Reconstructing a radiation: the chiton genus Mopalia in the north Pacific
Ryan P. Kelly A B C E and Douglas J. Eernisse DA Columbia University, New York, USA.
B American Museum of Natural History, New York, USA.
C Present address: Stanford University, Hopkins Marine Station, Pacific Grove, California, USA.
D California State University, Fullerton, California, USA.
E Corresponding author. Email: rpk@stanford.edu
Invertebrate Systematics 22(1) 17-28 https://doi.org/10.1071/IS06021
Submitted: 13 June 2006 Accepted: 13 December 2007 Published: 18 March 2008
Abstract
The chiton genus Mopalia Gray, 1847 is highly speciose despite showing little morphological differentiation. Many of the 24 extant species are conspicuous, large-bodied and ecologically important today, but pre-Pleistocene fossils for the genus are rare. Here, we use a combined analysis of four gene regions (16S and COI mtDNA, 18S and 28S rDNA) to estimate the phylogenetic relationships for Mopalia species and use the inferred phylogeny to analyse the group’s biogeography and patterns of speciation. We then use these molecular data to distinguish between two alternative interpretations of the fossil record, as there is a large temporal gap between the oldest fossils tentatively identified as Mopalia and the next oldest fossils (Miocene versus Plio-Pleistocene). Based on the estimated substitution rates from a wide variety of other marine animals, we conclude that the observed rates in Mopalia are consistent with a Miocene origin for the genus. Given this age for the group and assuming a molecular clock, most speciation events in Mopalia are inferred to have occurred on average ~5 Mya. The phylogenetic results indicate that most of the speciation events leading to extant species must have occurred along the western North American coast, though there appear to have been multiple spreading events across the Pacific. When considered along with results for the many other near-shore taxa that have similar distributions to Mopalia, our findings suggest the emergence of a coherent historical biogeography of the northern Pacific.
Additional keywords: Mollusca, neogene, phylogeny, polyplacophora, speciation.
Acknowledgements
RPK acknowledges R. Desalle for financial and intellectual support and for bench space at AMNH, and financial support provided by the Columbia University Department of Ecology, Evolution, and Environmental Biology, The National Science Foundation GK-12 Fellowship (No. DGE0231875 to Columbia University), The University of Washington’s Friday Harbor Marine Laboratories, The Lerner-Gray Grants for Marine Research, and the American Museum of Natural History. An earlier version of this study was first presented as part of RPK’s dissertation research, and the author gratefully acknowledges the support of his dissertation committee: Rob Desalle, Andrew Baker, Chris Raxworthy, Shahid Naeem, and DJE. DJE acknowledges support from the California State University, Fullerton and the Sea of Cortez Expedition and Education Project, Stanford University and the Instituto Nacional de Ecología in Mexico, for allowing participation as part of this expedition that made these Mexican comparisons possible, and likewise L. Lovell, Scripps Institution of Oceanography, for the loan of specimens dredged off the Pacific coast of Mexico. Much of the analysis and writing were completed while DJE was completing a sabbatical jointly supported by CSUF and a fellowship from the National Evolutionary Synthesis Center (NESCent), NSF #EF-0423641. We thank especially A. Draeger for helping to identify problematic specimens and for collecting all of our subtidal samples in the Monterey area. Others who have contributed to our collecting efforts for this study include B. Allen, A. Baldwin, R. Clark, C. Colt, G. Eckert, G. & S. Jeffrey, H. Keller, J. Manke, C. McGary, J. Pearse, M. Poulson, L. Rogers-Bennett, and T. Stebbins. Our studies benefited from the cooperation of the Reserve Managers at the Landels-Hill Big Creek, Kenneth S. Norris Rancho Marino, and Bodega Marine Reserves of the University of California Natural Reserve System. We thank H. Chaney, F. Hochberg, and P. Valentich-Scott who have greatly assisted us in curation of our specimen vouchers at the Santa Barbara Museum of Natural History, and J. Feinstein, Ambrose Monell Cryo Collection of the American Museum of Natural History, for help curating frozen tissue vouchers. DJE is grateful to several scientists consulted during revisions of this manuscript, including D. Zwickl for advice on phylogenetic analysis, C. Cunningham for help with molecular rates calculations, and G. Vermeij, L. Marinkovich, C. Powell, II, K. Amano, H. Saito, A. Oleinik, J. Barron, M. Lyle, L. Groves, and G. Kennedy for considerable help with interpreting the fossil record of Mopalia. We are grateful to J. Sullivan, M. Manier, E. Jacobs-Palmer, S. Palumbi, M. Whitlock, S. Otto, B. Sidlauskas, members of the NESCent Journal Club, and several anonymous reviewers, whose comments improved the manuscript significantly.
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