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tively). After moving, the larvae typically became inactive again, only feeding 27.8% in the laboratory and 25.5% in the field. However, in the field, encountering food is not guaranteed, and these percentages do not take into account distance traveled. Once a larva in the field had fed, it would either remain inactive or move with equal probability. However, larvae in the laboratory would remain inactive after feeding 81.3% of the time.

Larvae would eat the violet leaves rapidly during feeding bouts and, when not feeding, larvae would hide in clumps of grass or walk. McCorkle and Hammond (1988) observed similar feeding behavior for S. zerene hippolyta. The fifth instar larva that was released back to the prairie walked in a curved path (25.40 m over a 24 hour period) and fed only on violet plants that were in its path (although it walked past violet plants as close as a centimeter away and apparently did not perceive them). Wind direction did not seem to matter as the larvae walked equally close to violets up and down wind without noticing them. As the sun set, the larva walked and fed progressively less, until it became inactive at the base of a grass clump and remained there until the following morning. The larvae studied in the laboratory and greenhouse displayed a similar diurnal feeding pattern. The inactivity associated with nightfall may be due to a reduction in temperature, however temperature does not drop substantially as soon as the sun sets in eastern Kansas, leading us to believe that light is a more important cue for activity than temperature.

This study of a small sample of S. idalia larvae in Kansas tall-grass prairie indicates that they do not forage entirely at night. More study is needed covering a larger geographic region in order to determine how widespread this behavior is within S. idalia and the genus Speyeria.

Acknowledgments

We thank Alberto B. Broce and Matt R. Whiles, Department of Entomology, Kansas State University and William F. J. Parsons, Department of Entomology, University of Wisconsin for helpful comments on earlier versions of this manuscript. Voucher specimens (Lot # 076) are deposited in the KSU museum of Entomological & Prairie Arthropod Research. This is contribution No. 99-515-J of the Kansas Agricultural Experiment Station, Kansas State University, Manhattan, Kansas 66506.

Literature Cited

Debinski, D. M. & L. Kelly. 1998. Decline of Iowa populations of the regal fritillary (Speyeria idalia) Drury. J. Iowa Acad. Sci. 105:16-22.

Ehrlich, P. A. & A. H. Ehrlich. 1961. How to know the butterflies. Wm. C. Brown Co., Dubuque, Iowa. 262 pp.

Hammond, PC. & D. V. McCorkle. 1983. The decline and extinction of Speyeria populations resulting from human environmental disturbances (Nymphalidae: Argynninae). J. Res. Lepid. 22:217-224.

Holland, W J. 1931. The butterfly book. Second ed. Doubleday. New York, New York. 424 pp.

Kopper, B. J. 1997. Behavioral ecology of the regal fritillary, Speyeria idalia (Drury) (Lepidoptera: Nymphalidae), in Kansas tall-grass prairie: reproductive diapause, factors influencing ovipo-sition site selection, and larval foraging behavior. M.S. Thesis, Kansas State University, 105 pp.

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

McCorkle, D. V. & P. C. Hammond. 1988. Biology of Speyeria zerene hippolyta (Nymphalidae) in a marine-modified environment. J. Lepid. Soc. 42:184-195.

Royer, A. R. 1988. Butterflies of North Dakota. Minot State University, Minot, North Dakota. 192 pp.

Scudder, S. H. 1889. Butterflies of the eastern United States and Canada with special reference to New England. Published by the author. Cambridge, Massachusetts. Vol. 1, pp. 1-776; vol. 2, pp. 767-1774; vol. 3, pp. 1775-1958, pis. 1-89, 3 maps.

Brian J. Kopper,' 2 David C. Margolies, and Ralph E. Charlton, Department of Entomology, Kansas State University, Manhattan, Kansas 66506-4004, USA.

1 Present address: Department of Entomology, University of Wisconsin, 237 Russell Labs, Madison, Wisconsin 53706, U.S.A.

2 Corresponding author.

Submitted for publication 14 August 1999 , revised and accepted 25 July 2000.

Journal of the Lepidopterists' Society 54(3), 2001, 97-100

BIOLOGY OF ADELPHA MYTHRA FEEDING ON ASTERACEAE, A NOVEL PLANT FAMILY FOR THE NEOTROPICAL LIMENITIDINAE (NYMPHALIDAE), AND NEW DATA ON ADELPHA 'SPECIES-GROUP VII"

Additional key words: Adelpha syma, Adelpha cocala, life history, Rosaceae, Rubiaceae.

The Neotropical genus Adelpha Hiibner (Nymphalidae) includes about 85 species (Keith Willmott pers. comm.) spread from western USA to Uruguay, and occurring in a wide variety of habitats and vegetation types (Aiello 1984). Species determination is very difficult in some Adelpha groups, and the natural divisions of the genus are not yet fully resolved, although a number of species relationships have been proposed on the basis of the immatures (Aiello 1984). Unfortunately, immatures are known for only 32 species of Adelpha, solely 21 of which have some portion of the early stages illustrated. Thus, although a cladistic analysis of the genus is needed, it would be impossible at this time. Because information on additional species is essential to a better understanding of the genus (DeVries 1987; Aiello 1991), it is important that any new data about Adelpha immatures be reported (Otero & Aiello 1996).

This paper describes the immature stages of Adelpha mythra (Go-dart 1824) and A. syma (Godart 1824), reports their larval host plants, and discusses the position of both species within Adelpha, based on their immatures.

Study sites and methods. Adelpha mythra, a montane species in

southeast Brazil, is one of 16 species of Adelpha known in the Santa Genebra Forest Reserve (22°44'S, 47°06,W, altitude 600-630 m), a 250 ha fragment of semideciduous forest in Campinas, Sao Paulo State, SE Brazil (see additional information on the area in Morellato and Leitao-Filho 1995). In January 1999, a female A. mythra was observed there ovipositing on the scandent vine, Mutisia coccinea St. Hil. (Asteraceae). The egg did not hatch, so this very unusual "record" was thought to be an oviposition mistake of this female. However, from February to April 1999, A. mythra was reared from first to fourth instars collected on the same plant species and also on Bathysa meridionalis (Rubiaceae) in several parts of the Serra do Japi (23°11'S, 46°52'W), a mountain range (700-1300 m altitude) covered by semideciduous forest, in Jundiai, Sao Paulo State, SE Brazil (Brown 1992). Immatures of Adelpha syma were also found on Rubus (Rosaceae) in the Serra do Japi, and immatures of A. cocala were discovered feeding on a Rubiaceae in the Parque Ecologico do Voturua (46°22,W, 23°57'S, altitude 20-100 m), a 200 ha fragment of lowland subtropical rainforest in the city of Sao Vicente, coastal Sao Paulo State, SE Brazil.

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Journal of the Lepidopterists' Society

A. syma

LEG T1 T2_T3 A1 A2 /A3-6 A7 A8 A9 A10 PROLEG

Fig. 1. Fifth instar larvae of Adelpha syma (above) and A. mijthra (below)

SIMPL SCOLUS

ANAL -PROLEG

/^T1         T2        T3 A1        A2 /A3-6 A 7 A 8 A9 A10

LEG                                                          PROLEG

Fig. 2. Above, Chaetotaxy of first instar larva of Adelpha syma. Below, Distribution of scoli in a fifth instar larva of A. mijthra.

Larvae were reared in plastic cages cleaned daily, following Fre-itas (1991). Adults, head capsules and pupal skins are in the collection of the first author. Larval food plant vouchers, identified by Dr. Jorge Tamashiro, have been deposited in the herbarium of the Uni-versidade Estadual de Campinas.

Descriptions of immatures and host plants of Adelpha mythra. The only egg observed was greenish brown, sculptured with hexagonal pits, with spines arising from the pit junctions, consistent with eggs described for other species of Adelpha. The egg was placed on the upper surface of the leaf, near the apex. It was laid rapidly by a startled female, and did not hatch.

First and second instars were not described in detail. The third instar had conspicuous head scoli and a faintly visible variegated pattern that, with few changes, was maintained and intensified through the remaining larval stadia. The fifth instar (Fig. 1) was variegated green, with a pattern of oblique lateral stripes. The green areas changed to light orange as the larvae neared pupation. In the final larval stadium, the body scoli were short and thick, with a dense covering of spines; the scoli on A2 were arched posteriorly. The distribution of the scoli (Fig. 2) was the same as for most Adelpha species. The mature larva was about 25 mm long.

The first through fourth instars initiated feeding at the apex of a leaflet, leaving the midvein intact and extending it with fecula and silk to form "frass chains" (Aiello 1984). They rested upon these structures when not feeding. In addition, they attached dead leaf fragments and clumps of fecula to the base of the chains. When feeding on Bathysa meridionalis, larvae built "frass chains" on other parts of the same large leaf.

The pupa (Fig. 3A) showed an elongated general profile (about 17 mm long), with segment A2 produced and curved anteriorly, and segment T2 pointed and directed posteriorly. The head horns were pointed and shaped like tiny asymmetrical leaves, curving out from the sides of the head. The general color was brown, with dark lines on the wing pads and no reflective areas.

Mutisia coccinea (Asteraceae), a scandent shrub common in wet second growth habitats, was observed as the larval food plant of Adelpha mythra both in Santa Genebra and in the Serra do Japi. In the latter site, A. mythra was also reared on Bathysa meridionalis L. B. Smith & Downs (Rubiaceae), a plant with enormous leaves (up to lm in length and width), of montane habitats, where it occurs most often near watercourses. Bathysa meridionalis is used by many other Adelpha species in that site (Brown 1992 and AVLF pers. obs.).

Immatures and host plants of Adelpha syma. The egg was

greenish brown, with sculpturing and ornamentation as described for other species of Adelpha, placed on the upper surface of the leaf, near the apex. The r eight and diameter were about 0.8 mm; the duration was not determined.

The first instar was entirely brown, with pale body setae about 0.06 mm long, arranged as in Fig. 2. The head capsule was pale brown without ornamentation. The final body length was 4 mm. The second instar was entirely brown, with a spiny brown head. The body bore short stubby scoli. The final body length was 6 mm. The third and fourth instars showed a clearly visible variegated pattern, with head spines longer. The final lengths were 9 mm in the third and 15 mm in the fourth larval stadia. The fourth and fifth instars had the same general pattern. The fifth instar (Fig. 1) was variegated with green, cream and brown, showing a general pattern of oblique lateral stripes. The green areas change to light orange before pupation. The body scoli were short and thick, with dense spines, and the scoli of A2 arching posteriorly. The distribution of the scoli was the same as in A. mythra. The fully grown larva was 25 mm long. Data on head capsule widths for all instars are in Table 1. The first through fourth instars constructed frass chains and had a behavior similar to that described for A. mythra.

The pupa (Fig. 5B) showed an elongated general profile (about 18 mm long), with segment A2 projecting and curved anteriorly, and segment T2 pointec. and directed posteriorly. The head horns were very small and pointed, curved out from the sides of the head. The general color was brown, with dark lines on the wing pads and no reflective areas.

The host plant in the Serra do Japi was Rubus hrasiliensis Mart. (Rosaceae), a common blackberry of sunny second growth habitats, especially in montane sites. Rubus rosifolius Sm., an introduced species, is also used as larval ood in other montane sites in SE Brazil.

Positions within Adelpha. The scolus shape and the general pattern of the larvae, and the general form of the pupae, suggest

Table 1. Head capsule widths of Adelpha syma

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that both Adelpha mythra and A. syma belong to the Species-Group VII of Aiello (1984). The immatures of both species are very similar to those of A. cocala, the main difference being that the A2 process of the pupa is much longer in A. mythra and more arched in A. syma than in A. cocala (Fig. 3C). In A. mythra, the length of the A2 projection approaches the condition observed in A. phylaca (as described by Miiller 1886) (Fig. 3D), a species belonging to Group II of Aiello (1984). However, the T2 projections of A. mythra and A. syma are sloped posteriorly, and not curved upward as in Group II pupae, giving pupae in the two groups distinctive general appearances. The pupal head horns of A. mythra (Fig. 3A) are similar in shape to those of A. cocala (Fig. 3C), but are farther apart at their bases and are more tapered. The head horns of A. syma (Fig. 3B) are similar to those of A. phylaca (Fig. 3D), but are more curved.

Additional species of Adelpha need to be reared in order to clarify the scenario based on the morphology of the immatures and to make a cladistic analysis possible. Some species groups are based on only one, two or three species, and the immatures of many common species remain to be discovered, or their descriptions are not sufficiently detailed to permit assignment to a species-group. Additional descriptions of Adelpha immatures with figures are important, especially when they show apparent deviations from the eight known species groups (Aiello 1984, Otero & Aiello 1996).

Host plant use. Among the Nymphalidae, the association with Asteraceae as larval food plants is found in only a few groups (especially Melitaeinae and Acraeinae) (Ackery 1988, Freitas 1991).

I                                I

5 mm

Fig. 3. Pupae of Adelpha mythra (A), A. syma (B), A. cocala scale; the bar means 7.3 mm).

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The record of Asteraceae as a larval host of the Limenitidinae represents a new plant family for neotropical Adelpha. Larvae of Adelpha species have been recorded as feeding on Aquifoliaceae, Asteraceae, Bombacaceae, Caprifoliaceae, Combretaceae, Ericaceae, Fagaceae, Flacourtiaceae, Icacinaceae, Malpighiaceae, Melastom-ataceae, Moraceae, Ochnaceae, Piperaceae, Rosaceae, Rubiaceae, Tiliaceae, Ulmaceae, Urticaceae, Verbenaceae and Vochysiaceae (Jones & Moore 1883, Miiller 1886, Moss 1933, Biezanko et al. 1966, Aiello 1984, 1991, DeVries 1987, Ackery 1988, Brown 1992, Otero & Aiello 1996, Diniz & Moraes 1997, Constantino 1998, and this work). Some themes may be recognized in the different species-groups (Aiello 1984). Basically the species of Adelpha can be sorted into rubiaceous feeders and non-rubiaceous feeders. Four examples of species feeding on both Rubiaceae and other families have been reported (A. serpa, A. boreas tizona, A. syma and A. cocala), and A. mythra is the fifth recorded case. Although interesting, this pattern must be considered with caution, because some plant identifications need to be confirmed by additional field observations.

Acknowledgments

This study was conducted as part of a Ph.D. Thesis on Nymphalidae systematics and evolution. K. Willmott, Deane Bowers and an anonymous referee reviewed and made helpful critics and comments on the manuscript. This research was supported by fellowships from the Brazilian CNPq to AVLF and KSB.

and A. phylaca (D) (A. phylaca redrawn from Aiello, 1984, in a different

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Literature cited

Ackery, P. R. 1988. Hostplants and classification: a review of nymphalid butterflies. Biol. J. Linn. Soc. 33:95-203.

AlELLO, A. 1984. Adelpha (Nymphalidae): deception on the wing. Psyche 91:1-45.

Aiello, A. 1991. Adelpha ixia leucas: Immature stages and position within Adelpha (Nymphalidae). J. Lepid. Soc. 45(3):181-187.

Biezanko, C. M., A. Ruffinelli & C. S. Carbonell. 1966. Lepi-doptera del Uruguay. Notas complementarias. III. Bole tin Facul. Agron. Univ. Repub. (Montevideo) 91:1-53.

Brown, K. S. JR. 1992. Borboletas da Serra do Japi: Diversidade, habitats, recursos alimentares e variagao temporal, pp. 142-187, 18 figs. In L. P. C. Morellato (ed.), Historia natural da Serra do Japi. Ecologia e preservagao de uma area florestal no sudeste do Brasil. Campinas, Editora da Unicamp/Fapesp.

Constantino, L. M. 1998. Butterfly life history studies, diversity, ranching and conservation in the Choco rain forests of western Colombia (lnsecta: Lepidoptera). Shilap Revta. Lepid. (Madrid) 26 (101):19-39.

DeVries, P. J. 1987 The butterflies of Costa Rica and their natural history. Princeton University Press, Princeton, New Jersey.

Diniz, I. R. & H. C. Moraes. 1997. Lepidopteran caterpillar fauna of cerrado host plants. Biodiversity and Conservation 6:817-836.

Freitas, A. V. L. 1991. Variagao morfologica, ciclo de vida e sis-tematica de Tegosa claudina (Eschscholtz) (Lepidoptera,

Nymphalidae, Melitaeinae) no estado de Sao Paulo, Brasil. Revta. bras. Ent. 35:301-306.

Jones, E. D. & F. Moore. 1883. Metamorphoses of Brazilian Lepidoptera from San Paulo, Brazil, with nomenclature and descriptions of new forms. Second series. Proceedings of the Literary and Philosophical Society of Liverpool 37:229-259.

Morellato, L. P C. & H. Leitao-Filho. 1995. Introdugao, pp. 15-18. In L. P. C. Morellato & H. Leitao-Filho (eds.), Ecologia e preservagao de uma floresta tropical urbana. Reserva de Santa Genebra. Campinas, Editora da Unicamp.

Moss, M. 1933. Some generalizations on Adelpha, a neotropical genus of nymphalid butterflies of the group Limenitidi. Novit. Zool. (Tring) 39(1): 12-20.

Muller, W. 1886. Sudamerikanische Nymphalidenraupen: Versuch eines naturlichen Systems der Nymphaliden. Zoologische Jahrbucher (Jena) 1:417-678.

Otero, L. D. & A. AlELLO. 1996. Descriptions of the immature stages of Adelpha alala (Nymphalidae). J. Lepid. Soc. 50(4):329-336.

Andre Victor Lucci Freitas, Keith S. Brown, Jr., Museu de Historia Natural, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109 13083-970, Campinas, Sao Paulo, Brazil and Annette Aiello, Smithsonian Tropical Research Institute,Box 2072, Balboa, Ancon, Panama.

Received for publication 9 August J 999; revised and accepted 2 October 2000.