Classification, nomenclature and identification of lime swallowtail butterflies: a post-cladistic analysis (Lepidoptera: Papilionidae)
The taxonomy of the five species of ‘lime swallowtails’ of the Papilio demoleus group (P. demoleus, P. demodocus, P. erithonioides, P. grosesmithi, and P. morondavana) is re-examined. Sixty morphological characters are described, scored and illustrated based on features of the wings, antennae, and male and female genitalia. The results of cladistic analyses of these data are described and discussed in the light of a previous collaborative study that combined this morphological data set with DNA sequence data. While it is confirmed that the lime swallowtails form a monophyletic group, the internal relationships of the group, and the relationships of the group with other members of the genus, have yet to be established with any certainty. This implies that the current subgeneric classification of Papilio must be considered highly provisional. Dichotomous keys to the species and subspecies based on adults, together with a synonymic checklist, are presented. Implications of the current, possibly anthropogenic spread of two subspecies, P. d. demoleus and P. d. malayanus, are discussed. The nominal taxa P. demoleus flavosignatus Heydemann and P. demoleus libanius Fruhstorfer are both formally synonymised with P. demoleus demoleus (synn. nov.). Five on-line appendices accompany the paper. Appendix I gives full nomenclatural information, including diagnoses, synonyms and type data, for each species and subspecies. Lectotypes are designated here for 6 of the 20 nominal species-group taxa included in the lime swallowtails: Papilio morondavana Grose-Smith, P. grosesmithi Rothschild, P. erithonioides Grose-Smith, P. erithonius malayanus Wallace, P. demoleus novoguineensis Rothschild, and P. demoleus pictus Fruhstorfer. Type material or putative type material is located for the remaining nominal taxa, with the exception of P. erithoniusP. epius Fabricius, P. demoleus flovosignatus Heydemann and P. sthenelus Macleay. Appendix II presents a history of the application and misapplication of the names Papilio demodocus and Papilio demoleus, respectively, to the common afrotropical and oriental pest species. Appendix III summarises known distribution and natural history of the species and subspecies. The male and female genitalia of the various out-group species, as examined during this study, are described in Appendix IV. Appendix V provides an index to the principal entries for lime swallowtail taxa, including synonyms and infrasubspecific names. The five on-line appendices are completed by a comprehensive bibliography covering the entire work.
Key Words: higher-classification; species; subspecies; lectotype designations; type material; new synonymy; Papilio demoleus; P. demodocus; P. erithonioides; P. grosesmithi; P. morondavana; keys; distribution; natural history