Braz. J. Biol., 62(4A): 657-663, 2002 HOST-PLANT USE AMONG BUTTERFLY SPECIES 657 HOST PLANT USE AMONG CLOSELY RELATED Anaea BUTTERFLY SPECIES (LEPIDOPTERA, NYMPHALIDAE, CHARAXINAE) QUEIROZ, J. M. Departamento de Zoologia, Universidade Estadual de Campinas, C.P. 6109, CEP 13083-970, Campinas, SP, Brazil Correspondence to: Jarbas Mar��al Queiroz, Departamento de Ci��ncias Ambientais, IF, Universidade Federal Rural do Rio de Janeiro, Rod. BR 465, km 7, CEP 23850-230, Serop��dica, RJ, Brazil, e-mail: jarbas@ufrrj.br Received August 8, 2001 ��� Accept February 5, 2022 ��� Distributed November 30, 2002 (With 1 figure) ABSTRACT There is a great number of Charaxinae (Lepidoptera: Nymphalidae) species in the tropics whose larvae feed on several plant families. However the genus Anaea is almost always associated with Croton species (Euphorbiaceae). This work describes patterns of host plant use by immature and adult abun- dance on different vertical strata of sympatric Anaea species in a forest of Southeastern Brazil. Quantitative samples of leaves were taken in April/1999 and May/2000 to collect eggs and larvae of four Anaea species on C. alchorneicarpus, C. floribundus and C. salutaris in a semideciduous forest. Sampled leaves were divided into three classes of plant phenological stage: saplings, shrubs and trees. The results showed that the butterfly species are segregating in host plant use on two scales: host plant species and plant phenological stages. C. alchorneicarpus was used by only one Anaea species, whereas C. floribundus was used by three species and C. salutaris by four Anaea species. There was one Anaea species concentrated on sapling, another on sapling/shrub and two others on shrub/tree leaves. Adults of Anaea were more frequent at canopy traps but there were no differences among species caught in traps at different vertical positions. This work supplements early studies on host plant use among Charaxinae species and it describes how a guild of closely related butterfly species may be organized in a complex tropical habitat. Key words: Nymphalidae, Charaxinae, Anaea, Memphis, Croton. RESUMO Uso de plantas hospedeiras entre esp��cies proximamente aparentadas de borboletas Borboletas Anaea (Lepidoptera: Charaxinae) est��o freq��entemente associadas a esp��cies de Croton (Euphorbiaceae). Este trabalho descreve a utiliza����o de plantas hospedeiras por uma guilda de borboletas em uma floresta do Sudeste brasileiro. Folhas de tr��s esp��cies de Croton foram amostradas em 1999/2000 para a coleta de ovos e larvas de quatro esp��cies de Anaea. Paralelamente, foram colocadas armadilhas para adultos em duas diferentes posi����es verticais (1 m e 5 m) para compara����o do n��mero de indiv��duos capturados entre as esp��cies. Os resultados demonstraram que a guilda de borboletas segrega na utiliza����o de plantas hospedeiras em duas escalas: esp��cie hospedeira e est��gio fenol��gico das plantas. Croton alchorneicarpus foi utilizada como hospedeira apenas por A. ryphea C. floribundus foi utilizada por tr��s esp��cies de Anaea e C. salutaris, por quatro esp��cies. Anaea otrere foi mais abundante em plantas jovens, A. ryphea predominou em plantas jovens e arbustos e A. appias e A. arginussa apresentaram maiores densidades em folhas de arbustos e ��rvores de Croton. Os adultos de Anaea foram capturados com maior freq����ncia nas armadilhas postas na copa das ��rvores, todavia n��o houve diferen��a significativa entre as esp��cies. Este trabalho

Braz. J. Biol., 62(4A): 657-663, 2002 658 QUEIROZ, J. M. suplementa os dados sobre utiliza����o de plantas hospedeiras por Charaxinae e descreve como uma guilda de borboletas proximamente aparentadas pode estar organizada em um ambiente tropical complexo. Palavras-chave: Nymphalidae, Charaxinae, Anaea, Memphis, Croton. forest canopy (DeVries, 1988 DeVries et al. 1997 DeVries et al. 1999) but stratification of immatures was not investigated. This work describes patterns of host plant use by immatures of sympatric Anaea species that feed on Croton and compares the abundance of adults between understory and canopy strata in a forest of Southeastern Brazil. METHODS Three Croton species are commonly found along trails in the semideciduous forest of the Serra do Japi Reserve (23o16���S, 47o00���W) near Jundia��, S��o Paulo, Brazil, immediately after Japi. The place is a continuous forest of 300 km2 with an average canopy height ranging between 5 and 15 m above the ground. Croton floribundus Spreng is the most widespread and abundant species and C. salutaris Baill is the rarest species and occurs in discrete patches. A third species, C. alchorneicarpus Croizat, may be very abundant on some trails but is rare on others. Eggs of Anaea species (1.0 mm diameter) are frequently laid singly on the underside of leaves. Although they are very similar in appearance, there are some variations of color among species. Larvae from the first and second instar are quite similar to the naked eye, but from the third instar onward different species can be easily identified in the field. First to third instar larvae of Anaea species construct frass chains, adding feces with silk to the leaf vein, on the tip of which they rest. Fourth and fifth instar larvae may roll the leaf to form a tubular shelter, holding it closed with silk, and hide inside when not feeding (see Muyshondt, 1974, 1975a,b, and Caldas, 1994, for descriptions of immatures). I used a stratified sample of leaves in April/ 1999 and May/2000 to collect eggs and larvae of Anaea on the three Croton species in Japi. I took the samples at four different sites in the forest. These sites were as far as 4 km from each other. INTRODUCTION Groups of closely related butterfly species are almost always confined to related larval host plants (Gilbert & Singer, 1975). There is a great number of Charaxinae (Lepidoptera: Nymphalidae) species in the tropics whose larvae feed on several plant families (DeVries, 1987). Ackery (1988) showed that many species feeding on Euphorbiaceae belong to the Anaeini tribe. The genus Croton, pantropical in distribution and having more than 800 described species (Webster, 1994), is the main larval foodplant and hosts several species from Anaeini in the genera Hypna, Anaea, Polygrapha, Fountainea and Memphis (Ackery, 1988). The use of generic names in Charaxinae va- ries considerably among authors and I will follow Comstock���s (1961) nomenclature, which considered Memphis H��bner as a subgenus of Anaea H��bner (see Caldas, 1994). The genus Anaea includes most Neotropical Charaxinae (e.g., DeVries, 1987, 1988). Biological studies for several Anaea species can be found in Muyshondt (1974, 1975a,b) and Caldas (1991, 1994). Up to twelve Anaea (Memphis) species have been recorded feeding on Croton (see Ackery, 1988), but in Brazil only A. ryphea Cramer has been studied more extensively (see Caldas, 1991, 1994, 1995). Tropical adult butterflies may exhibit stratified distributions between canopy and understory (DeVries, 1988 DeVries et al., 1997, 1999 Beccaloni, 1997). In some cases, larvae of butterfly species may be more abundant on saplings than on adults of the same host plant species (see Karban, 1987). Several others immatures of herbivore species have shown vertical stratification on their host plants (e.g., Brown et al., 1997 and included references). In Ithomiinae the flight height of adults may be correlated to the occurrence of their larval host-plants (Beccaloni, 1997). Charaxinae species have often been trapped in the