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Early Tetrapod Relationships Revisited

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Biol. Rev. (2003), 78, pp. 251–345. f Cambridge Philosophical Society 251 DOI: 10.1017/S1464793102006103 Printed in the United Kingdom Early tetrapod relationships revisited MARCELLO RUTA1*, MICHAEL I. COATES1 and DONALD L. J. QUICKE2 1 The Department of Organismal Biology and Anatomy, The University of Chicago, 1027 East 57th Street, Chicago, IL 60637-1508, USA ([email protected]; [email protected]) 2 Department of Biology, Imperial College at Silwood Park, Ascot, Berkshire SL57PY, UK and Department of Entomology, The Natural History Museum, Cromwell Road, London SW75BD, UK ([email protected]) (Received 29 November 2001; revised 28 August 2002; accepted 2 September 2002) ABSTRACT In an attempt to investigate differences between the most widely discussed hypotheses of early tetrapod relation- ships, we assembled a new data matrix including 90 taxa coded for 319 cranial and postcranial characters. We have incorporated, where possible, original observations of numerous taxa spread throughout the major tetrapod clades. A stem-based (total-group) definition of Tetrapoda is preferred over apomorphy- and node-based (crown-group) definitions. This definition is operational, since it is based on a formal character analysis. A PAUP* search using a recently implemented version of the parsimony ratchet method yields 64 shortest trees. Differ- ences between these trees concern: (1) the internal relationships of aı¨stopods, the three selected species of which form a trichotomy; (2) the internal relationships of embolomeres, with Archeria crassidisca and Pholiderpeton scutigerum collapsed in a trichotomy with a clade formed by Anthracosaurus russelli and Pholiderpeton attheyi; (3) the internal relationships of derived dissorophoids, with four amphibamid species forming an unresolved node with a clade consisting of micromelerpetontids and branchiosaurids and a clade consisting of albanerpetontids plus basal crown-group lissamphibians; (4) the position of albenerpetontids and Eocaecilia micropoda, which form an un- resolved node with a trichotomy subtending Karaurus sharovi, Valdotriton gracilis and Triadobatrachus massinoti; (5) the branching pattern of derived diplocaulid nectrideans, with Batrachiderpeton reticulatum and Diceratosaurus brevirostris collapsed in a trichotomy with a clade formed by Diplocaulus magnicornis and Diploceraspis burkei. The results of the original parsimony run – as well as those retrieved from several other treatments of the data set (e.g. exclusion of postcranial and lower jaw data; character reweighting; reverse weighting) – indicate a deep split of early tetrapods between lissamphibian- and amniote-related taxa. Colosteids, Crassigyrinus, Whatcheeria and baphetids are progressively more crownward stem-tetrapods. Caerorhachis, embolomeres, gephyrostegids, Solenodonsaurus and seymouriamorphs are progressively more crownward stem-amniotes. Eucritta is basal to temnospondyls, with crown-lissamphibians nested within dissorophoids. Westlothiana is basal to Lepospondyli, but evidence for the monophyletic status of the latter is weak. Westlothiana and Lepospondyli form the sister group to diadecto- morphs and crown-group amniotes. Tuditanomorph and microbrachomorph microsaurs are successively more closely related to a clade including proximodistally: (1) lysorophids; (2) Acherontiscus as sister taxon to adelospondyls; (3) scincosaurids plus diplocaulids; (4) urocordylids plus aı¨stopods. A data set employing cranial characters only places microsaurs on the amniote stem, but forces remaining lepospondyls to appear as sister group to colosteids on the tetrapod stem in several trees. This arrangement is not significantly worse than the tree topology obtained from the analysis of the complete data set. The pattern of sister group relationships in the crownward part of the temnospondyl-lissamphibian tree re-emphasizes the important role of dissorophoids in the lissamphibian origin debate. However, no specific dissorophoid can be identified as the immediate sister taxon to crown-group lissamphibians. The branching sequence of various stem-group amniotes reveals a coherent set of internested character-state changes related to the acquisition of progressively more terrestrial habits in several Permo-Carboniferous forms. Key words: amniotes, characters, congruence, lissamphibians, parsimony ratchet, taxon exemplar, tetrapods, total-group. * Author for correspondence. 252 Marcello Ruta, Michael I. Coates and Donald L. J. Quicke CONTENTS I. Introduction ................................................................................................................................................ 253 II. Conflicting cladogram topologies ............................................................................................................. 255 III. Historical background ................................................................................................................................ 257 IV. Taxonomic definitions ............................................................................................................................... 259 V. Taxon exemplars ........................................................................................................................................ 261 (1) Devonian and Lower Carboniferous taxa ..................................................................................... 261 (a) Taxonomic sample ..................................................................................................................... 261 (b) Remarks ....................................................................................................................................... 262 (2) Acherontiscidae ................................................................................................................................. 262 (a) Taxonomic sample ..................................................................................................................... 262 (b) Remarks ....................................................................................................................................... 262 (3) Adelospondyli .................................................................................................................................... 262 (a) Taxonomic sample ..................................................................................................................... 262 (b) Remarks ....................................................................................................................................... 262 (4) Aı¨stopoda ............................................................................................................................................ 263 (a) Taxonomic sample ..................................................................................................................... 263 (b) Remarks ....................................................................................................................................... 263 (5) Baphetidae .......................................................................................................................................... 263 (a) Taxonomic sample ..................................................................................................................... 263 (b) Remarks ....................................................................................................................................... 263 (6) Colosteidae ......................................................................................................................................... 263 (a) Taxonomic sample ..................................................................................................................... 263 (b) Remarks ....................................................................................................................................... 263 (7) Diadectomorpha ................................................................................................................................ 264 (a) Taxonomic sample ..................................................................................................................... 264 (b) Remarks ....................................................................................................................................... 264 (8) Embolomeri and Eoherpetontidae ................................................................................................. 264 (a) Taxonomic sample ..................................................................................................................... 264 (b) Remarks ....................................................................................................................................... 264 (9) Gephyrostegidae ................................................................................................................................ 265 (a) Taxonomic sample ..................................................................................................................... 265 (b) Remarks ....................................................................................................................................... 265 (10) Lysorophia .......................................................................................................................................... 265 (a) Taxonomic sample ....................................................................................................................
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