Phylogenetics and divergence times of Papilioninae (Lepidoptera) with special reference to the enigmatic genera Teinopalpus and Meandrusa


Phylogenetic relationships of 18 genera of the swallowtail subfamily Papilioninae, four genera of Parnassiinae, and the monobasic Baroniinae are inferred based on 94 morphological characters and 5616 bp DNA from seven genes (16S, COI, COII, ND1, ND5, EF-1 alpha and wingless). Bayesian likelihood analyses show that Baroniinae are the sister of a clade comprising Parnassiinae and Papilioninae. Four Papilioninae tribes are recognized, Leptocircini, Teinopalpini, Papilionini and Troidini, with Leptocircini being the sister of the remaining tribes. Meandrusa and Teinopalpus are sister taxa and comprise the tribe Teinopalpini, which is the sister of a clade comprising Papilionini and Troidini. The tribe Troidini (pipevine swallowtails) comprises two subtribes: Battina (including only Battus) and Troidina. The endemic Madagascan genus Pharmacophagus is consistently placed as the sister to the remaining Troidina. The non-Pharmacophagus Troidina are tentatively divided into a Neotropical lineage and an Australasian lineage. Dispersal–vicariance analyses indicate that past dispersal events are most important for explaining current distribution patterns of Papilionidae. However, the division of the non-Pharmacophagus Troidina into a Neotropical lineage and an Australasian lineage is possibly due to the final break-up of southern Gondwana. A fossil-calibrated relaxed Bayesian molecular clock analysis confirms that the ages of the lineages fit this scenario. The basal lineages leading to the current subfamily-level diversity of Papilionidae probably arose around the K ⁄T boundary. Analyses of larval host-plant relationships within Papilionidae show very little phylogenetic pattern. However, Aristolochiaceae-feeding apparently evolved independently in non-Parnassiini parnassiines and Troidini. The Willi Hennig Society 2010. Swallowtail butterflies are the best-known family of Lepidoptera and may be the most well-known group of invertebrate animals. Numerous studies have been made on speciation, polymorphism, mimicry, sexual selection, host relations, chemistry, physiology, anatomy and historical biogeography. These insects are a flagship group for invertebrate conservation, and several large volumes have been devoted to their biology and evolution (e.g. Igarashi, 1979; Tyler et al., 1994; Scriber et al., 1995). Many studies have also addressed their systematic and phylogenetic relationships. Vane-Wright (2003, p. 480) noted that ‘‘measured by effort per species, more work has gone into trying to understand the interrelationships of the 600 or so species of Papilionidae than any other family of Lepidoptera’’, and yet ‘‘schemes abound, but we remain far from any consensus’’. One consensus widely agreed is the division of Papilionidae into three extant subfamilies: Baroniinae, *Corresponding author: E-mail address: Present address: Department of Entomology, Natural History Museum, Cromwell Road, London SW7 5BD, UK. The Willi Hennig Society 2010 Cladistics 10.1111/j.1096-0031.2010.00326.x Cladistics 26 (2010) 1–25 Parnassiinae and Papilioninae. Parnassiinae and Papilioninae are generally considered sister taxa, with the monobasic Baroniinae representing the stem lineage (but see Nazari et al., 2007; Michel et al., 2008). Although placed in Papilioninae, the phylogenetic positions of Meandrusa and Teinopalpus have been particularly problematic. Phylogenetic relationships In the following account, few works pre-dating 1980 are dealt with; Hancock (1983), Igarashi (1984) and Miller (1987a) gave reviews of these sources. Munroe and Ehrlich (1960) and Munroe (1961) summarized previous systematic evidence and recognized two tribes [Parnassiini and Luehdorfiini (=Zerynthiini)] within Parnassiinae, and four tribes (Leptocircini, Teinopalpini, Papilionini and Troidini) within Papilioninae. They considered Meandrusa to be a junior synonym of Papilio, placed Cressida and Euryades in Troidini, and followed Ford (1944a) in placing Teinopalpus in its own tribe, although a position within Leptocircini was also considered (Munroe and Ehrlich, 1960, p. 170; Munroe, 1961, p. 17). Hancock (1983) presented the first modern (cladistic) attempt to resolve the classification and phylogeny of Papilioninae (Fig. 1a), although this work was methodologically mixed (Miller, 1987a). Within Papilioninae, Hancock recognized three tribes: Leptocircini, Papilionini and Troidini. Igarashi (1984) presented a classification based on detailed studies of juvenile life stages. Although his phylogenetic reconstruction and classification were

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@inproceedings{Simonsen2010PhylogeneticsAD, title={Phylogenetics and divergence times of Papilioninae (Lepidoptera) with special reference to the enigmatic genera Teinopalpus and Meandrusa}, author={Thomas J. Simonsen and Evgeny V. Zakharov and Marie Djernaes and Adam M. Cotton and Richard I Vane-Wright and Felix A. H. Sperling and Nong Kwai and Hang Dong}, year={2010} }