Content area
Full Text
Key Words myrmecophily, parasitism, butterfly, symbiosis, biogeography
Abstract The estimated 6000 species of Lycaenidae account for about one third of all Papilionoidea. The majority of lycaenids have associations with ants that can be facultative or obligate and range from mutualism to parasitism. Lycaenid larvae and pupae employ complex chemical and acoustical signals to manipulate ants. Cost/benefit analyses have demonstrated multiple trade-offs involved in myrmecophily. Both demographic and phylogenetic evidence indicate that ant association has shaped the evolution of obligately associated groups. Parasitism typically arises from mutualism with ants, and entomophagous species are disproportionately common in the Lycaenidae compared with other Lepidoptera. Obligate associations are more common in the Southern Hemisphere, in part because highly ant-associated lineages make up a larger proportion of the fauna in these regions. Further research on phylogeny and natural history, particularly of the Neotropical fauna, will be necessary to understand the role ant association has played in the evolution of the Lycaenidae.
INTRODUCTION
The estimated 6000 species of Lycaenidae account for about one third of all Papilionoidea (1, 55, 208, 226). Full or partial life histories have been recorded for about 20% of these species, and of those whose full life histories are known, about 75% [(N=665 (Table 1)] associate with ants (95,139,181). These associations can be mutualistic or parasitic and range from loose facultative interactions in which larvae are only occasionally tended by several species of ants (about 45% of associations), to complex obligate associations in which larvae are always tended by ants, often by only a single species (30%) (Table 1). Even when lycaenids are not myrmecophilous, they may be protected against ant aggression by a suite of ant-associated adaptations. The Lycaenidae are additionally characterized by striking life history diversity. Herbivorous species consume an unusually wide array of different plant families (10, 80), and a small number of lycaenids (-3% of all associations or 12% of obligate ant associations) are parasitic or predatory (48,183).
The behavioral and ecological diversity of the Lycaenidae makes this group particularly amenable to comparative studies of life history evolution. Colonel John N. Eliot laid the groundwork for such research in 1973 by providing what he described as a "tentative arrangement" of the higher classification of the Lycaenidae (83). He later revised...