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Photographs included in the original manuscript have been reproduced xerographically in this copy. 35 mm slides or 6"X 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. [re Accessing the World’s Information since 1938 300 North Zeeb Road, Ann Arbor, Ml 48106-1346 USA Order Number 8820372 Revision of the genusPseudopolydesmus Attems, 1898 and its relationships to tlie North American genera of the family Polydesmidae Leach, 1815 Withrow, Charles Phillip, Ph.D. The Ohio State University, 1988 UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106 PLEASE NOTE: in all cases this material has been filmed in the best possible way from the available copy. Problems encountered with this document have been identified here with a check mark V 1. Glossy photographs or pages. 2. Colored illustrations, paper or _____print 3. Photographs with dark background____ 4. 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Other _____ ____ REVISION OF THE GENUS PSEUDOPOLYDESMUS ATTEÎ'ÎS, 1898 AND ITS RELATIONSHIPS TO THE NORTH AMERICAN GENEE^A OF THE FAMILY POLYDESMIDAE LEACH, 1815 Presented in partial fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Charles Phillip Withrow, B.S., M.S. ***** The Ohio State University 1988 Reading Committee; pproved By Barry D. Valentine John L. Crites Advisor David H. Stansbery Department of Zoology Charles A. Triplehom Copyright by Charles Phillip Withrow 1988 To My Parents ACKNOWLEDGEMENTS I would like to thank my advisor during my career at Ohio State University, B a n y D. Valentine, for his advice, course assistance, counseling, stimulation, and understanding. I would also like to thank Dr. Richard L. Hoffman, of Radford University for the ej^sure and introduction to the interesting field of diplopodology and for suggesting this topic. The stimulation, comradery, and help of graduate students in both departments of zoology and entomology has been well appreciated. I would like to thank the Department of Zoology and the Graduate School of Ohio State University for their continued financial stroport t h r o u ^ toy studies. Finally I would like to acknowledge the support of my parents, Robert C. and Jean P. Withrow, for their tolerance of ity early adventures into science. Their guidance and support of ïty decisions in life is unrepayable. 11 VITA March 13, 1955 ............... B o m - Gallipolis, Ohio 1977 ......................... B. S., Zoology, Marshall University, Huntington, West Virginia 1979 ......................... M.S., Biological Sciences, Marshall University, Huntington, West Virginia 1979 ......................... Teaching Associate, Department of Biological Sciences, Marshall University, Huntington, West Virginia 1981-87 ..................... Teaching Associate, The Ohio State University, Columbus, Ohio HJBIICATIONS 1978 Millipedes of Cabell County, West Virginia. W. Va. Acad. Sci. 38:48-50. FIELDS OF STUDY Major Field: Diplopoda Systzematics and Biogeography 1 1 1 TABLE OF CONTENTS Page DEDICATION........................................... i ACKNOWLEDGEMENTS..................................... ii VITA................................................. iii LIST OF TABLES....................................... vi LIST OF FIGURES...................................... viii LIST OF MAPS..........................................xii INTRODUCTION......................................... 1 MATERIALS AND METHODS................................ 6 Materials................................... 6 Generalized Methods.......................... 7 Phylogenetic and Classification Methods...... 10 RESULTS....... 15 Family Doratodesmidae........................ 16 Family Polydesmidae.......................... 19 Subfamily Mastigonodesminae.................. 21 Key to North American Genera................. 24 Subfamily Scytonotinae....................... 26 Speodesmus.................. 28 Scytonotus............................. 32 Utadesmus.............................. 37 Bidentoaon. ........................... 41 iv Coronodesmus........................... 45 Sipeorthus.............................. 48 Idahodesmus............................ 51 Siabfamily Polydesminae....................... 55 Polvdesmus............................. 57 Pseudopolydesmus........................ 64 Subfamily E^anerchodinae..................... 126 CLADISTIC ANALYSIS................................... 129 Character States...................... 129 Classification of Polydesmidae.................... 166 ZOOGBOGRAmY......................................... 167 The Vicariance Model.............................. 167 Distribution Patterns............................. 169 Historical Zoogeography........................... 172 BIBLIOGRAHiY......................................... 173 APPENDIX A: TABLES................................... 188 APPENDIX B: FIGURES.................................. 205 APPENDIX C: MAPS................ 254 APPENDIX D: IDCALETY DATA............................ 264 U S T OF TABLES Tables Page 1. Systematics of the Family Polydesmidae........ 189 2. List of generic character states for the Family Polydesmidae.......................... 190 3. Data set for the Family Polydesmidae and genera of Scytonotinae.............................. 192 4. Manhattan distance matrix for Family Polydesmidae........................ 194 5. Patristic distance matrix for the Family Polydesmidae................................ 195 6. Hamoplasy matrix for the Family Polydesmidae... 196 7. List of specific character states for the genus Pseudopolydesmus....................... 197 8. Data set for genus Pseudopolydesmus........... 198 9. Length and width measurements of Pseudopolydesmus species, males and females, with range, mean, standard deyiation, and number measured.............................. 199 10. W/L (width/length) and WL (width X length) measurements of Pseudopolydesmus species, males and females, with range, mean, standard deyiation, and number measured............... 200 11. Distance from telopodite tip to base of mesial and ectal (E^-^) processes as a percentage of total telopodite length........ 201 12. Manhattan distance matrix for the species in the genus Pseudopolydesmus................... 202 13. Patristic distance matrix for the species in the genus Pseudopolydesmus................... 203 yi 14. Hamoplasy matrix for the species in the genus Pseudopolydesmus............................. 204 Vll U S T OF FIGURES Figures Page 1-3. Dorsum, dorsal view, male. 1. Cerastelachvs cavemicola; 2. Mastigonodesmus vignai; 3. Epanerdhodus bidens.......................... 206 4-6. Dorsum, dorsal view, male. 4. Speodesmus echinourus; 5. Scytonotus granulatus; 6. Utadesmus henriensis......................... 207 7-9. Dorsum, dorsal view, male. 7. Bidentogon helferorum; 8. Coronodesmus bituberculatus; 9. Speorthus tugahbius......................... 208 10-12. Dorsum, dorsal view, male. 10. Idahodesmus dentatus; 11. Polvdesmus inconstans; 12. Pseudopolydesmus serratus.................... 209 13-15. R i ^ t gonopod, lateral view. 13. Cerastelaolivs cavemicola : 14. Mastigonodesmus vignai; 15. Epanerdhodus bidens.......................... 210 16-18. left gonopod of Speodesmus edhinourus. 16. medial view; 17. lateral view; 18. ventral view........................................ 211 19-20. Dorsal view. 19. Pseudopolydesmus erasus, male; 20. Scytonotus granulatus male and female 212 21-23. R i ^ t gonopod of Scytonotus granulatus. 21. medial view; 22. lateral view; 23. ventral view........................................ 213 24-26. R i ^ t gonopod of Utadesmus henriensis. 24. medial view; 25. lateral view; 26. ventral view........................................ 214 27-29. R i ^ t gonopod of Bidentogon helferorum. 27. medial view; 28. lateral view; 29. ventral view........................................ 215 30-32. R i ^ t gonopod of Coronodesmus bituberculatus. 30. medial view; 31. lateral