The ecology and evolution of ant association in the Lycaenidae (Lepidoptera)

Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, USA.
Annual Review of Entomology (Impact Factor: 13.02). 02/2002; 47(1):733-71. DOI: 10.1146/annurev.ento.47.091201.145257
Source: PubMed

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.

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    • "Many ant colonies are also inhabited by inquilines (i.e., social parasites) like Lycaenid caterpillars that compete with or eat host brood (Pierce et al. 2002). Similarly, avian brood parasites compete with or kill host offspring while exploiting adult hosts (Hauber 2003). "
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    ABSTRACT: Recent studies of inter-individual variation in behavior have focused primarily on its environmental causation and adaptive consequences, but commonly ignore questions regarding its proximate development. Here, we explore the effects of the late natal environment on the development of individual- and colony-level personalities in the acorn ant, Temnothorax longispinosus. This species is commonly parasitized by the slave-making ant Protomognathus americanus, and we predicted that rearing T. longispinosus with its brood parasite would alter the behavior of individual ants and the collective, colony-level personality of groups. Using a split-brood design (where brood from a single source colony is split equally across different rearing environments), we reared T. longispinosus in four conditions: their maternal queen, an unrelated conspecific queen, a slave-making queen, and a slave-making worker. Although individual aggressiveness and exploratory behavior did not differ between ants raised by their maternal queen or slave-making ants, ants raised by an unrelated conspecific queen showed increased aggressiveness 60 days after emergence. Further, groups of ants raised by slave-making workers were faster at locating new nest sites, a collective behavior, relative to groups reared by their maternal queen. Lastly, colonies containing T. longispinosus and their maternal queen had the greatest brood production at 60 days. Our results demonstrate that differences in individuals’ rearing environment can influence both individual- and colony-level personality in multi-level societies.
    Behavioral Ecology and Sociobiology 03/2015; 69(3):395-405. DOI:10.1007/s00265-014-1852-2 · 3.05 Impact Factor
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    • "Lycaenids have long been known for their interactions with mutualistic ants, and many species have developed mechanisms to recognize and select specific ant partners (Fiedler 1991; Pierce et al. 2002). In this context, our results (in addition to Kaminski et al. 2010) reveal that lycaenid oviposition mediated by ant presence may be more common than previously thought, even in lycaenids with a weak ant association , such as A. strophius. "
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    ABSTRACT: Ovipositing adult females of myrmecophilous lycaenids are expected to select plants based on ant presence in order to maximize the survivorship of immature stages. Usually, larvae feed ants with honey-like solutions and, in turn, ants ward off parasitoids. Nonetheless, a rarely investigated approach is whether ant partners can also extend their protective behavior towards lycaenids eggs. Here, we investigated the ant-related oviposition pattern of Allosmaitia strophius and Rekoa marius; then, we compared egg parasitism according to the presence of ants. Lycaenid oviposition and egg parasitism (in percent) were experimentally compared in ant-present and ant-excluded treatments. The study plant, Heteropterys byrsonimifolia, is an extrafloral nectaried shrub which supports several ant species. We sampled 280 eggs, of which 39.65 % belonged to A. strophius and 60.35 % to R. marius. Both lycaenids eggs were significantly more abundant on branches with ants, especially those with Camponotus crassus and Camponotus blandus, two ant species known to attend to lycaenids. A. strophius and R. marius parasitism was 4.5- and 2.4-fold higher, respectively, in ant-present treatments, but the results were not statistically significant. Our study shows that ant-mediated host plant selection in lycaenids might be much more widespread than previously thought, and not restricted to obligate myrmecophilous species. Tending ants may be inefficient bodyguards of lycaenid eggs, because unlike larvae which release sugared liquids, eggs do not offer obvious rewards to ants. Ants can ward off parasitoids of larvae, as observed elsewhere, but our findings show that positive ant-lycaenid interactions are conditional and depend on immature ontogeny.
    Naturwissenschaften 09/2014; DOI:10.1007/s00114-014-1232-9 · 1.97 Impact Factor
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    • "The association of ants with honeydew-producing Hemiptera (aphids, scale insects, membracids, etc.), or with nectarproducing larvae of lycaenid butterflies, is a well-studied phenomenon termed trophobiosis. Trophobioses are complex, 806 S. A. Schneider et al. typically mutualistic, relationships in which ants provide protection and other benefits to a partner species and procure a reliable food reward from this partner in exchange for their attendance (reviewed by Way, 1963; Hölldobler & Wilson, 1990; Gullan, 1997; Gullan & Kosztarab, 1997; Delabie, 2001; Pierce et al., 2002). The only ant–hemipteran trophobiotic relationship in which honeydew appears not to be a 'currency' of exchange involves ants of the genus Melissotarsus Emery (Hymenoptera: Formicidae: Myrmicinae: Melissotarsini) and certain armoured scale insects (Hemiptera: Diaspididae). "
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    ABSTRACT: The association between African armoured scale insects (Hemiptera: Coccoidea: Diaspididae) and ants belonging to Melissotarsus Emery (Hymenoptera: Formicidae: Myrmicinae) is the only trophobiosis known in which ants do not receive honeydew or nectar in exchange for protection and other services. Food reward for the ants in this mutualism remains unknown, despite repeated suggestions that diaspidids are consumed by the associated ants, thus serving as ‘domestic cattle’. We describe new observations on interactions between Melissotarsus emeryi Santschi and the diaspidid Morganella conspicua (Brain) from South Africa. Worker ants exhibited previously undescribed tending behaviours, most notably a ‘squeezing and licking’ performed on an adult female diaspidid and ‘culling’, in which a worker removed an adult female armoured scale from the host plant. These could represent the gathering of secretory products and the cultivation of an individual for consumption, respectively. An ant exclusion study over 12 days of isolation showed that adult female diaspidids and second-instar nymphs secreted no wax or exudates that attending ants would ordinarily collect. Workers of M. emeryi did not defend their nest against invading colonies of Crematogaster and other unidentified ants: we hypothesize that the primary mode of defence is maintenance of isolation within galleries. We describe three new ant-associated diaspidid species: Affirmaspis cederbergensis Schneider sp.n. from South Africa, Diaspis doumtsopi Schneider sp.n. from Cameroon, and Melissoaspis incola Schneider sp.n. from Madagascar. Melissoaspis formicaria (Ben-Dov) comb.n. is transferred from Morganella (Brain). Diagnostic characteristics for Melissoaspis Ben-Dov are revised, and additional taxonomic information defining this genus allows ease of identification. An updated identification key to the species of ant-associated diaspidids is provided. This published work has been registered in ZooBank,
    Systematic Entomology 10/2013; 38(4). DOI:10.1111/syen.12033 · 2.55 Impact Factor
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