Molecular phylogeny of the tribe Candalidini (Lepidoptera: Lycaenidae): systematics, diversification and evolutionary history
Authors: Braby Michael F. , Espeland Marianne , Müller Chris J. , Eastwood Rod , Lohman David J. , Kawahara Akito Y. , Maunsell Sarah C. , Pierce Naomi E. 
Published: June 2020

AbstractThe butterfly tribe Candalidini is geographically restricted to Australia and mainland New Guinea and its adjacent islands. With 60 species and subspecies, it represents a large radiation of Papilionoidea in the Australian region. Although the species‐level taxonomy is relatively well understood, the number of genera is uncertain, varying from two to eight. We reconstructed the phylogeny of the Candalidini based on a 13‐locus hybrid enrichment probe set (12.8 Kbp: COI, Thiolase, CAD, CAT, DDC, EF1‐a, GAPDH, HCL, IDH, MDH, RPS2, RPS5, Wingless), including all previously recognized genera and 76% (28/37) of the species‐level diversity of the tribe. Maximum likelihood analysis recovered the Candalidini as a strongly supported monophyletic group. In conjunction with morphological characters, the phylogeny provided a robust framework for a revised classification in which we recognize four genera, 37 species and 23 subspecies. The genusNesolycaenaWaterhouse & R.E. Turner is considered in synonymy withCandalidesHübner, and four other genera are not recognized, namely,HolochilaC. Felder,AdalumaTindale,ZetonaWaterhouse andMicroscenaTite. Of the four valid genera, theabsimilisgroup (23 species) is placed in the newly described genusEirmocidesBraby, Espeland & Müllergen. nov.(type speciesCandalides consimilisWaterhouse). Theerinusgroup (six species) is assigned toErinaSwainson, which is reinstated.Chrysophanus cyprotusOlliff is assigned toCyprotidesTite, which is also reinstated as a monotypic genus. The remaining seven species are placed inCandalides sensu stricto. Overall, we propose 47 new nomenclatural changes at the species and subspecies levels, including the synonymy ofHolochila biakaTite asEirmocides tringa biaka(Tite)syn. nov. et comb. nov.and recognition ofCandalides hyacinthinus gilesiM.R. Williams & Bollam as a distinct speciesErina gilesi(M.R. Williams & Bollamstat. rev. et comb. nov.A dated phylogeny using Bayesian inference in BEAST2 and biogeographical and habitat analyses based on the DEC model in BioGeoBEARS indicated that the ancestor of the Candalidini most likely evolved in rainforest habitats of the mesic biome in situ on the Australian plate of Southern Gondwana during the Eocene (c. 43 Ma). A major period of diversification occurred in the Miocene, which coincided with aridification of the Australian continent, followed by a further episode of radiation in montane New Guinea during the Plio‐Pleistocene.This published work has been registered on ZooBank by the authors: Michael Braby:http://zoobank.org/urn:lsid:zoobank.org:author:4D3A7605‐EBD0‐40F6‐A5F2‐7F67F59E3D60;Marianne Espeland:http://zoobank.org/urn:lsid:zoobank.org:author:00D6F9F9‐3902‐4A8B‐846F‐720AB32922A6;Chris Müller:http://zoobank.org/urn:lsid:zoobank.org:author:15FE5F26‐7596‐46C2‐9697‐1FD92A692D0D;http://zoobank.org/urn:lsid:zoobank.org:pub:47D5CA34‐C294‐4FBD‐84B6‐1C2A82B7CADF.