Palaeo island-affinities revisited--biogeography and systematics of the Indo-Pacific genus Cethosia Fabricius (Lepidoptera: Nymphalidae)

Abstract

The Indo-Pacific is a very complex region encompassing several micro-continents with unique tectonic and geomorphologic histories. Unsurprisingly, the biogeographic history of Indo-Pacific biota is generally poorly known, especially that of organisms found in the heart of the region, the biodiversity hotspot known as Wallacea. Here, we explore the biogeographic history of the Indo-Pacific butterfly genus Cethosia using all known species and many distinctive subspecies. Cethosia butterflies span the Indo-Pacific tropics, including several lineages with localized endemism that are critically important when reconstructing biogeographic history of the Indo-Pacific and, in particular, of Wallacea. A phylogenetic hypothesis is proposed, based on sequences of the mitochondrial genes cytochrome oxidase subunit I (COI) and NADH dehydrogenase 5 (ND5), and the nuclear wingless gene. Both Maximum Parsimony and Bayesian analyses showed that the genus is monophyletic and consistently recovered seven, generally very well supported, clades, namely the cydippe, leschenault, biblis, nietneri, hypsea, penthesilea and cyane clades. Species group relationships are largely concordant with general morphology (i.e., wing pattern and colouration) and, based on the phylogeny, we propose a revised systematic classification at the species level. The evolution of the genus appears associated with the inferred geological history of the region, in particular with respect to the expanding Wallacea theory, whereby ancient connected terranes were fragmented during the mid Miocene to early Pliocene, approximately 14-3 Mya. Recent diversification events in Cethosia were likely promoted by climatic fluctuations during the Pliocene and, to a lesser extent, the Pleistocene. Our results support the view that, while dispersal has been important for Cethosia throughout much of the region, the high levels of island endemism and the essentially allopatric radiations recovered in Cethosia in Wallacea are better explained by vicariant processes linked to the history of formation of micro-continent and associated palaeo islands.