The comparative osteology and phylogenetic relationships of lepidosirenid lungfishes
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Lepidosirenidae is a clade of freshwater lungfishes that comprise the South American Lepidosiren paradoxa and four African species of the genus Protopterus. These two genera have been geographically separated since the Early Cretaceous break-up of Gondwana, but they share similar biology and skeletal morphology. The lepidosirenid species traditionally were distinguished by a combination of features such as head-to-body ratios, the number of pairs of vertebral ribs, and the presence or absence of external gills, but there are no published discrete skeletal characteristics and no published comparative studies including all extant species. I used High Resolution X-Ray Computed Tomography (CT), X-Ray photography, and alcohol-preserved, cleared-and-stained, and dry skeletal specimens from museum collections to describe the skeletal morphology of all species of lepidosirenid lungfishes in a comparative context. I digitally disarticulated the bones in each CT scan to compile a comprehensive comparative atlas of the cranial and pectoral elements of all extant lungfish. I discovered that the anocleithrum in Lepidosiren paradoxa, which was previously thought to be lacking, is actually present. I also identified skeletal differences between species in the frontoparietal, parasphenoid, supraorbital, and suboperculum. I incorporated those characters into the first morphological phylogenetic analysis to determine the interrelationships of the lepidosirenids. I also used previously published molecular sequence data from the ribosomal RNA gene 16s to run combined morphological and molecular phylogenetic analyses. To generate phylogenetic hypotheses using different types of data and different methods of determining phylogeny, I employed the maximum parsimony, maximum likelihood, and Bayesian inference methods. Lepidosirenidae is monophyletic in almost all analyses, Protopterus is monophyletic in each analysis, and Protopterus annectens and Protopterus aethiopicus are sister taxa in every analysis. The phylogenetic positions of Protopterus dolloi and Protopterus amphibius are incongruent in many of the analyses, which indicates that further examination of the skeletal variation and addition of molecular sequences of different genes is needed. Based on the comparative morphological atlas and the phylogenetic analyses, questions of lepidosirenid biogeography, morphological variation within lungfish, and better identification of lungfish fossils can now be investigated in a more rigorous context.