside of each dorsal scolus, terminating at epicranial suture; dark brown stripe along curve from mid point of outside of each dorsal scolus, to sides of head just in front of subdorsal scolus; base of head dark brown from occiput to mandibles. Body: brown, except paler dorsally on abdominal segments A2-A6; broad subspiracular white stripe on A1-A7 with oblique brown intrusions from above, just posterior to each spiracle on A2-A7; four soft laterally flattened triangular middorsal projections, one on each of A3-A6; large oval middorsal spot, lying between the projections of A3 and A4, dark brown with a beige posterior-pointing arrow; caudae held separated, brown, broad.

Pupa (17 days, Fig. 2d). About 4.5 cm long, and 2 cm wide at widest point; beige with fine brown cryptic markings, giving the general appearance of a dried, curled and sun-bleached leaf; head ridged from center of eye to vertex; eye area adorned with stout dark brown upright setae; antennae with a median black stripe for their entire length, and with cross lines set approximately 0.5 mm apart; mesothorax mid-dorsally humped and keeled; a lateral keel begins near the thoracic spiracle, passes along the forewing, parallel to the inner margin, and, at a level with abdominal segment Al, smoothes to become a raised area following the forewing inner margin to the tornus; each forewing bears two white enameled triangles toward the base of the inner margin and just ventral to the wing keel; clear patches and a small dark triangle are found at the midpoint of each mesothoracic leg; on the abdomen a dark brown midventral line terminates at the tip of the cremaster, as does an oblique dark brown line that begins at the spiracle on abdominal segment A6; dorsum adorned with dark brown, upright setae from the mesothoracic hump through abdominal segment A8, on which the setae are somewhat appressed; spiracles narrowly elliptical, and that of A8 is obscure.

Diagnostic characters of early stage morphology. The mature larvae of Central American species of Caligo can be easily diagnosed by head and body color, and by the number and morphology of the head scoli. The mature larva of Caligo eurilochus sulanus Fruh-storfer has a dark tan head adorned with four pairs of

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Journal of the Lepidopterists' Society

Table 1. Larval food plants of the butterfly subfamily Brassolinae (Nymphalidae). Abbreviations: ARE = Arecales, BRO = Bromeliales, CYC = Cyclanthaceae, GEN = Gentianales, gym = gymnosperm, POA = Poales, ZIN = Zingiberales, ovp = oviposition record. References: Aiello, unpbl1; Aiello & Silberglied, 19782; Barcant, 19703; Biezanko et al., 19744; Burmeister, 18735; Casagrande, 19796; Condie, 19767; Cubero, 19858; d'Almeida, 19229; d'Araujo e Silva et al., 196810; DeVries, 198511, 198712; Fontaine, 191313; Harrison, 196314; Hayward, 196915; Moss, unpbl (in Ackery, 1988)16; Miiller, 188617; Rothschild, 191618; Small, unpbl19; Srygley, unpbl20; Srygley & Penz, unpbl21- Stauffer et al., 199322; Urich & Boos, 198123; Urich & Emmel, 199124; Yepez et al., 198525; Young, 197726, 198627; Young & Muyshondt, 197528, 198529.

Buttei"fly species

"palm"

Panicum lanatum Sw. (as capim amargoso)

Arecastrum (as Syagrus) romanzoffianum

(Cham.) Becc. Butia (as Syagrus) capitata (Mart.) Becc. "diversas especies de Palmae" "palmen" Arecastrum (as Cocos) romanzoffianum

(Cham.) Becc. (as geriva) Astrocaryum ayri Mart, (as brejauva) Bactris sp. Butia (as Cocos) eriospatha (C. Mart, ex Drude)

Becc. (as butiazeiro) Cocos nucifera L. (as baba de boi) Cocos nucifera L. (as coqueiro da Bahia) Cocos nucifera L. (as coqueiro anao) Copernicia cerifera Mart, (as carnaubeira) Livistona chinensis (Jacq.) R. Br.

(as pent-sao da China) Livistona rotundifolia (Lamarck) Mart. Phoenix dactylifera L. (as tamareira) Roy stoma (as Oreodoxa) oleracea (Jacq.)

O.F. Cook (as palmeira imperial) Roystonea (as Oreodoxa) regia (Kunth)

O.F. Cook (as palmeira real) Saccharum officinarum L. (as cana de agiicar) Musa sapientum L. (as bananeira) Chaemodora sp. Cocos nucifera L.

Acrocomia aculeata (Jacq.) Lodd. ex Mart. Archontophoenix alexanclrae (Muell.)

H. Wendl. & Drude Arecastrum romanzoffianum (Cham.) Becc. Astrocaryum, spp. Attalea sp. Bactris major Jacq. Bactris spp. Butia (as Cocos) eriospatha (C. Mart.

ex Drude) Becc. (as butiazeiro) Caryota mitis Lour, (as C. plumosa horticola) Caryota urens L.

Chrysalidocarpus lutescens H. Wendl. Chrysalidocarpus lutescens H. Wendl.

(as areca bambu) Cocos nucifera L. Cocos nucifera L. (as coqueiro da Bahia)

Cocos nucifera L. (as coqueiro anao) Copernicia cerifera Mart, (as carnauba) Desmoncus spp. Euterpe spp.

Hyophorbe lagenicaulis (L.H. Bailey) H.E. Moore Livistona chinensis (Jacq.) R. Br. (as pent-sao da China) Livistona sp. Mauritiaflexuosa L. f. Neodypsis decaryi Jumelle Orbignya spp. "palms"

Phoenix canariensis hort. ex Chabaud Phoenix dactylifera L. Phoenix dactylifera L. (as tamareira)

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Table 1. Continued.

Butterfly speci

5.  Caligo arisbe Hiibner

6.  Caligo atreus Kollar

Caligo atreus dionysos Fruhstorfer

7.  Caligo heltrao (Illiger)

8.  Caligo eurilochus (Cramer)

Caligo eurilochus brasiliensis (Felder)

Caligo eurilochus sulanus Fruhstorfer

9.  Caligo idomeneus (L.) 10. Caligo illioneus (Cramer)

Caligo illioneus oberon Butler

Caligo illioneus pampeiro Fruhstorfer 11. Caligo martia (Godart)

12. Caligo memnon (Felder & Felder)

POA Poaceae

ZIN ZIN ZIN ZIN ZIN ZIN ZIN

Cannaceae

Cannaceae

Marantaceae

Musaceae

Musaceae

Musaceae

Musaceae

,29

Phoenix reclinata Jacq.                                                 22

Pntchardia pacifica Seemann and Wendland           22

Ptychosperma macarthurii (A.A. Wendl.)                  22

G. Nicholson Roystonea (as Oreodoxa) oleracea (Jacq.)                  10 (nos 2364,

O.F. Cook (as palmeira imperial)                                2365)

Roystonea (as Oreodoxa) regia (Kunth)                     10 (nos 2364,

O.F. Cook (as palmeira real)                                       2365)

Roystonea oleracea (Jacq.) O.F. Cook                        25

Roystonea regia (Kunth) O.F. Cook                           22

Roystonea venezuelana L.H. Bailey                           22

Sabal mauritiiformis (Id. Karst.) Griseb.                    22

and H. Wendl. Sabal umbraculiferus Mart.                                         22

Scheelea macrocarpa Karsten                                     22

Washingtonia filifera (Linden ex Andre) H. Wendl. 22 Saccharum officinarum L. (as cana de acucar)          10 (no. 2365)

Musa sapientum L. (as bananeira)                              10 (no. 2365)

Ravenala madagascariensis Sonn.                              22

Strelitzia nicolai Regel & Korn.                                  22

Caere                                                                             10 (no. 2369)

Astervgyne martiana H. Wendl. (H. Wendl.)            11

ex Hemsl. Calathea sp. Heliconia spp. Cyclanths Heliconia sp. Musa sp. Canna indica L. Caete

Calathea zebrina (Sims) Lindl. Musa sapientum L. (as bananeira) "plusieurs Musaceae"

Hedychium coronarium J. Konig (as lirio do brejo) Calathea latifolia (Willd. ex Link.) Klotzsch Hedychium sp. Heliconia latispatha Benth. Heliconia latispatha Benth. Musa sapientum L. "plusieurs Masaceas (sic)" Euterpe edulis Mart, (as palmito) banana

Musa sapientum L. (as bananeira) Hedychium coronanum J. Konig (as lirio do brejo) Calathea sp. Heliconia sp. Musa sp. unidentified

Heliconia latispatha Benth. Musa sp.

Hedychium coronarium J. Konig (as lirio do brejo) Heliconia spp.

Musa sapientum L. (as bananeira) Saccharum spontaneum L. Heliconia sp. Musa sp. banana Echinochloa crus-galli (L.) P. Beauv.

(as capim canivao) Pennisetum purpureum Schumach.                            10 (no. 2372)

(as capim elefante) Canna indica L.                                                            1 (lot 85-57)

Canna sp.                                                                      7

Calathea latifolia (Willd. ex Link.) Klotzsch              20 (ovp)

Heliconia latispatha Benth.                                         20, 29

Heliconia spp.                                                               7, 11, 19

Musa sapientum L.                                                       20 (ovp)

Musa sp.                                                                         14, 29

7

11,

12

12

12

6

10(no. 2367)

6

10 (no. 2367)

9

10 (no. 2367)

20

17

1 (lot 80-26)

20 (ovp)

20 (ovp)

9

10 (no. 2368)

18

10 (no. 2368)

10 (no. 2368)

12

12

12

15

1 (lot 81-77)

16

10 (no. 2371)

11

10 (no. 2371)

21

12

12

18

10

no. 2372)

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Table 1. Continued.

Butterfly species

family

Caligo memnon memnon (Felder & Felder)

Caligo memnon telamonius (Felder & Felder)

13.  Caligo oberthurii oberthurii (Deyrolle)

14.  Caligo oileus (Felder & Felder) Caligo oileus scamander (Boisduval)

15.  Caligo placidianus Staudinger

16.  Caligo praxsiodus Fruhstorfer

17.  Caligo prometheus epimetheus

(Felder & Felder)

18.  Caligo teucer (L.)

Caligo sp.

19. Catoblepia amphirhoe (Hiibner)

20.  Catoblepia orgetorix championi Bristow

21. Dasyophthalma rusina (as geraensis) (Godart) Dasyophthalma rusina (Godart)

22. Dynastor darius (F)

Dynastor darius mardonius Fruhstorfer Dynastor darius stygianus Butler

23. Dynastor macrosiris (Doubleday)

24. Dynastor napoleon (Doubleday)

25. Eryphanis aesacus bubocula (Butler)

26. Eryphanis automedon (Cramer)

27. Eryphanis polyxena lycomedon

(Felder & Felder)

. Eryphanis reevesii (Doubleday)

(as bambu comum)

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Table 1. Continued.

Butterfly species

family

Eryphanis reevesii (Doubleday) (as Caligo rivesii)

29.  Narope cyllastros Doubleday Narope cyllastros cyllastros Doubleday

Narope cyllastros testacea Godman & Salvin

30.  Opoptera aorsa (Godart)

(as Opsiphanes aorosa)

31.  Opoptera staudingeri (Godman & Salvin)

Opoptera (as Opsiphanes) staudingeri (Godman h Salvin)

32.  Opoptera (as Opsiphanes) syme (Hiibner)

33.  Opsiphanes bogotanus Distant

Opsiphanes bogotanus bogotanus Distant 34. Opsiphanes cassiae (L.)

Opsiphanes cassiae cassiculus Stichel Opsiphanes cassiae lucullus Fruhstorfer

35. Opsiphanes cassina aiellae Bristow

Opsiphanes cassina fabricii (Boisduval)

POA    Poaceae

POA    Poaceae

POA    Poaceae

POA    Poaceae

POA    Poaceae

POA    Poaceae

POA    Poaceae

POA    Poaceae

POA    Poaceae

POA    Poaceae

POA     Poaceae

POA    Poaceae

POA    Poaceae

POA     Poaceae

ARE    Arecaceae

ZIN      Marantaceae

36. Opsiphanes invirae (Hiibner)

Bambusa vulgaris Schrad. ex J.C. Wendl.                  10 (no. 2383)

(as bambu comum)

Olyra latifolia L. (as taquarinha)                                 10 (no. 2383)

Pennisetum purpureum Schumach.                            10 (no. 2383)

(as capim elefante)

Bambusa sp.                                                                  17

Olyra latifolia L.                                                           17

Bambusa spp.                                                                15, 17

Bambusa (as Guadua) sp.                                            10 (no. 2384)

Bambusa sp. (as bambu)                                              10 (no. 2384)

Bambusa sp.                                                                  12

Bambusa vulgaris Schrad. ex J.C. Wendl.                  10 (no. 2387)

(as bambu comum)

Chusquea longifolia Swallen                                       8

Chusquea sp.                                                                 12

Chusquea sp.                                                                 11

Bambusa (as Guadua) sp.                                            10 (no. 2395)

palms                                                                             11, 12

Calathea inocephala (Kuntze) H.A. Kenn.                 1 (lot 81-41)

& Nicolson

Calathea latifolia (Willd. ex Link.) Klotzsch              20

banana                                                                           18

unidentified                                                                   13

Heliconia sp.                                                                  16 Heliconia sp. and "differentes plantes musacees" 5

Musa sapientum L.                                                       15

Musa sp.                                                                        3

banana                                                                           18

Musa sapientum L. (as bananeira)                              10 (no. 2389)

Cocos nucifera L.                                                          1 (lots 77-76,

82-1, 83-17, 87-3)

Livistona sp.                                                                  1 (lot 91-25)

palm                                                                               1 (lot 95-8)

Acrocomia vinifera Oerst.                                            11, 12

Bactris guineensis (L.) H.E. Moore (as                      28

Bactris minor)

Bactris sp.                                                                      11, 12

Cocos nucifera L.                                                          11, 12, 28

Erythea salvadorensis (H.Wendl. ex Becc.)               28

H.E.Moore (as Brahea saldorensis)

Roystonea regia (Kunth) O.F. Cook                           28 Arecastrum (as Cocos) romanzoffianum (Cham.) 10 (nos 2390,

Becc. (as geriva)                                                           2392)

Arecastrum (as Syagrus) romanzoffianum                 4

(Cham.) Becc.

Butia (as Cocos) eriospatha (C. Mart, ex                   10 (nos 2390,

Drude) Becc. (as butiazeiro)                                       2392)

Butia (as Syagrus) capitata (Mart.) Becc.                  4

Cocos nucifera L. (as coqueiro da Bahia)                   10 (no. 2390)

Copernicia cerifera Mart, (as carnauba)                     10 (no. 2390)

Livistona australis (R. Br.) C. Mart.                           4

Livistona australis (R. Br.) C. Mart.                           10 (nos 2390,

(as pent-sao austral)                                                     2392)

Livistona chinensis (Jacq.) R. Br.                                10 (no. 2392)

(as pent-sao chines)

Livistona chinensis (Jacq.) R. Br.                                4

Livistona rotundifolia (Lamarck) Mart.                     10 (no. 2390)

Palmeira de leque                                                         10 (no. 2392)

Phoenix canariensis hort. ex Chabaud                        4

Prestoea sp.                                                                   8

Raphia sp. (as palmeira ornamental)                          10 (no. 2391)

Roystonea (as Oreodoxa) oleracea (Jacq.)                  10 (no. 2390)

O.F. Cook (as palmeira imperial)

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Journal of the Lepidopterists' Society

Table 1. Continued.

Butterfly speci

family

Opsiphanes invirae amplificatus Stiche]

(as O. i. remoliatus) Opsiphanes invirae amplificatus (Stiche]) Opsiphanes invirae amplificatus Stichel

(as ampliplacita) Opsiphanes invirae cuspidatus Stichel

37.  Opsiphanes merianae Stichel

38. Opsiphanes quiteria (Cramer) Opsiphanes quiteria hadius Stichel Opsiphanes quiteria meridionalis Staudinger Opsiphanes quiteria meridionalis (as philon)

Staudinger

Opsiphanes quiteria quirinus Godman & Salvin

39. Opsiphanes tamarindi Felder & Felder Opsiphanes tamarindi sikyon Fruhstorfer

Opsiphanes tamarindi tamarindi Felder & Felder

Opsiphanes sp.

Cycas circinalis L. (as palmeira de jardim)              10 (no. 2390)

[dubious record]

Musa sapientum L. (as bananeira)                          10 (nos 2390,

2392)

"giriva and palms"                                                  18

Phoenix sp.                                                            15

Musa sapientum L. (as bananeira)                          10 (no. 2386)

Bactris major Jacq.                                                1 (lot 82-44)

palms                                                                    12

ornamental palm                                                   3

Arecastrum romanzoffianum (Cham.) Becc.            15

Bactris sp.                                                             1 (lot 81-74) Chrysalidocarpus (as Areca) lutescens H. Wendl. 10 (no. 2393)

Astrocaryum ayri Mart, (as brejauva)                     10 (no. 2394)

Euterpe edulis Mart, (as palmito)                            10 (no. 2394)

Chrysalidocarpus lutescens H. Wendl.                    8

Cocos nucifera L.                                                  26

Geonoma sp.                                                         8

palms                                                                    12

Prestoea allenii H.E. Moore                                   8

Musa sapientum L. (as bananeira)                          10 (no. 2396)

Musa sp.                                                                14, 17

Heliconia collinsiana Griggs                                   28

Heliconia latispatha Benth.                                    28

Musa sp.                                                               28

Canna indica L.                                                     1 (lot 84-9)

Heliconia latispatha Benth.                                    1 (lots 80-40,

80-43)

Heliconia sp.                                                         12

Musa sp.                                                                12

Cocos nucifera L. (as coqueiro anao)                      10 (no. 2385)

Cyrtostachys sp.                                                    20

Heliconia latispatha Benth.                                    1 (lot 82-18)

Musa sapientum L. (as bananeira)                          10 (no. 2385)

scoli, dark brown body with six middorsal projections (see figs. 6 and 7 in Malo & Willis 1961 p. 532). That of C. atreus dionysos Fruhstorfer has a tan colored head with fine vertical striations and three pairs of scoli,

with the dorsal pair enlarged at tip and curved outward (see fig. 32, El in DeVries 1987 p. 248), a tan colored body with many fine striations on dorsum, and five middorsal projections (DeVries 1987). The head cap-

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

Volume 53, Number 4

151

sule of the mature larva of C. memnon is banded with tan and dark brown with four pairs of scoli (see fig. 5 in Young & Muyshondt 1985 p.162; fig. 32, E2 in De-Vries 1987 p. 248), with the dorsal pair enlarged at tip. The body is light brown with a dark brown middorsal stripe, dark brown striations, and six middorsal projections (note that fig. 5 in Young & Muyshondt 1985 p. 162, and fig. 31 F in DeVries 1987 p. 248, do not portray the same body color pattern). The mature larva of C. illioneus has a head patterned in brown and creamy white adorned with three pairs of scoli plus a lateral tubercle, and the body is beige with a dark brown middorsal stripe and four middorsal projections (Fig. la). That of C. idomeneus has a beige head patterned with brown, three pairs of scoli with the dorsal pair enlarged at tip and curved outward (Fig. 2b). The body is brown with a lighter colored area dorsally and a large oval middorsal spot between A3 and A4, a broad white subspiracular stripe, and four middorsal projections (Fig. 2a). The pupae of all species are very similar, and those of C. illioneus and C. idomeneus seem to differ only in the size of the white triangular marking at the base of the wing (more prominent in C. idomeneus, Fig. 2d). Early stages of C. oileus have never been formally described.

Larval food plants of the Brassolinae. It is well known that brassoline immatures are restricted to monocotyledonous plants (Ehrlich & Raven 1965, Ackery 1988, Table 1), but little correlation has been found between plant use and brassoline classification (Ackery 1988). We found that larval food plants include four of the eight monocot superorders (Table 1), a distribution suggesting that brassolines are generalist monocot feeders. However, all food plant records together indicate that the majority of species feed on plants in the families Arecaceae, Musaceae, and Poaceae (Tables 1 and 2). Therefore, the apparent lack of correspondence between plant use and brassoline classification should be reexamined.

Available records are sufficient to show that brassoline genera vary both in diet breadth and their association with monocot families (Table 2; see Stichel 1909 and Bristow 1981, 1982, 1991 for taxonomic classification of the butterflies). For instance, although individual species of Caligo have been reported to feed on 1-4 plant genera in 1-4 families, collectively Caligo has a larval food plant range that includes 11 genera in 7 families (Table 2, see also Ackery 1988) suggesting multiple events of host colonization during its evolutionary history. Similar patterns occur in Eryphanes and Opsiphanes: species of Eryphanes typically feed on Poaceae, except for E. aesacus which has also been found on Arecaeae; and Opsiphanes tend to associate

with Arecaceae and Musaceae, except for O. bo-gotanus and O. tamarindi, whose food plant range also includes Marantaceae and Canaceae respectively Available information suggests that other brassolines are restricted to a single plant family (Blepolenis and Catoblepia on Arecaceae, Narope and Opoptera on Poaceae, Dynastor on Bromeliaceae). Noteworthy is that the food plant range of the putative basal genus Brassolis includes Arecaeae, Poaceae and Musaceae; the plant families upon which most brassolines feed as immatures. Although patterns of host association can be recognized at the generic level, their examination in an evolutionary context awaits a well supported phy-logeny for this group of butterflies.

We hope that the summary presented here encourages research aimed at furthering our understanding of the patterns of food plant utilization and evolution in brassoline butterflies.

Acknowledgments

We are grateful to: R. Kimsey, who brought in the C. idomeneus larva; C. Galdames, for identifying plants collected by R.B. Srygley and C.M. Penz; W. Hahn for guidance on palm nomenclature; P.J. DeVries, G. Lamas and A. Neild for their help and suggestions; and P. Ackery, P. J. DeVries and D. Jenkins for comments on the manuscript. We also acknowledge InReNaRe (now A.N.A.M.) for granting permission to conduct research and collect butterflies in Panama, and the Smithsonian Tropical Research Institute (STRI) for providing a vehicle to conduct research. Financial support was provided by STRI (to C. Penz), National Geographic Society (to R.B. Srygley), and National Science Foundation DEB98-06779 (to C.M. Penz).

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Received for publication 23 April 1999; revised and accepted 10 October 1999