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Systematic and Biological Studies of the Leopard Frogs (~Anapipiens Complex) of the United States

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MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 148 Systematic and Biological Studies of the Leopard Frogs (~anapipiens complex) of the United States BY j ANN E. PACE Department of Biology, Queens College, CUNY, Flushing, N. Y. (New address: James Ford Bell Museum of Natural History, University of Minnesota at Minneapolis) Ann Arbor MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN August 19, 1974 MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN FRANCIS C. EVANS, EDITOR The publications of the Museum of Zoology, University of Michigan, consist of two series-the Occasional Papers and the Miscellaneous Publications. Both series were founded by Dr. Bryant Walker, Mr. Bradshaw H. Swales, and Dr. W. W. Newcomb. The Occasional Papers, publication of which was begun in 1913, serve as a medium for original studies based principally upon the collections in the Museum. They are issued separately. When a sufficient number of pages has been printed to make a volume, a title page, table of contents, and an index are supplied to libraries and individuals on the-mailing list for the series. The Miscellaneous Publications, which include papers on field and museum techniques, monographic studies, and other contributions not within the scope of the Occasional Papers, are published separately. It is not intended that they be grouped into volumes. Each number has a title page and, when necessary, a table of contents. A complete list of publications on Birds, Fishes, Insects, Mammals, Mollusks, and Reptiles and Amphibians is available. Address inquiries to the Director, Museum of Zoology, Ann Arbor, hlichigan 48104. MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 148 Systematic and Biological Studies of the Leopard Frogs (~anapipiens complex) of the United States BY ANN E. PACE Department of Biology, Queens College, CUNY, Flushing, N. Y. (New address: James Ford Bell Museum of Natural History, University of Minnesota at Minneapolis) Ann Arbor MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN Aukgust 19, 1974 CONTENTS PAGE IN'TIIODUCTION ........................................ 1 ACKNOWLEDGMENTS ................................... 7 MATERIALS. METHODS. AND MISCELLANEOUS NOTES ........... 8 NOMENCLATURE ....................................... 11 Nomenclatural History ................................. 11 Species Characterizations and Synonymies .................... 14 VOCALIZATIONS ....................................... 30 Vocal Repertoires ..................................... 30 Functions of Vocalizations ................................31 Species Differences .................................... 34 VARIATION IN SECONDARY SEX CHARACTERS ................ 41 Vocal Sacs ......................................... 41 Mullerian Ducts ...................................... 46 Mullerian Ducts and Vocal Sacs ........................... 51 Tympana1 Size ....................................... 51 'Thumb Pads ......................................... 52 VARIATION IN CHARACTERS FOUND IN BOTH SEXES ........... 52 Dorsolateral Folds .......... ........................... 52 Tympana1 Spots ..................................... 55 Dorsal Spot Patterning and Snout Spot ...................... 58 Ventral Melanism ..................................... 61 Geographic Variation in Body Proportions .................... 63 Variation in other Morphological Characters ................... 66 VARIATION IN LEOPARD FROGS FROM THE SOUTHWEST ........ 66 INTERSPECIFIC DISTRIBUTIONAI. RELATIONSHIPS ............. 70 Parapatry ........................................... 70 Probable Cases of Species Range Disjunctions .................. 74 Apparent Range Disjunctions Omitted from Distribution Map of Rana pipiens ........................................... 78 Distributional Relationships in the Middle West ................. 79 R . pipiens and R . blairi in Nebraska ......................... 82 INTERSPECIFIC HYBRIDIZATION IN NATURE .................. 83 GROWTI3 RATES AND DEVELOPMENTAL TIME ................. 87 LIFE IllSTORIES OF LEOPARD FROGS .... .-. ................. 91 Rana pipiens ........................................ 91 Rana blairi ......................................... 93 Comparison of R . pipiens and R . blairi ...................... 94 Rarza utricz~laria ..................................... 95 DISRUPTIVE SELECTION AND PARAPATRY ................... 100 DISTRIRUTIONAL HISTORY AND RELATIONSIIIPS WITHIN THE COMPLEX .......................................... 101 R . pipiens and R . palustris ............................... 101 The Prairie Peninsula and Distributional Relationships in the Middle West ............................................. 103 Rana berlandieri and Florida Populations of R . z~tricularia ......... 104 SUMMr\RY AND CONCLUSIONS ............................ 106 APPENDIX I: LOCALITIES OF SPECIMENS EXAMINED ............ 107 APPENDIX 11: LOCALITIES OF FIELD AND CALL RECORDS ....... 130 APPENDIX 111: CATALOGUE NUMBERS OF FROGS WITH UNUSU/\I . DORSOLXTERAI, FOLDS ................................. 134 LITERATURE CITED ..................................... 135 ILLUSTRATIONS FIGURE P'AGE 1. Geographic distribution ofR . pipiens in the United States ....... 17 2. R. pipiens d, (a)view of vocal sac region and (b)dorsal view ....... 19 3. Reproduction of plate accompanying original description of R . pipiens .......................................... 4. Geographic distribution of R . utricularia ................... 5 . R . utricularia d from Kansas, (a)view of vocal sac region and (b)dorsal view .................................... 6. R . utricularia d from Florida, (a) view of vocal sac region and (b)dorsal view ..................................... 7 . Geographic distribution ofR . berlandieri in the United States ....... 8. Reproduction of illustration accompanying original description of R . berlandieri ..................................... 9. Geographic distribution of R . blairi ....................... 10. R . blairi d. (a)view of vocal sac region and (b)dorsal view ......... 11. Vocal response of (a)R . pipiens male and (b)R . palustris male to mating call of conspecific male ......................... 12. Relationship of pulse duration to pulse rate in mating call of R . utricularia ........................................ 13. Sonagrams of matingcalls ofR . utricularia (lowtemperatures) ...... 14. Sonagrams of mating calls of X . utricularia (hightemperatures) ...... 15. Sonagrams of mating calls of R . utricularia. R . pipi ns. and R . palustris (sympatric) and R. capito .....................I" 16. Relationship of pulse rate to temperature in mating calls (a) of R . utricularia and (b)of R . pipiens, R . palustris. and R . capito ...... 17. Relationship of pulse rate to temperature in mating calls of R . berlandieri and R . blairi .............................. 18. Geographic distribution of call types in the R . pipiens complex in the United States ................................... 19. Geographic distribution of call types in Indiana and Kentucky ..... 20 . Geographic distribution ofvocal sac types in the United States ...... 21 . Distribution of males with and without vestigial oviducts in the United States ..................................... 22 . Distribution ofFlorida males with and without vestigial oviducts ..... 23. Diagrammatic representation of dorsolateral fold types .......... 24. Geographic distribution of frogs with continuous and discontinuous dorsolateral folds ................................... 25 . Geographic variation in tympana1 spotting .................. 26 . Geographic variation in snout spotting ..................... 27 . Relationship between snout-urostyle length and head width for R . pipiens. R . utricularia. and R . blairi from Missouri and Iowa ........ 28 . Relationship between snout-urostyle length and head width for R . pipiens and R . utricularia from Indiana ................... 29 . Relationship between snout-urostyle length and head width for R . utricularia males from Florida with and without Mullerian ducts ..... 30. Distribution of male leopard frogs (excluding R . pipiens) with and without vestigial oviducts from southeastern Arizona ............ 31 . Distribution of male leopard frogs (excluding R . pipiens) with and without externalvocal sacsfromsoutheasternArizona ........... 32. Variation in size of adult male leopard frogs (excluding R . pipiens) from southeastern Arizona ............................. ILLUSTRATIONS Variation in dorsolateral folds of leopard frogs (excluding R. pipiens) from southeastern Arizona ............................. Geographic distribution of R. pipiens, R. utricularia, and R. blairi in Illinois ........................................ Geographic distribution of lactate dehydrogenase types in the United States ..................................... Relationship between size at seven weeks and age at metamorphosis of R. pipiens, R. utricularia, and their hybrids ................. Rclationship between size of metamorphosed frogs and age at meta- morphosis for R. pipiens and R. utricularia .................. Latitudinal variation in size of mature males of R. utricularia from the Atlantic Coastal Plain ............................. North-south variation in dorsal spot patterning in R. pipiens and R. palustris in Michigan ............................... Geographic distribution of R. palustris in the United States ......... INTRODUCTION Populations of leopard frogs occur over most of North and Central America from Great Slave Lake and
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