Book Chapters

Abstract The caterpillars of many Lepidoptera are neither attacked nor tended by ants but nevertheless appear to be obligately ant-associated and benefit from the enemy-free space created by ants. Obligate myrmecophiles that do not attract ants through stridulatory or chemical signaling are limited to habitats where ants are reliably present for other reasons, either among ant-attended hemipterans, on ant-plants, or around ant nests. Particularly in the tropics, obligate ant associates that passively coexist with ants are more diverse than previously recognized, including, for example, hundreds of African species in the lycaenid subfamily Poritiinae. Mutualists and parasites of ants have been reported in eleven families: Tineidae, Tortricidae, Cyclotornidae, Coleophoridae, Crambidae, Erebidae, Notodontidae, Hesperiidae, Pieridae, Lycaenidae, and Riodinidae. Altogether, myrmecophily has originated at least 30 times in Lepidoptera, and many groups may remain undiscovered. The butterfly families Lycaenidae and Riodinidae contain the vast majority of ant-associated species: larvae of at least 3841 (71%) of the \textasciitilde5390 described Lycaenidae and 308 (20%) of the \textasciitilde1562 described Riodinidae are known or inferred to be ant-associated, and both families possess specialized, convergently developed exocrine glands and stridulatory devices to communicate with ants. Many caterpillar-ant relationships previously characterized as mutualisms may actually be parasitic, as caterpillars can manipulate ants and ultimately exert a fitness cost. In the family Lycaenidae, highly specialized and obligate ant associations are found largely in the Old World tropics, Australia, and Southern Africa, where the stoichiometry of soil micronutrients, particularly sodium and phosphorus, climate, host plants, and geography may all selectively shape caterpillar-ant associations.