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Systematic Biology 2004 53(2):193-215; doi:10.1080/10635150490423403
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© 2004 Society of Systematic Biologists

Molecular Phylogeny, Historical Biogeography, and Divergence Time Estimates for Swallowtail Butterflies of the Genus Papilio (Lepidoptera: Papilionidae)

Evgueni V. Zakharov1, Michael S. Caterino2 and Felix A.H. Sperling1

1 Department of Biological Sciences, University of Alberta Edmonton Alberta T6G 2E9, Canada; E-mail: zakharov{at}ualberta.ca (E.V.Z.); E-mail: felix.sperling{at}ualbert.ca (F.A.H.S.)
2 Santa Barbara Museum of Natural History 2559 Puesta del Sol Road, Santa Barbara, California 93105, USA

Edited by Ted Schultz: Associate Editor


  Abstract

Swallowtail butterflies are recognized as model organisms in ecology, evolutionary biology, genetics, and conservation biology but present numerous unresolved phylogenetic problems. We inferred phylogenetic relationships for 51 of about 205 species of the genus Papilio (sensu lato) from 3.3-Kilobase (kb) sequences of mitochondrial and nuclear DNA (2.3 kb of cytochrome oxidases I and II and 1.0 kb of elongation factor 1{alpha}). Congruent phylogenetic trees were recovered within Papilio from analyses of combined data using maximum likelihood, Bayesian analysis, and maximum parsimony bootstrap consensus. Several disagreements with the traditional classification of Papilio were found. Five major previously hypothesized subdivisions within Papilio were well supported: Heraclides, Pterourus, Chilasa, Papilio (sensu stricto), and Eleppone. Further studies are required to clarify relationships within traditional "Princeps," which was paraphyletic. Several biologically interesting characteristics of Papilio appear to have polyphyletic origins, including mimetic adults, larval host associations, and larval morphology. Early diversification within Papilio is estimated at 55–65 million years ago based on a combination of biogeographic time constraints rather than fossils. This divergence time suggests that Papilio has slower apparent substitution rates than do Drosophila and fig-pollinating wasps and/or divergences corrected using best-fit substitution models are still being consistently underestimated. The amount of sequence divergence between Papilio subdivisions is equivalent to divergences between genera in other tribes of the Papilionidae, and between genera of moths of the noctuid subfamily Heliothinae.

Keywords: Character evolution; fossils; mimicry; molecular systematics; swallowtail butterflies; substitution rates

Received December 13, 2002; Revised April 2, 2003; Accepted November 15, 2003
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