The palatability and the ability of neotropical butterflies to escape after being detected, attacked and captured by wild kingbirds (Tyrannus melancholicus Vieillot), was investigated by the release of 668 individuals of 98 butterfly species close to the birds, during their usual feeding activities. Most of the butterflies were attacked and eaten. Only the troidine swallowtails (Parides and Battus; Papilionidae) were consistently rejected on taste and elicited aversive behaviours in birds. Most other aposematic and/or mimetic species in the genera Danaus and Lycorea (Danainae), Dione, Eueides and Heliconius (Heliconiinae), Hypothyris, Mechanitis and Melinaea (Ithomiinae), Biblis, Callicore and Diaethria (Limenitidinae) were generally eaten. Cryptic and non-mimetic species were always attacked and, if captured, they were also eaten. All Apaturinae, Charaxinae, Nymphalinae, Hesperidae, most Limenitidinae, Heliconiinae (Agraulis, Dryas, Dryadula and Philaethria) and Papilionidae (Eurytides, Heraclides and Protesilaus) were in this group. Results indicate that the learning process in kingbirds may demand a large mortality in prey populations, even among species generally accepted as unpalatable and aposematic. They also support the assertion that escaping ability and unpalatability evolved in butterflies as alternative strategies to avoid predation by birds. Mimetic relationships among several species are discussed. Evidence for the evolution of aposematism not related to unpalatability, but to escaping ability, was found for two hardto-catch Morpho species.

Bwbgical Journal ~f t t u Linnean society (1996), 59: 351-365. With 2 figures j-/ 7 4 PbA Palatablility and esc-ing ability in Neotropical butted&: tests with wild kingbirds (Tfiannus melan2holicu~Tyrannidae) 7 CARLOS E.G. PINHEIRO Department of elicited avers% beham3urs in birds. Most o G r a p o s a t i c and/or mimetic species in the gezra Danaur and Lycorea (Danainae), fitm,~uci& and Heliconiuc (Heliconiinae), H@Thyris, Mechanilir and M d s a (IthomGae), Bi?%is, Callipre an2 Dirulhria &en&dinaqwere g e n e a y e a t e n 2 P t i c and n&nimeticypecies weE always attacked a n c i f captured, they were also eaten. AU Apaturinae, Charaxinae, Nymphalinae, Hesperidae, most Limenitidinae, Heliconiins -&raulis, &as, & -& I .and F%i $&i a) and PaFdionidae (Eu-&h, Hcratlidw and%oorcsilnils) were in this group. Results indicze that thhearning p r o c e G kin&ir%may deEand a large-mortality in prey opulations, even among species gy accepted as unpetable and aposetmatic. Thr'o supp/ort -f i e assertion that esca&g ability and unpalsbility evolved in butterflies as alternative strategies to avoid predation by bi r r & Mimetic relationships among several specigare discussed. Evidence for the evolution of aposzatism not a a t e d to unpalatability, but to escaping ability, was found for two hardto-catch Morpho species. h -\ 01996 The Linnean Society of London ADDITIONAL

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The selective advantage of Miillerian mimicry in nature was investigated by releasing live mimetic and nonmimetic butterflies close to wild, aerial-hunting tropical kingbirds ( Grannw melancholzcus) and cliff-flycatchers (Hirundinea firruginea) in three Amazon habitats (rain forest, a city, and "canga" vegetation). Only mimetic butterflies elicited sight-rejections by birds, but protection conferred by mimicry was restricted to sites in which both predators and mimics co-occurred, as in the case of six mimicry rings at a forest site and two at a city site. Most other Miillerian mimics released at city and canga vegetation were heavily attacked and consumed by birds. These results appear to reflect the birds' previous experiences with resident butterfly faunas and illustrate how birds' discriminatory behavior varied among habitats that differed in butterfly species and mimicry ring composition.

Birds are able to recognize and learn to avoid attacking unpalatable, chemically defended butterflies after unpleasant experiences with them. It has also been suggested that birds learn to avoid prey that are efficient at escaping. This, however, remains poorly documented. Here, we argue that butterflies may utilize a variety of escape tactics against insectivorous birds and review evidence that birds avoid attacking butterflies that are hard to catch. We suggest that signaling difficulty of capture to predators is a widespread phenomenon in butterflies, and this ability may not be limited to palatable butterflies. The possibility that both palatable and unpalatable species signal difficulty of capture has not been fully explored, but helps explain the existence of aposematic coloration and escape mimicry in butterflies lacking defensive chemicals. This possibility may also change the role that putative Müllerian and Batesian mimics play in a variety of classical mimicry rings, thus opening new perspectives in the evolution of mimicry in butterflies.

1. This article reports the responses of wild, adult jacamars to butterflies with distinct coloration types in central Brazil. Fully aposematic species, i.e. those exhibiting bright and/or contrasting colours on both wing surfaces (= A/A), were predominantly sight‐rejected by birds and, with one exception, the few butterflies attacked and captured were taste‐rejected afterwards.2. Aposematic and cryptic butterflies, i.e. those exhibiting bright and/or contrasting colours on the upper and cryptic colours on the underwings (= A/C) were sight‐rejected while flying, when they show their conspicuous colours to predators. This suggests that birds associate butterfly colours with their difficulty of capture, as in the case of Morpho and several Coliadinae species. These butterflies, however, were heavily attacked at rest, when they are cryptic.3, Fully cryptic butterflies, i.e. those exhibiting cryptic colours on both wing surfaces (= C/C) did not elicit sight rejections by birds. Comparisons involving the number of attacks and the capture success of flying and resting individuals showed no significant differences in species more frequently observed like some cracker butterflies (Hamadryas feronia and H. februa) and Taygetis laches. Compared with the A/C Coliadinae, these butterflies showed a lesser, although not significantly different, ability to escape while flying, but a greater and significantly different ability to escape while at rest.4, A hunting tactic of jacamars, which consists of following flying A/C and C/C butterflies on sight, and waiting until they perch to locate and attack them, is described for the first time.

ABSTRACT. The skippers (Lepidoptera, Hesperioidea, Hesperiidae) of the Distrito Federal, Brazil. An updated list containing 335 species of skippers (Hesperiidae) found in the Distrito Federal (central Brazil), including data obtained from the literature, from several entomological collections, and from collecting of the authors, is presented. Species with presumed distribution in the region are not included. It is presented too, a supplementary list with 32 endemic species occuring in the Cerrado vegetation, from which 27 were found in the Distrito Federal.