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Systematic Biology 2004 53(3):363-383; doi:10.1080/10635150490445670
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© 2004 Society of Systematic Biologists

Phylogeny of the Nymphalidae (Lepidoptera)

André Victor Lucci Freitas and Keith S. Brown, Jr.

Museu de História Natural and Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas CP 6109, CEP 13083–970, Campinas, SP, Brazil; E-mail: baku{at}unicamp.br

Edited by Ted Schultz: Associate Editor


  Abstract

A generic-level phylogeny for the butterfly family Nymphalidae was produced by cladistic analysis of 234 characters from all life stages. The 95 species in the matrix (selected from the 213 studied) represent all important recognized lineages within this family. The analysis showed the taxa grouping into six main lineages. The basal branch is the Libytheinae, with the Danainae and Ithomiinae on the next branch. The remaining lineages are grouped into two main branches: the Heliconiinae-Nymphalinae, primarily flower-visitors (but including the fruit-attracted Coeini); and the Limenitidinae (sensu strictu), Biblidinae, and the satyroid lineage (Apaturinae, Charaxinae, Biinae, Calinaginae, Morphinae, Brassolinae, and Satyrinae), primarily fruit-attracted. Data partitions showed that the two data sets (immatures and adults) are very different, and a partitioned Bremer support analysis showed that the adult characters are the main source of conflict in the nodes of the combined analysis tree. This phylogeny includes the widest taxon coverage of any morphological study on Nymphalid butterflies to date, and supports the monophyly and relationships of most presently recognized subgroups, providing strong evidence for the presently accepted phylogenetic scheme.

Keywords: Adults; combined data; eggs; juveniles; larvae; morphology; Nymphalidae; phylogeny; pupae

Received March 26, 2002; Revised April 13, 2002; Accepted December 14, 2003
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