Phylogenetic Analysis of the Subtribe Ageroniina with Special Emphasis on Hamadryas (Lepidoptera, Nymphalidae) with an Identification Key to the Species of Hamadryas
A cladistic analysis of the genus Hamadryas was done in order to answer two questions: is the genus Hamadryas monophyletic and, what characters best define the different species groups of the genus? The computer programs WinClada ver. 1.00.08 and Nona ver. 2.8 for phylogenetic analysis were used. The 20 species of Hamadryas were analyzed together with Ectima erycinoides, Batesia hypochlora and Panacea procilla; these four genera together form the sub-tribe Ageronina of the subfamily Biblidinae. These 23 species with a total of 66 characters were included in the data matrix: 43 of external morphology, two from the fore wing spiral organ (responsible for sound production), and 20 of the male genitalia. Three different analyses, including all of the non-Hamadryas, using each species as the out-group, were done in order to compare results. A fourth analysis, using only Ectima erycinoides as an out group, was done in order to verify and compare the species groups of Hamadryas Also, three different phylogenetic attributes were mapped: biogeography, palatability and capacity to produce sound signals. An identification key to all the species of Hamadryas was prepared. All the analysis done combining Hamadryas with the other three genera suggests that Hamadryas as currently defined is polyphyletic. 1. Introduction The biodiversity on earth is so enormous that it would not be possible to study it, if it were not classified [1] and phylogenetic analyses of the species permit us to approach their evolutionary relationships in order to establish hypothetical species groups on which the classification can be based. Hamadryas Hübner (1806) is composed of twenty species ranging from the southern United States to northern Argentina [2]. According to DeVries [3] it is distinguished from other butterfly genera by their spotted “calico” pattern on the upper side. Jenkins [4] on the other hand states that Hamadryas is a taxonomically confused genus which has been in great need of revision and highlights the lack of a detailed study to find consistently valid distinguishing characters. This genus is well known for producing an audible clicking sound, an ability that has been reported for many but not all species. It has been suggested that this sound is used in courtship, defense, and territoriality [5–7]. It is produced through a modification of the internal walls of the forewing subcostal vein which consists of a spiral organ that ends in a tympanic membrane which in turn releases the sound into the air [7, 8]. In contrast to most of other nymphalid butterflies, this genus
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