Reconstructing the origins of praying mantises (Dictyoptera, Mantodea): the roles of Gondwanan vicariance and morphological convergence.

A comprehensive taxonomic sampling of Mantodea (praying mantises), covering virtually all higher-level groups, was assembled to reconstruct the phylogeny of the order. Sequence data were generated from five mitochondrial and four nuclear loci (12S rRNA, 16S rRNA, 18S rRNA, 28S rRNA, Histone III, Cytochrome Oxidase I & II, NADH dehydrogenase subunit 4, and Wingless) for 329 mantis exemplars along with seven cockroach and eight termite species. Only seven of 14 families, 14 of 33 subfamilies, and seven of 14 tribes were recovered as monophyletic, indicating that phylogeny is largely incongruent with classification. Mapping biogeographical regions on the phylogeny demonstrated that our results adhere closer to biogeographical distributions than to classification. Specific patterns in distribution suggest that major morphological convergences have confounded taxonomists' ability to reconstruct natural groups. A major revision of higher-level relationships is in order through a comprehensive investigation of morphology and molecular data. We found that major mantis lineages diverged prior to and during the isolation of geographical regions and subsequent ecomorphic specializations within these regions may have led to convergences in morphology. Divergence time estimation places the origin of Mantodea at the beginning of the Jurassic with most modern mantises originating on Gondwana in the Cretaceous. The first major divergence among modern mantises occurred as a result of the north-south splitting of South America and Africa. Subsequent divergences resulted from the breakup of Gondwana. The position of the Indian subcontinent appears to be central to the diversification of Afrotropical and Indomalayan mantises while Antarctica may have served as the conduit for the mantis invasions into South America and Australasia. When India separated from Antarctica and drifted north it distributed mantis lineages back into the Afrotropics and carried a diverse taxonomic assemblage to Asia.