Chris J Muller

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

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.

Phylogenetics of Coenonymphina (Nymphalidae: Satyrinae) and the problem of rooting rapid radiations.

We report a rapid radiation of a group of butterflies within the family Nymphalidae and examine some aspects of popular analytical methods in dealing with rapid radiations. We attempted to infer the phylogeny of butterflies belonging to the subtribe Coenonymphina sensu lato using five genes (4398bp) with Maximum Parsimony, Maximum Likelihood and Bayesian analyses. Initial analyses suggested that the group has undergone rapid speciation within Australasia. We further analyzed the dataset with different outgroup combinations the choice of which had a profound effect on relationships within the ingroup. Modelling methods recovered Coenonymphina as a monophyletic group to the exclusion of Zipaetis and Orsotriaena, irrespective of outgroup combination. Maximum Parsimony occasionally returned a polyphyletic Coenonymphina, with Argyronympha grouping with outgroups, but this was strongly dependent on the outgroups used. We analyzed the ingroup without any outgroups and found that the relationships inferred among taxa were different from those inferred when either of the outgroup combinations was used, and this was true for all methods. We also tested whether a hard polytomy is a better hypothesis to explain our dataset, but could not find conclusive evidence. We therefore conclude that the major lineages within Coenonymphina form a near-hard polytomy with regard to each other. The study highlights the importance of testing different outgroups rather than using results from a single outgroup combination of a few taxa, particularly in difficult cases where basal nodes appear to receive low support. We provide a revised classification of Coenonymphina; Zipaetis and Orsotriaena are transferred to the tribe Eritina.

Phylogenetics and biogeography of a spectacular Old World radiation of butterflies: the subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrini).

BackgroundButterflies of the subtribe Mycalesina (Nymphalidae: Satyrinae) are important model organisms in ecology and evolution. This group has radiated spectacularly in the Old World tropics and presents an exciting opportunity to better understand processes of invertebrate rapid radiations. However, the generic-level taxonomy of the subtribe has been in a constant state of flux, and relationships among genera are unknown. There are six currently recognized genera in the group. Mycalesis, Lohora and Nirvanopsis are found in the Oriental region, the first of which is the most speciose genus among mycalesines, and extends into the Australasian region. Hallelesis and Bicyclus are found in mainland Africa, while Heteropsis is primarily Madagascan, with a few species in Africa. We infer the phylogeny of the group with data from three genes (total of 3139 bp) and use these data to reconstruct events in the biogeographic history of the group.ResultsThe results indicate that the group Mycalesina radiated rapidly around the Oligocene-Miocene boundary. Basal relationships are unresolved, but we recover six well-supported clades. Some species of Mycalesis are nested within a primarily Madagascan clade of Heteropsis, while Nirvanopsis is nested within Lohora. The phylogeny suggests that the group had its origin either in Asia or Africa, and diversified through dispersals between the two regions, during the late Oligocene and early Miocene. The current dataset tentatively suggests that the Madagascan fauna comprises two independent radiations. The Australasian radiation shares a common ancestor derived from Asia. We discuss factors that are likely to have played a key role in the diversification of the group.ConclusionsWe propose a significantly revised classification scheme for Mycalesina. We conclude that the group originated and radiated from an ancestor that was found either in Asia or Africa, with dispersals between the two regions and to Australasia. Our phylogeny paves the way for further comparative studies on this group that will help us understand the processes underlying diversification in rapid radiations of invertebrates.

Polyura inopinatus Röber, 1940; a remarkable butterfly mystery resolved

Abstract The most distinctive species of Polyura, P. inopinatus, described from a single specimen said to be from North Sulawesi, Indonesia, has been a great mystery since it was first described by Röber, in 1940. The holotype, originally illustrated in monochrome in the journal Deutsche Entomologische Zeitschrift, Iris, was lost very soon after it was described, almost certainly destroyed during allied bombing of Dresden in the 1940s. No other specimen was known for almost eight decades. We suggest that the type locality (Sulawesi) is incorrect and that the holotype was more likely to have been collected in the Baining Mountains, East New Britain Province, Papua New Guinea. We report the recent discovery of several male P. inopinatus from West New Britain Province, and describe and illustrate specimens. A neotype is designated.

Taxonomy of the genus Udara (Lepidoptera: Lycaenidae) in New Caledonia and the Wet Tropics of Australia, with a review of butterfly endemism in these regions

The lycaenid genus Udara Toxopeus is recorded for the first time from New Caledonia where it is represented by the newly described species U. renevieri sp. nov. The species is figured, the male genitalia are illustrated, and the taxon is compared with U. cardia (C. Felder, 1860) (from Maluku through New Guinea to the Solomon Islands and Australia) to which it appears to be most closely related. The taxonomic status of U. cardia tenella syn. nov. and stat. nov. from the Wet Tropics biome of north‐east Australia is revised and treated as a subspecies of U. cardia based on comparative evidence of the morphology of the male genitalia and wing colour pattern elements. The holotypes of both Lycaena cardia C. Felder, 1860 and Lycaena tenella Miskin, 1891 are illustrated. Patterns of butterfly endemism in New Caledonia‐Loyalty Islands and the Wet Tropics are discussed in relation to the conservation importance of these biodiversity hot spots. Estimated levels of endemism are 21% for New Caledonia and 5% (17% if subspecies are included) for the rainforest component of the Wet Tropics.