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.

The ecology and evolution of ant association in the lycaenidae (lepidoptera)

Aphytophagy in butterflies : its relationship to myrmecophily

A facultative mutualism between the riodinid butterfly Thisbe ircnca and the ponerine ant Ectafomma ruidurn is described from Panama. Ants protect larvae against attacks of predatory wasps, but not against tachinid parasitoids. Several potential sources of ecological variation affecting the larval survival of Thisbe irenea are noted. A preliminary means of testing the ability of larvae to appease ants is described that may be applied to all butterfly-ant systems. Observations and literature records indicate that ant taxa which tend butterfly larvae are the same taxa that tend extrafloral nectaries and Homoptera. A general hypothesis for the evolution of myrmecophily among butterflies suggests that ant taxa which utilize secretions in their diet are major selective agents for the evolution of the larval ant-organs, and hence, ant-larval mutualisms. This idea is extended to suggest how appeasement of predaceous ant taxa through the use of larval ant-organs can influence an ant-larval relationship, eventually leading to mutualism.

Mutualism between Thisbe irenea butterflies and ants, and the role of ant ecology in the evolution of larval‐ant associations

Associations with ants, termed myrmecophily, are widespread in the butterfly family Lycaenidae and range from mere co-existence to more or less specific mutualistic or even parasitic interactions. Secretions of specialized epidermal glands are crucial for mediating the interactions. Transfer of nutrients (carbohydrates, amino acids) from butterfly larvae to ants plays a major role, but manipulative communication with the help of odour signals is also involved. By means of myrmecophily, lycaenid butterflies largely escape ant predation, and certain species gain protection through attendant ants or achieve developmental benefits from ant-attendance. Benefits to the ants range from minimal to substantial food rewards. While most lycaenid species maintain facultative relationships with a variety of ant genera, highly specific and obligatory associations have convergently evolved in a number of butterfly lineages. As a corollary, communication systems are largely unspecific in the former, but may be highly specialized in the latter. The sophisticated communication between obligate myrmecophiles and their host ants is tightly connected with the evolutionary rise of specialized life-cycles and thus is a source of augmenting diversity within the butterflies.

T. H. E. Jackson

Papers

The early stages of some african lycaenidae (lepidoptera), with an account of the larval habits