Type of Document	Dissertation
Author	Aschliman, Neil C
Author's Email Address	naschliman@bio.fsu.edu
URN	etd-06302011-041658
Title	The Batoid Tree Of Life: Recovering The Patterns And Timing Of The Evolution Of Skates, Rays And Allies (Chondrichthyes: Batoidea)
Degree	Doctor of Philosophy
Department	Biological Science, Department of
Advisory Committee	Advisor Name Title Gavin J.P. Naylor Committee Co-Chair Scott J. Steppan Committee Co-Chair Austin R. Mast Committee Member R. Dean Grubbs Committee Member William C. Parker University Representative
Keywords	elasmobranchs divergence times comparative morphology molecular systematics
Date of Defense	2011-06-02
Availability	unrestricted
Abstract Batoid fishes (skates, stingrays and allies) comprise the majority of species diversity and morphological disparity among chondrichthyans, one of the two primary divisions of extant jawed vertebrates. The largely recent and growing interest in batoid evolution has emphasized the need for a well-supported phylogeny against which evolutionary changes in traits can be interpreted. While batoids are morphologically well characterized and have an excellent fossil record, there is currently no consensus on the interrelationships of family-level taxa. Patterns of evolution within the two largest groups of batoids, skates and stingrays, also remain obscure. This dissertation presents novel frameworks for interpreting the patterns and timing of batoid evolution based on molecular data, morphology, and fossils. I recovered a resolved and time-calibrated phylogeny of the major extant groups of batoids using mitochondrial genomes, two independent nuclear markers, and fossil ages. Taxon sampling included 37 ingroup species from 22 of 23 families. Data partitioning schemes, potential biases in the sequence data, the relative informativeness of each fossil, and ancestral state reconstructions were explored. The molecular data set was then expanded with additional species in order to address questions at a finer taxonomic scale, in particular among skates and stingrays. Two nuclear and two mitochondrial markers were sequenced for 87 batoid species across 52 of 81 genera. A similar analytical approach was applied to this larger data set. I also performed a morphology-based phylogenetic analysis in order to interpret accurately coded morphological characters against the molecular frameworks. I updated the data set of McEachran and Aschliman (2004) with a number of corrections and modifications, and added new characters from the synarcual and other chondroskeletal structures. The molecular phylogenies indicate that the major lineages of batoids originated in relatively rapid sequence, followed by long periods of independent evolution. These trees corroborate morphology-based hypotheses in many respects, but have strongly divergent implications in others. Lineages inferred to be distantly related, such as skates and stingrays, or sawfishes and sawsharks, are indicated to have achieved very similar, specialized body plans through convergence. Skates and stingrays are unique among batoids in exhibiting a highly depressed disc supported to the apex by fin rays, and swim by passing waves along the lateral margin of the pectoral fin without additional propulsion by lateral motion of the tail and caudal fin. The plesiomorphic mode of locomotion and body plan for most batoid groups was probably not shark-like, as in sawfishes. Instead, it is inferred to be a combination of pectoral fin undulation and shark-like axial locomotion, which is correlated with a broader pectoral disc and reduced tail compared to the sawfish body plan. The higher-resolution molecular phylogeny is in many cases congruent with previous morphological studies or biogeography. Exceptions typically suggested a potential weakness in the molecular data, suggested that a morphology-based classification scheme is likely based on convergent characters or ignores a highly divergent morphotype nested within otherwise similar taxa, or helped resolve taxonomic confusion based on informal re-assignment by authority. Potentially non-monophyletic families and genera were identified for future study with expanded taxon sampling, additional sequence data and morphological re-evaluation. The origin of Batoidea is estimated to have occurred in the Late Triassic, with the major groups diverging throughout the Jurassic and possibly into the Cretaceous. Radiations of each major crown group are indicated to have occurred from the Late Cretaceous to the Cenozoic. The tree shape recovered for all major batoid groups, with long internal branches subtending subsequent radiations around the Cretaceous/Tertiary boundary, suggests that batoid standing diversity may be due in large part to lineage pruning and/or rapid radiation into vacated niche space. This is consistent with the fossil record, which suggests that batoids were more severely affected by the end-Cretaceous extinction event than were the other neoselachians. The updated morphological phylogeny included several key changes from earlier hypotheses and more closely approximates the molecular frameworks. There remain conflicts between morphological and molecular trees, such as the phylogenetic placements of skates and thornbacks, which currently appear to be difficult to reconcile. Future attempts to reconcile molecules and morphology will require expanded data sets and investigation of potential sources of error in each.
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