East Tennessee State University Digital Commons @ East Tennessee State University Electronic Theses and Dissertations Student Works 5-2015 A Systematic Review of the Soricimorph Eulipotyphla (Soricidae: Mammalia) from the Gray Fossil Site (Hemphillian), Tennessee Joshua Doby East Tennessee State University Follow this and additional works at: https://dc.etsu.edu/etd Part of the Biodiversity Commons, Evolution Commons, and the Zoology Commons Recommended Citation Doby, Joshua, "A Systematic Review of the Soricimorph Eulipotyphla (Soricidae: Mammalia) from the Gray Fossil Site (Hemphillian), Tennessee" (2015). Electronic Theses and Dissertations. Paper 2526. https://dc.etsu.edu/etd/2526 This Thesis - Open Access is brought to you for free and open access by the Student Works at Digital Commons @ East Tennessee State University. It has been accepted for inclusion in Electronic Theses and Dissertations by an authorized administrator of Digital Commons @ East Tennessee State University. For more information, please contact [email protected]. A Systematic Review of the Soricimorph Eulipotyphla (Soricidae: Mammalia) from the Gray Fossil Site (Hemphillian), Tennessee _______________________________________ A thesis presented to the faculty of the Department of Geosciences East Tennessee State University In partial fulfillment of the requirements for the degree Master of Science in Geosciences _______________________________________ by Joshua R. Doby May 2015 _______________________________________ Dr. Steven C. Wallace, Chair Dr. Jim I. Mead Dr. Blaine W. Schubert Keywords: Soricidae, Gray Fossil Site, Miocene, Paenelimnoecus , Phylogenetics, Venom ABSTRACT A Systematic Review of the Soricimorph Eulipotyphla (Soricidae: Mammalia) from the Gray Fossil Site (Hemphillian), Tennessee by Joshua R. Doby Due in part to the incompleteness of the Cenozoic fossil record in the eastern U.S., the evolution and immigration of shrews (Soricidae) is not well understood. A rich soricid fauna from the Gray Fossil Site (GFS), Washington County, TN, has enabled many new inferences to be made. There are 7 new species in 6 genera: Paenelimnoecus , “Blarinella”, Petenyia, Tregosorex, Crusafontina, and Gen et sp. nov. GFS species of the genera Paenelimnoecus , “Blarinella”, and Petenyia are the first occurrence of each genus in the New World. Tregosorex, Crusafontina, and the N.A. taxon Limnoecus all have their latest documented occurrence at the GFS, extending their temporal range by at least 1 million years. “Blarinella ” sp. nov. has a complete lateral groove in the inferior incisor, providing the earliest evidence for venom in soricids by at least 4 million years. GFS taxa also provide insight into the evolution of both Soricini and Blarinini. 2 ACKNOWLEDGEMENTS I would like to think the chair of my committee Dr. Steven (Wally) Wallace and my other committee members Dr. James Mead and Dr. Blaine Schubert for their support and guidance throughout this project. I would also like to thank: Shawn Haugrud for his work preparing the specimens used in this thesis and for discussions about the specimens, Sandy Swift for her work picking through the sediment and her initial identifications of the specimens as shrews, Brett Woodward for allowing me access to the specimens and providing them with new numbers when necessary, Dr. Joyner for his help with making a map, Dr. Rountre for access to specimens at the University of Michigan, and Dr. Reumer for his helpful comments about the systematics within the thesis. I would like to thank my fellow graduate students and Gray fossil Museum staff for all of their support during my time at ETSU. Finally, I would like to thank Connor Behrmann for his limitless patience and support over the last year of this project. 3 TABLE OF CONTENTS Page ABSTRACT ..................................................................................................................... 2 ACKNOWLEDGEMENTS ............................................................................................... 3 LIST OF FIGURES ......................................................................................................... 7 LIST OF ABBREVIATIONS .......................................................................................... 10 Chapter 1. INTRODUCTION .......................................................................................... 11 Background ............................................................................................. 11 Soricid Fossil Record .............................................................................. 12 European ...................................................................................... 12 Asian ............................................................................................ 12 North American ........................................................................... 13 2. MATERIAL AND METHODS ..................................................................... 15 Methods of Differential Diagnosis ......................................................... 15 Morphology ............................................................................................. 17 Locality (Gray Fossil Site) ....................................................................... 19 3. SYSTEMATIC PALEONTOLOGY ............................................................... 22 Soricidae .................................................................................................. 22 Allosoricinae ................................................................................ 22 Paenelimnoecus ............................................................... 23 Limnoecinae ................................................................................ 25 Limnoecus ........................................................................ 26 4 Soricinae .................................................................................................. 28 Blarinellini ................................................................................... 28 Tregosorex ....................................................................... 28 Petenyia ........................................................................... 32 “Blarinella ” ..................................................................... 36 Gen. et sp. nov. ............................................................... 39 Anourosoricini ......................................................................................... 46 Crusafontina .................................................................... 46 4. PHYLOGENETICS ......................................................................................... 50 Past Attempts .............................................................................. 50 Evolutionary Implications ....................................................................... 53 Relationship Between Blarinellini and Blarinini ........................ 53 Paenelimnoecus ........................................................................... 54 Relationship of “Blarinella” to Anchiblarinella wakeeneyensis ................................................... 55 Upper Antemolars as an Evolutionary Marker ........................... 56 Soricini ......................................................................................... 57 Discussions of Problems with Sorex ............................................ 58 Circular Reasoning in Phylogenies ........................................................ 61 5. DISCUSSION .................................................................................................. 63 Paleobiology ............................................................................................ 63 Venom ......................................................................................... 63 Paleoecology ............................................................................................ 65 5 Interpretations of GFS Soricid Associations .............................. 67 Arguments for Multiple Similar Species at Single Site .............. 67 Niche Partitioning ...................................................................... 67 Preservation Bias ........................................................................ 69 6. CONCLUSIONS ............................................................................................. 71 REFERENCES .................................................................................................................. 73 APPENDICES .................................................................................................................. 81 Appendix A: List of 40 Soricid Taxa Used in the Data Matrix of the Phylogenetic Analysis Including 45 Character States. ..................................................... 81 Appendix B: Issues with Particular Characters ................................................... 84 Appendix C: 17 Character states with Primitive State Based on Comparisons with Genus Domnina ............................................ 92 Appendix D: Translation of Ziegler’s Description of Antesorex ........................ 94 VITA .............................................................................................................................. 95 6 LIST OF FIGURES Figure Page 1. United States with location of the Gray Fossil Site, TN ............................................... 14 2. Sorex mandible with functionally integrated regions shaded Modified from Badyaev