This dissertation has been microfilmed exactly as received 68-12,857 McGRAW, James Carmichael, 1928- A STUDY OF COSMOCERCOIDES DUKAE (HOLL, 1928) WILKIE, 1930 (NEMATODA: COSMO- CERCIDAE) FROM AMPHIBIANS IN OHIO. The Ohio State University, Ph.D., 1968 Zoology University Microfilms, Inc., Ann Arbor, Michigan A STUDY OP COSMOCERCOIDES DUKAE (HOLL, 1928) WILKIE, 1930 (NEMATODA: COSMOCERCIDAE) PROM AMPHIBIANS IN OHIO DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University James Carmichael McGraw, A.B., M.S. ****** The Ohio S tate U niversity 1968 Approved by // /Adviser/Adviser D§partm£ntnt ofof ZoologyZoology and Entomology ACKNOWLEDGMENTS The writer wishes to thank Dr. John L. Crites of the Department of Zoology and Entomology at Ohio State University for furnishing nematodes from Ohio amphibians. Thanks are also due the following persons who helped to locate nematode specimens: Dr. Barry Valentine, Department of Zoology and Entomology at Ohio State University; Dr. Russell R. Williams, Waynesburg College; Dr. Paul D. Harwood, Hess and Clark; Dr. W. G. Inglis, Brisith Museum; W. W. Becklund, M.B. Chitwood and J.M. Humphrey, U.S.D.A.- Beltsville Parasitology Laboratory; Dr. Roy C. Anderson, University of Guelph; Dr. Robert Ph. Dollfus, Museum National d'Histoire Naturelle, Paris; Dr. J. E. Alicata, University of Hawaii-; V. N. Converse, University of Adelaide; M. Bravo Hollis, National University of Mexico; N. T. Talbot, Departmentcf Agriculture of the Territory of Papua and New Guinea; W. A. Reid of the University of Kansas; Dr. R. U. Gooding, University of Singapore; and the late A. C. Walton of Knox College. Thanks are due Celeste Taft for identification of terrestrial molluscs; Dr. C. J. Zenisek and Warren Dolfin for aid in the identification of amphibians; Mr. Igors Gortinskis and M. A. Ewert for translation of Russian; i i Dean Harold J. Perkins of the Faculty of Science and Mathematics at Plattsburgh State University College of Arts and Science for aid and encouragement; Dr. Russell Skavaril for statistical analyses; and Dr. Robert J. Shields of City College, City University of New York, for many suggestions. I would like to express my appreciation to Mrs. Dorothy Schreck and the library staff for easing the literature search. Special thanks go to Dr. Joseph N. Miller whose advice and criticisms throughout this study were very much appreciated. iii VITA March 20, 1928 Born - Martins Perry, Ohio 1951 .................... A.B., Oberlin College, Oberlin, Ohio 1955-1964 . Graduate Assistant, Department of Zoology and Entomology, The Ohio S ta te U n iv ersity , Columbus, Ohio 1957 ..................... M.S., The Ohio State University, Columbus, Ohio 1964-1968 . Assistant Professor of Biology, State University College of Arts and Science, Plattsburgh, New York FIELDS OP STUDY Major Field: Zoology Studies in Invertebrate Zoology. W. J. Kostir Studies in Medical Entomology. C. E. Venard Studies in Vertebrate Zoology. J. W. Price Studies in Parasitology. J. N. Miller i v CONTENTS Page ACKNOWLEDGMENTS ............................................................................................ l i VITA .................................................................................................................... iv TABLES...........................................................................................................................v ii INTRODUCTION ...................................................................................................... 1 HISTORICAL REVIEW ...................................................................................... 4 MATERIALS AND METHODS ............................................................................ 11 OBSERVATIONS AND RESULTS ....................................................................... 15 Occurrence of Cosmocercoid.es dukae ......................................... 15 Morphology of Cosmocercoides' dukae ......................................... 20 C u t i c l e ..................................................................................................... 21 Cephalic region ..... ........................................................ 23 E s o p h a g u s................................................................................................ 24 I n t e s t i n e ................................................................................................. 25 Female reproductive system ........................................................ 26 Male reproductive system ............................................................. 28 Special male anatomy ....................................................................... 29 Descriptive Summary of Cosmocercoldes dukae from Ohio amphibians ....................................................................... 30 Observations on the Holotype of Cosmocercoldes dukae (Holl, 1928) ............................................................................ 33 Comparison of Selected Groups of Cosmocercoldes dukae (Holl, 1928) ............................................................ 34 Observations on Cosmocercoldes pulcher W ilkie, 1930 ........................................................................................... 52 Additions to the Description of Cosmocercoldes p u lch er—W ilk ie, 1930 54 An Emended D escription of the Genus Cosmocercoides—W ilkie, 1930 ................................................... 57 Observations on the Life History of Cosmocercoides dukae (Holl, 1928 ......................................... 58 DISCUSSION......................... 68 SUMMARY............................................................................................................... v CONTENTS (Contd.) Page APPENDIX...................................................................... 82 LITERATURE CITED........................................................................................... 87 PLATES................................................................................................................... 91 v i TABLES T able Page 1. Cosmocercoldes dukae from Ohio Amphibians, A pril 19 6 3-July 1964 16 2. Measurements (in mm) of Cosmocercoides dukae from one American toad (Bufo amerlcanus) from Ohio and C. dukae reported by Anderson (I960) from terrestrial molluscs in Ontario . 36 3. Derived statistics (percentages) of C. dukae from one American toad from Ohio and C. dukae reported by Anderson (i 960 ) from terrestrial molluscs in Ontario .................................... 37 4. Male measurements, Group I .................................................... 39 5. Female measurements, Group I ............................................... 40 6 . Male derived s t a t i s t i c s , Group I ..................................... 41 7. Female derived statistics, Group I ............................... 42 8. Male measurements, Group I I ................................................... 43 9. Female measurements, Group I I ............................................... 44 10. Male derived s t a t i s t i c s , Group I I ..................................... 45 11. Female derived statistics, Group II ............................... 46 12. Male measurements, Group I I I ............................................... 47 13. Male derived s t a t i s t i c s , Group I I I ................................ 47 14. Female measurements, Group I I I .......................................... 48 15. Female derived statistics, Group III ........................... 48 16. Infection of snails with Cosmocercoides dukae from amphibians ............................................................................ 61 v i i TABLES (Contd.) T ab le Page 17. Infection of snails with Cosmocercoides dukae by ingestion of frog fe c e s ................................................ 63 18. A comparison of the species of the genus Cosmocercoides .............................................................................. 70 v i i i INTRODUCTION The nematode Cosmocercoides dukae (Cosmocercidae) has been found in a variety of hosts. In fact, the great diversity of animals that apparently serve as definitive hosts raises some doubt whether all such reports are con­ cerned with Cosmocercoides dukae or whether some of the specimens that have been assigned to this species may belong to a closely related one which has not been defined taxonomically. It must be admitted that a species such as C. dukae which has been reported from such diverse hosts as molluscs, as well as such vertebrates as amphibians and reptiles, is either very versatile in its acceptance of host animals or many closely related species of nematodes may be involved. On the other hand, since specimens of C. dukae from the same species of host exhibit such great variation in size and shape, it might be that there are fewer valid species in the genus Cosmocercoides than are now accepted, and that those species whose descriptions have been based on relatively few specimens may actually be synonyms of C. dukae. An attempt was made to identify specimens of what appeared to be Cosmocercoides dukae from the American Toad, Bufo americanus, and from the Eastern Spadefoot Toad, Scaphiopus holbrooki. Specimens from the former host were larger than those from the latter, and, seemingly more important, the nematodes from Bufo americanus contained more papillae than those from Scaphiopus holbrooki. Since vari­ ation in the number of papillae