Phylogeny of the Aphnaeinae: myrmecophilous African butterflies with carnivorous and herbivorous life histories

Abstract

The Aphnaeinae (Lepidoptera: Lycaenidae) are a largely African subfamily of 278 described species that exhibit extraordinary life-history variation. The larvae of these butterflies typically form mutualistic associations with ants, and feed on a wide variety of plants, including 23 families in 19 orders. However, at least one species in each of 9 of the 17 genera is aphytophagous, parasitically feeding on the eggs, brood or regurgitations of ants. This diversity in diet and type of symbiotic association makes the phylogenetic relations of the Aphnaeinae of particular interest. A phylogenetic hypothesis for the Aphnaeinae was inferred from 4.4 kb covering the mitochondrial marker COI and five nuclear markers (wg, H3, CAD, GAPDH and EF1 ) for each of 79 ingroup taxa representing 15 of the 17 currently recognized genera, as well as three outgroup taxa. Maximum Parsimony, Maximum Likelihood and Bayesian Inference analyses all support Heath's systematic revision of the clade based on morphological characters. Ancestral range inference suggests an African origin for the subfamily with a single dispersal into Asia. The common ancestor of the aphnaeines likely associated with myrmicine ants in the genus Crematogaster and plants of the order Fabales.

Male secondary sexual characters in Aphnaeinae wings (Lepidoptera: Lycaenidae)

“…The four aphnaeine genera (Aphnaeus, Cigaritis, Lipaphnaeus and Pseudaletis) possessing male secondary sexual character are paraphyletic in the hypothetical phylogenetic tree provided by the most recent paper on the subfamily (Boyle et al 2015). We note that these genera represent both of the main aphnaeine lineages being sisters to each other, and in one of the lineages there is a basal genus with androconia (Pseudaletis).…”

“…According to Boyle et al (2015) Bridges 1994) -if we take the hair tuft as an important character for classification. However, the uniform type of male genitalia as well as their facies mitigate against such splitting, and also there is a growing evidence that male secondary characters are not stable (see for example Robbins et al 2012and Martins et al 2016.…”

“…These specimens are indicated with RCS (= research collection of Szabolcs Sáfián, Sopron, Hungary). Genus-level classification follows Boyle et al (2015). Genera and species are listed in alphabetical order.…”

“…The subfamily Aphnaeinae was recently the subject of important studies (Heath 1997, Boyle et al 2015. These were inspired by Eliot who pointed out the peculiarity of this lycaenid assemblage and recorded morphological traits for his classification (Eliot 1973: 470 in footnote no.…”

Editorial

“…, ), Lycaenidae (Boyle et al. , ), Sphingidae (Ponce et al. , ) and Nymphalidae (Gemmell & Marcus, ).…”