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Why i 7 Per » 1, ah ' fee UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY VOLUME £5 = L9G hg966 EDITORS E. RayMonp Hatt, Chairman TuHeEoporE H. EaTon, JR. Henry S. FircH FRANK B. Cross MusEUM OF NATURAL HISTORY UNIVERSITY OF KANSAS LAWRENCE 1966 Museum oF NATURAL HISTORY UNIVERSITY OF KANSAS LAWRENCE PRINTED BY ROBERT R. (BOB) SANDERS, STATE PRINTER TOPEKA, KANSAS Lae ie) 15. CONTENTS OF VOLUME 15 The amphibians and reptiles of Michoacan, México. By William E. Duell- man. Pp. 1-148, pls. 1-6, 11 figs. December 20, 1961. Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962. A new species of frog (genus Tomodactylus) from western México. By Robert G. Webb. Pp. 175-181, 1 fig. March 7, 1962. Type specimens of amphibians and reptiles in the Museum of Natural His- tory, The University of Kansas. By William E. Duellman and Barbara Berg. Pp. 183-204. October 26, 1962. Amphibians and reptiles of the rainforests of southern El Petén, Guatemala. By William E. Duellman. Pp. 205-249, pls. 7-10, 6 figs. October 4, 1963. A revision of snakes of the genus Conophis (family Colubridae, from Mid- dle America). By John Wellman. Pp. 251-295, 9 figs. October 4, 1963. A review of the Middle American tree frogs of the genus Ptychohyla. By William E. Duellman. Pp. 297-349, pls. 11-18, 7 figs. October 4, 1963. Natural history of the racer Coluber constrictor. By Henry S. Fitch. Pp. 351-468, pls. 19-22, 20 figs. December 30, 1963. A review of the frogs of the Hyla bistincta group. By William E. Duell- man. Pp. 469-491, 4 figs. March 2, 1964. . An ecological study of the garter snake, Thamnophis sirtalis. By Henry S. Fitch. Pp. 493-564, pls. 23-25, 14 figs. May 17, 1965. Breeding cycle in the ground skink, Lygosoma laterale. By Henry S. Fitch and Harry W. Greene. Pp. 565-575, 3 figs. May 17, 1965. Amphibians and reptiles from the Yucatan Peninsula, México. By William E. Duellman. Pp. 577-614, 1 fig. June 22, 1965. . A new species of turtle, genus Kinosternon, from Central America. By John M. Legler. Pp. 615-625, pls. 26-28, 2 figs. June 20, 1965. . A biogeographic account of the herpetofauna of Michoacan, México. By William E. Duellman. Pp. 627-709, pls. 29-36, 5 figs. December 30, 1965. Amphibians and reptiles of Mesa Verde National Park, Colorado. By Charles L. Douglas. Pp. 711-744, pls. 37, 38, 6 figs. March 7, 1966. Index, pp. 745-770. 7. ae mie ne ee iat oy Se) ae ' = (i 1 pate (AE. & : oes s ae ae ee) Ta. » ge Dat =artthes tf r ot 1 e= (ona! Ss i lege? Ane Li tie wie Ga?U TT? eo serethG (Overy ¢ i) i | 7 7 . UNIVERSITY OF KANSAS PUBLICATIONS MuSEUM OF NATURAL HISTORY Volume 15, No. 1, pp. 1-148, pls. 1-6, 11 figs. Yee See rece neepeeomte December 20, 1961 ah a in "a BEND Lise, 4 a og $ ~ BM LaAsies LORELEI TICE rr henennnth steepest The Amiptabians and Reptiles of Michoacan, Mexico BY i eee WILLIAM E, DUELLMAN UNIVERSITY OF KANSAS LAWRENCE Y 1961 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Institutional libraries interested in publications exchange may obtain this series by addressing hie! eee Librarian, University of Kansas Library, Lawrence, Kansas. ones for individuals, persons working in a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of Kansas, Lawrence, Kansas. There is no provision for sale of this series by the University Library, which meets institutional requests, or by the Museum of Natural History, which meets the requests of individuals. However, when individuals request copies from the Museum, 25 cents should be included, for each separate number that is 100 pages or more in length, for the purpose ‘of defraying the costs of wrapping and mailing. * An asterisk designates those numbers of which the Museum’s supply (not the Library’s supply) is exhausted. Numbers published to date, in this series, are as follows: Vol. 1. Nos. 1-26 and index. Pp. 1-688, 1946-1950. *Vol. 2. (Complete) Mammals of Washing tom, By Walter W. Dalquest. Pp. 1-444, 140 figures in text. April 9, 1948. Vol. 8. *1, The _avifauna of Micronesia, its origin, evolution, and distribution. By Rol- lin H. Baker. Pp. 1-859, 16 figures in text. June 12, 1951. *2. A quantitative study of the nocturnal migration of Aig By George H. Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951. 8. Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 478- 580, 49 figures in text, 18 tables. October 10, 1951. 4. Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and Winter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, Index. _Pp. 651-681. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, $31 figures in text. December 27, 1951. . 5. Nos. 1-37 and index. Pp. 1-676, 1951-1958. *Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952. 7. ek, Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-308, 73 figures in text, 87 tables. August 25, 1952. 2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry vi ee and Lewis L. Sandidge. Pp. 805-338, 5 figures in text. August ' The silky pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker. Pp. 889-347, 1 figure in text. February 15, 1954. North American jumping mice (Genus Zapus). By vile H. Krutzsch. Pp. 849-472, 47 figures in text, 4 tables. April 21, 1954. Mammals from Southeastern Alaska. By Rollin H. Baker and James §. Findley. Pp. 478-477. April 21, 1954. Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479- 487. April 21, 1954. Subspeciation in the montane meadow mouse. Microtus montanus, in Wyo- ming and Colorado. By Sydney Anderson. Pp, 489-506, 2 figures in text. July 28, 1954. 8. A new subspecies of bat (Myotis velifer) from southeastern California and Arizona. By Terry A. Vaughan. Pp. 507-512. July 28, 1954. 9. Mammals of the San Gabriel mountains of California. By Terry A. Vaughan. Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954. 10. A new bat (Genus Pipistrellus) from northeastern Mexico. By Rollin H. Baker. Pp. 583-586. November 15, 1954. 11. A new subspecies of pocket mouse from Kansas. By E. Raymond Hall. Pp. 587-590. November 15, 1954 12. Geographic variation in the pucker gopher, OCU castanops, in Coa- huila, Mexico, By Robert J. Russell and Rollin H. Baker. Pp. 591-608. March 15, 1955. 18. A new cottontail (Sylvilagus jet naee fon northeastern Mexico. By Rollin H. Baker. Pp. 609-612. April 8, 1 14.. Taxonomy and Gare aeon of some nd a shrews. By James S. Findley. Pp. 618-618. June 10, De 15. The pigmy woodrat, Nodlonts goldmani, its distribution and systematic posi- tion. By Dennis G. Rainey and Rollin H. Baker. Pp. 619-624, 2 figures in text. June 10, 1955. Index. Pp. 625-651. (Continued on inside of back cover) ARIS UNIVERSITY OF KANSAS PUBLICATIONS MusEeuM OF NATURAL HIsTORY Volume 15, No. 1, pp. 1-148, pls. 1-6, 11 figs. December 20, 1961 The Amphibians and Reptiles of Michoacan, Mexico BY WILLIAM E., DUELLMAN UNIVERSITY OF KANSAS LAWRENCE 1961 UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HIsTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 15, No. 1, pp. 1-148, pls. 1-6, 11 figs. Published December 20, 1961 UNIVERSITY OF KANSAS Lawrence, Kansas | Tre q = = ny ; Le gt toe H EP j { i Bay BLS fear a foe 1 pe th & ieee Sa é | i} 12? } Gee tb % H LoS IRENA re rer A PRINTED IN THE STATE PRINTING PLANT TOPEKA, KANSAS 1961 The Amphibians and Reptiles of Michoacan, México BY WILLIAM E. DUELLMAN CONTENTS PAGE INTRODUCTION? cen cc sbeosehaci’ cecmerhsr eee ey ont Qekey NOOR SE Rae ore: 3 Acknowledgments, Fon 6 06 co hota ecard Sate ine cee rere Me 5 ETIStORICAL A COOUTIE Hes tat hol le iek ice ena at ASU oS CEE ere vA INA TURAL CIGAINDSCAPE eect ve boca ian chacs ites re ee ee Ee ier 9 GEOGRAPHY OF (THE) HIERPETOFAUNA |. a fos. tos See ene eee oe 13 ANNOTATED) [UIST OF SPEGIES: ; (4saiaag Goch nena eae ee eee 13 Amphibia s, sca. 4235 22 Sorel Bas cts dade ee oie Oe ee ee 14 CRIED oF cca Sete Bie nda ERE: A col SINS ves ORCC Pc 14 SalIOMELAy ca soteccs cesee toe te eee ae coos Le SEER ET A a ee 20 LEYS oT lag RR cathe Me rl ee ih yee tl a PR, PR aA Y sch end bin highs Uc 2 56 mestudines! 1. S)....0. Me eee Be ee, 56 Crocodilia: /}24ia2 ea), ee Aes OR ee eee 58 PS a ene HORE ECS ea ON Wa ME na aE es ae yr a sig aoe agra ake 59 Serpentes) 20.45 « s.ciyc nt Acyo.s ees Sracncuae (esau, Nise Gk eee aoe ee 88 SPECIES OF QUESTIONABLE. OCCURRENCE... 2). 5-0-5 2a) (ce ees a eee 124 GAZETTEER Soe es ee re eee ee tare Peer eas Ora OOS LO ere 129 SUNENEARY 25108 SS Te PO a PS, a he OE ee oT Re Ree 141 [LaTERATURE; ‘CITED (hoes SES oS ee Bn Be See 142 INTRODUCTION For almost 80 years North American herpetologists have been making extensive collections of reptiles and amphibians in México. Some parts of the country, because of their accessibility, soon be- came relatively well known; other regions lying off the beaten path were bypassed or inadequately sampled. Principally in the last decade herpetologists have been entering regions from which no collections previously were available in an attempt to fill gaps in known distributions and to discover unknown species of animals. In 1950 Dr. Donald D. Brand led an exploration party from the Uni- versity of Texas to the poorly explored and faunistically unknown region of southwestern Michoacan. James A. Peters accompanied Brand and collected amphibians and reptiles. In 1951 I welcomed the opportunity to accompany Brand on a second expedition to southwestern Michoacin. Such was the beginning of my interest (8) 4 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. in the herpetofauna of the region. I have been fortunate to return to Michoacan on four successive trips, all of which had as their purpose the accumulation of data on the herpetofauna that would result in a survey of the component species and an analysis of their distribution. My original intention was to amplify Peters’ (1954) study based on the collections made by him in 1950 and by me in 1951 in the Sierra de Coalcomaén. But it soon became evident that in order to understand the relationships of the herpetofauna of the Sierra de Coalcoman, the species inhabiting the Tepalcatepec Valley and ad- jacent mountain ranges would have to be studied. In the course of making that study I examined all specimens from Michoacan already in museums. There have been few detailed herpetofaunal studies in México. The first such study of any consequence was that by Bogert and Oliver (1945) on the herpetofauna of Sonora. In that paper the authors analyzed the fauna from a geographic view and showed the transition from tropical species in the southern part of the state to members of the Sonoran Desert assemblage to the north. Martin (1958) made a detailed study of the herpetofauna of the Gémez Farias region in southern Tamaulipas; he emphasized the ecological distribution of amphibians and reptiles in that region with special reference to cloud forests. Duellman (1958c) presented a prelimi- nary geographic analysis of the herpetofauna of Colima with special reference to the continuity of the species inhabiting the lowlands. Zweifel (1960) discussed in detail the herpetofauna of the Tres Marias Islands and commented on the derivation of the fauna. Duellman (1960d) provided a detailed account of the geographic distribution of the amphibians known to occur in the lowlands of the Isthmus of Tehuantepec and attempted to account for the pres- ent patterns of distribution. The present report is the first of two parts dealing with the her- petofauna of Michoacan. The purpose of this part is to present a full account of the species of amphibians and reptiles known to inhabit the state of Michoacan; the accounts of the species are ac- companied by a brief description of the natural landscape and of the various assemblages of species comprising the major faunistic groups within the region. A gazetteer of collecting localities is appended. The second part of the study, now in preparation, deals with the ecological and historical geography of the herpetofauna. Since the present part will be of interest primarily to systematic AMPHIBIANS AND REPTILES OF MICHOACAN 5 herpetologists, I have decided to separate it from the more general material of interest to biogeographers. One of the major problems that faces the worker undertaking a faunal study is the presence of species or genera of unsettled sys- tematic status. My work in Michoacan has been no exception; fifteen separate studies were undertaken in an attempt to solve sys- tematic problems in certain groups. Some systematic problems still remain but are of little consequence insofar as the entire faunal picture is concerned, or are so involved as to be impractical to undertake at this time. In accounts of species, such problems are mentioned in the hope that they will interest some worker who will be inclined to investigate them. ACKNOWLEDGMENTS While engaged in the study of the herpetofauna of Michoacan I have built up a debt of gratitude to many individuals, without whose aid my ambition to complete my study never would have been realized. I am especially grateful to those individuals who ac- companied me in the field; Lee D. Beatty, Richard E. Etheridge, Carter R. Gilbert, Fred G. Thompson, Jerome Tulecke, and John Wellman offered stimulating companionship and valuable assist- ance. On many occasions they suffered hardships on behalf of my interests. Studies of my own specimens have been augmented by material from other institutions. For permitting me to examine specimens in their care I am indebted to W. Frank Blair, Charles M. Bogert, Doris M. Cochran, William B. Davis, James R. Dixon, the late Emmett R. Dunn, Josef Eiselt, Alice G. C. Grandison, Norman Hartweg, Robert F. Inger, Arthur Loveridge, the late Karl P. Schmidt, Hobart M. Smith, Robert C. Stebbins, Margaret Storey, Edward H. Taylor, and Richard G. Zweifel. Several people have aided me in the study of specimens and in the analysis of data; I am grateful to Donald D. Brand, who first introduced me to Michoacan; since that time I have benefited much from his knowledge of the area. James A. Peters provided me with essential information concerning his field work in southern Michoa- can in 1950. James R. Dixon and Floyd L. Downs have permitted me to use freely the material and data that they accumulated in their recent field work in Michoacan. Norman E. Hartweg allowed me to use the specimens and data that he gathered in his survey of the herpetofauna in the region of Volcan Paricutin. L. C. Stuart, Charles 6 Unrversiry OF Kansas Pusts., Mus. Nat. Hist. F. Walker, and Richard G. Zweifel have helped in unraveling some of the systematic and distributional problems. I am especially grateful to my wife, Ann, who for six months helped me track down elusive species and explore new areas. Fur- thermore, she has stimulated me to carry this study to completion. Many people in Michoacan favored the field parties with quar- ters, transportation, and valuable information, which greatly fa- cilitated the field work. In this respect I am especially indebted to Ingeniero Ruben Erbina of Ingenieros Civiles Asociados, who not only let us use his home as our headquarters, but through a letter of introduction gave us the “key” to southern Michoacan. Ingeniero Pedro Tonda aided us in Arteaga and San Salvador. Ingeniero Anastacio Peréz Alfaro of the Comisién Tepalcatepec in Uruapan provided the latest maps of southern Michoacan and much essential information pertaining to travel conditions in the area. Sefor Nefty Mendoza gave us a home in Dos Aguas; this kindness allowed us to work in this interesting region during the height of the rainy season. Mr. and Mrs. Bob Thomas let us make use of their facilities at Hacienda Zirimicuaro. The naval officers at the Estacién Marina at Playa Azul made pleasant what might have been a dreadful stay in that small coastal village. To the managers and pilots of Lineas Aereas Picho in Uruapan I owe special thanks for going out of their way on more than one occa- sion to transport a stranded snake-hunter. Throughout the months of field work beginning in 1955 I constantly have been aided by the authorities and workers of the Comisién Tepalcatepec, a sub- division of the Secretaria de Caminos y Obras Publicas, and of the private corporation, Ingenieros Civiles Asociados. Much of the field work in Michoacan was made possible only through the co-operation of the natives who supplied mules, acted as guides, and aided in the collection of specimens. I have learned a great deal from these people. They will never see this report. Their work as guides, muleteers, and collectors greatly assisted me with the mountains of equipment that had to be piled on the backs of scrawny mules for transportation to places where the natives sel- dom trod. Their efforts in behalf of Don Guillermo never will be forgotten; I extend an especially hearty muchas gracias to Benjamin, Ignacio, Jesus, Lorenzo, Mariano, and Remigio. Much of the work on this report was done while I was associated with the Museum of Zoology at the University of Michigan. I thank Norman E. Hartweg and T. H. Hubbell for making available AMPHIBIANS AND REPTILES OF MICHOACAN if to me the facilities of the museum and for their numerous cour- tesies that aided me so much. My field work in Michoacan was supported by the Museum of Zoology at the University of Michigan (1951), by the Horace H. Rackham Schoo! of Graduate Studies of the University of Michi- gan (1955), by the Penrose Fund of the American Philosophical Society (1956), by the Bache Fund of the National Academy of Sciences (1958), and by the University of Kansas Endowment Association (1960). Permits for collecting specimens in México were provided by the Direccién General de Caza through the courtesy of Ing. Juan Lozano Franco and Luis Macias Arellano. Historical Account Unlike many parts of southern México and northern Central America, Michoacan received no attention from the collecting ex- peditions of the European museums in the last century. The earliest known herpetological specimens from Michoacan were ob- tained by Louis John Xantus, who was appointed U. S. Consul to Colima in 1859. In April, 1863, Xantus collected at Volcén Jorullo in Michoacan; in April and May of the same year he collected along the coast of Michoacan between the Rio Cachan and the Rio Nexpa. His small collection of 19 extant specimens is in the United States National Museum. Alfredo Dugés, a resident of Guanajuato, México, made early contributions to the knowledge of the herpeto- fauna of Michoacan. In 1885 he described Sonora michoacanensis, and in 1891 he described Eumeces altamirani; from what is known of the distribution of these species, he probably had collected in the Tepalcatepec Valley. During their biological survey of México, Edward W. Nelson and Edward A. Goldman spent a limited amount of time in Michoacan in 1892 and again in 1903 and 1904. Most of their collecting was done on the plateau in the north- central part of the state; their collections are in the United States National Museum. While collecting fishes in southern México, Seth E. Meek obtained some amphibians and reptiles from Lago de Patzcuaro in 1904; these are in the collections of the Chicago Natural History Museum. In 1908 Hans Gadow ventured into the then unexplored “tierra caliente” of the Balsas Valley and col- lected at Volcdn Jorullo and other localities in the valley. Later in the same year he collected at Guayabo, San Salvador, and Arteaga in the Sierra de Coalcoman and at Buena Vista and Co- & UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. fradia in the Tepalcatepec Valley. His collections were deposited in the British Museum (Natural History) and the Naturhistorisches Museum Wien. The first thirty years of the present century saw little more field work in Michoacan. In the 1930’s Edward H. Taylor and Hobart M. Smith collected throughout much of México. At various times they worked in Michoacan, principally along the road from México City to Guadalajara. In 1935 Hobart M. Smith spent a week at Hacienda El Sabino south of Uruapan; he revisted the locality again in 1936 and made a large and important collection of am- phibians and reptiles from the upper limits of the arid tropical scrub forest in the Tepalcatepec Valley. Specimens collected by Smith and Taylor were incorporated into the Edward H. Taylor- Hobart M. Smith collection, which subsequently was deposited in part in the Museum of Natural History at the University of Illinois and in part in the Chicago Natural History Museum. In 1939 Hobart M. Smith collected at Patzcuaro and between Uruapan and Apatzingan; these collections, made while he was a Walter Rath- bone Bacon Scholar of the Smithsonian Institution, are deposited in the United States National Museum. In 1940 and 1941 Frederick A. Shannon, who was a member of the Hoogstraal Expeditions under the auspices of the Chicago Natural History Museum, col- lected on Cerro de Tancitaro and at Apatzing4n; an account of the specimens collected there was published by Schmidt and Shannon (1947). The eruption of Volcan Paricutin in February, 1948, attracted the attention of many biologists, a group of which from the Museum of Zoology at the University of Michigan collected in the Cordillera Volcanica in 1945 and 1947. The amphibians and reptiles were col- lected and studied by Norman E. Hartweg. In 1950 James A. Peters accompanied Donald D. Brand on a preliminary exploration of the western part of the Sierra de Coalcoman and adjacent Pacific coast of Michoacan; in the same year Peters collected also on the Mexican Plateau and at Volcan Jorullo. His specimens are in the Museum of Zoology at the University of Michigan. Since 1950 many biol- ogists have collected in Michoacan in the course of work on certain groups of animals or in general surveys. In this way Raymond Alcorn, Robert W. Dickerman, James R. Dixon, Floyd L. Downs, Emmet T. Hooper, and Robert R. Miller have contributed to our knowledge of the herpetofauna. As stated previously, my own field work in Michoacan began in 1951, when I accompanied Donald D. Brand on an exploring AMPHIBIANS AND REPTILES OF MICHOACAN 9 expedition to the southern part of the state. In that year a short time was spent on the Mexican Plateau, principally in the area around Lago de Cuitzeo, and at Volcdn Jorullo. In July and August we made our headquarters at Coalcoman. From that town the field party travelled southward to Maruata on the Pacific coast and thence back over the mountains to Coalcoman. Later in that summer we travelled by mule from Coalcoman southeastward to the mouth of the Rio Nexpa. In 1955, accompanied by Lee D. Beatty, Carter R. Gilbert, and Fred G. Thompson, I collected in the Tepalcatepec Valley and at Coalcoman. We made a mule trip from Coalcoman to Cerro de Barolosa, where we made the first collections from the pine-fir forests in the Sierra de Coalcoman. Later in the same summer Carter R. Gilbert and I spent a week at Playa Azul on the Pacific coast. In March, April, and May, 1956, my wife and I collected for a short time in the Cordillera Vol- cA4nica and on the Mexican Plateau. In early April we moved into the Tepalcatepec Valley, where we collected intensively between Churumuco and Tepalcatepec. In May we collected on the Pacific coast between Boca de Apiza and La Placita. In July and August, 1956, accompanied by Richard E. Etheridge, we re- turned to Michoacan and again collected on the Mexican Plateau and in the Cordillera Volcdnica, before moving into the Tepal- catepec Valley. In an attempt to fill in gaps in the known distri- butions of many species and to sample the fauna in some previously uncollected areas, I returned to Michoacan in June, 1958. Ac- companied by Jerome B. Tulecke and John Wellman, I collected on the Mexican Plateau in the northwestern part of the state, on the southern slopes of the Cordillera Volcanica, and in the Tepal- catepec Valley. Most of our time was spent in the Sierra de Coal- coman, where we collected at Aguililla, Artega, and Dos Aguas. In 1960 two days were spent in Michoacan; a small collection was made in the eastern part of the Cordillera Volcanica. With the exception of the specimens collected in 1960, which are at the Mu- seum of Natural History at the University of Kansas, the speci- mens that I have collected in Michoacan are in the Museum of Zoology at the University of Michigan. NATURAL LANDSCAPE A proper understanding of the geographical distribution of ani- mals in a given region is possible only after a thorough acquaintance with the geography of the region. Likewise, in order to gain a knowledge of the ecological distribution and relationships of the 10 UnIversiry OF Kansas Pusts., Mus. Nat. Hist. components of the fauna, it is necessary to study the animals in their natural environments. In order to give the reader a picture of the physical features and the major animal habitats within the state of Michoacan, the following brief description is offered. Each of these facets mentioned below will be elaborated in detail in my final report on the herpetofauna of Michoacan. Physiography The state of Michoacan comprises an area of 60,093 square kilo- meters (Viv6, 1953). Within this area the rugged terrain has a total relief of nearly 4000 meters. There have been several attempts to classify the physiographic provinces of México; the classification used here is a slight modification of the scheme proposed by Ta- mayo (1949). I have tried to keep the system as simple as possible, but still useful in discussing the distribution of animals living in the region. For general purposes the state of Michoacan can be divided into lowlands and highlands as follows: LowLANDs Pacific Coastal Plain Balsas-Tepalcatepec Basin HicHLANDS Mexican Plateau Cordillera Volcdnica Sierra de Coalcoman Although the lowlands in the state are continuous, they are only narrowly connected and thus form two distinct physiographic and biotic areas. The Pacific Coastal Plain in Michoacdn extends for a distance of about 200 kilometers (airline) from the Rio Coahu- ayana to the Rio Balsas. The coastal plain is broad between the Rio Coahuayana and San Juan de Lima, and between Las Pefias and the Rio Balsas, where the hills rise some 12 kilometers inland from the sea. Between San Juan de Lima and Las Pefias the mountains extend to the sea; in this region rocky promontories form precipitous cliffs dropping into the sea. Between the promontories are small sandy or rocky beaches. Lying to the north of the Sierra de Coalcoman and the Sierra del Sur, but south of the Cordillera Volcanica, is a broad structural depression, the Balsas-Tepalcatepec Basin, The western part of this basin, which separates the Sierra de Coalcomaén from the Cor- dillera Volcanica, is the valley of the Rio Tepalcatepec, a major tributary of the Rio Balsas. The eastern part of the basin is the valley of the Rio Balsas. From the point of junction of the two rivers, the Rio Balsas flows southward through a narrow gorge, AMPHIBIANS AND REPTILES OF MICHOACAN 11 which separates the Sierra de Coalcoman from the Sierra del Sur, to the Pacific Ocean. In Michoacan the floor of the Balsas-Tepal- catepec Basin varies from 200 to 700 meters above sea level. The central part of México is a vast table-land, the Mexican Plateau, the southern part of which extends into northern Micho- acin. In this region the terrain is rolling and varies from 1500 to 1900 meters above sea level. Many small mountain ranges rise from the plateau and break the continuity of the rolling table-land. Located on the southern part of the Mexican Plateau in Michoacan are several lakes, the largest of which are Lago de Chapala, Lago de Cuitzeo, and Lago de Patzcuaro. Bordering the southern edge of the Mexican Plateau is a nearly unbroken chain of volcanos, the Cordillera Volcanica. The high- est peaks in Michoacan, Cerro San Andrés (3930 meters) and Cerro de Tancitaro (3870 meters), are in this range. Parts of the Cordil- lera Volcdnica in Michoacan are known by separate names; these are, from west to east: Sierra de los Tarascos, Sierra de Ozumatlan, and Serrania de Ucareo. Lying between the Tepalcatepec Valley and the Pacific Ocean, and east of the Rio Coahuayana and west of the Rio Balsas, is an isolated highland mass, the Sierra de Coalcoman. This mountain range rises to elevations of slightly more than 3000 meters. It has a length of about 200 kilometers and a width of about 80 kilometers. Except for a relatively low connection with the Cordillera Volcanica, the Sierra de CoalcomaAn is isolated from other mountain ranges in south- western México. CLIMATE The climates in Michoacan vary from tropical in the lowlands to cool temperate at high elevations in the Sierra de Coalcomén and Cordillera Volcanica. The highest temperatures are known in the Balsas-Tepalcatepec Basin, where at Churumuco the mean annual temperature is 29.3° C. and the range of monthly means is 3.5° C. (Contreras, 1942). Frosts occur sporadically on the Mexican Plateau, and in the winter snow falls on the highest mountains. Precipitation varies geographically and seasonally. Most of the rain falls between June and October. In the Balsas-Tepalcatepec Basin rainfall in the rest of the year is negligible. The annual aver- age rainfall at Coahuayana on the Pacific Coastal Plain is 871 mm. (Guzman-Rivas, 1957:52). In the Balsas-Tepalcatepec Basin rain- fall seldom exceeds 800 mm. per year. In the mountains precipita- tion is heavier and somewhat more evenly distributed throughout the 12 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. year, but still definitely cyclic. For example, Uruapan (elevation, 1500 meters ) receives an average annual rainfall of 1674 mm. (Con- treras, 1942). The prevailing winds are from the Pacific Ocean. The southern (windward) slopes of the Sierra de Coalcomén prob- ably receive more rain than any other part of the state. The Balsas- Tepalcatepec Basin lies in a rain shadow of the Sierra de CoalcomAén, and the Mexican Plateau lies in a somewhat less drastic rain shadow of the Cordillera Volcanica; these are the driest regions in the state. VEGETATION AND ANIMAL HABITATS For the purposes of this report I have adopted the classification of types of vegetation that seem to me most significant in terms of ecological distribution of reptiles and amphibians in Michoacdn. These types are as follows: TEMPERATE (1000-4000 meters) Fir Forest (2400-4000 meters ) Pine-oak Forest (1000-4000 meters ) Mesquite-grassland (1500-2100 meters ) TropicaL (0-1000 meters) Arid Tropical Scrub Forest (0-1000 meters ) Tropical Semi-deciduous Forest (150-600 meters ) The vegetation of the Pacific Coastal Plain and the Balsas-Tepal- catepec Basin consists of arid tropical scrub forest, composed of de- ciduous trees, which in many places are stunted and widely spaced. In the dry season there is little cover provided by this forest. In the rainy season there is a sparse growth of grasses and some shade pro- vided by the small leaves of the thorny trees. In Michoacan the rainfall is heaviest on the southern slopes of the Sierra de Coalcoman and somewhat less so on the southwestern slopes of the Cordillera Volcanica. At these relatively low eleva- tions (150 to 600 meters) there is tropical semi-deciduous forest, characterized by relatively dense shade throughout the year and by a leaf mulch on the ground. This type of forest forms the gallery forest along the larger streams in the Balsas-Tepalcatepec Basin and on the Pacific Coastal Plain. Rainfall also is heavy on the high mountain ridges, where tem- peratures are low. On these ridges, fir forest, often mixed with pine and oaks, is found. This habitat is characterized by a cool, moist climate, many rotting logs, and a moist ground cover of leaves and needles. Most of the mountains are covered with pine-oak forest, which in most places is decidedly subhumid, but where this forest occurs on the windward sides of high ridges, it sometimes is noticeably AMPHIBIANS AND REPTILES OF MICHOACAN 13 humid. In this forest the important animal habitats include the needle- and leaf-litter, and in some areas, bromeliads. The rolling terrain of the Mexican Plateau supports cacti, small leguminous trees, and grasses. Like the arid tropical scrub forest, this type of vegetation, the Mesquite-grassland association, is de- ciduous and thus provides little shelter in the dry season. Unlike the areas in which arid tropical scrub forest is developed, the Mesquite-grassland is found in areas having warm days and cool nights. GEOGRAPHY OF THE HERPETOFAUNA Although the main part of my final report on the herpetofauna of Michoacan will deal with the geographical and ecological patterns of distribution of the herpetofauna, a brief summary of the faunal assemblages is presented here. In Michoacan there are two major faunal assemblages, one in the lowlands, and one in the highlands. A large number of the species inhabiting the lowlands are wide-ranging species, such as Bufo marinus, Iguana iguana, and Boa constrictor. Sixty-three species are known to occur on the Pacific Coastal Plain; 41 of these, together with 36 others occur in the Balsas-Tepalcatepec Basin, a physio- graphic region to which several species of reptiles are endemic; for example, Enyaliosaurus clarki, Urosaurus gadowi, Cnemidophorus calidipes, and Eumeces altamirani. Generally speaking, the members of the highland faunal assem- blage have more restricted geographic ranges. The major exceptions are those species that are widely distributed on the Mexican Plateau, such as: Bufo compactilis, Sceloporus torquatus, and Salvadora bairdi. In the montane habitats of the Cordillera Volcdnica, 45 species of amphibians and reptiles are known; 34 species have been found in the Sierra de Coalcomdn. Fourteen species are known to occur in both ranges. Several species are known only from the Cordillera Volcanica and adjacent highlands, and three species are endemic to the Sierra de CoalcomAn. ANNOTATED LIST OF SPECIES In the following pages the 176 species and subspecies of amphibians and reptiles known to occur in the state of Michoacan are discussed in relation to their variation, life histories, ecology, and distribution in the state. Data have been gathered from 9676 specimens. I have not prolonged the accounts of species with information that has been presented elsewhere. Consequently, the length and completeness of the accounts are variable. I have given only the information that I consider a worthwhile contribution to our knowledge of the particular species. 14 UnIverSITY OF Kansas Pusts., Mus. Nat. Hist. The synonymies given at the beginning of each account include the first use of the trivial name by the original author, the first usage of the combination that I am using, and, if the circumstances make it necessary, additional names or combinations that have been proposed since the publication of the checklists of Mexican amphibians and reptiles by Smith and Taylor (1945, 1948, and 1950b). References cited only in the synonymies are not listed in the Litera- ture Cited. Preceding the discussion of each species is an alphabetical list of the localities in Michoacan from which specimens have been examined. The listing of a locality means that one or more specimens, as indicated, has been examined from that locality. Only for those specimens especially men- tioned in the text are catalogue numbers given. Abbreviations for the various museums and scientific collections are, as follows: AMNH American Museum of Natural History ANSP Academy of Natural Sciences of Philadelphia BMNH British Museum (Natural History ) CNHM Chicago Natural History Museum EHT-HMS Edward H. Taylor-Hobart M. Smith collection JRD James R. Dixon collection, College Station, Texas KU University of Kansas Museum of Natural History MCZ Museum of Comparative Zoology MVZ Museum of Vertebrate Zoology NMW Naturhistorisches Museum Wien SU Stanford University Museum of Natural History TCWC Texas Cooperative Wildlife Collection UIMNH _— University of Illinois Museum of Natural History UMMZ University of Michigan Museum of Zoology USNM United States National Museum UTNHC _ University of Texas Natural History Collection Throughout the accounts of the species all measurements are given in milli- meters; if the range of variation is given, the mean follows in parentheses. AMPHIBIA Caudata Ambystoma amblycephalum Taylor Ambystoma amblycephala Taylor, Univ. Kansas Sci. Bull., 26: 420, Novem- ber 27, 1940.—Fifteen kilometers west of Morelia, Michoac4n, México. Fifteen km. W of Morelia (19); 11 km. SSE of Opopeo (12); 8 km. § of PAtzcuaro; 24 km. S of Pdtzcuaro (2); Quiroga (20); Tacicuaro (167). Taylor and Smith (1945:580) presented data on 137 specimens collected at Tacicuaro on October 1, 1939; these are all larvae and metamorphosing individuals. Aside from these, the largest larva examined (UMMZ 104962 from 15 km. W of Morelia) has a snout-vent length of 70.0 mm. and a tail length of 53.5 mm. The larvae are pale pinkish tan above and somewhat paler below; there is a lateral row of cream colored spots. The tail-fin, which is deep- est at mid-length, extends to the back of the head and is flecked with brown. In small larvae the outer edge of the tail-fin is dark brown. The eyes are large. Two small metamorphosed specimens (UMMZ 98967) from 24 kilometers south of Patzcuaro are tenta- AMPHIBIANS AND REPTILES OF MICHOACAN 15 tively referred to this species. These specimens have body lengths of 49.0 and 45.0 mm. and tail lengths of 36.0 and 31.5 mm., respec- tively. They have 17-17 and 16-15 vomerine teeth arranged in a broad arch behind the choanae, 10 costal grooves, and 7 intercostal spaces between adpressed toes. The dorsal color is uniform brown; that of the venter is a dusty cream. Larvae were collected from shallow ponds near Quiroga and 15 kilometers west of Morelia; metamorphosed individuals were taken from beneath logs in pine and fir forests at elevations from 2800 to 2800 meters, Ambystoma dumerili dumerili (Dugés) Siredon Dumerili Dugés, La Naturaleza, 1:241, 1870—Lago de Patzcuaro, Michoacan, México Bathysiredon dimen: Dunn, Notulae Naturae, 36:1, November 9, 1939. Bathysiredon dumerilit dumerilii, Maldonado-Koerdell, Mem. y Rev. Acad. Nac. Cien., 56:199, 1948. Ambystoma (Bushyuveden) dumerili, Tihen, Bull. Florida State Mus., 8:3, June 20, 1958. Lago de Patzcuaro (22); ? Morelia. For many years this unusual salamander was known from only a few specimens mostly collected in the last century; Smith and Taylor (1948:7) stated: “It is presumed that this species is extinct owing to the introduction of exotic game and food fishes.” In 1951 and in 1955 I had been told that axolotls were sold in the market at Patz- cuaro; nevertheless, none was found on my visits there. In 1956 Charles M. Bogert obtained several large specimens at the market in Patzcuaro. These establish the continued existence of the sala- mander in Lago de Patzcuaro. On January 27, 1955, R. W. Dicker- man procured a specimen (KU 41573) in the market at Morelia. Since fish are brought to Morelia from Lago de Patzcuaro, the speci- men probably was from that lake. Nevertheless, the species may occur in other permanent bodies of water in Michoacan. Maldo- nado-Koerdell (1948) described Bathysiredon dumerili queretaren- sis from San Juan del Rio, Queretaro. This locality is about 200 air- line kilometers northeast of Lago de Patzcuaro and is in the Rio Moctezuma drainage. Ambystoma ordinarium Taylor Ambystoma ordinaria Taylor, Univ. Kansas Sci. Bull., 26:422, November 27, 1940.—Four miles west of El Mirador, near Puerto Hondo, Michoacan, México. Axolotl (56); Cerro San Andrés; 22 km. W of Mil Cumbres; 46 km. E of Morelia ( 34); 8 km. SE of Opopeo (5); Puerto de Garnica (8); Puerto Hondo (41); San Gregorio (16); San José de la Cumbre (20). 16 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Of 16 specimens (KU 51520-35) collected on June 18, 1955, near San Gregorio, 15 are adult females with swollen cloacae and minute ovarian eggs. Possibly these specimens had just recently deposited their mature eggs. In preservative the specimens are black above and dull creamy gray below. Measurements for the 15 females are: snout-vent length, 80.0-102.0 (92.5); tail length, 69.0-98.0 (84.2); head width, 15.8-20.5 (17.7); head length, 22.8-26.6 (24.4). A larval specimen with small gills has a snout-vent length of 72 mm. and a tail length of 62mm. Three specimens have 12 costal grooves; the other have 11. Of 20 specimens from San José de la Cumbre (UMMZ 112857 and 115143), 14 are neotenic adults; the others are larvae. In life the salamanders were blackish to olive-brown above with scattered cream-colored dots on the dorsum and flanks but in preservative are dull grayish black with indistinct pale spots and dark reticulations. The belly is pale gray with indistinct dark spots. Eleven females and three males have the following measurements, respectively: snout-vent length, 76.0-90.0 (80.7), 64.0-84.0 (74.3); tail length, 70.0-81.0 (75.0), 58.0-71.0 (66.7); head width, 19.5-28.5 (20.7), 17.5-20.5 (19.3); head length, 22.0-25.0 (23.0), 20.0-22.5 (21.5). The smallest larva has a snout-vent length of 43.0 mm. and a tail length of 38.0 mm. Two individuals have 12 costal grooves; the others have 11. All of the females contained eggs, the largest of which were 1.5 mm. in diameter. The stomachs of most of the specimens were distended with oligochaets, aquatic insect larvae, and small aquatic beetles. A series of 34 larvae (JRD 5904-37) from 46 kilometers east of Morelia are tentatively referred to this species. These specimens are olive-brown above with cream-colored spots on the flanks; the dorsal tail-fin does not extend onto the body. This species has been found only at elevations in excess of 2400 meters in pine and fir forests. At Rancho Axolotl James A. Peters collected larvae and neotenic individuals in a rocky stream and adults from beneath rocks and logs in the forest near the stream. Neotenic individuals and larvae were found in a clear stream in pine-fir forest at an elevation of 2700 meters near San José de la Cumbre; specimens were collected there in July, 1955, and again in July, 1956. The site was visited in April, 1956, at which time the stream consisted of only a few puddles; no salamanders were found. AMPHIBIANS AND REPTILES OF MICHOACAN i yé Ambystoma tigrinum velasci Dugés Ambystoma velasci Dugés, La Naturaleza, ser. 2, 1:142, 1888.—Laguna Santa Isabel, near Guadalupe Hidalgo, Distrito Federal, México. Ambystoma tigrinum velasci, Dunn, Copeia, no. 3:157, November 14, 1940. Patzcuaro (5); Tacicuaro (9). Definite specific assignment of these specimens, all larvae, can- not be made at this time. They have shovel-shaped heads and laterally compressed bodies with the dorsal tail-fin extending an- teriorly to the back of the head. The eyes are small. The body is pale tan with dark mottling on the tail and flanks. The average snout-vent length for nine specimens from Tacicuaro is 61.0 mm. The larvae from Tacicuaro (UMMZ 89255) were collected by Dyfrig Forbes in October, 1939; those from Patzcuaro, presumably Lago de Patzcuaro (BMNH 1914.1.28-247-8 and CNHM 948), were collected by Hans Gadow and Seth Meek in 1908. Pseudoeurycea belli (Gray) Speieres belli Gray, Catalogue Batrachia Gradientia British Museum, p. 46, 1850.—México. Type locality restricted to 2 miles east of Rio Fri rio, Puebla, México, by Smith and Taylor (1950a:341). Pseudoeurycea bellii, Taylor, Univ. Kansas Sci. Bull., 30:209, June 12, 1944. Axolotl (2); Carapan; Cerro Tancitaro (84); Macho de Agua; 22 km. W of Mil Cumbres; Opopeo; Patzcuaro (8); Puerto Hondo (2); San José de la Cumbre; San Juan de Parangaricutiro (42); Uruapan (5); Zacapu (4). This salamander seems to reach its greatest abundance in Micho- acan in the Sierra de los Tarascos between Patzcuaro and Tancitaro, where it is found at elevations from 1500 to 2900 meters. It is found less commonly in the eastern part of the Cordillera Volcanica in Michoacan, where it sometimes occurs in association with Pseudo- eurycea robertsi. On June 22 and 28, 1955, four clutches of eggs of this species were found beneath adobe bricks and rocks on the volcanic ash that has buried the village of San Juan de Parangaricutiro. The eggs were un- stalked and separate, but adherent in clumps of three or four (PI. 2, Fig. 1). The outer membranes were covered with fine particles of ash. The ash beneath the stones where the eggs were found was only slightly moist; one clump of eggs was partially desiccated. Three complete clutches have 20, 23, and 34 eggs; one clutch of 15 eggs was being eaten by beetles (Tenebrionidae: Eleodes sp.). The eggs vary in size from 4.6 to 6.5 mm. and average 5.8 mm. in di- ameter. They are unpigmented. Surrounding the embryo is a vitelline membrane, an inner, and an outer envelope (Fig.1). In an 2—7817 18 UnIvERSITY OF Kansas Pusts., Mus. Nat. Hist. average-sized egg having an embryo 4 mm. in length, the diameter of the outer membrane is 5.3 mm., the inner membrane 5.0 mm., and the vitelline membrane 4.6 mm. All of the eggs contained embryos in which the limb buds were developed; in about half of these the eyes were distinctly visible. Fic. 1, Diagram of an egg of Pseu- doeurycea belli from San Juan de Parangaricutiro, Michoacan. xX 10. The first heavy rain of the season occurred on the night of June 22, 1955. Thus, at least sometimes, Psuedoeurycea belli lays its eggs be- fore the onset of the rainy season. A female having a snout-vent length of 110 mm., collected on June 22, 1955, contained 36 ovarian eggs having diameters from 3.0 to 3.5 mm. The fact that small juveniles were collected on the same date indicates that this sala- mander lays eggs over a period of several weeks in late spring and early summer. The smallest juvenile examined has a snout-vent length of 17.0 mm. and a tail length of 7.5mm. Twelve juveniles from the vicinity of San Juan de Parangaricutiro have an average snout-vent length of 19.4 mm. and an average tail length of 9.7 mm. In juveniles the ad- pressed limbs either touch or overlap by one intercostal space; in adults there are two or three intercostal spaces between adpressed toes. Therefore the greatest number of intercostal spaces between adpressed limbs is found in the largest specimens. A similar rela- tionship between adpressed limbs (= length of limbs) and snout- vent length was shown for Plethodon richmondi by Duellman (1954a). The number of vomerine teeth is variable; the number of teeth seems to be closely correlated with the size of the sala- mander (Fig. 2). A similar correlation between the number of AMPHIBIANS AND REPTILES OF MICHOACAN 19 maxillary teeth and body length was reported for Chiropterotriton multidentatus by Rabb (1958). In 12 juvenile Pseudoeurycea belli there are 6-13 (8.8) vomerine teeth, and in 11 adults having snout- vent lengths greater than 90 mm. there are 39-49 (44.0) vomerine 50 VOMERINE TEETH ° 10 20 30 40 50 60 70 60 30 100 ile) 120 SNOUT—VENT LENGTH (IN MM) Fic. 2. Correlation between the number of vomerine teeth and snout-vent length in 79 Pseudoeurycea belli from Michoacan. teeth. The coloration of the juveniles resembles that of the adults (PIT): The differences between this species and Pseudoeurycea gigantea are minor. Taylor (1939a) distinguished gigantea from belli by the larger size, fewer intercostal spaces between adpressed limbs, more vomerine teeth, and absence of occipital spots in gigantea. Taylor and Smith (1945) stated that in life the spots in gigantea are orange instead of red as in belli. Five specimens of Pseudoeurycea belli from Michoacan, including one juvenile, lack occipital spots. In the 34 living individuals that I have seen from Michoacan the spots varied from deep red to orange. Therefore, of the characters listed by Taylor (op. cit.) to diagnose Pseudoeurycea gigantea, only the over-all larger size and smaller number of intercostal spaces between adpressed limbs (= relatively longer limbs) are useful in separating Pseudoeurycea belli and gigantea. Pseudoeurycea robertsi (Taylor) Oedipus robertsi Taylor, Univ. Kansas Sci. Bull., 25:287, July 10, 1939.— Nevado de Toluca, México. Pseudoeurycea robertsi Taylor, Univ. Kansas Sci. Bull., 30:209, June 12, 1944. Atzimba (3); Macho de Agua (9); Puerto Lengua de Vaca (14). Previously this species has been recorded only from the type lo- cality. In July, 1956, individuals referable to this species were found at two sites in pine-fir forest immediately to the east of Macho de Agua and in pine-oak-fir forest at Atzimba. On August 20, 1958, 20 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. a series was collected in pine-fir forest at Puerto Lengua de Vaca. These localities are between 2900 and 3000 meters in the Cordillera Volcanica in eastern Michoacan. In life the coloration of these salamanders was highly variable. The belly and undersurfaces of the tail and hind limbs were pale gray, with or without silvery white flecks; the chin was a cream-color and flecked with silvery white in some specimens. The middorsal area was brown, orange-brown, or dull grayish yellow. The flanks and lateral surfaces of the tail were black with yellowish flecks or streaks on the flanks and yellowish or orange-brown flecks on the tail. The iris was golden brown. Measurements of eight males and two females are, respectively: snout-vent length, 42.5-56.0 (49.5), 54.0- 60.0 (57.0); tail length, 42.0-56.0 (48.1), 52.0-55.0 (53.5). The smallest juvenile has a snout-vent length of 28.0 mm. and a tail length of 23.0 mm. Of the 26 available specimens, six have 12 costal grooves, and the others have 11. In comparison with 86 topotypes, the specimens from Michoacan have a less striking dorsal color pattern; none has a well-defined dorsal reddish brown area or bold reddish mottling on the tail. Furthermore, the specimens from Michoacan have paler venters than do topotypic specimens. Salientia Rhinophrynus dorsalis Duméril and Bibron Rhinophrynus dorsalis Duméril and Bibron, Erpétologie générale, vol. 8:758, 1841.—Veracruz, Veracruz, México. Mouth of the Rio Balsas (10). These specimens (BMNH 1914.1.28.181-90) were collected by Gadow in 1908 and reported by him (1930:72): “Whilst this very sluggish termite-eating toad is common enough in the sweltering hot country of the state of Vera Cruz, up to an elevation of 1500 feet, it was unknown on the west side of the Isthmus until I found it in great numbers near the mouth of the Balsas River, in and near freshwater pools, where it attracted attention by its loud peculiar voice during the pairing season in the month of July.” Subsequently, Peters (1954:3) verified the identification of these specimens. Although torrential rains fell during the week in July, 1955, that I spent at Playa Azul near the mouth of the Rio Balsas, the distinctive voice of Rhinophrynus was not heard. Elsewhere on the Pacific coast of México adult Rhinophrynus have been reported only from Tehuan- tepec and a few localities on the coastal lowlands of Chiapas. Taylor (1942b:37) found on the coast of Guerrero a tadpole that was re- AMPHIBIANS AND REPTILES OF MICHOACAN Path ferred to the genus Rhinophrynus by Orton (1943). In the summer of 1960 adults of Rhinophrynus were collected near Acapulco, Guerrero (Fouquette, in litt.). These recent collections verify the existence of the species along the Pacific lowlands of México at least as far north as Michoacan. Scaphiopus hammondi multiplicatus Cope Scaphiopus multiplicatus Cope, Proc. Acad. Nat. Sci. Philadelphia, 15:52, June 8, 1863.—Valley of México. Scaphiopus hammondi multiplicatus, Kellogg, Bull. U.S. Natl. Mus., 160:22, March 81, 1932. Angahuan (5); Cuitzeo (4); Cusefio Station (2); Jiquilpan (9); Morelia (7); Patzcuaro (3); Quiroga; Tarécuaro; Uruapan (24); Zacapu. This small toad has been found at elevations between 1500 and 2500 meters on the Mexican Plateau and associated mountain ranges; it occurs in mesquite-grassland and in pine forests. Calling males and females laden with eggs have been collected in the rainy season in the months of July and August. The call is a medium- pitched snore. In living individuals the dorsal ground color varies from pale brown to gray with dark brown or olive-brown markings. In many individuals the tips of the small dorsal pustules are red. Bufo coccifer Cope Bufo coccifer Cope, Proc. Acad. Nat. Sci. Philadelphia, 18:130, 1866.— Arriba, Costa Rica. Apatzingan (27); Lombardia; Nueva Italia (5). In life the dorsal color pattern consists of a yellowish tan ground color with dark brown spots; the middorsal stripe is deep yellow or cream color. The venter is a dusty cream color, and the iris is pale gold. Males have dark brown horny nuptial tuberosities on the thumb. The following measurements are of 21 males and four females, respectively: snout-vent length, 43.5-51.7 (48.1), 55.6-62.6 (59.1); tibia length, 16.6-18.8 (17.6), 18.8-20.3 (19.3); head width, 16.7-19.7 (18.4), 20.6-22.2 (21.4); head length, 13.8-16.6 (14.8), 16.5-18.2 (17.3). The specimens from the Tepalcatepec Valley differ slightly from specimens from southeastern México and Central America. Those from Michoacan have low and narrow cranial crests; in about one- half of the specimens the occipital crest exists only as a row of tubercles, and in some the postorbital and suborbital crests are barely discernible. Specimens from the southern part of the range, Costa Rica and Nicaragua, have much higher and thicker cranial crests; in these the occipital crest is well defined and extends pos- 22 Untversiry OF Kansas Pusts., Mus. Nat. Hist. teriorly to a point back of the anterior edge of the parotid gland; the postorbital and suborbital crests are well marked. Of 48 speci- mens from Esquipulas, Guatemala, all have high crests, but these are not so well developed as in ten specimens from Matagalpa, Nicaragua, and three from various localities in Costa Rica. Six specimens from Tehuantepec, Oaxaca, have cranial crests that are lower than those in specimens from Guatemala. In three of the specimens from Tehuantepec the occipital crests are reduced to a series of tubercles. Of six specimens from Agua del Obispo, Guer- rero, four have poorly developed occipital crests. These observa- tions suggest the presence of a cline in the development of the cranial crests; specimens have higher crests in the southern part of the range than in the northern part. In México Bufo coccifer has been collected only in semi-xeric habitats, but to the south, from Guatemala to Costa Rica, it has been found in more upland and humid habitats. Southern speci- mens are darker than those from the north, a possible correlation with the differences in habitat. These toads probably range throughout the Tepalcatepec Valley, but they are unknown from the coast of Michoacan. Breeding choruses were found after heavy rains on June 24, 1955, and on August 2, 1956. The first was in a muddy ditch; the second was in a flooded grassy field. The call is a high-pitched, but not loud, “whirrr.” Males were calling from the edge of the water or from clumps of grass in the water. Clasping pairs were in the water; amplexus is axillary. Bufo compactilis compactilis Wiegmann Bufo compactilis Wiegmann, Isis von Oken, 26:661, 1833.—Meéxico. Type locality restricted to Xochimilco, Distrito Federal, México, by Smith and Taylor (1950a:3380). Bufo compactilis compactilis, Smith, Herpetologica, 4:7, September 17, 1947. Cuitzeo (2); Emiliano Zapata (20); Jiquilpan (5); La Palma (5); Morelia; Tupataro. The southwestern terminus of the range of this species is on the Mexican Plateau in Michoacan. All specimens from the state have spotted venters. In living toads the dorsal ground color was gray or grayish tan with olive green spots. The vocal sac was brownish gray; the iris was a bright golden color. On June 11, 1958, many individuals were calling from shallow water in a flooded field at Emiliano Zapata. The call is a slow trill, in which the individual notes are discernible. AMPHIBIANS AND REPTILES OF MICHOACAN 93 Bufo marinus (Linnaeus) Rana marina Linnaeus, Systema naturae, ed. 10, 1:211, 1758.—America. Bufo horribilis Wiegmann, Isis von Oken, 26:654, 1833.—Misantla and Vera- cruz, Veracruz, México. Taylor and Smith, Proc. U.S. Natl. Mus., 95: 551, January 30, 1945. Bufo angustipes Taylor and Smith, Proc. U.S. Natl. Mus., 95:553, January 80, 1945.—La Esperanza, Chiapas, México. Aguililla; Apatzingan (3); Barranca de Bejuco; Capirio; Charapendo; Chichi- huas; Coahuayana (2); Coalcoman (7); Cofradia (2); 25 km. S of Cuatro Caminos; El Sabino (10); Huahua, La Playa (18); Ojos de Agua de San Telmo; Ostula; Playa Azul (2); Pémaro (2). This large toad is characteristically found in areas supporting tropical scrub forest to elevations of about 1000 meters. The species is much more abundant than the numbers listed above suggest. In the dry season individuals have been observed in patios, along streams, and by irrigation ditches. In the rainy season the loud, rattling call of the males is heard at night throughout the Tepalca- tepec Valley and the coastal lowlands. Taylor and Smith (1945:552) revived Wiegmann’s Bufo horribilis for the large toads of México that are here referred to B. marinus. Their action was based upon the supposition that the “species marinus” is composite. Although probably true, this supposition has yet to be proved. Until the large, and apparently related, species of Bufo inhabiting tropical America have been studied systematically as a unit, the recognition of segments of the population as either species or subspecies is meaningless. Taylor and Smith (op. cit.:553) based the description of a new species, Bufo angustipes, on one rather emaciated, formalin-hardened female from La Esperanza, Chiapas. The type (USNM 116518), when compared with nu- merous specimens of Bufo marinus from throughout the range of the species in México and northern Central America, displays no com- bination of characters to set it off from the others. Therefore, I suggest that Bufo horribilis Wiegmann and Bufo angustipes Taylor and Smith be placed in the synonymy of Bufo marinus (Linnaeus ) until future systematic study of the genus and this species in par- ticular establishes the existence of recognizable taxa. Bufo marmoreus Wiegmann Bufo marmoreus Wiegmann, Isis von Oken, 26:66, 1833.—Veracruz, Vera- cruz, México. Barranca de Bejuco; Coahuayana (11); El Diezmo (2); La Placita (9); La Orilla (12); Motin del Oro; Ostula (9); Playa Azul (5); Pémaro (15); Sali- tre de Estopilas; San Pedro Naranjestila. In Michoacan this species is confined to elevations of less than 1000 meters on the coast and foothills of the Sierra de Coalcoman. 24 UnrIversiry OF Kansas Pusts., Mus. Nat. Hist. In this region in the months of June and July, breeding congregations have been found in temporary pools and along streams. Smith and Taylor (1948:39), in their key to the Mexican species of Bufo, placed emphasis on the nature of the supraorbital and post- orbital crests (whether they form a curve or a sharp angle) in dis- tinguishing Bufo marmoreus from Bufo perplexus. In the original description of perplexus, Taylor (1943a:347) characterized the species as follows: supraorbital and postorbital crests forming a sharp angle, instead of a curve as in marmoreus; supratympanic crest smaller than in marmoreus; diagonal lateral stripe lacking in females; concentration of dorsal tubercles as found in marmoreus lacking in males. The discovery of specimens in which the crests form a curve and others in which the crests form an angle in both the Tepalcatepec Valley and in the coastal lowlands prompted an investigation of these characters and others throughout the ranges of the species. An examination of 410 specimens has resulted in the following con- clusions. TABLE 1.—VARIATION IN THE SHAPE OF THE SUPRAORBITAL AND POSTORBITAL CRANIAL CRESTS IN BUFO MARMOREUS AND B. PERPLEXUS. Locality N Curved Intermediate Angular Tepaleatepec Valley....| 50] 10 (20.0%) 17 (34.0%) | 23 (46.0%) MorelossAiccehs oct sneer 12 2 (16.6%) 5 (41.7%) 5 (41.7%) Izucar, Puebla......... 4 2 (50.0%) 0 (0.0%) 2 (50.0%) Southern Sinaloa....... 1 1 (100.0%) 0 (0.0%) 0 (0.0%) Puerto Vallarta, Jalisco 2 2 (100.0%) 0 (0.0%) 0 (0.0%) Colm atch tor ions cr acnen: AS 25: (5d- 09) AS | 40209,) 2 ( 5.0%) Coast of Michoacdn....| 55 | 35 (63.6%) | 17 (80.9%) 3. (Ga.5%) Acapulco, Guerrero... .. 7 7 (100.0%) 0 (0.0%) 0 (0.0%) Chilpancingo, Guerrero 10 1 (10.0%) 4 (40.0%) 5 (50.0%) Pochutla, Oaxaca...... 13 6 (46.2%) 6 (46.2%) LCG 9) Tehuantepec, Oaxaca...| 177 | 81 (45.8%) | 67 (37.8%) | 29 (16.4%) Tonold4, Chiapas....... 1 0 (.0.0%) 0 (0.0%) 1 (100.0%) Weracrugeree nie eee 33 | 26 (78.8%) 6 (18.2%) 1 C320) TOE Se vace ieee 410 | 198 (48.3%) | 140 (84.2%) | 72 (17.5%) 1. Although the highest percentage of individuals having the supraorbital and postorbital crests forming a sharp angle is from localities in the Balsas-Tepalcatepec Basin, numerous individuals from throughout the range of marmoreus have the crests forming an angle (Table 1). 2. In all samples of ten or more specimens, some toads have the supraorbital and postorbital crests forming a sharp angle, some have the crests form- ing a curve, and some have an intermediate condition. 8. The relative size of the supratympanic crest is highly variable in all sam- ples examined. AMPHIBIANS AND REPTILES OF MICHOACAN Fic. 3. Adult male of Bufo perplexus from Apatzingan, Michoacan, 2a—7817 S< ale 25 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Fic. 4. Adult male of Bufo marmoreus from Pémaro, Michoacan. Wz \ 1.5. REA a Hatchling of Pseudoeurycea belli from San Juan de Parangaricutiro, Michoacan. 8. PLATE +4 Fic. 1. Adult male of Tomodactylus nitidus nitidus from Tuxpan, Michoacan. x 4. Fic. 2. Adult male of Tomodactylus nitidus orarius trom Tecolapa, Colima. x 4. PLATE 5 Fic. 1. Adult male of Tomodactylus nitidus petersi from Apatzingan, Michoacan. x 4. Fic. 2. Adult male of Tomodactylus rufescens from Dos Aguas, Michoacan. 4. PLATE 6 bo Fic. 1. Adult male of Hypopachus caprimimus from Tuxpan, Michoacan. 2%. Adult male of Hypopachus oxyrrhinus ovis from Tangamandapio, Michoacan. ™ 3. AMPHIBIANS AND REPTILES OF MICHOACAN oT 4. A distinct, pale-colored, diagonal lateral stripe is found in females only from localities outside of the Balsas-Tepalcatepec Basin; females from the basin have a spotted dorsum. 5. Males from the Balsas-Tepalcatepec Basin usually have a broad middorsal line that is yellow or pale tan; those from outside the basin have either a narrow middorsal line or none. 6. Males from the Balsas-Tepalcatepec Basin have low, scattered dorsal tubercles (Fig. 3); males from outside the basin have a concentration of tubercles in a broad band on the back (Fig. 4). Therefore the nature of the cranial crests is of little value in sep- arating two populations, but the color pattern of the females and the nature of the dorsal tubercles of the males do show distinct differ- ences. Furthermore, certain differences in size and proportion are evident; Bufo marmoreus is a slightly larger toad and has a rela- tively longer tibia and longer head than perplexus (Table 2). TABLE 2.—COMPARISON OF CERTAIN MEASUREMENTS AND PROPORTIONS IN BuFO MARMOREUS AND B. PERPLEXUS. (MEANS ARE GIVEN IN PARENTHESES BELOW THE RANGES.) Tibia length | Head length Snout-vent }———__—_ SPECIES Sex N length Snout-vent | Snout-vent length length B. marmoreus........ of 15 61.5-72.5 | 35.9-41.6 | 28.3-33.3 (65.2) (39.0) (81.6) BS Der plexcie’s:. atv fou 20 50.0-59.0 | 33.7-38.1 | 26.4-31.1 (54.9) (36.4) (29.5) B. marmoreus........ Q cf 68.0-76.0 | 33.0-36.8 | 26.8-32.6 (70.7) (34.7) (29.6) B. perplexus......... f°) 6 64.1-69.8 | 32.4-36.9 | 25.1-29.0 (66.8) (35.5) (27.5) Taylor (1943a:347) described Bufo perplexus from Mexcala on the Rio Balsas in Guerrero. Among the many paratypes are speci- mens from Tonola, Chiapas, and Tehuantepec, Oaxaca. These ap- parently were referred to perplexus solely on the nature of the cranial crests. All of the specimens examined during the course of the present study from the lowlands of Veracruz and from the Pacific lowlands from Sinaloa southward to Chiapas are referable to Bufo marmoreus; those from the Balsas-Tepalcatepec Basin are referable to Bufo perplexus, as defined above. Ten specimens from Chilpancingo, Guerrero (UNMZ 115352), do not readily fit either species. Perhaps there is gene exchange between the inland and 8—7817 28 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. coastal populations through the relatively low pass at Chilpancingo, at the mouth of the Rio Balsas, and near the convergent headwaters of the Rio Coahuayana and Rio Tepalcatepec in southern Jalisco. If this can be demonstrated, then Bufo perplexus would have to be considered as a subspecies of Bufo marmoreus, instead of an allo- patric species. Bufo perplexus Taylor Bufo perplexus Taylor, Univ. Kansas Sci. Bull., 29:347, October 15, 1943.— Balsas River near Mexcala, Guerrero, México. Aguililla (2); Apatzingan (42); Buena Vista (5); Capirio (3); La Playa (25); Lombardia (6); Nueva Italia (9); Rio Cancita, 14 km. E of Apat- zingan; Rio Tepalcatepec, 27 km. S of Apatzingan; San Salvador (4); Tzi- tzio; Volcan Jorullo. Bufo occidentalis Camerano Bufo occidentalis Camerano, Atti R. Accad. Sci. Torino, 14:887, Decem- ber 31, 1878.—México. Type locality restricted to Guanajuato, Guana- juato, México, by Smith and Taylor (1950a:330). Firschein, Copeia, no. 3:220, September 15, 1950. Bufo simus, Smith and Taylor, Bull. U. S. Natl. Mus., 194:42, 1948. Barranca Seca (82); Cerro de Barolosa (4); Cerro Tancitaro, 3 km. E of Apo (2); Cerro Tancitaro, 19 km. E. of Apo (10); Charapendo; Coal- coman (7); Dos Aguas (4); Jacona, Jaramillo (2); Las Tecatas; Los Reyes (181); Tancitaro (10); Uruapan (8). This toad is an inhabitant of pine and oak forests between 900 and 2400 meters. Near Charapendo on the slopes of the Sierra de los Tarascos and at Coalcomén it apparently reaches its lowest alti- tudinal limits. At both of these localities the pine-oak forest is re- placed by arid tropical scrub forest on the lower slopes. Twenty-four tadpoles were collected on May 3 in a quiet section of a fast stream near Barranca Seca. The tadpoles have a robust body, broadest about two-thirds the distance from the snout to the posterior edge of the body, half again as broad as deep. Eyes dorsolateral; nostrils dorsal, somewhat directed forward, and about three-fifths the distance from the tip of the snout to the eye; spiracle sinistral and lateral, located at about midbody; anus median; tail long and slender; tail-musculature extends nearly to tip of tail; depth of tail-musculature at mid-length about one-third total depth of tail; dorsal tail-fin not extending onto body (Fig. 5); average body length of ten tadpoles having small hind limb buds, 14.4 mm.; average tail length, 22.0 mm. Mouth ventral, nearly terminal, about one-third as wide as widest part of body; anterior lip has no papillae; lower lip bordered by two rows of papillae and lateral lips by one row of papillae; beaks moderately well developed, the upper forming a broad arch and AMPHIBIANS AND REPTILES OF MICHOACAN 29 finely denticulate; tooth rows %, the upper rows extending to the edge of the lips, subequal in length, and slightly longer than lower rows, which also are subequal in length; inner upper tooth row broken medially; inner lower tooth row sometimes broken (Fig. 6). The body is black dorsally and laterally, and bluish gray ventrally; the tail musculature is brown and stippled with darker brown. The fins are transparent and stippled with brown, the stippling being most pronounced on the posterior two-thirds of the upper tail-fin. Fic. 5. Tadpole of Bufo occidentalis (UMMZ 94269) from Barranca Seca, Michoacan. xX 8. Cones na: noe ee eesed renened etnateny EW, 7 “a PANN OM RHO IR AAG Cry, ~ s ca eT nT Mere Lore : ww MARA yn yay, % i“ Q Np, oS Pe nL Wig AE NEAT may nett iziny {7 PIMC THI Ry a OCU ANAT m1, WttMn ” QA ae» tee (afr Fic. 6. Mouthparts of larval Bufo occidentalis (UMMZ 94269 ) from Barranca Seca, Michoacan. X 20. Forty recently metamorphosed individuals average 18.9 mm. in snout-vent length. The relationships of this toad seem to be with Bufo bocourti Brocchi, an inhabitant of pine and oak forests in the uplands of Chiapas and Guatemala. In Bufo occidentalis the tympanum usually is indistinct and sometimes completely covered, and it is absent in bocourti. Bufo occidentalis has a broader interorbital area and relatively shorter and more rounded parotid glands than bocourti. The tadpoles of the two species are nearly identical (see Stuart, 1943:12). 30 UNIVERSITY OF Kansas PuBis., Mus. Nar. Hist. Leptodactylus labialis (Cope) Cystignathus labialis Cope, Proc. Amer. Philos. Soc., 17:90, 1877.—No type locality designated; type locality restricted to Potrero Viejo, Veracruz, México, by Smith and Taylor (1950a:350). Leptodactylus labialis, Brocchi, Mission Scientifique au Mexique et dans l’Amerique Centrale, pt. 3, sec. 2, livr. 1:20, 1881. Apatzingén (26); Capirio (5); Cofradia (9); El Sabino (4); Lombardia; Rio Tepalcatepec, 27 km. S of Apatzingan (2). In the Tepalcatepec Valley this frog reaches the northernmost known limit of its range in western México. Although the species is abundant in the valley, it apparently is absent from the coastal lowlands. In the Tepalcatepec Valley Leptodactylus melanonotus seems to be more abundant than labialis. In the rainy season both species have been heard calling from the same ponds and flooded fields. There are only slight differences in size between the sexes; measurements of 20 males and eight females are, respectively: snout-vent length, 32.3-39.5 (35.1), 34.1-39.2 (87.2); tibia length, 14.3-17.0 (15.4), 14.9-16.8 (15.8); head width, 11.0-18.6 (12.0), 12.2-13.2 (12.6); head length, 12.8-15.1 (13.8), 12.8-14.6 (13.7). Leptodactylus melanonotus (Hallowell) Cystignathus melanonotus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 12:485, 1861.—Nicaragua. Type locality restricted to Recero, Nicaragua, by Smith and Taylor (1950a:320). Leptodactylus melanonotus, Brocchi, Mission Scientifique au Mexique et dans l’Amerique Centrale, pt. 3, sec. 2, livr. 1:20, 1881. Apatzing4n (103); Capirio; Charapendo (7); Coahuayana; Cofradia (10); El Sabino (21); La Playa (3); Lombardia (5); Maruata; Nueva Italia (7); Ostula (9); Playa Azul (11); Rio Marquez, 10 km. S of Lombardia; Rio Marquez, 13 km. SE of Nueva Italia (6); Rio Tepalcatepec, 27 km. S of Apatzingan. This species is widespread in the lowlands of the state; it has been collected up to elevations of 1050 meters in the Tepalcatepec Val- ley. In the dry season individuals were discovered beneath rocks along streams and in damp arroyos; in the rainy season they were found wherever there was water. Males were heard calling from flooded fields, ditches, rocky streams, and small puddles. The call is a series of individual notes: “woink, woink, woink.” Adult males are noticeably smaller than females; measurements for 20 males and ten females from Apatzingan are, respectively: snout-vent length, 29.6-34.6 (32.3), 36.8-44.1 (40.8); tibia length, 12.6-15.1 (14.0), 16.5-19.0 (17.8); head width, 10.8-11.9 (11.3), 12.6-14.8 (18.7); head length, 11.2-13.2 (11.9), 13.1-14.8 (14.0). Brownish yellow ventral glands are present in some juveniles and in some adults collected in the dry season as well as in the rainy season. AMPHIBIANS AND REPTILES OF MICHOACAN 31 Leptodactylus occidentalis Taylor Leptodactylus occidentalis Taylor, Trans. Kansas Acad. Sci., 39:349, 1987.— Tepic, Nayarit, México. Five km. W of Tangamandapio. On the night of June 11, 1958, this species was calling from a hya- cinth-choked ditch. Although numerous individuals were heard, only one specimen was obtained. The frogs were calling from the tangled mat of hyacinths along with Hyla eximia, Hypopachus oxyrrhinus ovis, and Rana pipiens. Taylor (1936a:352) characterized this species as follows: “The narrow head, small maximum size (38 mm. for females, 33 mm. for males ), the character of the postaxillary and postfemoral glands, the narrower groups of vomerine teeth, clearly distinguish this western Mexican form from the more robust, larger melanonotus to the south. The call is likewise fainter and different in quality.” Concerning the glands, Taylor (loc. cit.) remarked: “There is a possibility that the horny excrescence covering the glands may appear only during the breeding season. This character is quite as strongly marked in females as in males.” Bogert and Oliver (1945:324) concluded that the population of Leptodactylus in northwestern México could not be distinguished from melanonotus in other parts of the country and thus synonymized Leptodactylus occidentalis with melanonotus. Bogert and Oliver (op. cit.: 824) stated that the extent as well as the presence or absence of ventral glands was highly variable in all samples examined by them. Upon seeing numerous living individuals of Leptodactylus melan- onotus from many parts of its range in México and individuals of the population of Leptodactylus in northwestern México (Nayarit and Sinaloa), I was immediately impressed not so much by the differ- ences in the development of the ventral glands, but by the color of the glands. The differences in color are apparent in freshly pre- served specimens. With the exception of Leptodactylus from north- western México, specimens of melanonotus from throughout México and northern Central America have yellow or yellowish brown glands. Specimens from northwestern México have black or brown- ish black glands that are conspicuously darker than those found in melanonotus. Examination of 653 preserved specimens of Lepto- dactylus melanonotus from México and Guatemala has failed to reveal specimens with black ventral glands, like those found in specimens from northwestern México, to which the name Lepto- dactylus occidentalis has been applied. Furthermore, in melanono- tus the glands are less distinct and more extensive than in occi- 32 UNIvERSITY OF Kansas Pusts., Mus. Nar. Hist. dentalis; in the latter species glands are absent from the throat and midventral area, where they often are present in melanonotus (Fig. 7). In some individuals of both species collected in the dry season and in some collected in the rainy (breeding) season the glands are absent; the development of these glands, therefore, does not seem to be correlated with breeding. Likewise, the glands are present or absent in either sex, and often as not they are present in juveniles. Presence of the glands, therefore, cannot be corre- lated either with sexual or ontogenetic development. Since the Fic. 7. Diagrammatic view of ventral surfaces of Leptodactylus melanonotus (A) and Leptodactylus occidentalis (B), showing usual position and size of glandular areas. Approx. natural size. glands are found in individuals from all parts of the range, it is unlikely that there is a correlation between the development of the glands and the environment. Aside from the differences in the ventral glands, the call is dif- ferent in the two populations. The call of Leptodactylus occi- dentalis is a rather harsh “wack, wack, wack” as contrasted with the more nasal “woink, woink, woink” of melanonotus. Sound spectographs are needed to analyze the differences in calls. None of the specimens of occidentalis examined approaches in size the largest individuals of melanonotus; possibly the size of the frogs is another valid character for separating the species. On the basis of the above data it is evident that the frogs in northwestern AMPHIBIANS AND REPTILES OF MICHOACAN 33 México show certain characters that distinguish them from Lepto- dactylus melanonotus, as it is known throughout the rest of México. It is not known for certain that melanonotus and occidentalis are sympatric. Several series of old, poorly preserved specimens from Nayarit and Sinaloa cannot be placed in either species, for none has visible ventral glands. Leptodactylus melanonotus is known from Acaponeta, Nayarit (AMNH 43913-25), and the following localities in Jalisco: Barro de Navidad (UMMZ 118098), La Con- cepcién (UMMZ 118081), La Resolana (UMMZ 102104), and Tenachitl4n (UMMZ 113045-6). Records for Leptodactylus occi- dentalis are: Alamos, Sonora (AMNH 51356-65); Culiacan (AMNH 49511-9), Chele (UMMZ 110914), and Rosario (UMMZ 113062) in Sinaloa; Ixtl4n del Rio (UMMZ 102108), San Blas (UMMZ 112814, 112994, 110892, 115543), and Tepic (UMMZ 115544) in Nayarit; Ameca (UMMZ 102106-7) and La Cofradia on the south shore of Lago de Chapala (UMMZ 102105) in Jalisco; and Tanga- mandapio, Michoacan (UMMZ 119145). From these scattered records it appears that Leptodactylus occidentalis in the southern part of its range stays in the uplands, whereas melanonotus is con- fined to the lowlands. Microbatrachylus hobartsmithi (Taylor) Eleutherodactylus hobartsmithi Taylor, Trans. Kansas Acad. Sci., 39:355, 1987.—Uruapan, Michoacan, México. Microbatrachylus hobartsmithi Taylor, Univ. Kansas Sci. Bull., 26:501, November 27, 1940. Cascada Tzararacua (6); 21 km. W of Ciudad Hidalgo; 29 km. E of Morelia; Puerto Hondo; San José de la Cumbre (13); Uruapan (2); Zita- cuaro. Of six specimens from Cascada Tzararacua, five are colored like typical M. hobartsmithi, having the anterior and posterior surfaces of the thighs and the upper arms pale pink in life and a grayish brown dorsum in preservative. The other specimen (UMMZ 94231) has in preservative a dark brown dorsolateral line on each side en- closing a pale tan area that extends from the snout to the vent. One specimen from 29 kilometers east of Morelia (UIMNH 40338) and 13 specimens from San José de la Cumbre (UMMZ 102111) do not have the prominent tarsal tubercles characteristic of M. hobartsmithi. Also, in these fourteen specimens the palmar tubercles are larger, and the dark anal patch more distinct, than in typical M. hobart- smithi. Possibly these specimens, which are from the high mountains 34 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. in the eastern part of Michoacan, represent another species of Micro- batrachylus. However, Taylor (1940d:501) reported a series of M. hobartsmithi from the mountains 10 miles west of Villa Victoria in the western part of the state of México. The largest specimen from Michoacan is a gravid female (UIMNH 16104) having a snout-vent length of 23.5 mm. Microbatrachylus hobartsmithi has been found in rocky ravines along streams in the Cordillera Volcanica and the southwestern escarpment of these mountains at elevations from 1450 to 2750 meters. Microbatrachylus pygmaeus (Taylor) Eleutherodacytylus pygmaeus Taylor, Trans. Kansas Acad. Sci., 39:352, 1937.—1 mile north of Rodriguez Clara, Veracruz, México. a aiae pygmaeus Taylor, Univ. Kansas Sci. Bull., 26:500, Novem- er 2 Microbatrachylus albolabris Taylor, Univ. Kansas Sci. Bull., 26:502, Novem- ber 27, 1940.—2 miles west of Cordoba, Veracruz, México. Microbatrachylus minimus Taylor, Univ. Kansas Sci. Bull., 26:507, Novem- ber 27, 1940.—Agua del Obispo, Guerrero, México. Microbatrachylus imitator Taylor, Univ. Kansas Sci. Bull., 28:70, May 15, 1942.—La Esperanza, Chiapas, México. Arteaga (328). This large series (UMMZ 119247-8) was collected on June 22 and 23, 1958, before the onset of the heavy summer rains. The frogs were found in a shaded ravine at the north edge of Arteaga; they were obtained during the day, at which time they were actively moving about in the leaf litter along a small stream. These frogs are all referred to M. pygmaeus, because this is the earliest name available for frogs showing the variation in character- istics displayed by this large series. The characters used by Taylor (1936a, 1940d, 1941a, and 1942b) and Smith and Taylor (1948) to distinguish the various species of Microbatrachylus include color pattern, relative length of the hind limb, presence and position of dorsal dermal folds or pustules, relative size of inner and outer meta- tarsal tubercles, and the number of palmar tubercles. All specimens from Arteaga have two palmar tubercles; the inner and outer meta- tarsal tubercles are subequal in size. Furthermore, aside from sexual difference, there is little variation in the relative length of the hind limbs (Table 3). However, many color patterns do exist in the series; each of these color patterns is described below. AMPHIBIANS AND REPTILES OF MICHOACAN 35 TABLE 3.—SNOUT-VENT LENGTH EXPRESSED AS A PERCENTAGE OF TIBIA LENGTH IN ANIMALS OF SIx CoLoR PATTERNS OF MICROBATRACHYLUS PYG- MAEUS. (LETTERS REFER TO THE VARIANTS HAVING THE COLOR PATTERN DiscussED IMMEDIATELY BELOw ) Twice re Number Range standard P Cer Sex of of Mean error ACTEEN specimens variation of mean A rol 25 51.4-57.5 55.2 3.34 2 25 49 .3-54.9 51.6 3.12 B ou 20 51.0-57.1 55.4 2.44 2 21 47 .3-54.9 §1.2 3.52 C rot 6 54.5-56.2 QB ches ar eee gees fe) 6 50.0-52.9 SUG Ree ees D of 17 52.9-58.2 55.4 2.64 g 14 48 .5-56.6 §2.1 4.16 E ou 10 50.9-56.9 55.1 3.40 2 7 49.6-54.5 7 I Te ae Sse F Q 2 51.9-52.6 BZ. nee erties A.—225 specimens: Dorsum mottled brown and cream, usually with a dark spot between the eyes and one or two dark V-shaped marks with the apex anteriorly on the back; 55 of these have a narrow cream-colored line from the tip of the snout to the vent and thence onto the posterior surfaces of the thighs. All are pustulate above; in most specimens the pustules form no pattern, but in some they tend to form a V in the scapular region. B.—41 specimens: Dorsum pale tan or cream-color with brown mottling on flanks; a brown interorbital bar and a brown chevron in scapular region. Dorsum irregularly pustulate; in some specimens the pustules tend to form a V in the scapular region. C.—12 specimens: Dorsum colored like “A”, but having a broad yellow stripe narrowly bordered by black from the tip of the snout to the vent; in some specimens there is a narrow yellow stripe on the posterior surfaces of the thighs. The dorsum is irregularly pustulate. D.—81 specimens: Dorsum variably streaked with cream-color or pale tan and brown; usually a broad cream-colored stripe from eyelid to groin bordered laterally by a somewhat narrower brown stripe; middorsal area cream-color and separated from dorsolateral cream-colored stripe by a brown stripe, or middorsal area brown with a cream-colored or yellow, narrow stripe from tip of snout to vent; a dark stripe from tympanum to flank; dorsal surfaces of heels creamy white to pale orange; anal patch brown. A dermal ridge from posterior edge of eyelid to rump; another ridge extends posteromedially from the eyelid; scattered pustules on the dorsum in some specimens. E.—17 specimens: A narrow dark stripe from snout, through nostril and eye, over tympanum, to vent, enclosing a unicolor dorsum (reddish tan to yellowish tan in life); heels pale tan or yellow above; anal patch black. A faint dermal ridge from posterior edge of eyelid to rump, or part way to rump. F.—2 specimens: Mottled brown and cream-color above; upper lips and 36 UNIVERSITY OF KANSAS PUBLS., Mus. Nat. Hist. upper arms white. A dermal fold from posterior edge of eyelid to rump; scattered pustules on dorsum. Some of these color variants are assignable to names proposed by Taylor: “A” and “B” undoubtedly are M. pygmaeus (Taylor, 1936a); “C” probably is M. pygmaeus; “D” is referable to M. mini- mus (Taylor, 1940d) in most characteristics, although the colora- tion is more nearly like that of M. lineatissimus (Taylor, 1941a), a larger species characterized by a relatively long hind limb; “E” apparently is M. imitator (Taylor, 1942b); “F” is M. albolabris (Taylor, 1940d). Examination of series of these frogs from other parts of México shows a similar composition of color variants. Of 78 specimens from the Rio Sarabia and the village of Sarabia in Oaxaca (UMMZ 115428-37), 57 are “A,” six are “D,” three are “E,” and 12 are “F”; of 22 specimens from Teapa, Tabasco (UMMZ 113829), 11 are “A,” five are “D,” two are “E,” and four are “F’; of 33 specimens from Potrero Viejo, Veracruz (USNM 115447-58, 115461-71, 116840-2, 116864-70), ten are “A,” 18 are “E,” and ten are “F”’; of 31 specimens from La Esperanza, Chiapas (USNM 115477-9, 116827-39, 116849-63), 28 are “A” and four are “F.” It is highly doubtful if these color variants are actually distinct species. Goin (1950 and 1954) in his studies of inheritance of color pattern in West Indian species of the genus Eleutherodactylus has shown that similar color pattern variants come from the same clutch of eggs; furthermore, Goin has worked out the genetic ratios of certain of these variants. Heathwole (in litt.) obtained “normal” specimens and individuals having a broad middorsal stripe (“C” in figure 9) from a clutch of eggs of Eleutherodactylus gollmeri. The presence of a broad middorsal yellow stripe is com- mon in Eleutherodactylus rugulosus. Perhaps the most interesting aspect of variability in color pat- tern in Mexican eleutherodactylids is the parallelism between mem- bers of the Eleutherodactylus rhodopis-group and some members of Microbatrachylus. In the former group there are white-lipped individuals (Eleutherodactylus beatae Boulenger ), individuals hav- ing a unicolor reddish or yellowish dorsum (E. dorsoconcolor Tay- lor), and individuals having a dorsal pattern of irregular longi- tudinal brown and cream-colored streaks (E. venustus Giinther). In the humid forests of southern Veracruz, northern Oaxaca, and Chiapas members of both groups occur sympatrically. A proper understanding of the evolutionary significance of these variants in the two groups, as well as proper allocation of the presently recog- nized species, must await experimental evidence based on studies AMPHIBIANS AND REPTILES OF MICHOACAN 37 of the inheritance of color pattern. Nevertheless, at present it is apparent that certain characters, especially the nature of the der- mal folds and pustules, and the color pattern, are of little taxo- nomic value in distinguishing “species” of Microbatrachylus. The data derived from a study of the large series from Arteaga, together with that from the other series examined, suggests that Microba- trachylus albolabris, imitator, minimus, and pygmaeus are morpho- types of one species. Of these names, pygmaeus is the oldest. Consequently Microbatrachylus pygmaeus has been used here for the series from Arteaga. Although Microbatrachylus hobartsmithi, a species distinguished from all of the above by the presence of tubercles on the outer edge of the tarsus, is known from Michoacan northward into Nayarit, Microbatrachylus pygmaeus previously has not been known north of Guerrero, where it occurs in habitats similar to that in which it was collected at Arteaga. Eleutherodactylus augusti cactorum Taylor Eleutherodactylus cactorum Taylor, Univ. Kansas Sci. Bull., 25:391, July 10, 1989.—20 miles northwest of Tehuacdan, Puebla, México. Eleutherodactylus augusti cactorum, Zweifel, Amer. Mus. Novitates, 1813:20, December 28, 1956. Cheran; Coalcoman; Uruapan. The few specimens indicate that this species occurs at moderate to high elevations in the state. The specimens from Cheran and Urua- pan were obtained in pine forests; the specimen from Coalcoman was found on a rocky hillside covered with dense forest and located about 100 meters below the lower limits of the pine forest in the area. A specimen from Rancho Reparto (elevation 1850 meters ) on the west slope of Cerro Barolosa was lost. The specimen from Coalcoman (UMMZ 104728) is a juvenile having a snout-vent length of 25.0 mm. In life it was tan above, mottled with olive-green. The ventral surfaces were gray; the hind limbs were distinctly barred with yellow and brown, and the lips were barred with yellow and black. Eleutherodactylus occidentalis Taylor Eleutherodactylus occidentalis Taylor, Proc. Biol. Soc, Washington, 54:91, July 31, 1941.—Hacienda El Florencio, Zacatecas, México. Arteaga (2); Cascada Tzararacua; Coalcoman (2); 19 km. SW of Coire (3); La Placita (7); Los Reyes; Ostula (4); Pémaro (2). The locality records for this species suggest that it is a member of a group of animals, the distribution of which includes the western part of the Mexican Plateau and the Pacific lowlands. In Michoacdn 88 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. this frog has been collected in pine-oak forest at Cascada Tzararacua and at Los Reyes, in arid scrub forest at Arteaga and Coalcomén, and in tropical semi-deciduous forest on the lower Pacific slopes of the Sierra de Coalcoman. On July 5, 1950, James Peters (1954:6) found calling males at La Placita. Most of the specimens are immature; four adult males have snout-vent lengths of 30.9-33.0 (32.2) mm. In all specimens the first finger is noticeably longer than the second; the inner meta- tarsal tubercle is large, flat, and cream-colored, contrasting with the dark brown sole of the foot. When the hind limbs are ad- pressed, the heels broadly overlap. Characteristically, a dark line extends from the snout, through the eye, above the tympanum, to a point above the insertion of the forelimb. Usually there is a dark bar behind the tympanum, two dark brown bars from the eye to the mouth and thence onto the lower jaw, and another dark bar on the upper lip between the eye and nostril. One adult from Arteaga, an adult and a juvenile from La Placita, and one juvenile each from Coire, Ostula, and Pémaro, have the lower lip barred with dark brown and white, and have a white stripe extending the length of the upper lip. In life the dorsum varies from dark gray or olive-brown to tan or reddish brown. This species belongs to a group containing two other populations that are currently recognized as species—calcitrans, known only from Omiltemi, Guerrero, and mexicanus, reported from the moun- tains of Oaxaca. Another apparently undescribed member of this group has been collected in the mountains of northern Puebla. The locality records indicate that the group inhabits the mountains on the periphery of the Mexican Plateau, except in western México, where Eleutherodactylus occidentalis extends to the Pacific low- lands. Eleutherodactylus rugulosus vocalis Taylor Eleutherodactylus vocalis Taylor, Univ. Kansas Sci. Bull., 26:401, Novem- ber 27, 1940.—Hacienda E] Sabino, Michoacan, México. Arteaga (10); El Sabino (8); Salitre de Estopilas (3); Tumbiscatio (2); Tzitzio (2). The distributional data on this frog in Michoacan indicate that it inhabits riparian situations in arroyos and canyons in the lower slopes of the Cordillera Volcdnica and the Sierra de Coalcoman, where it has been taken at elevations only below 1100 meters. The dorsal color of living individuals from Arteaga varied from dark gray and olive brown to tan and reddish brown. The iris AMPHIBIANS AND REPTILES OF MICHOACAN 39 was grayish brown. In contrast, individuals from Agua del Obispo, Guerrero, had pale golden eyes; specimens from Matias Romero, Oaxaca, had gold eyes heavily flecked with gray; and individuals from Volcan San Martin, Veracruz, had bronze eyes. The use of the trinomial here is arbitrary. Frogs of the Eleuthero- dactylus rugulosus group in México (rugulosus, avocalis, and vo- calis) exhibit only slight differences in size, proportions, and colora- tion (Duellman, 1958c:6). Furthermore, the named populations are allopatric. Eleutherodactylus rugulosus vocalis, as defined by Duellman (loc cit.), occurs in the foothills of the Sierra Madre Occidental and associated ranges from central Sinaloa southward into Michoacan. Tomodactylus angustidigitorum Taylor Tomadoctylus angustidigitorum Taylor, Univ. Kansas Sci. Bull., 26:494, November 27, 1940.—Quiroga, Michoacan, México. Angahuan (6); Apo; Carapan (21); 19 km. S of Carapan (13); Cerro Tancitaro (12); Cheran; Corupu (14); Cusefio Station (14); Opopeo (3); Paracho (11); Paricutin (2); Patzcuaro (3); Quiroga (59); San Juan de Parangaricutiro (16); Tancitaro (25); Uruapan (8); Zacapu (11). This species is indigenous to the pine-oak forests on the south- em rim of the Mexican Plateau, and has been collected at eleva- tions from 1500 to 2500 meters. Males have been observed to call from rocks, rock fences, clumps of grass, and low bushes; the call is a single “peep.” At San Juan de Parangaricutiro numerous specimens were found in the daytime beneath adobe bricks and lava on the volcanic ash derived from Volcan Paricutin; at Paracho individuals were found by day beneath rocks in a pine forest. In most specimens the dorsum is dark reddish brown, and the prominent inguinal glands are cream-color or pale orange (PI. 8, Fig. 1). Of eight individuals collected at Paracho, one was reddish brown, two were pinkish tan, three were dark brown, and two were black. Tomodactylus fuscus Davis and Dixon Tomodactylus fuscus Davis and Dixon, Herpetologica, 11:157, July 15, 1955.—1.5 miles southeast of Huitzilac, Morelos, México. Los Cantiles (2); 28 km. E of Morelia. The range of this species includes the Sierra Ajusco in México and Morelos and thence westward to the Serrania Ucareo in Michoacan. The specimen from 28 kilometers east of Morelia was found in an oak forest on a steep hillside at an elevation of 2100 meters. One from Los Cantiles was calling from a steep cliff at 40 UNIvERsITY OF KANSAS PuBLs., Mus. Nat. Hist. an elevation of 2200 meters in pine-oak forest. This specimen (UMMZ 119156) in life had a pale olive-brown dorsum with ir- regular dark brown mottling and transverse bars on the limbs. The interorbital bar, the upper arms, and the tips of the dorsal pustules were pale orange; the iris was pale grayish gold (Pl. 3, Fig. 2). ' Tomodactylus nitidus nitidus (Peters) Liuperus nitidus Peters, Monats. Akad. Wiss. Berlin, p. 878, 1869.— Izicar de Matamoras, Puebla, México. Tomodactylus amulae Giinther, Biologia Centrali-Americana, Reptilia and Batrachia, p. 219, April, 1900.—Amula, Guerrero, México. es nitidus nitidus, Dixon, Texas Jour. Sci., 9:385, December, 1957. Copuyo (15); Tuxpan (8); Tzitzio (11). One specimen from Tzitzio (UMMZ 99155) was referred to Tomodactylus nitidus petersi by Dixon (1957:390). A re-examina- tion of this specimen, and examination of ten others from the same locality (UMMZ 121571) reveals that the relatively small size of the tympanum and absence of dense ventral spotting place these specimens closer to T. nitidus nitidus than to T. nitidus petersi. The specimens from Tuxpan (UMMZ 114808-4) had in life a gray to olive tan ground color with dark olive-green markings, bright yellow thighs with olive-green transverse bands, yellowish tan shanks with olive-green bars, yellow groin, white inguinal glands with black markings, grayish white belly with scattered brownish black spots in some specimens, and a deep golden iris (PI. 4, Fig. 1). These specimens were found calling from bushes in a rocky field at an elevation of 1800 meters. The call is a high-pitched “pee-ee-eep.” Tomodactylus nitidus orarius Dixon Tomodactylus nitidus orarius Dixon, Texas Jour. Sci., 9:392, December, 1957. —4.5 miles southwest of Tecolapa, Colima, México. La Placita (8); Pémaro. These specimens, referred to Tomodactylus petersi by Duellman (1954b:5), were included in T. nitidus orarius by Dixon (1957:392). Color notes based on living individuals from Tecolapa, Colima (UMMZ 114312 and 116922), are: gray above mottled with brown; venter dirty white; anterior and posterior surfaces of thighs bright yellow; iris pale golden (Pl. 4, Fig. 2). The call is a soft “braa” usually followed by three high notes: “braaa-eep-ee-eep.” In Mi- choacin this subspecies has been found only in the coastal region and the lower foothills of the Sierra de Coalcom4n, an area in which AMPHIBIANS AND REPTILES OF MICHOACAN 4l it replaces Tomodactylus nitidus petersi. This is the only Tomodac- tylus known to inhabit coastal lowlands. Tomodactylus nitidus petersi Duellman Tomodactylus petersi Duellman, Occ. Pap. Mus. Zool. Univ. Michigan, 560:5, October 22, 1954.—Coalcoman, Michoacan, México. Tomodactylus nitidus petersi, Dixon, Texas Jour. Sci., 9:390, December, 1957. Aguililla; Apatzingan (8); Cascada Tzararacua; Charapendo (5); Coalcoman (5); 18 km. E of Dos Aguas (6); El Sabino (5); La Playa (2); Jiquilpan; Uruapan (2); Volcan Jorullo; Zamora. In life, specimens from Apatzingin (UMMZ 114308-9) varied in dorsal color from grayish tan to pale brown; the dorsal markings were olive green. The thighs and groin were yellowish orange; the iris was pale golden, and the vocal sac was purplish gray (PI. 5, Fig. 1). Measurements for 13 adult males from the Tepalcatepec Valley are: snout-vent length, 21.9-26.8 (24.3); tibia length, 8.4-9.9 (9.3); head width, 7.2-9.2 (7.8); head length, 7.6-8.7 (8.2). At Apatzingén and Charapendo in the Tepalcatepec Valley males were found calling from rocks and bushes in open arid tropical scrub forest. The call, a triple note “peep-ee-eep,” is repeated once every 90 to 135 seconds. Tomodactylus nitidus petersi probably ranges throughout the Tepalcatepec Valley and surrounding foothills. Dixon (1957:392) referred the specimens from Zamora, Jiquilpan, and Uruapan to this subspecies. Uruapan is near the lower limits of the pine forest on the slopes of the Cordillera Volc4nica; Zamora and Jiquilpan are on a low part of the Mexican Plateau southeast of Lago de Chapala. Tomodactylus rufescens Duellman and Dixon Tomodactylus rufescens Duellman and Dixon, Texas Jour. Sci., 11:78, March, 1959.—Dos Aguas, Michoacan, México. Dos Aguas (14); 18 km. E of Dos Aguas (6). Fourteen specimens from the pine-oak forests around Dos Aguas (UMMZ 118508-10, 121498-9) have reddish brown dorsal color and a narrow cream-colored middorsal line (Pl. 5, Fig. 2). Twelve of these specimens are adult males having snout-vent lengths of 20.7 to 24.6 (22.5) mm. One female has a snout-vent length of 24.8 mm., and one juvenile has a snout-vent length of 14.5 mm. Six specimens are from a region of mixture of pine-oak forest and arid tropical scrub forest at 18 kilometers east of Dos Aguas (UMMZ 121497, 121500). All are males having snout-vent lengths of 18.0 to 22.6 (20.7) mm. The dorsum is tan marked with black; the thighs are yellowish orange. 42 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist. The specimens from 18 kilometers east of Dos Aguas were found on July 22, 1960, by Floyd L. Downs and John Winklemann, who collected calling males of Tomodactylus rufescens and Tomodac- tylus nitidus petersi at the same locality. Downs (personal com- munication) stated the call was a single note. At Dos Aguas I heard T. rufescens give two calls, one a single “peep,” the other a triple note—“pee-ee-eep.” In the higher parts of the Sierra de Coalcoman Tomodactylus rufescens seems to fill the same niche as T. angustidigitorum does in the Cordillera Volcanica. At lower elevations in their respective mountain ranges the species occur sympatrically with T. nitidus petersi. Diaglena reticulata Taylor Diaglena reticulata Taylor, Univ. Kansas Sci. Bull., 28:60, May 15, 1942.— Cerro Arenal, Oaxaca, México. Nueva Italia (3); Ostula (7). Until recently frogs of the genus Diaglena were known only from a few specimens from southern Sinaloa (Diaglena spatulata) and from the Pacific lowlands of the Isthmus of Tehuantepec (Diaglena reticulata). Peters (1955a) reported specimens from Ostula, Michoac4n, and compared these specimens with one D. reticulata from Tehuantepec, Oaxaca, and four D. spatulata from Sinaloa. This comparison showed that the specimens from Michoa- can, although showing some minor differences from D. reticulata, are closer to that species than to D. spatulata. Subsequent to Peters’ work, series of both species of Diaglena, including additional speci- mens from Michoacan and from Colima, have been collected, and a more qualified comparison is now possible. In comparing specimens of D. spatulata from southern Sinaloa (UMMZ 115322) with specimens of D. reticulata from Tehuante- pec, Oaxaca (UMMZ 115321), the differences noted by Taylor (1942c:60) were found to be constant. But specimens from Ostula, Michoacan (UMMZ 104418), and five individuals from Colima (TNHC 26379-83) were found to be intermediate in certain charac- ters. The skin of the dorsum in D. reticulata is granular; that in D. spatulata is smooth. The skin in specimens from Ostula and Colima is slightly granular. The dorsal ground color of D. reticulata is yellowish brown with dark reticulations; the dorsal ground color of D. spatulata is olive-green. Specimens from Ostula and Colima most closely resemble those from Tehuantepec in coloration, but the reticulations are more coarse, and the ground color has an olive- AMPHIBIANS AND REPTILES OF MICHOACAN 43 green tint. Diaglena reticulata also differs from D. spatulata in having a larger over-all size, slightly broader head, a narrower interorbital distance, and a more pointed snout with a deeper labial shelf (Table 4). The specimens from Ostula and Colima are inter- mediate between D. reticulata from Oaxaca and D. spatulata from Sinaloa in body proportions. Of three specimens from the Tepalcatepec Valley (JRD 5991-3), only two are suitable for measuring. These specimens are smaller than adults from the coastal areas and have broader heads and snouts, but narrower interorbital distances, than specimens in the other samples (Table 4). The texture of the skin is like that of specimens from Ostula and Colima. The coloration resembles that of D. reticulata, but the reticulations are bold and form indistinct bands on the hind limbs. TABLE 4.—CoOMPARISON OF Four CHARACTERS IN FIVE SAMPLES OF DIAGLENA. (Att Data ARE FOR MALES; MEANS GIVEN IN PARENTHESES BELOW RANGES. ) Head ies Internarial L purer Snout-vent eladehy distance eens) OCALITY e) length specimens Snout-vent Hoad Head length widtli width OS. COL IRA 9 71.1-87.5 | 25.4-29.1 | 63.0-71.4 | 11.9-13.8 (80.7) (27.9) (67.1) (12.9) Coast of Michoacén 5 72.0-79.2 | 24.3-27.2 | 67.0-73.8 | 13.7-14.4 (74.8) (25.6) (71.4) (14.1) Colima 2/3. 23. +t 71.7-79.6 | 26.1-28.6 | 70.5-75.3 | 16.0-17.9 (74.8) (27.4) (72.0) (16.6) Tepalcatepec Valley...... 2 63.0-65.4 | 28.3-32.2 | 57.3-62.4 | 17.0-20.2 (64.2) (30.3) (59.9) (18.6) DINSIOR acces « 11 71.9-81.3 | 24.0-27.3 | 70.5-78.1 | 15.0-17.3 (77.3) (25.7) (73.4) (16.1) All specimens from Michoacan and Colima more closely approach Diaglena reticulata than D. spatulata. The acquisition of additional specimens, especially from the area between Sinaloa and Colima and from Guerrero, is necessary to determine the relationships among the various populations known at present. Both species of Diaglena inhabit tropical scrub forest; none has been found in the more humid and tropical semi-deciduous forests. Humid forest replaces the scrub forest in the lowlands of southern Nayarit and 4—7817 44 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. northern Jalisco; possibly this forest acts as a barrier to the distri- bution of Diaglena and thus serves as a divider between the ranges of D. spatulata to the north and D. reticulata to the south. Pternohyla fodiens Boulenger Pternohyla fodiens Boulenger, Ann. Mag. Nat. Hist., ser. 5, 10:326, 1882.— Presidio, Sinaloa, México. Nueva Italia (2). These specimens (JRD 5994-5) were found on the road near Nueva Italia during a heavy rain on the night of August 25, 1960, by James R. Dixon. Both are females having snout-vent lengths of 64.0 and 59.0 mm. They are typical of the species as it is known from Sinaloa, Nayarit, Jalisco, and Colima. These specimens constitute the southernmost record for the species, which ranges in semiarid habitats from southern Arizona southward along the Pacific lowlands of México to Colima and in- land on the Mexican Plateau in Jalisco. Phyllomedusa dacnicolor Cope Phyllomedusa dacnicolor Cope, Proc. Acad. Nat. Sci. Philadelphia, 16:181, September 30, 1864.—Colima, Colima, México. Funkhouser, Occ. Pap. Nat. Hist. Mus. Stanford Univ., 5:37, April 1, 1957. Agalychnis alcorni Taylor, Copeia, no. 2:31, June 2, 1952.—Rio Tepal- catepec, 17 miles south of Apatzingan, Michoacan, México. Agalychnis dacnicolor, Duellman, Herpetologica, 18:29, March 30, 1957. Phyllomedusa alcorni, Funkhouser, Occ. Pap. Nat. Hist. Mus. Stanford Univ., 5:80, April 1, 1957. Aguililla (18); Apatzingan (7); Charapendo; Coahuayana (8); Coalcoman (54); El Sabino; Huetamo Road (2); La Orilla; La Placita; Nueva Italia (4); 32 km. E of Neuva Italia (2); Rio Cancita, 14 km. E of Apatzingan; Rio Tepalcatepec, 27 km. S of Apatzingdn; Salitre de Estopilas (2); Tzi- tzio (4). This large tree frog has been found only in the lowlands below elevation of about 1000 meters, usually in arid tropical scrub forest. Calling males were heard on rainy nights throughout the rainy season; in nearly every instance both males and females were found in low trees and bushes. On summer nights when there had been no rain, adults were found sitting on bushes in the scrub forest. At Coalcoman on July 1, 1955, a chorus was heard at midday. About forty Phyllomedusa dacnicolor were found in one guayava bush at the edge of a recently dried pond. Individual males were calling; clasping males were silent. The call is a barking groan. Fifteen individual egg masses were hanging from branches and leaves in tear-drop fashion. Each egg mass contained 100 to 350 pale green eggs, located only in the exterior part of the clear gela- AMPHIBIANS AND REPTILES OF MICHOACAN 45 tinous mass. Two composite egg masses appeared to have been made up by egg deposition on the part of three to five females (PI. 2, Fig. 2). As shown by Duellman (1957a), the characters used by Taylor (1952) to diagnose Phyllomedusa alcorni are sexually dimorphic. Funkhouser (1957) apparently was unaware of this sexual di- morphism, for she recognized P. alcorni and P. dacnicolor as distinct species. Phrynohyas inflata (Taylor) Acrodytes inflata Taylor, Univ. Kansas Sci. Bull., 30:64, June 12, 1944.— La Venta, Guerrero, México. Phrynohyas inflata, Duellman, Misc. Publ. Mus. Zool. Univ. Michigan, 96:19, February 1, 1956. Phrynohyas corasterias Shannon and Humphrey, Herpetologica, 13:15, March 30, 1957.—4.8 miles east of San Blas, Nayarit, México. Barranca de Bejuco. One specimen of this large species was collected in 1951; it was found on a low branch in tropical semi-deciduous forest at an eleva- tion of 65 meters. In life there were olive-gray blotches on a pale gray dorsum; the iris was a dark golden color. This species, which is known from only a few specimens, seems to be restricted to the coastal lowlands and low foothills from Guerrero northward to Nayarit. Shannon and Humphrey (1957) described Phrynohyas corasterias from Nayarit. Their description was based on a small female having a snout-vent length of 34.4 mm. The new species was diagnosed as differing from P. inflata in having less webbing on the feet, a poorly developed supratympanic fold, a more pustulate dorsum, and marked differences in dorsal pattern, color, and nature of antebrachial banding. The significance of the webbing was questioned by Shannon and Humphrey. The nature of the supratympanic fold and dorsal pustules changes with age (Duellman, 1956a:31). Phrynohyas inflata is known to attain a snout-vent length of 95 mm. Dermal structures that undergo onto- genetic change are of little importance in comparing a juvenile with a large adult. The only significant difference in color pattern be- tween P. inflata and P. corasterias is the presence of wide transverse bands on the limbs of the latter. In this respect P. corasterias ap- proaches P. latifasciata, a species known only from two specimens from southern Sinaloa. The acquisition of additional specimens from Jalisco, Nayarit, and Sinaloa may show that P. inflata and P. latifas- ciata are conspecific, as suggested by Duellman (1956a:21). None- theless, the specimen on which the description of P. corasterias was 46 UNIVERSITY OF Kansas PuBts., Mus. Nat. Hist. based is not sufficiently different from the known specimens of P. inflata to warrant specific recognition. Hyla arenicolor Hyla arenicolor Cope, Jour. Acad. Nat. Sci. Philadelphia, ser. 2, 6:84, July, 1866.—Northern Sonora, México. Type locality restricted to Santa Rita Mountains, Pima County, Arizona, by Smith and Taylor (1950a:354). Agua Cerca; Cascada Tzararacua (3); Chinapa; Cojumatlin; Dos Aguas; El Sabino (25); El Espinal; Lago de Camécuaro; Lombardia (2); Tupataro; Zinapécuaro. Altitudinally this frog ranges from 500 to 2100 meters; although the environments in which it has been found vary from open arid tropical scrub forest to pine forest, it usually is found near rocky streams in these habitats. There is great disparity in size between specimens from the mountains and those from the Tepalcatepec Val- ley. Seven males from elevations in excess of 1400 meters have an average snout-vent length of 34.7 mm.; nine from elevations below 1000 meters have an average snout-vent length of 49.1 mm. In life a male collected at night at Lombardia (UMMZ 112846) had dark brown spots on a grayish brown dorsum; the groin, anterior and posterior surfaces of the thighs, and ventral surfaces of the hind limbs and palms were yellowish orange. The belly and tips of digits were white; the vocal sac was purplish brown, and the iris was dark grayish gold. In contrast, a specimen obtained in the daytime at Chinapa (UMMZ 119204) had indistinct gray spots on a pale ashy gray dorsum; the flash colors were yellow. After dark the spots were dark olive-brown on a grayish brown dorsum. Two males were found calling from a rocky stream near Lom- bardia on July 12, 1955. The call is a nasal “ah-ah-ah-ah.” Hyla baudini Duméril and Bibron Hyla baudinii Duméril and Bibron, Erpétologie générale, vol. 8:564, 1841. —Meéxico. Type locality restricted to Cérdoba, Veracruz, México, by Smith and Taylor (1950a:3846). Aguililla (5); Apatzingan (80); Arteaga; Buena Vista; Charapendo; Coa- huayana; Cofradia (4); El Sabino (12); La Placita; La Playa; Maruata; Nueva Italia (3); 82 km. E of Nueva Italia (2); Ostula (4); Rio Tepal- catepec, 25 km. S of Apatzing4n (3); Salitre de Estopilas; San José de la Montafia (2); Tumbiscatio; Tzitzio. This tree frog is widespread in the coastal lowlands and in the Tepalcatepec Valley up to elevations of about 1200 meters. It is found in numbers in the early part of the rainy season, at which time males were heard calling from bushes and trees along ditches and temporary ponds. The call is a loud nasal “waank-waank- waank.” One individual that was emitting a long and unusually AMPHIBIANS AND REPTILES OF MICHOACAN 47 high-pitched call was found to have one hind limb engulfed by a Leptodeira maculata. When active at night these frogs usually are pale tan to reddish brown above with dark brown markings. A specimen found sitting on a maguey plant in the daytime was pale ashy gray with a pale green upper lip. Hyla bistincta Cope Hyla bistincta Cope, Proc. Amer. Philos. Soc., 17:87, 1877.—Veracruz, México. Type locality restricted to Acultzingo, Veracruz, México, by Smith and Taylor (1950a:846). Cerro San Andrés; Dos Aguas (2); Los Conejos (8); Uruapan (50). In the Parque Nacional at Uruapan this species was found in abundance during the day. The frogs hide in an entanglement of vines and vegetation overhanging several small spring-fed streams. Tadpoles were in the rocky streams, and metamorphosing young were on vegetation at the edges of the streams. In life the dorsum is greenish tan with brown mottling; in some indivduals the entire dorsum is dark chocolate brown. The flanks are pale lemon yellow barred with lavender-brown. Notes on the color of a living frog from Dos Aguas (UMMZ 119193) are: Dorsal ground color a medium shade of brown with dark brown flecks; flanks black with silvery white and pale yellow spots; belly pale yellowish white; throat mottled with grayish brown; iris pale copper color. Fic. 8. Tadpole of Hyla bistincta (UMMZ 115231) from Uruapan, Michoacan. x 2. Description of Tadpole: Body somewhat depressed; maximum width of body slightly more than one-half of body length. Nostrils placed dorsolaterally and directed anteriorly, situated about mid- way between tip of snout and eye. Eyes of moderate size, dor- solateral in position and directed upwards. Tail about twice as long as body, thrice as long as deep, and tapering gradually to a rounded tip. Tail-musculature not extending to tip of tail fin. Spiracle sinistral, lateral, and situated at midbody. Vent dextral; 48 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. the cloacal tube extending along ventral part of tail for a distance equal to about one-eighth of body length (Fig. 8). Average body length of six tadpoles with small hind limb buds, 19.5 mm.; tail length, 38.3 mm. Mouth ventral, its width equal to about two- thirds of greatest width of body. Lips bordered by two rows of small papillae; row of larger papillae between upper lip and outer upper tooth-row, similar row between lower lip and outer lower tooth-row; laterally these rows degenerating into numerous small papillae. Horny beaks well developed; upper beak moderately arched and deeply indented; lower beak slightly indented. Ser- rations of beaks blunt and peglike, moderately developed on both beaks, but slightly stronger on lower one. Tooth-rows 2/3; upper rows nearly equal in length and slightly longer than lower rows, which are subequal in length; inner upper tooth row interrupted medially by rounded notch; inner lower tooth-row turned down- ward laterally; teeth in all rows about equal in size, but decreas- ing in length laterally (Fig. 9). een caress oy g aS! HURL ay Boa (>>, ww lay hy ee os Re ~ go ssn: Snug Bie ee at wo Pal “etary Z ne ves bo: ». LTT te «sre OMI, = 7 Sree ety, one het em nanan JPN, L278 AAR AA dA Zi si nen OOO ASATIISAY Fic. 9. Mouthparts of larval Hyla bistincta (UMMZ 115281) om Uruapan, Michoacan. x 15. Color in formalin: pale grayish brown dorsally and laterally; pale gray ventrally; tail-musculature brown; tail-fin translucent with scattered melanophores most numerous on upper fin. In most details these tadpoles resemble those of Hyla robertsorum described by Rabb and Mosimann (1955). Four metamorphosing young have snout-vent lengths of 23.0- AMPHIBIANS AND REPTILES OF MICHOACAN 49 23.5 (23.2); a completely metamorphosed individual has a snout- vent length of 24.8 mm. In Michoacan this stream-breeding hylid occurs at elevations of 1,600 to 2,400 meters in the Sierra de Coalcoman and in the moun- tains rising from the Mexican Plateau. Hyla eximia Baird Hyla eximia Baird, Proc. Acad. Nat. Sci. Philadelphia, 7:61, October 20, 1854.—Valley of México. Type locality restricted to Coyoacan, Distrito Federal, México, by Smith and Taylor (1950a:329). Hyla microeximia Maslin, Herpetologica, 13:81, July 10, 1957.—8 miles northwest of Jocotepec, Jalisco, México. Ciudad Hidalgo (36); Cuitzeo; 29 km. NW of Jacona; Jiquilpan (2); Lago de Camécuaro (2); Lago de Patzcuaro (129); Los Reyes; Morelia; Sahuayo (8); San Gregorio (63); Tangamandapio (4); Temazcal (26); Tupataro; Tuxpan (15); Undameo (2); Uruapan (20); Zacapu; Zamora (27); Zina- pécuaro (10). More than 80 per cent of the specimens from Michoacan have brown spots between the lateral and dorsolateral dark stripes, and more than 50 per cent have spots between the dorsolateral stripes, at least posteriorly. In comparison with specimens from the Valley of México, those from Michoacan have more distinct dorsolateral stripes that extend farther anteriorly, sometimes to the eyelid, and in this respect are more nearly like those from Jalisco and Nayarit (Taylor, 1989b:425). Some specimens from the western part of Michoacan possess certain characters used by Maslin (1957:81) to distinguish Hyla microeximia from H. eximia; nevertheless, the variation is such that two species cannot be distinguished in Michoa- cin. Four series of freshly preserved specimens have been studied in detail; in the discussion below they are arranged from west to east; the measurement is for snout-vent length of ten males from each sample: Zamora.—Twenty-two specimens (UMMZ 102083), 24.0-27.6 (26.1) mm. Dorsolateral dark stripe, or row of dashes, present in all specimens; dark spots in lateral and dorsal green fields; lateral dark stripe confluent with dorsolateral stripe posteriorly in 18 specimens; white line not extending to groin. Temazcal.—Thirty-five specimens (UMMZ 119162), 26.5-31.1 (28.2) mm. Dorsolateral dark stripe of row of spots present only posteriorly in most; both dorsolateral stripes and dorsal spots lacking in four specimens; heavy spotting dorsally in three others; lateral and dorsolateral dark stripes confluent posteriorly in three; lateral white stripe extending to groin in 16 specimens. Cuidad Hidalgo.—Thirty-six specimens (UMMZ 119168), 26.4-30.9 (28.2) mm. Dorsolateral dark stripe or row of spots present only posteriorly in most; no brown spots in the green fields of many specimens; large brown inguinal spot in most specimens; heavy spotting dorsally in four; lateral and dorsolateral 50 UNnIvERSITY OF Kansas Pusts., Mus. Nat. Hist. dark stripes confluent posteriorly in five; lateral white line extending to groin in most specimens. Tuxpan.—Fifteen specimens (UMMZ 115227), 28.7-33.0 (30.5) mm. Dor- solateral dark stripe or row of dashes in all specimens; dark spots in lateral green fields, at least posteriorly in most; dark spots posteriorly in the dorsal green field in five; lateral dark stripe separated from dorsolateral stripe in all specimens; lateral white line extends to the groin in all specimens. As can be seen from the above descriptions, the distinguishing characters of Hyla microeximia—confluence of lateral and dorso- lateral dark stripes posteriorly, extent of lateral white stripe, and distribution of dark spots dorsally—are found in individuals from all of the populations sampled. In the samples from western Michoacan there is a higher incidence of microeximia-like frogs than in those from other parts of the state. Hyla eximia is a wide- ranging species varying greatly geographically and individually. A thorough review of the species and related members of the Hyla eximia-group is necessary before certain populations can justifiably be segregated as subspecies or species. In Michoacan Hyla eximia has been collected in mesquite-grass- land, pine-oak forest, and cultivated areas on the Mexican Plateau from 1500 to 2300 meters; apparently it is absent from the Sierra de Coalcoman. This is the most abundant frog on the southern part of the Mexican Plateau; in the rainy season breeding choruses are found in temporary pools and in the marshes adjacent to the permanent lakes. Hyla lafrentzi Mertens and Wolterstorff Huy lafrentzi Mertens and Wolterstorff, Zool. Anz., 84:235, August 25, 1929.—Desierto de los Leones, Distrito Federal, México. Cerro San Andrés (26); Opopeo (9). In March, 1949, James A. Peters collected this species at eleva- tions of 2400 to 2800 meters on the west slope of Cerro San Andrés. The frogs were found beneath logs and rocks in a damp canyon in coniferous forest. Among the juveniles in this series is a com- pletely transformed individual (UMMZ 102093) having a snout- vent length of 14.5 mm. Five adults have snout-vent lengths of 36.2-39.5 (38.0) mm. Hyla lafrentzi has noticeably longer hind limbs than H. eximia; in the former, when the hind limb is brought forward along the body, the tibiotarsal articulation extends to the snout. There are dark transverse bands on the hind limbs; the dorsolateral stripe is broken into an anterior and a posterior seg- ment, and the latter is narrowly bordered by white in most specimens. AMPHIBIANS AND REPTILES OF MICHOACAN 51 Hyla lafrentzi occurs at higher elevations than any other frog in Michoacan; the locality records from throughout the range in- dicate that it is restricted to pine and pine-fir forests. In these habitats it replaces Hyla eximia, which inhabits the lower pine- oak forests and mesquite-grassland on the Mexican Plateau. Ponds are absent at places where Hyla lafrentzi has been collected; pos- sibly the eggs are laid in streams. Hyla smaragdina Taylor Hyla smaragdina Taylor, Copeia, No. 1:18, March 30, 1940.—6 kilometers east of Cojumatlan, Michoacan, México. Hylella azteca Taylor, Proc. Biol. Soc. Washington, 56:49, June 16, 1943.— Tepoztlin, Morelos, México. Cojumatlin (30); Copuyo (7); 18 km. E of Dos Aguas (22); Ostula (8); Pémaro (3); Sahuayo; Salitre de Estopilas (7). Taylor (1940a:18) diagnosed this species as having few or no vomerine teeth, no vocal sac, a rather broad and flat head, two large tubercles below the anus, a granular venter, and a green dorsum in life. The specimens on which the description was based are either immature or non-breeding individuals; all were collected from bromeliads growing on cacti near Cojumatlén. Another small, flat- headed hylid from Tepoztlan, Morelos, was described and diagnosed by Taylor (1943b:49) as differing from Hyla smaragdina in having a vocal sac and a broader head. This specimen was named Hylella azteca. Specimens from the coastal region of Michoacan and Colima were referred to Hylella azteca by Peters (1954:7) and Duellman (1958c:8). Comparison of topotypic Hyla smaragdina and the holotype of Hylella azteca (UIMNH 25044) with the several series of specimens from Michoacan has resulted in the conclusion that all pertain to only one species. Although the type series of Hyla smaragdina con- sists of immature specimens, the males in that series do possess vocal sacs. Since these were not breeding individuals, the sacs are not well developed. The characters of the anal tubercles and the relative width of the head are of no value in separating the two species. The apparently aestivating individuals comprising the type series of Hyla smaragdina, and the type of Hylella azteca, which also was found in a bromeliad, were green in life. Of the calling males found on the coast of Michoac4n, most were yellowish tan when found; two were pale green, but soon changed to pale tan. Calling males from Copuyo and Dos Aguas were pale yellowish tan. Therefore the color of the dorsum is of little significance in distinguishing the two named populations. 52 University oF Kansas Pusts., Mus. Nat. Hist. Males of Hyla smaragdina have been found calling in the months of June and July from rocky streams; the call is a nasal “haah-haah- haah,” repeated quickly and constantly for as long as 30 seconds. As pointed out by Duellman (1958c:9), this breeding behavior is un- like that suggested by Taylor (1943b:51). In Michoacan Hyla smaragdina has been found in tropical semi-deciduous forest, oak forest, and mesquite-grassland at elevations from 150 to 1500 meters. Hyla smithi Boulenger Hyla smithi Boulenger, Zool. Rec. Reptilia and Batrachia, 38:33, 1902.— Cuernavaca, Morelos, México. Aguililla (14); Apatzingan (104); Arteaga; Charapendo (5); Coalcoman (11); El Sabino (44); La Playa (6); Lombardia (2); Nueva Italia (8); Playa Azul; Salitre de Estopilas (2). This small hylid is abundant in the Tepalcatepec Valley to eleva- tions of about 1000 meters; it was found infrequently on the coastal lowlands. Males call from bushes in and around flooded fields and ditches, from grasses and small herbs in the water and from vegeta- tion overhanging small streams. The call consists of a series of short, high notes, somewhat reminiscent of a katydid’s song. In the dry season occasional males were heard calling from irrigated fields near Apatzingan. In the daytime individuals were found in the axils of leaves of the elephant-ear plants (Xanthosoma). In living individuals the dorsal color usually is uniform pale yel- low; often the lateral white stripe is barely visible. The vocal sac is bright yellow, and the iris is pale gold. In some individuals there are scattered dark brown spots or flecks on the back and upper surfaces of limbs. Twenty males from Apatzingén have the follow- ing measurements: snout-vent length, 22.8-26.0 (25.0) mm., tibia length, 10.7-18.6 (12.6) mm.; head width, 7.2-8.0 (7.6) mm., head length, 7.1-8.1 (7.7) mm. Hypopachus caprimimus Taylor Hypopachus caprimimus Taylor, Univ. Kansas Sci. Bull., 26:526, November 27, 1940.—Agua del Obispo, Guerrero, México. Buena Vista; Copuyo (6); Charapendo (3); Cofradia; Jaramillo; Jungapeo; San Salvador; Tuxpan. Specimens of Hypopachus from the Balsas drainage in Michoacan have characters consistent with topotypic H. caprimimus. Eleven specimens from the southern edge of the Mexican Plateau all have the flanks darker than the dorsum, a distinct and continuous dark stripe from the occiput to the groin, a large dark spot in the inguinal region, and a pair of dark transverse stripes on the thigh and shank AMPHIBIANS AND REPTILES OF MICHOACAN 53 (Pl. 6, Fig. 1). With the exception of three specimens from Chara- pendo, all have a predominantly brown venter with round, cream- colored spots. Peters (1954:8) referred specimens from Buena Vista and San Salvador to Hypopachus oxyrrhinus. He stated that the specimen (BMNH 1914.1.28.150) from San Salvador had flanks much darker than the dorsum and a well-defined continuous stripe from the occi- put to the groin; this specimen has the characters of H. caprimimus. The specimen (BMNH 1914.1.28.151) from Buena Vista resembles H. oxyrrhinus in some characters, but it is not like H. oxyrrhinus ovis on the Mexican Plateau in Michoacan. The specimen has paired transverse stripes on the hind limbs as does H. caprimimus, and is here referred to that species. In Michoacan this species has been collected in arid tropical scrub forest at elevations of 200 to 1800 meters in the northern foothills of the Sierra de Coalcoman, the Tepalcatepec and Tuxpan valleys, and on the lower slopes of the Cordillera Volcanica. Calling males have been found along streams. One specimen from Charapendo was regurgitated by a Leptodeira maculata. Hypopachus oxyrrhinus ovis Taylor Hypopachus ovis Taylor, Univ. Kansas Sci. Bull., 26:520, November 27, 1940.—Tepic, Nayarit, México. Hypopachus oxyrrhinus ovis, Shannon and Humphrey, Herpetologica, 14:89, July 23, 1958. Emiliano Zapata; 80 km. NW of Jacona (2); 10 km. NE of P&tzcuaro (2); Tangamandapio (16); 24 km. W of Zamora (16). Thirty-seven specimens from the Mexican Plateau in northwest- ern Michoacan agree well with the diagnosis of Hypopachus oxyr- rhinus ovis by Shannon and Humphrey (1958). With the excep- tion of one specimen from Tangamandapio, all have dark bellies extensively mottled or spotted with cream-color. Most of the specimens have some form of an irregular, usually broken, dark line from the occiput to the groin. In eight specimens there is no line or linear arrangement of spots; instead the dorsum is spotted or flecked with dark brown. The ground color of the dorsum and flanks varies from dull reddish brown to grayish brown; cream- colored spots are evident on the flanks and posterior surfaces of the thighs in all specimens (PI. 6, Fig. 2). In comparison with 14 specimens from Queseria, Colima (UMMZ 80001-2), individuals from the Mexican Plateau have a 54 UnIversITY OF Kansas Pusts., Mus. Nat. Hist. darker venter with bolder markings, and a more mottled dorsum. In Michoacan this species has been taken between 1500 and 2200 meters on the Mexican Plateau, where it inhabits mesquite-grass- land and cultivated areas. Rana dunni Zweifel Rana dunni Zweifel, Copeia, no. 2:78, July 15, 1957.—Lago de Patzcuaro, Michoacan, México. Lago de Patzcuaro (23); Rio de Morelia, near Undameo (8). Aside from the type series of this species, there are in the Museum of Zoology at the University of Michigan six specimens taken from “tanks” at the limnological station at Patzcuaro by Paul S. Martin in 1948, and eight specimens found in shaded ditches along the Rio de Morelia by Robert R. Miller on April 4, 1957. The Rio de Morelia flows into Lago de Cuitzeo; this drainage is separated from Lago de Patzcuaro by a chain of hills about 2400 meters in elevation. Dr. Richard G. Zweifel has examined these specimens and has informed me that, although they differ slightly from typical Rana dunni, they are much closer to that species than to Rana montezumae. Rana megapoda Taylor Rana megapoda Taylor, Univ. Kansas Sci. Bull., 28:310, November 12, 1942.—Chapala, Jalisco, México. La Palma (8). These specimens (USNM 118998-114005) are from the marshes along the southeastern shore of Lago de Chapala. Five females have snout-vent lengths of 124.0-188.1 (181.5), and one male has a snout-vent length of 110.2 mm. Two juveniles have snout- vent lengths of 49.7 and 56.3 mm. The coloration of the juveniles is more bold than that of the adults. The body proportions of these specimens agree with those presented by Zweifel (1957:80). Rana montezumae Baird Rana montezumae Baird, Proc. Acad. Nat. Sci. Philadelphia, 7:61, October 20, 1854.—Mexico City, Distrito Federal, México. La Palma; 8 km. NW of Maravatio (10); Sahuayo; Tupataro (7). This species probably is more abundant and widespread than is indicated by the few specimens listed above. It has been found only in the vicinity of permanent water on the Mexican Plateau and the mountains rising from the plateau at elevations of 1500 AMPHIBIANS AND REPTILES OF MICHOACAN 55 to 2000 meters. Its apparent absence from Lago de Patzcuaro cannot be explained, unless Rana dunni replaces it there. Rana pipiens Schreber Rana pipiens Schreber, Der Naturforscher, Halle, 18:185, 1782.—Raccoon, Gloucester County, New Jersey. Aguililla (2); Apatzingén (13); Arteaga; Axolotl (16); Camachines (33 Capirio; Cascada Tzararacua (3); Cerro San Andrés (6); Charapendo 3f Ciudad Hidalgo; Coalcomén (17); Cuitzeo (3); El Sabino (10); Jacona (8); 29 km. NW of Jacona (8); Jiquilpan; La Orilla (3); La Palma (5); La Playa (er Lago de Chapala (8); Lago de Patzcuaro (6); Lombardia; Los Conejos 67); Los Reyes (7); Macho de Agua; Maravatio; Morelia (5); Opopeo (3); Patzcuaro (9); 26 km. S of P&tzcuaro (52); Puerto Hondo (3); Rio Duero, 14 km. E of Zamora (18); Rfo Tepalcatepec, 27 km. S of Apatzingan (2); San Gregorio (38); San José de la Cumbre (5); Tangamandapio; Zacapu; 18 km. W of Zamora (35). Except on the Pacific lowlands, this species is abundant throughout the state. It has been collected from sea level to 2800 meters, the greatest altitudinal range of any amphibian in Michoacan. It has been found frequently in the Tepalcatepec Valley; it is not a dis- tinctly highland species in southern Michoacdn, as stated by Peters (1954:9). One specimen from Aguililla (UMMZ 119257) is an albino. In this specimen there is a faint pattern on the hind limbs; otherwise the entire body is creamy white; the eyes are pink. Rana pustulosa Boulenger Rana pustulosa Boulenger, Ann. Mag. Nat. Hist., ser. 5, 11:348, 1883.— Ventanas, Durango, México. Arteaga (4); 21 km. S of Arteaga; Cascada Tzararacua (3); Coalcomén (3 8 km. ENE of Dos Aguas (3); El Sabino (53); Los Reyes (3); Tzitzio (4 ruapan. Although Rana pustulosa seems to be absent from the Mexican Plateau in Michoacan, it has been collected at elevations of 850 to 2150 meters on the slopes of the Cordillera Volcdnico and in the Sierra de Coalcoman. Usually the frogs are found along rocky streams, but at Coalcoman they were found in a hyacinth-choked old river channel, and at El Sabino, in irrigation ditches. In most specimens the dorsum is dark olive-brown; in some it is pale olive-tan with dense dark brown mottling on the back and dark transverse bands on the hind limbs. Thirteen tadpoles (UMMZ 94271) taken from a seepage pool by a stream near Uruapan closely resemble the description of tadpoles of this species given by Taylor (1942b). . > 56 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. REPTILIA Testudines Chelonia mydas (Linnaeus) Testudo mydas Linnaeus, Systema naturae, ed. 10:197, 1758.—Type lo- cality restricted to Ascension Island by Mertens and Miiller (1928:23). Chelonia mydas, Brongniart, Bull. Sci. Soc. Philom., 2:89, 1800. Beach between Rio Motin and Rio Colotlan (2); Maruata; Playa Azul (4). Green sea turtles are abundant along the coast of Michoacan. Laying females and fresh nests were found on August 6-12, 1950, July 14-16, 1951, and July 8-10, 1955. The general account of sea turtles on the coast of Michoacan that was given by Peters (1957) is supplemented here by my field notes on the actions of one female observed on the night of July 14, 1951, near Maruata by Donald D. Brand and I. Because of a full moon, visibility was excellent. In the course of the day several Chelonia were seen in the surf; shortly after dark the first turtle was observed on the beach. Several were observed to come out on the beach and crawl nearly to the strand line, only to return to the sea. At 10:20 p.m. one turtle was seen about 15 meters from the water. We watched this turtle from some distance and observed that by 10:26 p.m. she had moved about ten meters to a bank of sand about two meters high. Ten minutes later she had climbed the bank and disappeared over the top into the brush. We moved closer and remained hidden below the bank. Although we could not see the turtle, we could hear her movements. Between 10:37 and 10:57 p.m. the turtle dug, often flipping the dry sand for a distance of about two meters. When this energetic digging ceased, we moved up the bank to see that she was facing inland and sitting in a depres- sion about one and one-half meters in diameter and 30 centimeters in depth. She had cleaned out this depression in the past 20 min- utes. Between 11:00 and 11:36 p. m. she dug the nest hole by first scooping sand with one hind flipper and then with the other; when sand was thrown by one flipper, there was a similar, but weaker, motion by the other flipper. At 11:36 p.m. she stopped digging. By crawling up behind the turtle we were able to examine the nest cavity, which measured 21 centimeters across the top and 38 centi- meters deep. The diameter of the bottom of the hole was estimated to be about 50 centimeters. At 11:40 p. m. she released the first egg; a minute later she dropped the second. At 11:42 p. m. the third and fourth eggs were released; these were coherent, as were the fifth and sixth eggs released at 11:43 p.m. After this, as many as three eggs AMPHIBIANS AND REPTILES OF MICHOACAN 57 were dropped at atime. After laying about 60 eggs, she paused for a minute and then continued laying. By 11:55 p.m. she had laid 98 eggs; after this, the process of deposition slowed considerably. She dropped a fragment of an egg followed by normal eggs. At midnight she deposited a miniature egg about 20 mm. in diameter. This terminated the deposition. Immediately she began to cover the nest. Within ten minutes after the last egg was deposited the nest had been covered. The turtle first had been seen at 10:20 p. m.; judging from its speed and its distance from the water, the turtle probably had been on land for about ten minutes. About 25 minutes were used in crawling from the water to the nesting site. One hour and 33 minutes were spent at the nesting site; of this time twenty minutes were taken for egg deposition. The turtle was not followed back to the water, but if the return trip took approximately the same amount of time as required to travel from the ocean to the nesting site, the total elapsed time from departure to return to the water was about two and one-half hours. We collected the eggs as they were deposited. There were 106 eggs, each having a diameter of about 40 mm., plus one small egg and a fragment of another. The turtle had a carapace about one meter in length. From our limited observations of sea turtles and their tracks on the beaches, and from the accounts of these animals by the resi- dents of the coastal region, great numbers of sea turtles use these relatively uninhabited beaches for nesting grounds. However, the turtles do not go unmolested. The natives capture turtles and col- lect their eggs. Opened and emptied nests also showed signs of predatory activity on the part of other mammals. In the vicinity of Playa Azul several turtles were killed by dogs. Kinosternon hirtipes hirtipes Wagler Cinosternon hirtipes Wagler, Naturl. Syst. Amph., p. 87, 1830.—México. Type locality restricted to Mazatlan, Sinaloa, México, by Smith and Taylor (1950b:25). Kinosternon hirtipes hirtipes, Schmidt, Check list N Amer. Amph. Rept., ed. 6, p. 89, 1953. Eight km. W of Ciudad Hidalgo; Jiquilpan; La Palma; Lago de Camécuaro (4); Lago de Cuitzeo (3); Lago de Patzcuaro (8); 14 km. E of Zamora (4). One specimen from eight kilometers west of Ciudad Hidalgo (UIMNH 24707) is from the Rio Tuxpan, a tributary of the Rio Balsas; this is the only record for the species from the Balsas drain- age. All others are from the lakes or rivers flowing into the lakes on the southern part of the Mexican Plateau. This species exists 58 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. in Lago de Patzcuaro to the apparent exclusion of the abundant and widespread Kinosternon integrum. Kinosternon integrum LeConte Kinosternon integrum LeConte, Proc. Acad. Nat. Sci. Philadelphia, 7:183, 1854.—México. Type locality restricted to Acapulco, Guerrero, México, by Smith and Taylor (1950b:25). Agua Cerca (3); Aguililla; Arteaga (8); Apatzing4n (7); Barranca de Herradero; Buenavista (20); Capirio (2); Charapendo (3); Chupio; Coa- huayana (2) Coalcoman (169); Copuyo (4); El Sabino (8); Jacona; Jiquilpan (12); La Orilla (2); La Playa (2); Lago de Cuitzeo (27); Las Higuertas; Lombardia (3); Los Reyes (5); Morelia; Ojos de Agua de San Telmo; San Pedro Naranjestila; Tacicuaro. Excepting Lago de Patzcuaro, Kinosternon integrum occupies all permanent and temporary ponds, lakes, and streams below 2200 meters throughout the state. At Coalcomaén the species was in roadside ditches, small puddles, flooded fields, a hyacinth-choked ox-bow of the Rio Coalcoman, as well as in the Rio Coalcoman and its tributaries. Specimens from Arteaga and Barranca de Herradero were found in clear rocky streams; the one from Las Higuertas was found in a small muddy pond in pine-oak forest. On August 26, 1960, James R. Dixon found a copulating pair in a pool at Capirio. The large series from Coalcoman contains juveniles and adults; these turtles formed the basis for the study of relative growth of plastral scutes in this species by Mosimann (1956). Geoemyda rubida perixantha Mosimann and Rabb Geoemyda rubida perixantha Mosimann and Rabb, Occ. Pap. Mus. Zool. Univ. Michigan, 548:1, November 9, 1953.—Eight kilometers south of Tecoman, Colima, México. Apatzing4én (2); Coahuayana; La Placita; Punta San Juan de Lima. These specimens have been discussed in detail by Mosimann and Rabb (19538). All are from the arid tropical scrub forest; those from the coastal regions were collected at elevations of less than 40 meters, and those from the Tepalcatepec Valley were col- lected at an elevation of 335 meters. Crocodilia Crocodylus acutus acutus Cuvier Crocodylus acutus Cuvier, Ann. Mus. Hist. Nat. Paris, 10:55, 1807.—Santo Domingo. Crocodylus acutus acutus, Miiller and Hellmich, Ibero-Amerik. Stud., 13: 128, 1940. Boca de Apiza (2); Playa Azul (2). The crocodile or “caiman” is abundant in the brackish lagoons along the cost of Michoacan; three large adults and several juveniles AMPHIBIANS AND REPTILES OF MICHOACAN 59 were observed at Estero Pichi at Playa Azul; others were seen at Mexiquillo and Maruata. Residents of the Balsas-Tepalcatepec Basin frequently have reported “caimanes” in the Rio Balsas and Rio Tepalcatepec, but the existence of the crocodile in these rivers has not been verified by specimens. Sauria Phyllodactylus duelimani Dixon Phyllodactylus duellmani Dixon, Southwest Nat., 5:37, April 15, 1960.— Rancho E] Espinal, Michoac4n, Mexico. Fourteen km. SSW of Apatzing4n; Capirio; Cafradia (3); El Espinal (3). This species is known only from the Tepalcatepec Valley, where it has been found in open arid situations from 180 to 500 meters. Specimens were found in the daytime in stumps, dead cacti, and the hollow branches of the legume, Apoplanesia paniculata. In life adults were pale gray or grayish tan above and creamy white below. A juvenile having a snout-vent length of 18 mm. had a pale orange tail with gray cross-bands. In the adults the tail was colored like the body. The specimen from 14 kilometers south-southwest of Apatzingan (KU 29764) and those from Cofradia (BMNH 1914.1.28.28-30) were not listed by Dixon (1960). Phyllodactylus homolepidurus Smith Phyllodactylus homolepidurus Smith, Univ. Kansas Sci. Bull., 22:121, No- vember 15, 1985.—Five miles southwest of Hermosillo, Sonora, México. El Ticuiz (2); La Placita; Ostula (2); Pémaro; San Pedro Naranjestila. These specimens have been referred to Phyllodactylus homolepi- durus by James R. Dixon (in litt.), who is currently studying the American members of the genus. Geckos of this species have been found in tropical semi-deciduous forest in the coastal lowlands to elevations of 500 meters. Most specimens were found beneath the bark of standing dead trees or stumps. Two individuals from El Ticuiz (UMMZ 115102) in life were dark gray above with brownish tubercles; the belly was a dusty cream-color. Apparently this species does not enter the Tepalcatepec Valley, where Phyllo- dactylus lanei is abundant. Phyllodactylus lanei Smith Biullodactyius lanei Smith, Univ. Kansas Sci. Bull., 22:125, November 15, 1985.—Tierra Colorado, Guerrero, México. Apatzingan (18); 21 km. S of Arteaga; El] Sabino (53); La Playa; Ostula ); Rio Marquez, 10 km. S of Lombardia (8); 16 km. N of Tafetan. This widespread species has been taken at elevations of less than 1100 meters in the Balsas-Tepalcatepec Basin, where it occurs in 5—7817 60 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. riparian situations in the foothills. Specimens have been collected in tropical semi-deciduous forest at Ostula and in oak forest south of Arteaga; both of these localities are on the Pacific slopes of the Sierra de Coalcoman, a region inhabited by Phyllodactylus homo- lepidurus. Both species have been collected at Ostula. A juvenile from 21 kilometers south of Arteaga (UMMZ 118933) had alternating black and white bands on the tail. In life most of the lizards are dull ashy gray or grayish tan above and white below. According to Dixon (in litt.), one specimen from Apatzingén (UMMZ 115102) resembles Phyllodactylus magnus in scutellation, but it lacks the distinctive yellow venter of that species. Apparently Phyllodactylus lanei is restricted to rather mesic en- vironments in the Balsas-Tepalcatepec Valley and surrounding foot- hills; in the more open arid environments on the floor of the valley it seems to be replaced by Phyllodactylus duellmani. Phyllodactylus paucituberculatus Dixon Phyllodactylus paucituberculatus Dixon, Southwest. Nat., 5:40, April 15, 1960.—Rio Cupatitzio (= Rio Marquez), 6.5 miles south of Lombardia, Michoacan, México. Rio Marquez, 10 km. S of Lombardia (6). Two of these specimens (UMMZ 112692-3) were discussed in de- tail by Dixon (1960:40) in his description of the species. On August 25, 1960, Dixon collected four additional specimens at the type locality, a conglomerate cliff along the Rio Marquez. These will be reported by him in his forthcoming study of the genus. Anolis dunni Smith Anolis dunni Smith, Copeia, no. 1:9, May 10, 1936.—Agua del Obispo, Guerrero, México. Arteaga (3); 19 km. S of Arteaga. Three females from Arteaga (UMMZ 119075) have snout-vent lengths of 41, 41, and 44mm. In life the pale grayish brown dorsum was marked with dark brown; the belly was white, and the throat was pale pink. All have a dark interorbital bar and dark vertical bars on the upper labials. In two specimens there are only scattered dark flecks on the dorsum; in the third there is a dark postorbital stripe, a dark lateral stripe, and four narrow transverse bands on the body. A male from 19 kilometers south of Arteaga (UMMZ 119076) having a snout-vent length of 49 mm. had in life a tan dorsum, a broad white stripe from the ear to the groin, scattered small white spots on the dorsum, and indistinct pale cream-colored spots on the posterior surfaces of the thighs. This male has the dark labial bars, AMPHIBIANS AND REPTILES OF MICHOACAN 61 but lacks the dark interorbital bar, found in the females. The large rose-pink throat fan extends to about the middle of the belly. In all of the specimens the middorsal scales are keeled and much smaller than the smooth pavementlike or slightly imbricate ventrals. All have two gulars in contact with the mental, five scales between the nasals, five scales (not including the first labials) in contact with the rostral, and four rows of loreals. In these characters these specimens agree well with Anolis dunni from Guerrero, as diagnosed by Davis (1954b). Previously Anolis dunni has been reported only from the vicinity of Agua del Obispo, Guerrero, a locality situated at an elevation of about 900 meters in pine-oak forest in the Sierra del Sur. All known close relatives of Anolis dunni occur only in Guerrero: A. taylori Smith and Spieler from Acapulco, A. gadowi Boulenger from Tierra Colorado, A. liogaster Boulenger, and A. omiltemanus Davis from Omiltemi. The present specimens from elevations of about 900 meters in riparian stream vegetation and oak forest represent the northern known limits of this group of Anolis. Anolis nebulosus (Wiegmann) Dactyloa nebulosa Wiegmann, Herpetologia Mexicana, p. 47, 1834.— México. Type locality restricted to Mazatlan, Sinaloa, México, by Smith and Taylor (1950b:66). Anolis nebulosus, Bocourt, Mission Scientifique au Mexique et dan |’Amer- ique Centrale. Reptiles, livr. 2:77, 1873. Acahuato (8); Agua Cerca; Apatzingan (4); Araparicuaro (3); 29 km. S of Ario de Rosales (3); 20 km. S of Arteaga (2); Barranca de Bejuco; Cascada Tzararacua (5); Cerro Tancitaro (13); Cheran; Chupio (5); Coalcoman (10); Cofradia; Dos Aguas (10); 18 km. E of Dos Aguas (3); El Diezmo; El Sabino (48); El Ticuiz; Jiquilpan (2); La Orilla; La Placita; La Playa (8); Los Conejos (2); Los Pozos; Nogueleras (2); Ostula; 8 km. W of Patzcuaro (2); 8 km. NE of P&tzcuaro; Playa Azul (3); Rio Cachan; Rio Marquez, 10 km. S of Lombardia; Rio Tepalcatepec, 27 km. S of Apat- zingan; San Juan de Lima (6); San Pedro Naranjestila; Temazcal; Tuxpan (2); Tzitzio; Uruapan (74); 11 km. N of Uruapan (2); Volcan Jorullo; 16 km. E of Zacapu (2); 18 km. W of Zamora; Ziracuaretiro. Even with the abundance of material the assignment of a specific name to these anoles is only tentative, for definite determination between Anolis nebulosus Wiegmann and A. nebuloides Bocourt is uncertain. Bocourt (1873:75) distinguished A. nebuloides from A. nebulosus by the following characters: (1) head scales keeled, not smooth; (2) snout narrower; (3) ear opening larger; (4) supraorbital semicircles separated by a row of small scales and not in contact; (5) dorsal scales larger and subequal in size to the belly scales. Boulenger (1885:77) used the same characters; Smith and Taylor (1950b:58) in their key to the Mexican species 62 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. of Anolis stated that the dorsal scales are slightly smaller than the ventrals in A. nebulosus and markedly smaller in A. nebuloides. Smith (in litt.) stated that the characters of the relative sizes of the dorsal and ventral scales were incorrect in that key. The application of the above criteria to specimens from Micho- acan has not resulted in the recognition of two species. The ma- jority of the specimens have the supraorbital semicircles separated by at least one small scale; the head scales, with the exception of those on the snout in a few individuals, are smooth; the dorsal scales are only slightly smaller than the ventrals. In other char- acters of scutellation the specimens are highly variable. The males in life have an orange throat fan. Anoles of this kind have been found in Michoacan, Colima, Jalisco, Nayarit, and southern Sinaloa. Near Oaxaca, Oaxaca, specimens were collected that superficially resemble those from Michoacan and farther north. These have low keels on the snout scales, dorsals somewhat larger than the ventrals, and a pink throat fan. In ten males from Oaxaca the size of the dorsal scales relative to that of the ventrals is 1.00: 0.88; the same ratio for 25 males from Michoacan is 1.00: 1.08. In both samples there are specimens in which the dorsal and ventral scaies are about equal in size. Investigations by Richard E. Etheridge on the osteology of Anolis, including those species here being considered, have revealed rela- tively constant differences in the parasternalia and in the caudal vertebrae. The application of Etheridge’s findings to anoline sys- tematics must await the completion of his study. The carination of the scales on the snout versus smooth scales there seems to be the only significant character given by Bocourt that dis- tinguishes A. Nebuloides from A. nebulosus. The difference in the color of the throat fan, which is apparent only in living individuals, is more striking. Obviously more than one species is represented, as is borne out by the differences in the color of the throat fan and in the osteology, but there is uncertainty about the correct name for each species. On the strength of Bocourt’s diagnosis of keeled snout scales in A. nebuloides, I am applying that name to the population in Oaxaca and A. nebulosus to the specimens from Michoacan. As arranged here, the two species can be distinguished, as follows: A. nebulosus.—Dorsal scales only slightly smaller than the ventral scales; snout scales usually smooth; throat-fan bright orange in adult males. A. nebuloides.—Dorsal scales somewhat larger than the ventral scales; snout scales having a low keel; throat-fan pink in adult males. AMPHIBIANS AND REPTILES OF MICHOACAN 63 With respect to geographic distribution, A. nebulosus has been collected from southern Sinaloa southward to Michoacan. The lizards here referred to A. nebuloides have been taken only in pine- oak forest on the mountain slopes near Oaxaca City. Zweifel and Norris (1955:283) reported anoles with pink throat-fans from south- ern Sonora; possibly those specimens are A. nebuloides; I have not examined them. I have seen several preserved specimens from the vicinity of Tehuantepec, Oaxaca. Although they probably belong to this group, those specimens differ from both A. nebulosus and A. nebuloides in their larger size, relatively larger head, and much larger throat fan. Aside from the minor variation in scutellation, specimens of Anolis nebulosus from Michoacan vary greatly in coloration. Usually the females have some form of a broad middorsal pale-colored band. In life this is dull yellow, tan, or orange. Two females from Dos Aguas are strikingly different; one (UMMZ 119521) has a broad middorsal orange stripe that is scalloped laterally and bordered by gray. The other (UMMZ 119081) has a narrow middorsal cream- colored line. Males usually are unicolor brown or olive-tan; some- times the middorsal region is darker. Some individuals have dark cross-bands or chevrons on the dorsum. One male from Dos Aguas (UMMZ 119080) has a cream-colored lateral stripe. In Michoacan Anolis nebulosus occurs from sea level to elevations slightly in excess of 2100 meters, usually in areas of dense cover, whether this be herbaceous, viney, or woody, ordinarily on the ground as well as in bushes and trees. One was in a bromeliad grow- ing about ten meters above the ground. In the arid Tepalcatepec Valley anoles of this species are most frequently found in the tangled growth along streams. Above Uruapan they were found in pine-oak forest, and on the Mexican Plateau between Zamora and Zacapu they were found in a bunch grass-scrub oak association. Anolis schmidti Smith Anolis schmidti Smith, Publ. Field Mus. Nat. Hist., zool. ser., 24:21, January 80, 1939.—Manzanillo, Colima, México. wee Placita; San Juan de Lima. Peters (1954:11) reported on the specimen from La Placita; an- other was secured at San Juan de Lima in 1956. The latter (UMMZ 115078) is a male having a snout-vent length of 43.0 mm. and a tail length of 70.5 mm. The dorsal ground color is pale tan; there are five pairs of irregular dark brown dorsolateral blotches. In life the throat fan was pale orange. These specimens agree with those from 64 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Colima described by Duellman (1958c:10). The distribution of Anolis schmidti seems to be restricted to the coastal lowlands from Michoacan to Nayarit. Basiliscus vittatus Wiegmann Basiliscus vittatus Wiegmann, Isis von Oken, 21:373, 1828.—México. Type rosa a to Veracruz, Veracruz, México, by Smith and Taylor 1950b:72). Apatzingin (9); Capirio; Coahuayana (5); El Cerrito; El Sabino (2); El Ticuiz; La Placita (3); Maruata (2); Motin del Oro; Ostula; Playa Azul (3). This species has been found only on the coast and in the low Tepalcatepec Valley. In the latter area it is restricted to riparian situations along the larger streams. The lizard is abundant in the mangrove swamps bordering the brackish lagoons on the coast. In July, 1955, scores of individuals were seen around Estero Pichi at Playa Azul. Adults, especially the large males, are exceedingly wary and difficult to collect. At all localities where they were found, the lizards were most often seen in dense bushes, where they are well camouflaged. Individuals of all sizes were observed to run across the surface of the ponds. Iguana iguana rhinolopha Wiegmann Iguana rhinolopha Wiegmann, Herpetologia Mexicana, p. 44, 1834.—México. Type locality restricted to Cérdoba, Veracruz, México, by Smith and Tay- lor (1950b:72). Iguana iguana rhinolopha, Van Denburgh, Proc. Acad. Nat. Sci. Philadelphia, 1897:461, January 18, 1898. Apatzingan (8); Capirio (3); El Cerrito; El Ticuiz (2); La Placita; La Playa (2); Maruata; Playa Azul; Rio Cachan. Like the preceding species, this lizard is always found near water. It does not ascend the foothills of the Sierra de Coalcoman, but in the Balsas Basin it reaches elevations of 800 meters at La Playa. Large adults are often seen in the large trees making up the gallery forests along rivers. From high perches the lizards drop into the water with a terrific splash. Bright green juveniles were abundant in bushes along the Rio Tepalcatepec in July, 1955. Ctenosaura pectinata (Wiegmann) Cyclura pectinata Wiegmann, Herpetologia Mexicana, p. 42, 1834.—México (by inference). Type locality restricted to Colima, Colima, México, by Bailey (1928:25). Ctenosaura pectinata, Gray, Catalogue of the lizards . . . British Mu- seum, p. 191, 1845. Apatzingan (27); between Ario de Rosales and La Playa; Barranca de Bejuco; Capirio (2); Coalecomén (4); El Espinal; El Sabino (2); El Ticuiz; Jazmin (2); La Huacana; La Placita (8); LaPlaya (8); Limoncito; Lombardia; Motin del Oro; Playa Azul; Rio Cancita, 12 km. E of Apatzingan (2); Rio Marquez, 10 km. S of Lombardia (2); ? Uruapan; Volcan Jorullo. AMPHIBIANS AND REPTILES OF MICHOACAN 65 Ctenosaura pectinata is a common lowland species that ascends the slopes of the Sierra de Coalcoman and the Cordillera Volcdnica to elevations of about 1050 meters (approximating the lower limits of the oak forest). The record from Uruapan (USNM 10234, col- lected by Dugés) is doubtful. These large lizards are most easily observed on rock fences along roads. Near Apatzingin innumerable individuals can be seen in mid-morning. Later in the day, as the sun rises higher in the sky, the lizards retreat to the shade of the crevices in the fences. The abundance of these lizards in the Tepalcatepec Valley, together with evidence gathered from the natives of the valley, indicates that these lizards are seldom used for human consumption there. On the other hand, several people in Coalcoman consider the “iguana negra” (local name for Ctenosaura) to be a delicacy and serve it at every opportunity. In early July, 1951, brilliant green young of the year were collected at La Playa and at Coalcoman. Enyaliosaurus clarki (Bailey) Ctenosaura clarki Bailey, Proc. U.S. Natl. Mus., 73:44, September 26, 1928. —Ovopeo (= Oropeo), Michoacan, México. Enyaliosaurus clarki, Duellman and Duellman, Occ. Pap. Mus. Zool. Univ. Michigan, 598:1, February 16, 1959. Twelve km. SSW of Apatzingan; Capirio (7); Cofradia (3); El Espinal (2); 32 km. E of Huetamo; Jazmin (5); Oropeo (10); Rancho Nuevo; Rio Can- cita, 12 km. E of Apatzingan (8); Tepalcatepec (3); Zicuiran (6). This species is known only from the low areas of the Balsas-Te- palcatepec Basin between elevations of 200 and 510 meters. It is commonly found in the open arid tropical scrub forest dominated by Prosopsis sp., Apoplanesia paniculata, and Cercidium plurifoliola- tum. Continued collecting in the Tepalcatepec Valley has borne out the suggestions of Duellman and Duellman (1959) concerning the distribution and abundance of this lizard. Also, continued collect- ing in Colima and on the Pacific coast has failed to reveal the pres- ence of Enyaliosaurus there. Phrynosoma asio Cope Phrynosoma asio Cope, Proc. Acad. Nat. Sci. Philadelphia, 16:178, Septem- ber 30, 1864.—Colima, Colima, México. Apatzingan (4); San Salvador. In Michoacan this species has been obtained only in the Tepal- catepec Valley and on the northern slopes of the Sierra de Coal- coman between 300 and 700 meters. Apparently the lizard is ab- sent from the coastal lowlands of Michoacan and Guerrero. The distribution of this species, therefore, is discontinuous. One popu- 66 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. lation inhabits the lowlands of Colima and the Balsas-Tepalcatepec Basin inland to northern Guerrero and Morelos; a southern popula- tion inhabits the Plains of Tehuantepec in Oaxaca. A juvenile from Apatzing4n (USNM 47789) has a snout-vent length of 40.0 mm. and a tail length of 19.5 mm. Sceloporus aeneus aeneus Wiegmann Sceloporus aeneus Wiegmann, Isis von Oken, 21:370, 1828.—México. Type locality restricted to Tres Cumbres, Morelos, México, by Smith and Taylor (1950b:187). Sceloporus aeneus aeneus, Smith, Occ. Pap. Mus. Zool. Univ. Michigan, 361:6, December 15, 1937. Angahuan; Araparicuaro (2); Capacuaro (2); Carapan (11); Cherdn (11); 18 km. WNW of Ciudad Hidalgo (10); Cusefio Station; Jera4huaro; Los Conejos (386); Macho de Agua (7); Opopeo; Paracho (2); Patzcuaro Tah Pino Gordo; 18 km. W of Quiroga (2); Tancitaro (49); Uruapan (14); 16 km. NW of Zacapu (5); between Zacapu and Zamora (2); 13 km. E of Zinapécuaro; 14 km. SE of Zitacuaro (14). This smal] terrestrial species inhabits the pine and fir forests of the Cordillera Volcanica between elevations of 1850 and 3100 meters; apparently it is absent from the Sierra de Coalcoman. It seems to prefer rather open coniferous forests in which there is a more or less continuous cover of grasses on the ground. On warm sunny days the lizards can be observed scurrying about in the grass; in the early hours of the day, or on cold days, they are found beneath stones, logs, or dead clumps of bunch grass. Sceloporus asper Boulenger Sceloporus asper Boulenger, Proc. Zool. Soc. London, 1897:497, October, 1897.—La Cumbre de los Arrastrados, Jalisco, México. Apatzingan (3); 10 km. E of Dos Aguas; Uruapan (41). This strictly arboreal lizard is abundant in the mixed broad- leafed forest near Uruapan. The lizards are exceedingly wary and can be approached only with difficulty. In life males have pale blue bellies; the throat is pale pink. The pale gray dorsum marked with irregular darker gray blotches blends well with the color of the tree trunks on which the lizard lives. The one specimen from Dos Aguas was found on a pine tree; it provides the only record for the species from the Sierra de Coalcoman. Sceloporus bulleri Boulenger Sceloporus bulleri Boulenger, Proc. Zool. Soc. London, 1894:729, April, 1895.—Las Cumbre de los Arrastrados, Jalisco, México. Acuaro de las Lleguas (18); Barolosa (9); Dos Aguas (61); 10 km. NE of Dos Aguas (5). Heretofore this species has been known only from a few speci- mens from scattered localities in the Sierra Madre Occidental in AMPHIBIANS AND REPTILES OF MICHOACAN 67 southwestern Jalisco and Sinaloa. The collection of a large series of these lizards in virgin pine forest at elevations of more than 2000 meters in the Sierra de Coalcoman now makes possible an analysis of variation in the species. Superficially S. bulleri resembles S. torquatus, but S. bulleri is smaller, has more dorsal scales, fewer scales in the dark collar, and fewer femoral pores. In 88 specimens of S. bulleri there are 36-41 (38.7) dorsal scales and 2 or 3 (2.6) middorsal scales in the collar, as compared with 28-31 (29.3) dorsal scales and 3 or 4 (8.4) mid- dorsal scales in the collar of 26 specimens of S. torquatus from Uruapan. In 20 adult males of S. bulleri there are 18-15 (14.3) femoral pores, and 18-16 (14.4) in 11 females; 13 males of S. tor- quatus have 14-21 (17.3) femoral pores, and 13 females have 15-21 (16.7). Seventeen adult males of S. bulleri have snout-vent lengths of 72-91 (82.0); ten females, 71-87 (75.7). In comparison, 18 adult males of S. torquatus have an average snout-vent length of 88.9 mm., and 18 females, 88.5 mm. In S. bulleri there is little variation in the head scales. The frontal is in contact with the interparietal in 63, and not in 24, specimens; the median frontonasal is in con- tact with the frontal in 13, and not in 74, specimens. In 39 speci- mens there are two canthals, and in 48 there is one; in 29 speci- mens there are three preauriculars, and in 58 there are four. In life adult males have a pale blue tail, bright blue belly patches, a purplish blue throat, and pale blue lines on the sides of the head and neck. This species was obtained at four localities in the high moun- tains of the Sierra de Coalcoman. In this mountain range Scelo- porus bulleri apparently replaces S. torquatus, a species that is widespread in the Cordillera Volcanica and on the Mexican Plateau. At Dos Aguas and at Acuaro de las Lleguas the lizards were abun- dant in the tall pine forest, where they were found on standing pine trees, on pine logs, and on rock outcroppings. Sceloporus dugesi intermedius Dugés Sceloporus intermedius Dugés, La Naturaleza, 4:29, 1877.—La Noria, near Zamora, Michoacan, México. Sceloporus dugesii intermedius, Smith, Univ. Kansas Sci. Bull., 24:663, February 16, 1938. Cojumatlan (6); Jiquilpan (11); Lago de Camécuaro; Lago de Chapala; Morelia (23); Patzcuaro (84); Quiroga (85); Sahuayo (4); Tacicuaro (2); Tangamandapio (17); Tangancicuaro (9); Zacapu (4); Zamora (11); Zinapécuaro (9). This lizard is strictly an inhabitant of the Mexican Plateau, where it is found in rocky places, sometimes in pine-oak forest, 68 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist. but more frequently in mesquite-grassland. It is a terrestrial species, and is most often seen on rock fences at elevations of 1500 to 2200 meters. This species differs from S. bulleri and S. torquatus in having two rows of supraoculars, instead of one; also it has more dorsal scales. Twenty-six specimens of Sceloporus dugesi intermedius from Tan- gamandapio and Tangancicuaro have 44-48 (45.7) dorsal scales, as compared with an average of 38.7 in S. bulleri and 29.3 in S. torquatus. In life Sceloporus dugesi intermedius has a dull green- ish gray dorsum; in males the belly patches are bright blue bor- dered medially by black, and the throat is bluish gray. The larg- est specimen examined is a male having a snout-vent length of 80 mm. Sceloporus gadowae Boulenger Sceloporus gadoviae Boulenger, Proc. Zool. Soc. London, 1905, 2:246, Octo- ber 7, 1905.—Mezquititlan, Guerrero, México. Chupio; El Sabino (77); La Playa (6); Rio Marquez, 10 km. S of Lom- bardia (11). Although this species has a rather extensive range in the Balsas- Tepalcatepec Basin in the state of Michoacan, Guerrero, Morelos, and Puebla, it is only locally abundant in that area. Usually these lizards are found on rocky cliffs in which there are many crevices for cover. Sceloporus gadowae is abundant on a conglomerate cliff along the Rio Marquez south of Lombardia. Although the closely related S. pyrocephalus is abundant in the stream valley and in the hills above the cliff, S. gadowae has been found only on the cliff; few individuals of S. pyrocephalus have been observed on the cliff. A similar situation was discovered on a much more extensive con- glomerate cliff along the Rio Balsas near Mexcala, Guerrero. Near Tehuitzingo, Puebla, where S. pyrocephalus was not found, S. gado- wae was found on conglomerate cliffs. Probably there is strong competition between the two species; possibly this has resulted in the restriction of S. gadowae to isolated cliff-habitats within the extensive range of the more widespread S. pyrocephalus. In Michoacan Sceloporus gadowae has been found along the lower slopes of the Cordillera Volcdnica at elevations from 250 to 1050 meters. All of the localities from which this lizard is known lie in the arid tropical scrub forest. AMPHIBIANS AND REPTILES OF MICHOACAN 69 Sceloporus grammicus microlepidotus Wiegmann Sceloporus microlepidotus Wiegmann, Herpetologia Mexicana, p. 51, 1834.— México. Type locality restricted to México, Distrito Federal, by Smith and Taylor (1950b:120). Sceloporus grammicus microlepidotus, Smith and Laufe, Trans. Kansas Acad. Sci., 48:332, December, 1945. Angahuari; Apo (10); Atzimba (3); Carapan (5); Cerro San Andrés (17); Cerro Tancitaro (18); Corupu; Cusefio Station (2); Jacona; Jerahuaro (10); Macho de Agua; Mil Cumbres; 46 km. E of Morelia; 60 km. E of Morelia (2); Opopeo (14); Patzcuaro (30); Puerto Hondo (19); San Geeeario (41 }; San José de ja Cumbre (8); Sierra Patamba; Tancitaro (233); pataro; Undameo; Uruapan (180); between Zacapu and Zamora; 24 ae SE of Zitacuaro; between Zurumbeneo and Cerro Garnica. This small species of Sceloporus is an ubiquitous inhabitant of the coniferous forests from 1550 to 8100 meters in the Cordillera Vol- canica. Usually it is seen on tree trunks, but occasionally on the ground. Near the lower limit of the altitudinal distribution of the species, as at Uruapan, individuals sometimes are found on broad- leafed trees. Apparently Sceloporus heterolepis replaces S. grammi- cus microlepidotus in the Sierra de Coalcoman. Sceloporus heterolepis Boulenger Sceloporus heterolepis Boulenger, Proc. Zool. Soc. London, 1894:731, April, 1895.—La Cumbre de los Arrastrados, Jalisco, México. Araparicuaro; Cerro Barolosa (6); Dos Aguas (13); Los Conejos; 11 km. N of Uruapan (8). Although Michoacan has not previously been included in the range of this lizard, it was first collected in the state by Gadow in 1908 (BMNH 1914.1.28.69 from Araparicuaro). The description of S. heterolepis given by Smith (1939:197) can be supplemented by data on the 23 specimens now in the collections of the Museum of Zoology at the University of Michigan. All have two canthals; there are 55 to 71 (63.6) scales in the middorsal row; 1 to 3 rows middorsally are somewhat enlarged and bordered on either side by a row of larger scales bearing high keels. There are 14 to 20 (16.2) femoral pores. Eight adult males have snout-vent lengths from 49 to 61 (58.0) mm. and tail lengths from 57 to 74 (66.0) mm.; four adult females have snout-vent lengths from 52 to 57 (55.2) mm. and tail lengths from 60 to 66 (63.5) mm. The smallest of eight juveniles has a snout-vent length of 28 mm. and a tail length of 82 mm. The dorsum in adults is pale grayish brown; there are three irregular chevron-shaped dark marks and a triangular dark brown mark above the insertion of the hind limbs; on the tail are dark brown rings. There are scattered faint blue flecks on the flanks and narrow transverse dark lines on the lower limbs. Males 70 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. have pale bluish green belly patches and an orange-salmon-colored throat; the belly in females is pale orange-tan. The juveniles have a more contrasting color pattern; the dark chevrons on the dorsum are bordered posteriorly by pale gray. In Michoacan this species has been obtained in pine and pine-fir forests from 1800 to 2700 meters. On Cerro Barolosa and at Dos Aguas, both in the Sierra de Coalcoman, the lizards were found beneath the bark of dead, standing pines. In the Sierra de Coal- coman Sceloporus heterolepis seems to fill the niche of the small arboreal Sceloporus in the coniferous forest in southwestern México, a position held by S. grammicus microlepidotus in the Cordillera Volcanica; the latter species does not occur in the Sierra de Coal- coman. Five specimens of Sceloporus heterolepis are known from the Cordillera Volcanica, whereas 603 of S. grammicus microlepi- dotus have been collected there. The ecological relationships that exist between the two species in the Cordillera Volcanica are not known. Insofar as is known, Sceloporus heterolepis reaches the southern limits of its range in the Sierra de Coalcoman and in the western part of the Cordillera Volcanica. Other records for the species are from the Sierra Madre Occidental in Jalisco. Langebartel (1959) described Sceloporus shannonorum from the mountains near the Durango-Sinaloa border; the single specimen of S. shannonorum differs significantly from S. heterolepis only in having fewer dorsal scales (48). The acquisition of additional material, especially from Nayarit and northern Jalisco, probably will provide a basis for showing that these two populations are conspecific. Sceloporus horridus oligoporus Cope Sceloporus oligoporus Cope, Proc. Acad. Nat. Sci. Philadelphia, 16:177, Sep- tember 30, 1864.—Colima, Colima, México. Sceloporus horridus oligoporus, Taylor, Univ. Kansas Sci. Bull., 24:520, Feb- ruary 16, 1988. Charape Apatzingan (50); Arteaga (2); Capirio (2); Cascada Tzararacua; Conevende (4); Coahuayana (8); Coalcomin (32) 19 km. S of Corralito; km. E of Dos Aguas; El Sabino (55); El Ticuiz; Huetamo (2); Jazmin pacer (2); La Orilla (2); La Placita; pinotnS (3); Playa Azul ( “(5), Tzitzio (8); Uruapan (4); Volcan Jorullo (2); Ziracuaretiro; Zirimicuaro All of the specimens from Michoacan seem to be typical S. hor- ridus oligoporus; none has more than six femoral pores. Characteristically this species is found in open arid scrub forest; it reaches its greatest abundance in rocky areas in which there are scattered leguminous trees and bushes. It has been found in these AMPHIBIANS AND REPTILES OF MICHOACAN 71 low trees and bushes almost as frequently as it has been found on the ground; none has been seen in large trees or far above the ground. Altitudinally, this species ranges from sea level to about 1600 meters. Sceloporus melanorhinus calligaster Smith Sceloporus melanorhinus calligaster Smith, Proc. U.S. Natl. Mus., 92:360, November 5, 1942.—Acapulco, Guerrero, México. Aguililla; Apatzing4n (18); Barranca de Herradero; Capirio (19); Coahuay- ana (4); Coalcoman (2); Cofradia (4); El Cerrito; El Sabino (38); El Ticuiz (8); La Placita (6); Lombardia (4); Playa Azul; Rio Marquez, 10 km. S of Lombardia (2); Rio Marquez, 13 km. SE of Nueva Italia (4); Salitre de Estopila; San Juan de Lima (2); Santa Ana; Tzitzio; Ziracuaretiro. Smith (1942a:360) diagnosed Sceloporus melanorhinus calligaster as having fewer femoral pores than the other subspecies of S. mel- anorhinus and as having the lateral belly patches in the males con- fluent in the midline. Examination of forty specimens from the Tepalcatepec Valley and the coastal regions of Michoacan does not substantiate this diagnosis. The number of femoral pores varies from 15 to 22 (18.9); 14 individuals (85%) had 20 or more femoral pores. Smith (loc. cit.) stated that S. melanorhinus in Oaxaca had 18 to 27 (21.6) femoral pores and that 77 per cent of the specimens had more than 20 femoral pores. Of the 24 males examined from Michoacan, 18 have the lateral belly patches separated in the mid- line. Usually they are separated by no more than one scale, but in some individuals they are separated by two or more scales. Although the above data minimize certain differences between the northern and southern populations of this species, certain of the color pattern characters seem to be diagnostic of the subspecies inhabiting the Pacific lowlands from Guerrero to Nayarit. Large adults of S. m. calligaster have only a faint dorsal pattern, which in the subspecies melanorhinus and stuarti consists of a series of large, dark, inter- connected triangles on the back. This pattern is present in young and small adults of S. m. calligaster; furthermore, in this subspecies the ventral coloration of the males differs from that found in the more southern populations. Adult males of S. m. calligaster have a black throat, that changes to brilliant blue posteriorly, and a large white spot medially on the chin. This spot is present in some speci- mens from Oaxaca and Chiapas, but, if present, it is much smaller and less distinct than in specimens from Michoacan. In S. m. calli- gaster the chest and midventral area are orange to salmon-color. A male from Lombardia in life was colored as follows: Dorsum grayish tan bearing faint bluish gray flecks; chest deep salmon- 72 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist. orange, this color continuing down midventral area to the somewhat paler groin; belly patches pale blue fading to pale green laterally; throat black anteriorly enclosing a white spot; throat blue posteriorly and bluish green posterolaterally. Individual lizards were observed to change in dorsal color from a pale ashy gray to a rather dull brown. Normally, inactive indi- viduals and those observed on overcast days were dull brown. Sceloporus melanorhinus calligaster is found in trees in riparian situations in the lowlands to elevations of about 1500 meters. It does not inhabit the arid tropical scrub forest in the Tepalcatepec Valley or on the coast, but in those areas is found in the gallery forests along streams and rivers. The lizards are wary and live high in the trees; they are especially difficult to locate in the rainy season, when the trees are in full leaf. Sceloporus pyrocephalus Cope Sceloporus pyrocephalus Cope, Proc. Acad. Nat. Sci. Philadelphia, 16:177, September 80, 1864.—Colima, Colima, México. Acahuato (2); Apatzingan (142); Arteaga (4); 26 km. S of Arteaga (4); Capirio (6); Chinapa; Chupio; 19 km. S of Corralito (5); El Sabino (220); Jazmin (3); La Placita (8); La Playa (14); La Salada (6); Lombardia (5); Nueva Italia (14); Ojos de Agua de San Telmo (2); Oropeo (3); Ostula; Punta de San Telmo (8); Rio Cancita, 14 km. E of Apatzingan (13); Rio Marquez, 10 km. S of Lombardia (10); Rio Marquez, 13 km. SE of Nueva Italia (3); San Juan de Lima (2); Santa Ana (2); Tafetan (2); Tepalcatepec (2); Tzitzio (6); Volcan Jorullo (3). This smal] species is extremely common in the Tepalcatepec Valley and noticeably less so on the coast. It is usually found on the ground in rocky areas, but males frequently have been seen on the trunks of low trees in the scrub forest. Altitudinally, it ranges from sea level to slightly more than 1000 meters. The sexes are readily distinguished in the field (Oliver, 1937; Smith, 1939; Duellman, 1954b). In the dry season only males were ob- served in the Tepalcatepec Valley, but in the rainy season both sexes were found in approximately the same numbers. Sceloporus scalaris scalaris Wiegmann Sceloporus scalaris Wiegmann, Isis von Oken, 21:370, 1828.—México. Type locality restricted to México, Distrito Federal, by Smith and Taylor (1950b:137 ). Sceloporus scalaris scalaris, Smith, Occ. Pap. Mus. Zool. Univ. Michigan, 861:2, December 15, 1937. Carapan (2); Cheran; Ciudad Hidalgo; Huingo (3); Jacona (8); Jiquilpan (2); Lago de Camécuaro (2); Lago de Chapala; Lago de Cuitzeo (5); Morelia (4); Patzcuaro (4); Queréndaro; Quiroga; Tacicuaro (5); Tarécu- aro; Zacapu (4); Zamora (4); Zinapécuaro (11). This small terrestrial species does not seem to be abundant any- where in the state. It sometimes is found in open pine, oak, or AMPHIBIANS AND REPTILES OF MICHOACAN 73 pine-oak forest, but usually it is cbserved in areas supporting bunch grass. In such places the lizards sun and forage on the open ground and quickly take refuge in the large clumps of grass. Altitudinally, the species ranges from 1550 to 2300 meters. Although Sceloporus scalaris scalaris has been found in association with S. dugesi inter- medius, S. spinosus, and S. torquatus, it does not seem to form any close ecological association with any of these species. In the pine forests of the Cordillera Volcanica S. s. scalaris is replaced by Sceloporus aeneus aeneus, another small terrestrial species that occurs in great abundance throughout the coniferous forests of the Cordillera Volcanica. Sceloporus siniferus siniferus Cope Sceloporus siniferus Cope, Proc. Amer. Philos. Soc., 11:159, 1869.—Pacific side of the Isthmus of Tehuantepec. Type locality restricted to Tehuan- tepec, Oaxaca, México, by Smith and Taylor (1950b:134). Sceloporus siniferus siniferus, Smith and Taylor, Bull. U. S. Natl. Mus., 199:134, October 26, 1950. Twenty-six km. S of Arteaga; Barranca de Bejuco (2); Coahuayana; El Ticuiz (2); La Mira; La Orilla (2); La Placita (9); Maruata; Ojos de Agua de San Telmo; Ostula (4); Playa Azul (6); Pémaro (2); Puerto de las Higuerita; Santa Ana (3). This small terrestrial species inhabits the dense arid tropical scrub forest on the coast and lower foothills of the Sierra de Coalecoman to elevations of about 150 meters. It also occurs in the lower Balsas Valley, but it has not been found in the scrub forest of the broad Tepalcatepec Valley. Perhaps the large number of Sceloporus sini- ferus on the coastal lowlands is responsible for the small number there of S. pyrocephalus, another terrestrial species of about the same size. The latter is abundant in the Tepalcatepec Valley, where S. siniferus siniferus has not been found. Sceloporus siniferus siniferus is a fast runner and difficult to collect; consequently, the small number of specimens available is not indicative of its abun- dance. Sceloporus spinosus spinosus Wiegmann Sceloporus spinosus Wiegmann, Isis von Oken, 21:370, 1828.—México. Type locality restricted to Puebla, Puebla, México, by Smith and Taylor (1950b:116). Sceloporus spinosus spinosus, Martin del Campo, Anal. Inst. Biol. México, 8:262, 1987. Cojumatlan (2); Huetamo Road; Lago de Cuitzeo (4); Maravatio (8); Tupataro (2). Although this species is widespread on the southern part of the Mexican Plateau, it is uncommon in Michoacan. It has been col- 74 UNIVERSITY OF KANsAs PuBLs., Mus. Nat. Hist. lected only in rather open situations in the mesquite-grassland on the plateau between 1500 and 2800 meters, where it has been found in association with Sceloporus dugesi intermedius and S. scalaris scalaris. Most specimens of Sceloporus spinosus spinosus have been observed on rock fences. In this habitat the species is the larger member of a pair of species, the smaller of which is Sceloporus dugesi intermedius. Sceloporus torquatus torquatus Wiegmann Sceloporus torquatus Wiegmann, Isis von Oken, 21:369, 1828.—México. Type locality restricted to México, Distrito Federal, by Smith and Taylor (1950b:126). Sceloporus torquatus torquatus, Cope, Proc. Amer. Philos. Soc., 22:402, Angahuan (31); Araparicuaro; Capdcuaro (3); Carapan (11); Cerro Tan- citaro; Cheran; Ciudad Hidalgo; Cojumatlan; Copandaro (2); Corupu (4); Cuseno Station (9); El Alamo; Jacona (6); Jiquilpan (2); Jungapeo He Lago de Camécuaro; Lago de Chapala; Lago de Cuitzeo (3); La Palma (2); Los Conejos (8); Los Reyes (3); Maravatio (9); Morelia (17); Paracho (3); Patzcuaro (27); Pino Gordo; Queréndaro (2); Quiroga; Sahuayo (3); San José de la Cumbre; San Juan de Panangaricutiro; Tacicuaro (10); Tan- citaro (200); Tangamandapio; Tangancicuaro (3); Temazcal (2); Tupataro (5); Uruapan (186); Zacapu; Zinapécuaro (10); Zirimicuaro (12); Zita- cuaro. This large species inhabits the Mexican Plateau and the Cordil- lera Volcanica, but not the Sierra de Coalcoman, where apparently it is replaced by Sceloporus bulleri. Sceloporus torquatus tor- quatus usually is found in pine or pine-fir forests at elevations be- tween 1450 and 3000 meters. In many places it is almost entirely arboreal, but in areas where there are many fallen trees or rock fences and rock piles, many individuals have been found on the ground near the rocks or logs. In the coniferous forests this species is associated with S. grammicus microtepidotus and S. aeneus aeneus. The distinction made by Smith (1938:572) between the sub- species S. torquatus torquatus and melanogaster is slight. Indi- viduals with pale bluish spots are found throughout the range of the species in Michoacan; spotting is especially evident in the young. Individuals having an incomplete nuchal collar have been found at Maravatio and at Zinapécuaro in the northern part of the state; in this character these specimens resemble S. torquatus melanogaster, which is found to the north from Guanajuato to Zacatecas and San Luis Potosi. AMPHIBIANS AND REPTILES OF MICHOACAN 745) Sceloporus utiformis Cope Sceloporus utiformis Cope, Proc. Acad. Nat. Sci. Philadelphia, 16:177, September 30, 1864.—Colima, Colima, México. Nineteen km. S of Arteaga (2); Cascada Tzararacua (17); Coahuayana (3); Coalcoman (6); El Sabino (2); El Ticuiz (2); Ostula (3); Pémaro; Rio Cachan; San Juan de Lima; Uruapan (26). In Michoacan the range of this species is discontinuous. It has been found between 1050 and 1550 meters on the slopes of the Cordillera Volcanica, and on the coast and seaward slopes of the Sierra de Coalcoman up to an elevation of 900 meters. It is absent from the Tepalcatepec Valley. At Uruapan and at Cascada Tzar- aracua this lizard was found on the ground in oak forest or in open pine-oak forest; on the coast and foothills of the Sierra de Coal- coman it was found on the ground in the gallery forests along streams, and not in the scrub forest. Urosaurus bicarinatus tuberculatus (Schmidt) Uta tuberculata Schmidt, Amer. Mus. Novitates, 22:4, December 1, 1921.— Colima, Colima, México. Urosaurus bicarinatus tuberculatus, Mittleman, Bull. Mus. Comp. Zool., 91:169, September, 1942. Twenty-six km. S of Arteaga; Cascada Tzararacua (2); Chupio; Coahuay- ana; Coalcoman (8); El Sabino (2); Jungapeo; La Orilla (2); La Placita (4); Playa Azul (4); Pémaro (2); San Salvador (16); ? Tupataro; Uruapan (12); Tzitzio; Zamora. The known distribution and geographic variation of Urosaurus bicarinatus in southwestern México presents a confused picture. In general rugosity, specimens from the coastal region of Micho- acan (Coahuayana, La Orilla, La Placita, Playa Azul, and Pomaro) resemble U. bicarinatus tuberculatus to the north along the Pacific coast. Furthermore, specimens from the coast have less stippling in the gular region than do those from the Sierra de Coalcoman and the slopes of the Cordillera Volcanica. Specimens from the mountains have greatly carinate enlarged dorsals, large lateral tubercles, and heavily stippled throats; in these characters they resemble specimens from Morelos, Guerrero, and Oaxaca. As mentioned by Peters (1954:14), some specimens from La Orilla and San Salvador are like U. bicarinatus bicarinatus in certain charac- ters, and one specimen has the blue ventral patches restricted to the sternal area, a characteristic of U. bicarinatus anonymorphus of Oaxaca and eastern Guerrero. Examination of all available speci- mens from Michoacan indicates that the nature of the dorsal scales is of little value in separating the subspecies. The specimens from Michoacan are here provisionally referred to U. bicarinatus tuber- 6—7817 76 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. culatus, because cursory examination of specimens from several localities between Nayarit and Oaxaca shows that there are only minor differences between the named populations. Individuals from the northern part of the range are more rugose and have larger blue ventral patches and less gular stippling than those from the south. In Michoacan Urosaurus bicarinatus tuberculatus is found in wooded areas, not in open scrub forest, in the coastal area to ele- vations of about 900 meters, and along the slopes of the Cordillera Volcanica and the southern edge of the Mexican Plateau at eleva- tions from 1000 to 1700 meters. The record for Tupaétaro probably is erroneous, for no other specimens of this species are known from the central plateau. Essentially, the distribution of this species parallels that of Sceloporus utiformis, a strictly terrestrial species. Urosaurus bicarinatus tuberculatus lives on tree trunks. Below 1000 meters in the Tepalcatepec Valley Urosaurus bicarinatus tuber- culatus is replaced by Urosaurus gadowi. Urosaurus gadowi (Schmidt) Uta gadovi Schmidt, Amer. Mus. Novitates, 22:3, December 1, 1921.— Cofradia, Jalisco, México (in error) = Cofradia, Michoac4n, México (Duellman, 1958b:49). Urosaurus gadovi, Mittleman, Bull. Mus. Comp. Zool., 91:154, September, 1942. Acahuato (2); Apatzingan (56); 12-16 km. S of Apatzingain (12); Buena- vista (7); Capirio (23); Cofradia (21); El Sabino (13); Guayabo; Jazmin; La Playa; La Salada (3); Nueva Italia (7); Rancho Nuevo; Rio Cancita, 14 km. E of Apatzingan (5); Rio Marquez, 10 km. S of Lombardia (2); Rio Marquez, 13 km. SE of Nueva Italia (3); San Salvador (2); Santa Ana; Tepalcatepec; Volcan Jorullo (3); Zicuiran (2); Ziracuaretiro. Although individuals of this species have been collected at eleva- tions slightly exceeding 1200 meters on Volcan Jorullo and at 1100 meters at Ziracuaretiro on the southern slopes of the Cordillera Vol- cAnica, for the most part these lizards are found at elevations of less than 800 meters, where they inhabit the open arid scrub forest of the Tepalcatepec Valley, a region to which this species is endemic (Duellman, 1958b:49). These small lizards usually are found on the trunks and main branches of the small trees in the scrub forest; in this habitat they are associated with Sceloporus horridus oligo- porus, a much larger species. Males have a pale orange spot on the throat and a pale blue belly; females have immaculate venters. A specimen from Guayabo on the northern slopes of the Sierra de Coalcomén was referred to Urosaurus irregularis (Fischer) by Peters AMPHIBIANS AND REPTILES OF MICHOACAN The (1954:15). I have studied this specimen (BMNH 1914.1.28.110), a female having a snout-vent length of 46 mm., and agree with Peters that it closely resembles Fischer’s description and figure (1882: pl. 17, fig. 1). This specimen and those seen of Urosaurus gadowi all have pavementlike enlarged dorsal scales that are complete across the vertical line. In U. gadowi the enlarged dorsals usually are in four to six irregular rows; in the specimen from Guayabo the dorsals are in tworows. Although none of the other specimens of U. gadowi examined has only two rows of enlarged dorsals, I prefer to consider the specimen from Guayabo as an aberrant individual of that species, rather than U. irregularis. Guayabo is in the known range of U. gadowi. Urosaurus irregularis is known only from the type specimen in the Bremen Museum; the type locality, according to Fischer (1882:232), is “Aus dem Hochlande von Mexico.” If an examina- tion of the type specimen of U. irregularis shows it to be identical with U. gadowi, then U. irregularis would be the name for the lizards here referred to U. gadowi. Mabuya brachypoda Taylor Mabuya brachypoda Taylor, Univ. Kansas Sci. Bull., 38 (1):308, December 0, 1956.—Four kilometers east-southeast of Los Angeles de Tilardn, Guanacaste, Costa Rica. El Sabino (42); La Placita; Playa Azul; Tzitzio (8). Previously this species has been reported from La Placita as Mabuya mabouya alliacea by Peters (1954:15). Webb (1958:1311) provided evidence that Mexican specimens were conspecific with Mabuya brachypoda, as described from Costa Rica by Taylor (1956: 808). The large series in the Taylor collection studied by Webb and listed by him as being from Uruapan actually is part of a series collected by Hobart M. Smith at El] Sabino at an elevation of 1050 meters, 30 kilometers south of Uruapan. This species probably ranges throughout the coastal region of the state; individuals from La Placita and Playa Azul were taken in dense scrub forest near sea level. Scincella assata taylori (Oliver) Leiolopisma assatum taylori Oliver, Occ. Pap. Mus. Zool. Univ. Michigan, 860:12, November 20, 1937.—Santiago, Colima, México. Scincella assata taylori, Mittleman, Herpetologica, 6:20, June 5, 1950. Twenty-one km. S of Arteaga; Ostula. The specimen from Ostula was obtained in semi-deciduous broad- leaf forest at an elevation of 120 meters; that from 21 kilometers south of Arteaga was taken in oak forest at an elevation of 830 meters. 78 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. Both localities are on the coastal slopes of the Sierra de Coalcoman. Probably the species inhabits the heavy forests on the lower slopes of these mountains. The specimen from south of Arteaga (UMMZ 119117) in life had a tan dorsum and a bright orange-pink tail. Eumeces altamirani Dugés Eumeces altamirani Dugés, La Naturaleza, ser. 2, 1:485, 1891.—Apatzing4n, Michoacan, México. Twelve km. E of Apatzing4n; El Sabino (4). One specimen of this rare species was found beneath a rock in the open scrub forest 12 kilometers east of Apatzingan on July 3, 1955. Another skink, presumably of this species, was seen at Ca- pirio. The specimen from east of Apatzingén is a male having a snout-vent length of 97 mm. and an incomplete tail. In most re- spects it compares favorably with accounts of the species given by Taylor (1936b:55 and 1936c:102). The frontal is divided by a transverse suture; the enlarged dorsal scales are arranged in 11 pairs anteriorly, followed by 48 unpaired enlarged scales. The head and middorsal area are brown; there is a pale tan stripe on the edges of the vertebral and paravertebral rows, bordered by a dark brown stripe on the paravertebral row, which, in turn, is bordered by a pale tan stripe on the lateral edge of the paravertebral scale row and the median edge of the adjacent scale row. The stripes extend from the neck to the base of the tail. The flanks are mottled with brown and cream-color; the labials are cream-color barred by brown; the venter is a pale cream-color. Dugés (1891:485) described Eumeces altamirani from “las re- giones cAlidas del Estado de Michoacan” and subsequently (1896: 480) gave Apatzingén as a locality for the species. Presumably he had only one specimen. In 1935 Hobart M. Smith collected the species at E] Sabino on the lower slopes of the Cordillera Volcanica bordering the Tepalcatepec Valley. All of the known specimens are from this valley and the adjacent slopes, an area to which the species apparently is endemic. Eumeces colimensis Taylor Eumeces colimensis Taylor, Publ. Field Mus. Nat. Hist., zool. ser., 20:77, May 15, 1935.—Colima, Colima, México. Coalcoman; Salitre de Estopila. The species was reported by Peters (1954:16); no additional material has been discovered. The species is known only from foothills and low mountains at elevations between 130 and 950 meters in Michoacan and Colima. AMPHIBIANS AND REPTILES OF MICHOACAN 79 Eumeces copei Taylor Eumeces copei Taylor, Proc. Biol. Soc. Washington, 46:1338, June 5, 1933.— 10 miles southeast of Asuncidn, México, México. Cerro Tancitaro (3); Zacapu. This member of the Eumeces brevirostris-group has been found only in pine or pine-fir forests at elevations from 1800 to 2700 meters. It probably ranges throughout the high mountains of the state north of the Tepalcatepec Valley; its apparent absence in other parts of the Cordillera Volcanica, other than on Cerro Tancitaro, is surprising. The species has been taken near Asuncién in the state of México and at Lagunas de Zempoala in Morelos. In this species the lateral pale yellow stripe, which is bordered below by dark brown, extends to the groin and onto the base of the tail. The dorsolateral stripe is separated from the copper-colored middorsum by a narrow brown stripe. Eumeces dugesi Thominot Eumeces Dugesii Thominot, Bull. Soc. Philom. Paris, ser. 7, 7:138, 1883.— Guanajuato. Type locality restricted to Guanajuato, Guanajuato, México, by Smith and Taylor (1950b:169). Carapan (6); Cheran (5); Opopeo (2); 17 km. S of Patzcuaro (3); San José de la Cumbre (2); Tancitaro (2); Tangancicuaro; Uruapan; Zacapu. Individuals of this species frequently have been found beneath rocks and logs in pine-oak, pine, or fir forests from elevations of 1550 to 1850 meters. To judge from specimens available, E. dugesi probably is the most abundant and widespread species of skink in the state. In this species the lateral yellow stripe is indistinct and is per- sistent only in the axilla; the dorsolateral stripes terminate anterior to the hind limbs and are not separated from the tan dorsum. Eumeces indubitus Taylor Eumeces indubitus Taylor, Univ. Kansas Sci. Bull., 21:257, November 27, 1934.—Near Cuernavaca, Morelos, México. Puerto Hondo. The one specimen of this species from Michoacan was collected by Edward H. Taylor in pine forest at Puerto Hondo, near Zit4- cuaro, at an elevation of about 2750 meters (Taylor, 1935:466). The species is known from the high mountains of eastern Michoacan, western México, and northern Morelos. 80 UNIVERSITY OF Kansas Pustis., Mus. Nat. Hist. Eumeces parvulus Taylor Eumeces parvulus Taylor, Proc. Biol. Soc. Washington, 46:175, October 26, 1938.—Tepic, Nayarit, México. El Ticuiz; La Placita; Pémaro (2); San Pedro Naranjestila (3). Aside from the specimens reported by Peters (1954:17), one other specimen was obtained at El Ticuiz. It has 22 scale rows, 3 supra- oculars in contact with the frontal, 2 postlabials, and a unicolored olive-tan dorsum. In life the anterior dorsolateral stripes were pale pinkish tan, the labials cream color, the throat white, and the tail pale blue. All specimens were found in semi-deciduous broadleaf forest at elevations of less than 500 meters on the seaward slopes of the Sierra de Coalcoman. Ameiva undulata sinistra Smith and Laufe Ameiva undulata sinistra Smith and Laufe, Univ. Kansas Sci. Bull., 31 (1):59, May 1, 1946.—Manzanillo, Colima, México. Apatzing4n (9); 19 km. S of Arteaga (3); Barranca de Bejuco (2); Coahu- ayana (6); Coalcoman (8); El Ticuiz (10); La Placita (2); Limoncito (3); Ostula (2); Playa Azul; Salitre de Estopila; San Juan de Lima (2); San Pedro Naranjestila (4). Six males and six females from the Tepalcatepec Valley have more femoral pores than do 16 males and nine females from the coastal lowlands; the ranges and average number of femoral pores in the former are 40-50 (44.8) for males and 38-40 (38.6) for fe- males; males from the coast have 34-44 (89.2), and females have 32-40 (86.2) femoral pores. In all specimens the number of lamel- lae beneath the fourth toe varies from 26 to 83 (29.7). In life juveniles have a pale olive-tan dorsum and a dorsolateral dark band, superimposed on which is a row of darker brown spots. The dorso- lateral band is bordered below by a narrow cream-colored stripe. The tail is tan above and grayish white below; the belly is pale bluish white. Adult males are brilliantly colored in life. A male having a snout-vent length of 108 mm. had a rusty brown dorsum and bright blue bars on the flanks separated by dark brown inter- spaces. The side of the head was pale green, and the chin and throat were golden yellow. In some specimens the throat is orange. Juveniles and subadults have dark flecks on the brown or tan mid- dorsal area, but these are absent in the largest males. This species inhabits the heavily wooded areas in the lowlands to elevations of about 950 meters. In the Tepalcatepec Valley it has been found only in gallery forests along streams. In both the Tepal- catepec Valley and the coastal lowlands there is a noticeable absence of large adults in the dry season. AMPHIBIANS AND REPTILES OF MICHOACAN 81 Cnemidophorus calidipes Duellman Cnemidophorus calidipes Duellman, Occ. Pap. Mus. Zool. Univ. Michigan, 574:1, December 23, 1955.—Capirio, Michoacan, México. eee at (56); 12-20 km. S of Apatzingan (5); 19 km. E of Apatzing4n 5); km. S of Arteaga; Capirio (57); El Espinal (18); Jazmin (9); 11 km. S ot Lombardia; Nueva Italia. This small, distinctive species of the sexlineatus-group of Cnemi- dophorus was discovered in the Tepalcatepec Valley in 1955 (Duell- man, 1955); subsequent field studies showed it to be widespread in the valley (Duellman, 1960c). One specimen (KU 29747) is from the relatively arid, low Pacific slope of the Sierra de Coalcoman, 25 kilometers south of Arteaga. All other specimens have been taken at elevations of 200 to 650 meters in the Tepalcatepec Valley, where the species characteristically inhabits the open scrub forests of the valley floor, especially the Cercidium-Prosopis-Apoplanesia associations, where there is a sparse growth of grasses. In this habitat it is most frequently seen in association with Cnemidophorus costatus zweifeli and C. deppei infernalis. Aside from the charaters given in Table 5, Cnemidophorus cali- dipes differs from other species of Cnemidophorus in Michoacan by possessing a complete (or nearly so) supraorbital semicircle- series of granules; in other species the granules seldom extend anteriorly beyond the posterior border of the frontal. Cnemidophorus communis communis Cope Cnemidophorus communis Cope, Proc. Amer. Philos. Soc., 17:95, 1877.— No type locality given; type locality restricted to Colima, Colima, México, by Zweifel (1959a:74). Cnemidophorus communis communis, Zweifel, Bull. Amer. Mus. Nat. Hist., 117:74, April 27, 1959. Aguililla (2); Apatzing4n (6); 13 km. S of Arteaga (2); 19 km. S of Arteaga (3); Capirio (3); Coahuayana (3); Coalcoman (44); El Ticuiz; between El Ticuiz and Ojos de Agua de San Telmo; Le Flecits (6); Pémaro (2); Rio Cachan; Salitre de Estopila; San Juan de Lim The specimens from Coalcoman and the wet localities were referred to Cnemidophorus sacki copei by Peters (1954:18) and Duellman (1954b:12). Zweifel (1959a) referred these specimens to Cnemidophorus communis communis and pointed out the prob- able sympatry of C. communis and C. costatus (= sacki of Zweifel) in the Tepalcatepec Valley. There is considerable geographic variation in the number of dorsal granules around the midbody. Sixteen specimens from the coastal regions of Michoacan have 129-146 (186.3) granules; nine from the Tepalcatepec Valley have 124-137 (128.8), and 44 from Coalcomén at an elevation of 950 meters in the Sierra de Coalcoman, intermedi- UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. 82 “UL CG “Wau OOT ‘UU 86 ‘WU FS ‘UI £6 “utd GET "WU 64, Ygsue] quoA-jnous UIMNUIXB yuid -dsUBI() yorlq puv ¥uIg yoryq pus yurd-ysinig = Radi pee id: SALES youd yods an[q red seoee era Uld eikea Fe | Utd Bo) (05) yeoryL BP iey. yaep ut sjyods uey fsodiys weedd yourysip xIg sjods [v10}R] en,q /40UuTy -SIp sjeiqoyiaavied fadiys [esioppru peoig S[B1IGe} -19A pasny Ayperyjszed snjd sodiys yourjstp yysIT syuBy uO sieq [BoTZIOA fads [esiop -pltr MOT[IA YIM posny sadijzs [viqo}IoAvIe | syuey ystppor ‘adiys [vsoye] WeeId pvoiq ‘sodizys [v10}8[Osiop pus [viqe}eAvied user) sjods ysinjq JO MOJ O4UI UayoIq adt14s [B.109R] ‘sodtys ]e10}e[Os10p pus jeiqoyleavied usaly UINSIOPpIw UseIs eed YIM posny s[Biqo} -1aAbred /8}0ds JO SMOJ [8.19] B[OSIOP PUB [B19}'T A410 -110380d syods ojed pu ApIOLIayUv sIBq-ssO1D syods moyjaA Jo SMO XI8 YJIM UWNsIOp Weer) sjods puw SIVG ONG [VoIPAGA YIM WNsIOp UMOIG 44ST u1oyyed 10[09 4,Npy (G€) 1h-ZE (8£) 8F-ZE (Gg) LE-ZE (6€) L¥-ZE (98) €F-TE (28), Ses28 (If) 6P-ZE (6) €P-LE (bh) Ze-8E (68) Lb-1€ soiod [B1OWI9 (98) 26-08 SMa ee davieisiien sHegeh'e) oa" op hima S2.UD]D98 (SFI) F9I-OZT | snpriy snuassiyneur (IZ) OZI-LIT | snwassanauy snwissyneur) (ZZ1) OFT-SOT |'° °° SNISILOLI SNULISSIDAUY (OL0ZIK16= | Sa SS sypusafur raddap (9TT) LII-9I1 Oo Vedeoeredso Osc. OD taddap vaddap OOD: I=6is|2- = Yyaframz snyo}soo (66) ZOI-26 (FI) 9FI-SOI $2)p]Uapr990 $1]0}809 crak Gea STUNUUULOD STUNUULOD oheuete VetegeRot eu eile is, anoveniete sadipypo (¢2) 98-99 sopnuBis 1es10q] salogddg (XINQ NYOVOHOIJ, WOUA SNAWIONdG YOA AVY SLNNOD ATVOS) NYOVOHOIJ NI SAUOHAOGIWANT) JO SAIOAUSANS ANV SAIOAdS NAY, FHL AO NOSIUVAWOD—'G ATAV I, AMPHIBIANS AND REPTILES OF MICHOACAN 83 ate geographically between the coast and the Tepalcatepec Valley, have 105-144 (119.7). The number of granules in specimens from the coast of Michoacan compares favorably with the range of 118-154 (137.8) for 34 specimens from Colima, Colima (Zweifel, 1959a:107). Aside from the characters given in Table 5, C. communis communis can be distinguished from other members of the Cnemidophorus sexlineatus-group (calidipes, costatus, and scarlaris) by its relatively small post-antebrachial scales. Although this is the largest species of Cnemidophorus in Michoa- can (adult males attain a snout-vent length of 135 mm. ), it is neither widespread nor abundant. On the coastal lowlands it occurs pri- marily with Cnemidophorus lineatissimus lividus. In the coastal lowlands there is little open scrub forest, a type of habitat that seems to be preferred by C. communis communis. In the Tepalcatepec Valley, C. communis communis occurs in the open scrub forest with the more abundant large species C. costatus (subspecies zweifeli). Only in the scrub forest in the Coalcoman Valley, where no other species of Cnemidophorus occurs, is C. communis communis abundant. Cnemidophorus costatus occidentalis Gadow Cnemidophorus communis occidentalis Gadow, Proc. Zool. Soc. London, 1906, 1:339, August 23, 1906.—Type locality restricted to Ixtlan, Nayarit, México, by Smith and Taylor (1950b:182). Cnemidophorus costatus occidentalis, Zweifel, Copeia, No. 1:98; March 17, 1961. Jiquilpan (4). Only four specimens from the extreme northwestern part of the state are referable to this subspecies. These have 97 to 102 dorsal granules at midbody and lack the blue gular band or spot charac- teristic of the subspecies in the Tepalcatepec Valley. Probably C. costatus occidentalis ranges throughout the Chapala depression, but to the east it is replaced by Cnemidophorus scalaris scalaris. Cnemidophorus costatus zweifeli Duellman Cnemidophorus sacki zweifeli Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 10:589, May 2, 1960.—Capirio, Michoacan, México. Apatzingan (107); Buenavista (3); Capirio (31); Charapendo (12); Chi- napa (2); 19 km. S of Corralito (3); Jazmin (2); between La Playa and Volcan Jorullo (2); Limoncito (3); 14 km. S of Lombardia (11); Nueva Italia (15); Rio Marquez, 10 km. S of Lombardia (2); Rio Marquez, 13 km. SE of Nueva Italia; Tafetan (18); 14 km. E of Tepalcatepec (2); Tzitzio (11); 19 km. S of Tzitzio; Volcan Jorullo (5); Ziracuaretiro; Zirimicuaro. These lizards were referred to Cnemidophorus sacki copei by Duellman (1954b:12 and 1955:6); Duellman (1960a) described the subspecies zweifeli and assigned it to Cnemidophorus sacki. Zwei- 84 UNIVERSITY OF Kansas PuBts., Mus. Nat. Hist. fel (1961:98) used the specific name C. costatus for the whiptails on the southwestern part of the Mexican Plateau (C. c. occiden- talis). Since occidentalis and zweifeli are conspecific, the combina- tion C. costatus zweifeli is used here for the population inhabiting the Tepalcatepec Valley. This lizard is abundant in the Tepalcatepec Valley, where it lives in open and dense scrub forest, usually at elevations of less than 1000 meters. Throughout the valley it is found in association with Cnemidophorus deppei infernalis, and in the lower parts of the val- ley it also is associated with Cnemidophorus calidipes. Observations made in the dry season indicate that large adults are not active at that time. On the coastal lowlands and in the valleys in the Sierra de Coalco- man Cnemidophorus costatus zweifeli is replaced by C. communis communis. To the east in the Balsas Basin C. costatus zweifeli inter- grades with C. costatus costatus. Cnemidophorus deppei deppei Wiegmann Cnemidophorus deppei Wiegmann, Herpetologia Mexicana, p. 29, 1834.— México. Type locality restricted to Tehuantepec, Oaxaca, México, by Smith and Taylor (1950b:179). Cnemidophorus deppei deppei, Cope, Trans. Amer. Philos. Soc., 17:31, 1892. Salitre de Estopila; San Pedro Naranjestila. This small species, which is extremely abundant on the coastal lowlands of Guerrero, seems to be rare on the coast of Michoacan, where it has been taken at elevations of 130 and 500 meters in open situations in otherwise forested areas. Duellman and Well- man (1960:25) discussed these specimens in relation to their sub- specific assignment. They were referred to Cnemidophorus deppei lineatissimus by Peters (1954:18). Cnemidophorus deppei infernalis Duellman and Wellman Cnemidophorus deppei infernalis Duellman and Wellman, Misc. Publ. Mus. Zool. Univ. Michigan, 111:32, February 10, 1960.—Mexcala, Guerrero, México. Acahuato; Apatzing4n (227); Capirio (3); El Sabino; Jazmin; La Playa (6); Lombardia (6); Nueva Italia (4); Rio Marquez, 10 km. S of Lom- bardia (6); Rio Marquez, 13 km. SE of Nueva Italia (10); south of Tan- citaro; Volcan Jorullo (3). This is one of the most abundant and widespread lizards in the Tepalcatepec Valley. Throughout its range it is ecologically asso- ciated with Cnemidophorus costatus zweifeli, which ranges to elevations somewhat higher than the 1050 meters known for C. deppei infernalis. This small lizard reaches its greatest abundance AMPHIBIANS AND REPTILES OF MICHOACAN 85 in grassy areas on the floor of the Tepalcatepec Valley, where in the Cercidium-Prosopis-Apoplanesia associations it occurs with Cnemidophorus calidipes. Duellman and Wellman (1960) discussed the variation and re- lationships of Cnemidophorus deppei, of which the subspecies infernalis is restricted to the Balsas-Tepalcatepec Basin. Cnemidophorus lineatissimus exoristus Duellman and Wellman Cnemidophorus lineatissimus exoristus Duellman and Wellman, Misc. Publ. Mus, Zool. Univ. Michigan, 111:44, February 10, 1960.—Rancho Santa Ana, four kilometers northeast of San Salvador, Michoacan, México. Thirteen to 25 km. S of Arteaga (18); Capirio (19); Limoncito (13); Santa Ana (22). As in Cnemidophorus calidipes, the distribution of this subspecies seems to be restricted to the Tepalcatepec Valley, except in the vicinity of Arteaga, where it occurs on the southern slope of the Sierra de Coalcoman. As pointed out by Duellman and Wellman (1960:46), the specimens from south of Arteaga are like those from the Tepalcatepec Valley in scutellation and coloration, and not like Cnemidophorus lineatissimus lividus from the geographi- cally closer coastal lowlands. In the Tepalcatepec Valley Cnemidophorus lineatissimus exor- istus inhabits gallery forests along the larger streams; in this habitat it is associated with Ameiva undulata sinistra. From the other species of Cnemidophorus in Michoacan, C. lineatissimus exoristus can be distinguished by the possession of seven longitudinal stripes in adults and by the characters of scutellation given in Table 5. Cnemidophorus lineatissimus lineatissimus Cope Cnemidophorus lineatissimus Cope, Proc. Amer. Philos. Soc., 17:94, 1877.— Colima and Guadalajara. Type locality restricted to Colima, Colima, México, by Smith and Taylor (1950b:179). Cnemidophorus lineatissimus lineatissimus, Duellman and Wellman, Misc. Publ. Mus. Zool. Univ. Michigan, 111:41, February 10, 1960. Boca de Apiza (4). These specimens have 117 to 126 dorsal granules at midbody, a noticeably lower count than that for Cnemidophorus lineatissimus lividus on the coast of Michoacan, which has 126 to 164 (148). Apparently these specimens represent immature C. lineatissimus lineatissimus; the differences between these and C. lineatissimus lividus from nearby localities indicate that possibly the populations are distinct species and not subspecies, as suggested by Duellman and Wellman (1960:41). 86 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. Cnemidophorus lineatissimus lividus Duellman and Wellman Cnemidophorus lineatissimus lividus Duellman and Wellman, Misc. Publ. Mus. Zool. Univ. Michigan, 111:50, February 10, 1960.—Maruata, Michoacan, México. Barranca de Bejuco (4); Boca de Apiza (2); Coahuayana (6); El Ticuiz (7); La Placita (11); Maruata (7); Motin del Oro; Ostula (5); Playa Azul (4); Playa Cuilala (2); Pémaro (2); Salitre de Estcpila (2); San Pedro Naranjestila. This is the most abundant and widespread species of Cnemi- dophorus on the coastal lowlands of Michoacan, where it ranges from sea level to elevations of about 500 meters. In this area it inhabits dense arid scrub forest and semi-deciduous broad-leafed forest. Both of these habitats are continuous, or nearly so, along the lowlands and foothills of the Sierra de Coalcoman. This in itself may explain the abundance of Cnemidophorus lineatissimus and the relative scarcity of C. deppei and C. communis in the coastal area, for C. deppei and C. communis usually inhabit more open arid scrub forest, as occurs in the Tepalcatepec Valley. Living in the dense scrub forest with C. lineatissimus is Ameiva undulata sinistra. Cnemidophorus scalaris Cope Cnemidophorus gularis scalaris Cope, Trans. Amer. Philos. Soc., 17:47, 1892.—Chihuahua, Chihuahua, México. Cneiac nner scalaris, Zweifel, Bull. American Mus. Nat. Hist., 117:72, 9. a. (2); Jacona; Lago de Cuitzeo (42); Morelia; 21 km. N of Morelia Zweifel (1959a:72) assigned the small species of Cnemidophorus having a relatively low number of dorsal granules and inhabiting the southern part of the Mexican Plateau to C. scalaris, which he diagnosed as rarely exceeding 100 mm. in snout-vent length and always having an average of less than 100 dorsal granules at mid- body and usually less than 90. Forty-two specimens from the south shore of Lago de Cuitzeo (UMMZ 119558) have 80-91 (85.8) dorsal granules. Four specimens from 21 kilometers north of Norelia (UIMNH 6952 and UMMZ 104743) have 89, 78, 92, and 84 granules; one from Morelia (UMMZ 104742) has 78; two from Araro (UMMZ 119522) have 80 and 87; one from Jacona (UIMNH 24703) has 88. Since no large adult males are present in the series from Micho- acan, an adequate comparison of coloration between these and populations on the northern part of the Mexican Plateau cannot be made. Cnemidophorus scalaris is a name applied to the lizards inhabiting the Mexican Plateau from Chihuahua south to Puebla by Zweifel (1959a:72). It is doubtful if all of the popula- AMPHIBIANS AND REPTILES OF MICHOACAN 87 tions assigned to this subspecies belong there; possibly more than one species is involved, but the paucity of material prevents further analysis at this time. Heloderma horridum horridum (Wiegmann) Trachyderma horridum Wiegmann, Isis von Oken, 22:421, 1829.—México. Type locality restricted to Huajintlan, Guerrero, México, by Smith and Taylor (1950b:193). Heloderma horridum horridum, Bogert and Martin del Campo, Bull. Amer. Mus. Nat. Hist., 109:20, April 16, 1956. Apatzingan; Coalcoman; La Placita; Oropeo; Pardcuaro. This species is known from elevations of less than 1000 meters in the Tepalcatepec Valley, the Sierra de Coalcoman, and the coastal lowlands. Specimens from Coalcoman, La Placita, and Paracuaro came from areas of dense woods; those from Apatzingan and Oropeo might have come from patches of dense woods in the otherwise open scrub forest of the Tepalcatepec Valley. Gerrhonotus imbricatus imbricatus Wiegmann Gerrhonotus imbricatus Wiegmann, Isis von Oken, 21:379, 1828.—México. Type locality restricted to México, Distrito Federal, by Smith and Tay- lor (1950b:201). Gerrhonotus imbricatus imbricatus, Dunn, Proc. Acad. Nat. Sci. Philadelphia, 88:475, October 20, 1986. Acuaro de las Lleguas (9); Cerro Barolosa (4); Cerro Tancitaro (36); Dos Aguas (22); Paracho; Sierra Patamba; Tinguidin; Zacapu. Specimens from the Sierra de Coalcoman are noticeably different from those inhabiting the mountains rising from the Mexican Plateau. Of 45 specimens from Cerro Tancitaro and adjacent areas on the Mexican Plateau and in the Cordillera Volcdnica, 15 have twelve longitudinal rows of dorsal scales and 30 have fourteen rows. Of seven specimens from the state of México, 5 have twelve rows and 2 have fourteen; of nine specimens from central Veracruz, 8 have twelve rows and one has fourteen; of six specimens from Hidalgo, 5 have twelve rows and one has sixteen; of two specimens from Guanajuata, one has fourteen and the other has sixteen rows. All of the 35 specimens from the Sierra de Coalcoman have sixteen rows. Furthermore, these specimens have the superciliary row ex- tended anteriorly, so that the anterior superciliary is in broad contact with the loreal. Specimens from Cerro Tancitaro have a shorter su- perciliary row, so that the anterior superciliary is not in broad con- tact with the loreal. These characters were used by Tihen (1949:220) to distinguish Gerrhonotus imbricatus ciliaris from G. imbricatus imbricatus. According to Tihen, the subspecies G. imbricatus 88 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. ciliaris ranges from Guanajuato and Hidalgo northward to Chihua- hua and Coahuila, whereas the nominal subspecies occurs from Michoacan and Hidalgo southward to Oaxaca. Specimens from the Sierra de Autlan in Jalisco are like those from Cerro Tancitaro; consequently, there seems to be no connection between the popula- tions of G. imbricatus ciliaris in the mountains of the northern part of the Mexican Plateau with the ciliaris-like individuals found in the Sierra de Coalcoman. The acquisition and study of additional ma- terial from throughout the range of the species is necessary to clarify the picture of geographic variation. Until then, I prefer to consider all of the specimens from Michoacan as Gerrhonotus imbricatus im- bricatus. The largest specimen is a male having a snout-vent length of 136 mm. Two juveniles collected in July 24, 1960, have snout-vent lengths of 86 and 42 mm. A specimen having a snout-vent length of 127 mm. and a tail length of 145 mm. was regurgitated by a Crotalus pusillus, which had a body length of 550 mm. Gerrhonotus imbricatus imbricatus is an inhabitant of coniferous forests. In the Cordillera Volcanica it occurs from 1500 to 8500 meters at the top of Cerro Tancitaro. In the Sierra de Coalcoman it occurs from 2100 to 2700 meters. On July 4, 1955, a pair was found in copu- lation beneath a pine log at 2700 meters on Cerro Barolosa. The male was lying on top of the female and was holding her head firmly in his jaws; the male’s tail was curled under the female’s tail, so that the cloacae were in contact. Serpentes Typhlops braminus (Daudin) Eryx braminus Daudin, Hist. . . . des reptiles, 7:279, 1803.—Vaza- gapatam, India. Typhlops braminus, Cuvier, Régne animal, ed. 2, 2:73, 1829. Apatzingan; Arteaga. Both specimens known from Michoacan were collected by Gadow in 1908. Peters (1954:20) remarked that the specimen from Arteaga probably does not indicate a rapid spreading of the species, which most likely was introduced into México at the time that vessels were stopping at Acapulco from the Philippines (Taylor, 1940b:444), but instead may indicate that pack trains from Acapulco passed through the Sierra de Coalcoman. The occurrence of this snake along a long- used camino substantiates this belief. AMPHIBIANS AND REPTILES OF MICHOACAN 89 Leptotyphlops bressoni Taylor Leptotyphlops bressoni Taylor, Copeia, No. 1:5, March 9, 1939.—Hacienda El Sabino, Michoacan, México. E] Sabino. This species still is known definitely only from the type specimen collected on the lower slopes of the Cordillera Volcanica at the north- ern edge of the Tepalcatepec Valley. A specimen (now lost) re- ported from Aguililla by Cope (1887:63) possibly represents this species (see Smith and Taylor, 1945:21, and Peters, 1954:20). Leptotyphlops gadowi Duellman Leptotyphlops gadowi Duellman, Copeia, No. 2:93, May 29, 1956.—Apatzin- gan, Michoacan, México. Apatzingan. No additional specimens of this species have been collected since the species was described by Duellman (1956b:93). Data given with the specimen by Gadow indicate that it came from his camp above Apatzingaén at an elevation of about 800 meters. Although the exact position of this camp is unknown, the lower slopes of the Cordillera Volcanica above Apatzing4n usually support arid scrub forest at elevations below 1000 meters. Therefore, this species prob- ably is an inhabitant of the arid scrub forest. Leptotyphlops phenops bakewelli Oliver Leptotyphlops bakewelli Oliver, Occ. Pap. Mus. Zool. Univ. Michigan, 860:16, November 20, 1937.—Paso del Rio, Colima, México. Leptotyphlops phenops bakewelli, Smith, Proc. U.S. Natl. Mus., 93:445, October 29, 1943. La Placita (4); La Salada; Ostula. The five specimens from the coastal lowlands are from elevations of less than 150 meters; these were collected by Peters (1954:20); the specimen from La Salada is from an elevation of 580 meters in the Tepalcatepec Valley. Peters (loc. cit.) remarked that the rostral and the tip of the tail that were described as white by Oliver (1937: 17) actually are sulphur-yellow in life. Loxocemus bicolor Cope Loxocemus bicolor Cope, Proc. Acad. Nat. Sci. Philadelphia, 18:77, June 30, 1861.—La Union, El Salvador. Loxocemus sumichrasti Bocourt, Ann. Sci. Nat., ser. 6, 4:1, 1876.—Tehuan- tepec, Oaxaca, México. Apatzingan (6); La Orilla; Lombardia. As noted by Peters (1954:21), this species was not recorded from Michoacan by Smith and Taylor (1945:27), but Gadow (1930:30) collected a specimen at La Orilla in 1908. This specimen (BMNH 90 Unrversirty OF Kansas Pusts., Mus. Nat. Hist. 1914.1.28.124) is a male having 235 ventrals and 47 caudals, a dark brown dorsum, and cream-colored labials and venter. The anterior chin-shields are considerably longer than the scales bordering the chin-shields. In these characters this specimen agrees with the diag- nosis of Loxocemus bicolor given by Taylor (1940c:447), who re- vived Loxocemus sumichrasti Bocourt. Of the six specimens from Apatzingan in the Tepalcatepec Valley, three males have 248 to 253 (246.6) ventrals and 44 to 45 (44.3) caudals; three females have 238 to 247 (244.0) ventrals and 42 to 44 (43.0) caudals. Certain char- acters of scutellation utilized by Taylor for separating L. bicolor and L. sumichrasti are inconsistent in this series. The chin-shields are longer than the adjacent scales, like those illustrated in L. bicolor by Taylor (op. cit., fig. 1). The relative lengths of the prefrontal and internasal sutures are subequal, or the prefrontal suture is slightly longer. Thus, in these characters of scutellation these snakes are like L. bicolor, but in coloration they are like L. swmichrasti; the dor- sal color in life was an iridescent dark bluish gray, and the belly was pale gray or bluish gray. The supposed differences in scutellation between L. bicolor and L. sumichrasti have been questioned by Woodbury and Woodbury (1944:360); these authors treated L. sumichrasti as a subspecies of L. bicolor. As pointed out by Zweifel (1959b:5), such an arrange- ment is not tenable, for, although individuals with each kind of color pattern have not been collected together at any one locality, the over-all geographic picture is one of sympatric distribution. Only snakes having the coloration of L. sumichrasti have been collected in the Balsas-Tepalcatepec Basin. I agree with Zweifel (loc. cit.) that on the basis of morphological similarities and sympatric dis- tribution, L. bicolor and L. sumichrasti seem to be dimorphic phases of the same species, showing no more striking differences in colora- tion than Lampropeltis getulus californiae, a now classical example of pattern dimorphism in snakes. In Michoacan, as in other parts of its range, Loxocemus bicolor in- habits arid scrub forest environments at low elevations. Boa constrictor imperator Daudin Boa imperator Daudin, Hist. nat. . . . des reptiles, 5:150, 1803.—Mé- xico. Type locality restricted to Cérdoba, Veracruz, México, by Smith and Taylor (1950a:347). Boa constrictor imperator, Forcart, Herpetologica, 7:199, December 31, 1951. Apatzingan (4); Coalcom4n; El Sabino (2); La Placita; La Playa (2); Lom- bardia; Nueva Italia (2); Rio Cachan; Rio Marquez, 13 km. SE of Nueva Italia; Rio Nexpa; Volcan Jorullo. These specimens have come from a variety of habitats from eleva- tions of less than 1,000 meters. The species seems to be equally AMPHIBIANS AND REPTILES OF MICHOACAN 91 abundant in the broad-leafed semi-deciduous forests of the coastal foothills and in the arid Tepalcatepec Valley. In the latter area most of the specimens were collected at night. Coniophanes fissidens dispersus Smith Coniophanes fissidens dispersus Smith, Proc. U. S. Natl. Mus., 91:106, No- vember 13, 1941.—El] Limoncito, Guerrero, México. Arteaga. Further collecting in southern Michoacan has failed to add addi- tional material of this species, which is known in the state from the one specimen collected by Gadow in 1908. The species possibly ranges throughout the coastal foothills of the Sierra de Coalcoman. Peters (1954:21) described the specimen from Arteaga. Coniophanes lateritius lateritius Cope Coniophanes lateritius Cope, Proc. Acad. Nat. Sci. Philadelphia, 13:524, March 31, 1862.—Guadalajara, Jalisco, México. Coniophanes lateritius lateritius, Smith and Grant, Herpetologica, 14:20, April 25, 1958. Nineteen km. S of Arteaga. The one specimen available from Michoacan of this apparently rare species was discussed by Wellman (1959:127), who pointed out that although the specimen was geographically intermediate be- tween the subspecies C. I. lateritius (Jalisco and Nayarit) and C. l. melanocephalus (Morelos and Puebla), the specimen (UMMZ 118954) was like C. 1. lateritius in scutellation and in color pat- tern differed from other known specimens of the species in hav- ing had in life a pale orange, instead of a brick-red, dorsum. Addi- tional specimens from the Sierra de Coalcoman will be required in order to determine whether this specimen is a representative of an orange-colored population or merely is aberrant in coloration. The present specimen is from an elevation of 900 meters in oak forest on the southern slopes of the Sierra de Coalcoman; other lo- cality records for the species indicate that it inhabits broad-leafed forest in foothills from Nayarit to Puebla. Conophis vittatus vittatus Peters Conophis vittatus Peters, Monats. Akad. Wiss. Berlin, p. 519, 1860.—No type locality given. Type locality restricted to Laguna Coyuca, Guerrero, México, by Smith and Taylor (1950a:381). Conophis vittatus vittatus, Smith, Jour. Washington Acad. Sci., 31:119, March 17, 1941. Arteaga; Coalcomaén (4); La Playa; 19 km. S of Tzitzio. All specimens of this terrestrial snake have been collected in areas of scrub forest between 800 and 1100 meters above sea level. Since 7—7817 92 University OF Kansas Pusts., Mus. Nat. Hist. the species is known from the coastal regions of Guerrero and Colima, its absence from the cost of Michoacan is unexplainable; probably the lack of specimens from these areas is due solely to inadequate collecting. Conopsis biserialis Taylor and Smith Conopsis biserialis Taylor and Smith, Univ. Kansas Sci. Bull., 28 (2):333, November 12, 1942.—Ten miles west of Villa Victoria, México, México. Capdcuaro (5); Cerro San Andrés; Cherian; Ciudad Hidalgo; Macho de Agua 735 Patzcuaro (8); Tancitaro (24); Uruapan (9); 24 km. SE of Zitacuaro (1 This species is abundant in the coniferous forests at elevations from 1550 to 2800 meters throughout the Cordillera Volcanica; ap- parently it does not occur in the Sierra de Coalcoman. On August 1, 1956, a copulating pair was found beneath a rock at Capacuaro. One of the best characters to distinguish this species from Toluca lineata, which occurs with Conopsis throughout its range in Michoa- can, is the presence of large, black ventral blotches in Conopsis biserialis, as contrasted with the two rows of small black spots in Toluca lineata. Conopsis nasus Giinther Conopsis nasus Giinther, Catalogue . . . snakes . . . British Mu- seum, p. 6, 1858.—California (in error). ‘Type locality restricted to Guanajuato, Guanajuato, México, by Smith and Taylor (1950a:330). Carapan (2); Erongaricuaro; Maravatio (3); Morelia (2); Nahuatzén; Patzcuaro (7); Tacicuaro (8); Tancitaro. This species has been collected in oak, pine-oak, and fir forests at elevations of 1900 to 2450 meters on the mountains rising from the Mexican Plateau. It does not seem to be so abundant as Conopsis biserialis. Sufficient ecological data to determine differences in habi- tat between the two species have not been compiled. Diadophis dugesi Villada Diadophis punctatus dougesii Villada, La Naturaleza, 3:226, 1875.—Potreros de Balbuena, Distrito Federal, México. Diadophis dugesii, Blanchard, Bull. Chicago Acad. Sci., 7:51, December 30, 1942. Morelia (2); P4tzcuaro; Quiroga. Apparently this snake is uncommon in Michoacan. It has been found only at elevations of 1900 to 2200 meters in pine and pine-oak forests on the mountains rising from the Mexican Plateau. AMPHIBIANS AND REPTILES OF MICHOACAN 93 Dryadophis melanolomus stuarti Smith Dryadophis melanolomus stuarti Smith, Proc. U.S. Natl. Mus., 93:418, Octo- ber 29, 1943.—Acapulco, Guerrero, México. Coahuayana; El Ticuiz; La Placita (3); Punto San Juan de Lima; Punto San Telmo. The few specimens indicate that in Michoacan, as elsewhere on the Pacific coast of México, this species is restricted to forested re- gions on the coastal plain. It does not occur in the Tepalcatepec Valley. The coloration, in life, of a juvenile (UMMZ 114604) is as follows: The dorsum is uniform pale grayish tan on posterior one-third of body and on tail; anteriorly there are pale grayish tan middorsal blotches separated by grayish white interspaces, which are about one-half the length of the blotches. Posteriorly the blotches are less distinct, fading into the uniform grayish tan ground color of the posterior part of the body. The blotches extend laterally onto the fourth and fifth scale rows. Large squarish lateral intercalary blotches of darker brown interconnect with the dorsal blotches. The top of the head is pale olive-brown; a dark brown postorbital stripe extends from the eye to the posterior edge of the last upper labial. The labials, chin, and ventrals 1-30 are creamy white, changing to a dusty cream-color posteriorly; the chin and ventrals 1-30 are heavily spotted with dark brown. The iris is a cream-color above and chocolate brown below; the tongue is blue. Drymarchon corais rubidus Smith Drymarchon corais rubidus Smith, Jour. Washington Acad. Sci., 31:474, No- vember 11, 1941.—Rosario, Sinaloa, México. Apatzingin (5); Arroyo El Salto; Arteaga; Capirio; El Sabino (7); La Palma; La Placita; Ostula; San Juan de Lima. Not all of the specimens from Michoacan are typical in color pat- tern of this subspecies, as defined by Smith (1941a:475). All speci- mens from the Tepalcatepec Valley are uniformly black above; they have reddish or cream-colored chins and the anterior two-thirds of the belly salmon-pink or reddish buff. Individuals from the Sierra de Coalcoman (Arteaga and Arroyo El Salto) are like those from the Tepalcatepec Valley. Three specimens from the coastal low- lands differ noticeably in color pattern: UMMZ 104504, adult male (Ostula).—Pale brown above flecked with black anteriorly; at midbody, flecks form narrow transverse bands that become progressively wider posteriorly, until on tail no brown pigment evident, all ventrals reddish buff, except last eight, which are black. 94 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. UMMZ 104602, adult female (La Placita ).—Black above, reddish cross-bands and flecks on all of body; dorsal and ventral surfaces of tail black; chin cream-color and entire belly reddish buff. UMMZ 114626, adult male (San Juan de Lima).—Black above; dull rust-colored cross-bands on anterior half of body; chin white; belly rust-colored on anterior two-thirds of body and black pos- teriorly. One specimen from La Palma on the Mexican Plateau (KU 29275) has the top of the head an olive-color, the entire dorsum black, the chin and ventrals 1-42 a cream-color, remainder of venter black, and all of the labials heavily barred with black. A juvenile from Capirio in the Tepalcatepec Valley (UMMZ 114627) is black above and has pale olive-colored flecks on the anterior one-third of the body; the top of the head is dark olive-brown, and the sides of the head are somewhat paler. Anteriorly the belly is a cream-color; posteriorly it is black. The specimens from the Tepalcatepec Valley are typical of Dry- marchon corais rubidus. Those from the coastal lowlands differ in having large areas of brown or red pigment on the dorsum, a condi- tion not mentioned by Smith in his description of the subspecies. The specimen from La Palma, like many others from various locali- ties on the Mexican Plateau, resembles in certain characters D. corais orizabensis (Smith, op. cit.:477). Our knowledge of the geographical variation in coloration in this species is incomplete; many populations have been assigned to subspecific rank without justification. In Michoacan this species is found from sea level to 1350 meters in the Sierra de Coalcoman and to 1800 meters at La Palma on Lago de Chapala. It has been collected in scrub forest, semi-deciduous broad-leafed forest, and oak forest. Drymobius margaritiferus fistulosus Smith Drymobius margaritiferus fistulosus Smith, Proc. U. S. Natl. Mus., 92:383, November 5, 1942.—Miramar, Nayarit, México. Apatzingan (3); Coahuayana; Coalcomén (3); El Sabino (3); El Ticuiz; 12 km. § of Tzitzio. This snake is abundant in the lowlands of the state; the few speci- mens listed above are indicative not of the rarity, but rather of the speed and agility, of this diurnal snake. It most frequently is found near water, where there is a dense growth of vegetation. One indi- vidual was observed in a large pool inhabited by several small Rana pipiens, and another was seen along the bank of a hyacinth-choked AMPHIBIANS AND REPTILES OF MICHOACAN 95 river channel. A third individual was captured while it was in pur- suit of a Cnemidophorus. This species has been collected on the coastal lowlands and sea- ward foothills of the Sierra de Coalcoman and in the Tepalcatepec Valley to elevations of 1150 meters. Elaphe triaspis intermedia (Boettger) Pityophis intermedius Boettger, Ber. Offen. Vereins. Naturk., 22:148, 1883. —México. Type locality restricted to Hacienda El Sabino, Michoacan, México, by Dowling (1960:74). Elaphe triaspis intermedia, Mertens and Dowling, Senckenbergiana, 33:201, November 15, 1952. Twenty-four km. E of Apatzing4n; Chupio; El Sabino (4); 11 km. E of Emiliano Zapata. Dowling (1960) has shown that specimens from the Balsas-Tepal- catepec Basin have fewer ventrals and caudals than those from the Sierra del Sur or the coast. All specimens from Michoac4n were collected in open forest, either scrub or oak forest. They were found in drier situations than those described for the species in southern Tamaulipas by Martin (1958:69). In Michoacan Elaphe triaspis intermedia is known from the Tepalcatepec Valley, the lower slopes of the Cordillera Volcdnica, and the western edge of the Mexican Plateau at an elevation of 1850 meters. It probably occurs in the lower parts of the Sierra de Coalcoman and along the Pacific coast, for it is known from the coastal lowlands of Guerrero and Colima. In August, 1951, I saw a snake that probably was this species in Barranca de Bejuco. Enulius unicolor (Fischer) Geophis unicolor Fischer, Abh. Nat. Ver. Bremen, 7:227, 1882.—México. Type locality restricted to Chilpancingo, Guerrero, México, by Smith and Taylor (1950a:831). aus unicolor, Taylor and Smith, Univ. Kansas Sci. Bull., 25:247, July 10, 1939. Between Ario de Rosales and La Playa; Coalcoman; Jungapeo (4); between Zitacuaro and Tuxpan. This small snake has been collected from beneath rocks in brushy areas and broad-leafed forest between 900 and 1800 meters; it has not been found in coniferous forest. The limited ecological data suggest that the species inhabits the transition zone between the tropical scrub forest and the temperate hardwood forest. All of the specimens have 17 rows of scales; four males have 169-178 (174.2) ventrals and 102-111 (106.8) caudals; two females have 192 and 195 ventrals and 96 and 87 caudals. Three individuals have one postocular on one side and two on the other; in the other 96 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. specimens there are two postoculars on each side. The largest male has a body length of 232 mm. and a tail length of 130 mm.; the largest female has a body length of 274 mm. and a tail length of 119 mm. Geagras redimitus Cope Coeeas redimitus Cope, Jour. Acad. Nat. Sci. Philadelphia, ser. 2, 8:141, San Juan de Lima (2). Previously this species was known definitely only from the Plains of Tehuantepec, Oaxaca. Sphenocalamus lineolatus was descried by Fischer (1883:5) from Mazatlan; this name has been placed in the synonymy of Geagras redimitus Cope. Although Fischer gave the type locality only as “Mazatlan” and did not designate the state, it is probable that the type originated from Mazatlan, Sinaloa. The present specimens are from a locality almost midway between Tehuantepec and Mazatlan and support the possibility that Geagras ranges along the Pacific coast of México from Oaxaca to Sinaloa. The two specimens from Michoacan (UMMZ 114446-7), both males, have 118 and 122 ventrals, 31 and 83 caudals, body lengths of 108 and 81 mm., and tail lengths of 20 and 15 mm. Both have 1-1 preoculars, 1-1 postoculars, 1-2 temporals, 6-6 upper labials, and 5-5 lower labials. In life, the dorsum was pale tan; the top of the head and the middorsal and lateral stripes were dark brown; the belly was white. The occipital spots were pale pinkish tan. Both speci- mens were found beneath rocks in tropical semi-deciduous forest at an elevation of 15 meters on the coastal plain. Geophis dugesi Bocourt Geophis dugesii Bocourt, Miss. Scientifique au Mexique et dans l’Amerique Centrale, Rept., livr. 9:573, 1883.—Tangancicuaro, Michoacén, México. Carapan; Tangancicuaro; Zacapu. Aside from the three specimens listed above, there are two (SU 4407-8) bearing the data “Michoacan.” Bocourt (1883:574) stated that the type specimen from Tangancicuaro had six or seven pale cross-bands on the anterior part of the body. An illustration, pre- sumably of the same specimen, by Dugés (1884:PI. 9) shows five distinct and one indistinct cross-bands. Of the four specimens that I have examined, none has more than three pale cross-bands, and one has only one indistinct cross-band. Two females have 154 and 158 ventrals and 88 and 37 caudals; two males have 150 and 151 ventrals and 43 and 42 caudals. This species is known only from elevations between 1750 and 2050 AMPHIBIANS AND REPTILES OF MICHOACAN 97 meters on the southwestern edge of the Mexican Plateau in the state of Michoacan. Geophis incomptus Duellman Geophis incomptus Duellman, Occ. Pap. Mus. Zool. Univ. Michigan, 605:3, May 29, 1959.—Dos Aguas, Michoacan, México. Dos Aguas (15). This species, which seems to be related to Geophis maculiferus, is known only from the pine-oak forest in the vicinity of Dos Aguas (elevation 2100 meters) in the Sierra de Coalcoman. Aside from the five specimens comprising the type series, there are ten other specimens in the Museum of Zoology at the University of Michigan collected by Floyd L. Downs in July, 1960. Data from these speci- mens and those comprising the type series show that in this sample seven males have 146-153 (149.3) ventrals and 35-37 (36.0) caudals; eight females have 150-154 (152.4) ventrals and 29-84 (82.5) caudals. The largest specimen is a female with a body length of 344 mm. and a tail length of 53 mm. Geophis maculiferus Taylor Geophis maculiferus Taylor, Univ. Kansas Sci. Bull., 27:119, December 30, 1941.—Near Cicio [sic] = Tzitzio, Michoacan, México. Tzitzio. The type and only known specimen of Geophis maculiferus (UIMNH 25078) is a female having 140 ventrals and 30 caudals, dorsal scales in 15 rows, one postocular, and an anterior temporal. Only one other species in México has dorsal scales in 15 rows and has an anterior temporal; that species is G. incomptus, which differs from G. maculiferus in having six or seven lower labials, instead of five, and in having the edges of the ventrals dark, instead of a uni- formly cream-colored belly. The locality from which the specimen was obtained lies at an elevation of 1630 meters on the southern slope of the Cordillera Volcanica. At that elevation there is an interdigitation of arid tropical scrub forest and pine-oak forest; probably Geophis macu- liferus inhabits the pine-oak forest. Geophis nigrocinctus Duellman Geophis nigrocinctus Duellman, Occ. Pap. Mus. Zool. Univ. Michigan, 605:1, May 29, 1959.—Dos Aguas, Michoacan, México. Dos Aguas (8). The three specimens comprising the type series of the species were found beneath logs and in a stump in pine-oak forest at an elevation of 2100 meters. A discussion of the variation in these 98 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. specimens and of probable relationships of the species was given by Duellman (1959). Floyd Downs spent several days at Dos Aguas in July, 1960; although he found ten specimens of Geophis incomp- tus, no further specimens of G. nigrocinctus were obtained. Geophis petersi Boulenger Geophis petersii Boulenger, Catalogue Snakes . . . British Museum, 2:321, September 23, 1894.—Mexico City. Type locality restricted to Patzcuaro, Michoacan, México, by Smith and Taylor (1950a:335). Cheran; Coalcoman; Morelia; Patzcuaro (6). This seems to be the most widespread species of Geophis in Michoacan. It has been found at elevations between 950 and 2350 meters, chiefly in pine or pine-oak forest. Boulenger (1894:321) described Geophis petersi from a specimen stated to be from Mexico City, a locality which probably is in error. The only localities from which the species is definitely known are those listed in this account. Three males and five females from the Mexican Plateau and the Cordillera Volcanica have respectively 140-144 (141.7) and 148-151 (146.0) ventrals and 39-41 (40.0) and 29-35 (33.2) caudals. All have dorsal scales in 15 rows, 1 postocular, no anterior temporal, and a relatively small triangular supraocular. The specimen from Coalcoman (UMMZ 104698) was referred to Geophis nasalis by Peters (1954:22). This specimen is abnormal in several characters; in five places there is a fusion and separation of the vertebral and paravertebral scale rows, producing a change from 17 to 15 rows of dorsal scales. Fusion of the three rows takes place at the level of the 8th, 41st, 47th, 54th, and 65th ventrals. Furthermore, there is a small secondary postocular on each side of the head. In other characters the specimen is like G. petersi; the resemblances to that species are greater than to G. nasalis, which has been recorded from Guatemala and southern Chiapas. Geophis tarascae Hartweg Geophis tarascae Hartweg, Occ. Pap. Mus. Zool. Univ. Michigan, 601:1, May 4, 1959.—Uruapan, Michoacan, México. Uruapan (8). A female of this species was collected in the Parque Nacional at the north edge of Uruapan in 1899, and a male was taken there in 1947; these specimens were used by Hartweg in his description of the species. Floyd L. Downs obtained another specimen in the Parque Nacional on July 19, 1960. It has 164 ventrals and 46 caudals; in life, the ground color of the neck was brown with a purplish tint; the dorsal markings were black; the chin was a cream- AMPHIBIANS AND REPTILES OF MICHOACAN 99 color, and the belly was white. This specimen is distinguished from those of all other species of Geophis in Michoacan in that it has dark irregular cross-bars on the dorsum and a row of dark spots on the venter. Hypsiglena torquata ochrorhyncha Cope Hypsiglena ochrorhyncha Cope, Proc. Acad. Nat. Sci. Philadelphia, 12:246, November 15, 1860.—Cape San Lucas, Baja California, México. Hypsiglena torquata ochrorhyncha, Bogert and Oliver, Bull. Amer. Mus. Nat. Hist., 83:378, March 80, 1945. Tupataro. The systematic status of the geographic variants of Hypsiglena in México and southwestern United States has been commented on by several authors. Tanner (1944) considered H. torquata and H. ochrorhyncha to be distinct species; Bogert and Oliver (1945:879) and Duellman (1957b:238) presented evidence indicating that H. torquata and H. ochrorhyncha intergrade in Sinaloa and southern Sonora. In Hypsiglena the scutellation, including the numbers of labials, dorsals, ventrals, and caudals, seem to vary in a clinal man- ner. Nevertheless, these snakes can be divided into two distinct populations on the basis of the nuchal color pattern, consisting of an ochrorhyncha-type (a broad dark nape-band, the lateral edges of which extend anteriorly and fuse with a postorbital stripe, and a narrow nape stripe extending from the posteromedian edges of the parietals to the dark nape band) and a torquata-type (a somewhat narrower dark nape-band bordered anteriorly by a pale nuchal area, and no dark nape stripe). Snakes having the ochrorhyncha-type of nuchal pattern are found on the Mexican Plateau from Michoacan northward into the desert regions of Sonora and the southwestern United States. Snakes having the torquata-type of pattern are found on the coastal lowlands and adjacent slopes of the Sierra Madre Occidental from southern Sinaloa to Colima and thence inland in the Balsas-Tepalcatepec Basin to Morelos and Guerrero. An excep- tion is Hypsiglena torquata dunklei from Forlén and San Fernando, Tamaulipas; it has the torquata-type of nuchal pattern. The dis- tributional picture is somewhat complicated because some individ- uals having the torquata-type of nuchal pattern also have a faint nape stripe. If these are taken as exceptions, the general picture of dis- tribution in México is H. t. torquata on the Pacific lowlands from Sinaloa southward to the Balsas Basin and H. t. ochrorhyncha on the Mexican Plateau. Smith (1943:433) resurrected Hypsiglena jani Dugés for the snakes of the ochrorhyncha-type on the southern part of the Mexican 100 UnIvERSITY OF Kansas Pusts., Mus. Nat. Hist. Plateau. He stated that the southern specimens differed from north- ern ones in having a nuchal spot 9 or 10 scales in length, as compared with a spot 2 to 6 scales in length in northern specimens. A cursory examination of specimens from the areas between Arizona and Michoacan showed that there is a gradual increase in the size of the spot from north to south. If no other characters can be found to distinguish the populations, they should be considered as a single subspecies. Hypsiglena affinis differs from H. torquata in possessing 19 instead of 21 rows of dorsal scales. Additional material is needed from the western slopes of Jalisco and the Barrancas in Zacatecas and Du- rango, before definite allocation of affinis can be made. Bogert and Oliver (1945:379) discussed the status of certain named populations in Baja California and concluded that only one species occurs there, and that the species probably is conspecific with H. torquata. A careful review of the genus Hypsiglena might show that there is only one species. The one specimen from Michoacan (USNM 46518) is from an elevation of about 2300 meters near the southern edge of the Mexican Plateau. Hypsiglena torquata torquata (Giinther) Leptodeira torquata Giinther, Ann. Mag. Nat. Hist., ser. 3, 5:170.—Laguna Island, Nicaragua (in error). Hypselena forauste torquata, Taylor, Univ. Kansas Sci. Bull., 25:371, July 10, 1939. Apatzingan; Capirio; Cofradia. Specimens from the three mentioned localities have the dark nuchal spot bordered anteriorly by a pale blotch. In life the speci- men from Capirio (UMMZ 114424) had rich reddish brown dorsal spots; the dorsal ground color was grayish white above and some- what more gray laterally. The pale nuchal area was a cream-color, and the iris was grayish red. All of the specimens were found in the arid scrub forest in the Tepalcatepec Valley at elevations between 200 and 350 meters. Imantodes gemmistratus gracillimus (Giinther) Dipsas gracillima Giinther, Biol. Centrali-Americana, Rept., p. 177, July, 1895.—southern México. Type locality restricted to Acapulco, Guerrero, México, by Smith and Taylor (1950a:331). Imantodes gemmistratus gracillimus, Zweifel, Amer. Mus. Novitates, 1961:12, September 16, 1959. La Orilla. The specimen from La Orilla was reported by Peters (1954:23) as Imantodes gemmistratus oliveri; Zweifel (1959c) showed that I. g. AMPHIBIANS AND REPTILES OF MICHOACAN 101 oliveri did not range west of Tehuantepec and that the snakes in- habiting the coastal lowlands of Guerrero, Michoacan, and Colima were assignable to the subspecies gracillimus. It may be assumed that this subspecies ranges throughout the coastal lowlands and foot- hills of the Sierra de Coalcoman. Imantodes gemmistratus latistratus (Cope) Dipsas gemmistrata latistrata Cope, Bull. U.S. Natl. Mus., 32: 68, 1887.— Southern Jalisco. Type locality restricted to Guadalajara, Jalisco, México, by Smith and Taylor (1950a:334). Imantodes gemmistratus latistratus, Zweifel, Amer. Mus. Novitates, 1961:3, September 16, 1959. El! Sabino. The one specimen from Michoacan was collected near the upper limits of the scrub forest on the slopes of the Cordillera Volcdnica. Zweifel (1959c:10) stated that in certain aspects of coloration this specimen was like I. gemmistratus gracillimus, but in scutellation and other features of coloration it was like I. g. latistratus. There are too few specimens of this species to define the ranges of the various subspecies with any degree of accuracy, but from the limited num- ber of specimens available, it seems that I. gemmistratus gracillimus occurs on the Pacific lowlands from Guerrero northward to Colima. Northward on the Pacific lowlands from Colima to Sinaloa and in the Balsas-Tepalcatepec Basin is found I. gemmistratus latistratus. Lampropeltis doliata (Linnaeus) Coluber doliatus Linnaeus, Systema naturae, ed. 12, 1:379, 1766.—Charles- ton, South Carolina. Lampropeltis doliata, Klauber, Copeia, No. 1:11, April 15, 1948. Coalcomin (3); El Sabino; 24 km. W of Morelia; Rio Nexpa; Uruapan. The few specimens of this species from Michoacdn show a wide range of variation; furthermore, the present systematic status of the subspecies of Lampropeltis doliata portrays an incongruous pattern of distribution. Specimens from the Sierra de Coalcoman have rela- tively narrow red bands that are not interrupted dorsally by exten- sions of the black rings; the scales in the red bands have black tips. The specimen from El] Sabino (EHT-HMS 5253) and the one from the Rio Nexpa on the coast (USNM 31491) have broader red bands; the scales in the red bands do not have black tips. A specimen from 24 kilometers west of Morelia (UIMNH 17782) and one from Urua- pan (UMMZ 121508) have the red bands interrupted dorsally by extensions from the black rings. Specimens from the Sierra de Coalcoman were referred to L. doliata blanchardi by Peters (1954:24), who noted that in some 102 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. characters these snakes were like L. d. nelsoni and in others like L. d. polyzona. The individual from El] Sabino was referred to L. d. nel- soni by Taylor (1940c:465); the one from 24 kilometers west of Morelia was referred to L. d. arcifera by Smith (1942c:198). If these assignments are correct, three subspecies of Lampropeltis doliata occur in Michoacan: blanchardi in the Sierra de Coalcoman, nelsoni on the coast and in the Tepalcatepec Valley, and arcifera on the Mexican Plateau and in the Cordillera Volcanica. Such a distribu- tion is plausible, but the few specimens and our general lack of knowledge of the variation and relationships of the different popula- tions do not permit a definite assignment at this time. Lampropeltis ruthveni Blanchard Lampropeltis ruthveni Blanchard, Occ. Pap. Mus. Zool. Univ. Michigan, 81:8, April 28, 1920.—P4tzcuaro, Michoacan, México. Morelia; Patzcuaro; Tacicuaro. At the present time this species is known definitely from only three localities on the Mexican Plateau in Michoacén. An incomplete skin from El] Sabino (EHT-HMS 5438) was referred to this species by Taylor (1940c:465); the specimen cannot be found, so verification of the identification cannot be made at this time. Leptodeira latifasciata (Giinther) Hypsiglena latifasciata Giinther, Biologia Centrali-Americana, Reptilia, p. 138, October, 1894.—Southern México. Type locality restricted to Huajintlan, Morelos, México, by Smith and Taylor (1950a:3381). Henedss latifasciata, Dunn, Proc. Natl. Acad. Sci., 22:696, December, Apatzingan; El Sabino; La Playa; 82 km. E of Nueva Italia. This nocturnal snake apparently ranges throughout the arid Balsas-Tepalcatepec Valley to elevations of about 1050 meters. It has been collected only in the arid scrub forest. Aside from the specimens listed by Duellman (1958a:93), there is one (UMMZ 120223) having eight body blotches, a body length of 510 mm. and a tail length of 108 mm. Leptodeira maculata (Hallowell) Megalops maculatus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 13:488, March 31, 1862.—“Tahiti.” Type locality restricted to Manzanillo, Colima, México, by Duellman (1958a:54). Leptodeira maculata, Duellman, Bull. Amer. Mus. Nat. Hist., 114:53, February 24, 1958. Aguililla (2); Apatzingan (24); Arteaga (2); Capirio (3); Charapendo (2); Coahuayana (3); Cofradia; Cuatro Caminos; La Placita (3); Lom- bardia (69); Nueva Italia (29); Pémaro; Rio Marquez, 10 km. S of Lom- bardia (2); Salitre de Estopila; Tafetan (2); Volcan Jorullo. AMPHIBIANS AND REPTILES OF MICHOACAN 103 This snake is abundant in the arid Tepalcatepec Valley; most of the specimens have been collected in arid scrub forest at eleva- tions of less than 500 meters. With the onset of the rains in late June and early July, large numbers of these snakes can be found around temporary pools, where they feed on small frogs and toads. In the dry season few individuals were found, and all of those were beneath cover. Specimens from the coast have more body- blotches than do those from the Tepalcatepec Valley (Duellman, 1958a:56); otherwise the snakes show little variation. Leptodeira splendida bressoni Taylor Leptodeira bressoni Taylor, Univ. Kansas Sci. Bull., 25:321, July 10, 1939.—Hacienda El Sabino, Michoacan, México. Leptodeira splendida bressoni, Duellman, Bull. Amer. Mus. Nat. Hist., 114:84, February 24, 1958. Coalcoman (3); El Sabino (3); Uruapan (5). The range of Leptodeira splendida bressoni apparently does not overlap that of Leptodeira maculata; the latter is restricted to the lower reaches of the arid scrub forest, whereas L. s. bressoni in- habits the upper limits of the arid scrub forest and the lower part of the pine-oak forest. Specimens have been collected between 950 and 1630 meters on the slopes of the Cordillera Volcanica and at 950 meters in the Sierra de Coalcoman. At Uruapan individuals were found beneath rocks along a stream and in a stone fence. Leptodeira duellmani, which was described from Coalcoman by Peters (1954:25), is an aberrant individual of L. s. bressoni (Duell- man, 1958a:56). Leptophis diplotropis (Giinther) Ahaetulla diplotropis Giinther, Ann. Mag. Nat. Hist., ser. 4, 9:25, 1872.— Tehuantepec, Oaxaca, México. Leptophis diplotropis, Bocourt, Mission scientifique au Mexique et dans l’Amerique Centrale, Reptiles, livr. 15:835, 1897. Between Aguililla and Dos Aguas; Arteaga; Coalcom4n; El Diezmo; El Sa- bino (5); La Playa; Ocorla. Most specimens of this species have been collected in tropical semi-deciduous forest at elevations of less than 1000 meters. In the Sierra de Coalcoman one was taken in pine-oak forest at an elevation of 1700 meters near Ocorla; another was found in broad-leafed forest between Aguililla and Dos Aguas at an elevation of 1600 meters. Most individuals have been seen in trees or bushes. The absence of broad-leafed forest in the Tepalcatepec Valley probably accounts for the absence of this snake in that area. 104 Universiry OF Kansas Pusis., Mus. Nat. Hist. Manolepis putnami (Jan) Dromicus putnami Jan, Elenco sistematico degli Ofidi, p. 67, 1863.—San Blas, Nayarit, México. Manolepsis putnami, Smith and Taylor, Bull. U.S. Natl. Mus., 187:92, Octo- ber 5, 1945. La Placita (3); Maquili; Ostula. In Michoacan the species has been found only in tropical semi- deciduous forest on the lower slopes of the Sierra de Coalcoman. From the observations made by Peters (1954:28), this snake is diurnal and feeds on teiid lizards. Masticophis striolatus striolatus Mertens Coluber striolatus Mertens, Zoologica (Stuttgart), 32:190, 1934.—Substitute name for Coluber lineatus Bocourt, a secondary homonym of Coluber lineatus Linnaeus = Lygophis lineatus. Type locality restricted to Presidio de Mazatlan, Sinaloa, México, by Smith and Taylor (1950a:843). Masticophis striolatus striolatus, Zweifel and Norris, Amer. Midl. Nat., 54: 2492, July, 1955. Apatzingin (4); Arteaga; Coalcoman (3); El Sabino; Jiquilpan; La Palma; La Playa (3); Lombardia; Nueva Italia; Rio Cachan; Santa Ana; Uruapan (2); Vole4n Jorullo; Ziracuaretiro. This large diurnal species inhabits open scrub forest and culti- vated terrain from sea level to about 1650 meters. On the Mexican Plateau it is known from the area around Lago de Chapala, to which it possibly gained access through the valleys in the headwaters of the Tepalcatepec drainage. Specimens from southern Michoacan have been reported previously by Peters (1954:28) and Duellman (1954b: 16) as Masticophis flagellum lineatus. Masticophis taeniatus australis Smith Masticophis taeniatus australis Smith, Jour. Washington Acad. Sci., 31:390, September 11, 1941.—Guanajuato, Guanajuato, México. Tacicuaro (2); Zamora. This species reaches the southern limit of its distribution in the state of Michoacan. The limited ecological data available suggest that the species inhabits the open mesquite grassland of the Mexican Plateau. Oxybelis aeneus auratus (Bell) Dryinus auratus Bell, Zool. Jour., 2:324, 1825.—México. Type locality re- en to Tehuantepec, Oaxaca, México, by Smith and Taylor (1950a: 340). Oxybelis aeneus auratus, Bogert and Oliver, Bull. Amer. Mus. Nat. Hist., 83:381, March 80, 1945. Coahuayana; El Sabino (4); between Las Tecatas and Las Higuertas; be- tween Los Pozos and La Ciénega; Playa Azul; Pémaro (2); between Pémaro and Maruata (2); Punto San Telmo; Rio Tizupan. On the basis of the number of specimens seen and collected on the seaward slopes of the Sierra de Coalcoman, this is a common snake AMPHIBIANS AND REPTILES OF MICHOACAN 105 there. Most specimens were collected in tropical semi-deciduous forest; others were collected in oak forest to an elevation of 1700 meters. Apparently Oxybelis does not inhabit the lower parts of the Tepalcatepec Valley; the only specimens from the inland area are four from El Sabino, which is situated at about 900 meters on the slopes of the Cordillera Volcinica. One individual was seen in gallery forest near Limoncito at an elevation of 730 meters on the northern slopes of the Sierra de Coalcoman. Pituophis deppei deppei (Duméril) Elaphis deppei Duméril, Mem. Acad. Inst. France, 23:453, 1835.—Meéxico. Type locality restricted to San Juan Teotihuacdn, México, México, by Smith and Taylor (1950a:334). Pituophis deppei deppei, Stull, Occ. Pap. Mus. Zool. Univ. Michigan, 250:1, October 12, 1932. Carapan (2); Morelia; Tacambaro; Tacicuaro; Zacapu. Duellman (1960b) showed that the widespread species Pituophis deppei was composite and that the “lined subspecies” actually represented another species, Pituophis lineaticollis. Pituophis dep- pei occurs only on the Mexican Plateau; in Michoacan it inhabits mesquite grassland and oak-bunch grass associations between 1900 and 2200 meters. Pituophis lineaticollis lineaticollis (Cope) Arizona lineaticollis Cope, Proc. Acad. Nat. Sci. Philadelphia, 13:300, December 28, 1861.—Southern Mexican Plateau. Type locality re- stricted to 24 kilometers northwest of Oaxaca, Oaxaca, México, by Duellman (1960b:607). Pituophis lineaticollis lineaticollis, Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 10:607, May 2, 1960. Acuaro de las Lleguas; Dos Aguas (8); Morelia; Tancitaro (5). This species reaches the northern limits of its range in the Sierra de Coalcoman and on the Mexican Plateau in Michoac4n. On the plateau it has been collected in mesquite grassland at eleva- tions between 1500 and 2000 meters. In the Sierra de Coalcoman individuals were found in open pine-oak forest at 2100 meters elevation and in a meadow surrounded by pine-oak forest at 2300 meters. Pseudoficimia frontalis (Cope) Toluca frontalis Cope, Proc. Acad. Nat. Sci. Philadelphia, 16:167, Septem- ber 30, 1864.—Colima, Colima, México. Pseudoficimia frontalis, Giinther, Biologia Centrali-Americana, Reptilia, p. 96, May, 1893. Apatzingan; Coalcoman (6); El Sabino (2). Most specimens were found beneath rocks in grassy areas near the upper limits of the arid scrub forest, both in the Sierra de 106 UNIVERSITY OF KANnsAS PuBLs., Mus. Nat. Hist. Coalcoman and on the southern slopes of the Cordillera Volcanica; all are from elevations of less than 1100 meters. One specimen was found on a road at night near Apatzingan. This species has been found in similar habitats near Huajintlan, Guerrero, and in arid scrub forest at lower elevations in Colima. It is unknown from the coast of Michoacan. Pseudoficimia pulcherrima Taylor and Smith Pseudoficimia pulcherrima Taylor and Smith, Univ. Kansas Sci. Bull., 28:246, May 15, 1942.—Huajintlan, Guerrero, México. Apatzingan. This specimen (CNHM 39208) was reported by Schmidt and Shannon (1947:81); they stated that it was a paratype of P. pul- cherrima. However, Taylor and Smith (1942a:246) did not men- tion the specimen; aside from the type (EHT-HMS 5497), the only other specimen they designated as belonging to the type series was UMMZ 85711 from Chilpancingo, Guerrero. The taxonomic validity of Pseudoficimia pulcherrima remains doubtful, for only minor characters distinguish it from P. frontalis. Furthermore, all known specimens of P. pulcherrima are from within the geographic range of P. frontalis. Rhadinaea hesperia hesperia Bailey Rhadinaea hesperia Bailey, Occ. Pap. Mus. Zool. Univ. Michigan, 412:8, May 6, 1940.—Omilteme and Sierra de Burro, Guerrero. Type locality eas to Omilteme, Guerrero, México, by Smith and Taylor (1950a: 832). Rhadinaea hesperia hesperia, Smith, Proc. Biol. Soc. Washington, 55:185, December 81, 1942. Arteaga (8); Coalcom4n; El Sabino (2); Uruapan; Volcan Jorullo (2). One specimen from Volcan Jorullo (UMMZ 104494), three from Arteaga (UMMZ 119281), and one from Uruapan (UMMZ 92342) are typical of the subspecies R. h. hesperia in possessing a lateral cream-colored line on the sixth and parts of the fifth and seventh dor- sal scale rows and in lacking a dark line on the second scale row. The specimens from E] Sabino (EHT-HMS 5441 and UIMNH 18933) and one from Coalcoman (UMMZ 104502) have the cream- colored line on the sixth and adjacent parts of the fifth and seventh scale rows and have a dark line on the second scale row. Another individual from Volcan Jorullo (UMMZ 104682) has cream-colored lines like the others, but it possesses two lateral dark lines, one on the second scale row, and one on the third. AMPHIBIANS AND REPTILES OF MICHOACAN 107 Smith (1942d:186) diagnosed Rhadinaea hesperia hesperioides as differing from the nominal subspecies in having the cream-colored line on the fourth and fifth scale rows and in possessing a dark line on the second scale row. The specimens seen all have the lateral cream-colored line centered on the sixth scale row, as is character- istic of R. h. hesperia. Although many of the specimens also possess a dark line on the second scale row, these specimens are here assigned to R. h. hesperia. Additional specimens are necessary to define accurately the subspecies and their ranges. Peters (1954:29) assigned the specimens from Coalcoman to R. h. hesperioides. In life the specimens from Arteaga had bright cream-colored tem- poral stripes and dorsolateral stripes on the anterior part of the body. The chin and anterior one-sixth of the belly was white; posteriorly the venter was bright orange-red. In Michoacan this snake has been found in tropical semi-decid- uous forest, arid scrub forest, and pine-oak forest at elevations from 850 to 1500 meters. Rhadinaea laureata (Giinther) Dromicus laureatus Giinther, Ann, Mag. Nat. Hist., ser. 4, 1:419, 1868.— Mexico City. Rhadinaea laureata, Boulenger, Catalogue Snakes . . . British Museum, 2, p. 179, September 23, 1894. op ee Carapan (8); Cherian (3); Paracho (2); Patzcuaro; Tancitaro This snake is abundant in the Cordillera Volcadnica, but it is un- known in the mountains to the northeast of Morelia or in the Sierra de Coalcoman. Most specimens were found beneath volcanic rocks imbedded in the ashy soil in pine forest between 1800 and 2300 meters. Rhadinaea taeniata (Peters) Dromicus taeniatus Peters, Monats, Akad. Wiss. Berlin, p. 275, 1863.— México. Rhadinaea taeniata, Bailey, Occ. Pap. Mus. Zool. Univ. Michigan, 412:14, May 6, 1940. Tancitaro (2). This species, which is known only from a small region in the mountains of Jalisco and central Michoacan, is represented by two specimens (CNHM 37130 and 890380) collected at Tancitaro (see Schmidt and Shannon, 1947:80). 8—7817 108 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Salvadora bairdi Jan Salvadora Bairdii Jan. Icon. gener. ophid., livr. 2, pl. 3, fig. 2, 1860.— México. Type locality restricted to Acambaro, Guanajuato, México, by Smith and Taylor (1950a:830). Barranca Seca; Carapan; Cerro San Andrés; Cojumatlin (2); Jiquilpan; Morelia; Patzcuaro (4); Quiroga; Sahuayo (2); Tacicuaro (12); Tan- citaro (56); Uruapan (2); Zacapu (2); between Zitacuaro and Tuxpan (3). This species is abundant on the Mexican plateau, where it inhabits the more grassy areas in the mesquite grassland and cutover land in the pine forests from 1550 to 2500 meters. Davis and Dixon (1957:21) described a specimen from Zacapu as having two dark paravertebral stripes diverging on the temporals and extending through the eye onto the loreal, a characteristic of Salvadora lineata. On the basis of this specimen, Davis and Dixon suggested that Salvadora bairdi and S. lineata were subspecifically related. The examination of the large number of specimens from Michoacan has revealed this kind of coloration in only one other specimen, an individual from Tacicuaro, in which the stripes diverge, but do not extend through the eye onto the loreal, Data on scutella- tion for the large series from Tancitaro were given by Schmidt and Shannon (1947:78), and for the series from Tacicuaro by Smith (1943:466). Salvadora mexicana (Duméril, Bibron, and Duméril) Zamenis mexicanus Duméril, Bibron, and Duméril, Erpétologie générale, 7 (pt. 1), p. 695, 1854.—Cape Corrientes, Jalisco, México. Salvadora mexicana Giinther, Ann. Mag. Nat. Hist., ser. 3, 12:349, 1863. Apatzingan (12); Capirio (2); El Sabino (5); Huetamo; La Placita; La Playa (4); Lombardia; Nueva Italia; Ojos de Agua de San Telmo; Oropeo; Rio Cancita, 14 km. E of Apatzingan; Santa Ana. This is one of the most abundant snakes in the arid lowlands of the Tepalcatepec Valley; observations indicate that it probably is equally abundant on the coastal lowlands. Near Apatzingin as many as five of these snakes have been seen in one-half hour. The snakes seem to be equally abundant and active in the dry season and in the rainy season. Most individuals were seen on the ground, but two were found in low trees. On several occasions Salvadora mexicana was observed in pursuit of lizards on the ground. Cap- tured individuals regurgitated Cnemidophorus costatus zweifeli, Cnemidophorus deppei infernalis, Sceloporus horridus oligoporus, Sceloporus pyrocephalus, and Urosaurus gadowi. Salvadora mexicana inhabits only the arid scrub forest at eleva- tions from sea level to about 1000 meters. AMPHIBIANS AND REPTILES OF MICHOACAN 109 Sibon nebulatus (Linnaeus) Coluber nebulatus Linnaeus, Systema naturae, ed. 10, 1, p. 222, 1758.— rica (in error). Type locality restricted to Jicaltepec, Veracruz, México, by Smith and Taylor (1950a:349). Sibon nebulatus, Taylor, Univ. Kansas Sci. Bull., 26:473, November 27, 1940, Aquila. The one specimen from Michoacan was collected by Peters (1954: 30) in tropical semi-deciduous forest on the coastal foothills of the Sierra de Coalcomin. As presently known, the range of this species in western México extends from Chiapas to Nayarit. Throughout this region the species avoids scrub forest; this may explain its absence in the Balsas-Tepalcatepec Valley. Sonora michoacanensis michoacanensis (Dugés) Contia michoacanensis Dugés, in Cope, Proc. Amer. Philos. Soc., 22:178, 1885.—Michoacan. Type locality restricted to Apatzingan, Michoacdn, México, by Smith and Taylor (1950a:335). Sonora michoacanensis michoacanensis, Stickel, Proc. Biol. Soc. Washing- ton, 56:116, October 19, 1943. Apatzingan (3); Coalcoman (8); 12 km. S of Tzitzio. These specimens, together with all known specimens from the Sierra del Sur in Guerrero (KU 28790-1, MVZ 45123) and the upper Balsas Basin in Puebla (UIMNH 41688), are referable to S. m. michoacanensis. The dorsal pattern consists of a highly variable number of cross-bands of red, white, and black. In the specimens from Michoacan there are as many as 17 red cross-bands on the body. One specimen from Apatzingan (CNHM 37141) has just behind the head a white band, bordered on either side by a narrow black band; posteriorly the body is uniform red. Two specimens from Coalcoman (UMMZ 109905-6) have respectively 11 and 13 red cross-bands and 20 and 17 white cross-bands, and the posterior part of the body is devoid of red color. Other specimens from these localities have red, black, and white cross-bands throughout the length of the body. Sonora michoacanensis michoacanensis is distinguished from S. michoacanensis mutabilis by the presence of cross-bands on the tail in the latter (Stickel, 1943:116). One specimen from Coal- coman (UMMZ 109904) has one narrow band on the tail; all others from Michoacan have uniformly red tails. Apparently Sonora michoacanensis michoacanensis ranges in semi- arid and arid habitats from the upper Balsas Basin in Puebla west- ward to the lower slopes of the Sierra de Coalcoman, whereas S. m. 110 UNIVERSITY OF KANsAsS Pusts., Mus. Nat. Hist. mutabilis lives in foothills of the Sierra Madre Occidental from southern Jalisco to Nayarit. Zweifel (1959b:6) presented evidence to show that specimens of S. m. mutabilis supposedly from “Distrito Federal” probably bear erroneous locality data. Tantilla bocourti (Giinther) Homalocranium bocourti Gimnther, Biologia Centrali-Americana, Reptilia, p. 149, 1895.—Guanajuato, Guanajuato, México. Tantilla bocourti, Cope, Amer. Nat., 30:1021, December, 1896. Carapan; Patzcuaro (2); between Zitaécuaro and Rio Tuxpan (11). This small snake is an inhabitant of the coniferous forests and the pine-oak forests on the Cordillera Volcdnica. Data on the series from between Zitacuaro and the Rio Tuxpan were given by Taylor (1940c:481). Tantilla calamarina Cope Tantilla calamarina Cope, Proc. Acad. Nat. Sci. Philadelphia, 18:320, Feb- ruary 13, 1867.—Guadalajara, Jalisco, México. Apatzingan; La Placita. Although this snake has been collected at high elevations along the rim of the Mexican Plateau in Nayarit, Jalisco, México, and Puebla, the specimens from Michoacan are from arid scrub forest at elevations of less than 400 meters. The species has been found in similar habitats in Colima (Oliver, 1937:24) and in Sinaloa and the Tres Marias Islands (Zweifel, 1960:110). Toluca lineata lineata Kennicott Toluca lineata Kennicott, in Baird, Report on the United States and Mexican boundary survey, 2, Reptiles, p. 28, 1859.—Valley of México. Toluca lineata lineata, Taylor and Smith, Univ. Kansas Sci. Bull., 28:348, May 15, 1942. Capacuaro; Carapan (12); Cheran (23); Cojumatlin; Los Reyes; Morelia (2); Nahuatzen; Paracho (10); Patzcuaro (17); Uruapan (2). This small snake is an inhabitant of the coniferous forests between elevations of about 1550 and 2800 meters. Not infrequently, indi- viduals have been found in pine-oak forest within these elevations. The generic status of Toluca is unsettled. Taylor and Smith (1942b) separated Toluca from Conopsis by the presence of en- larged and grooved posterior maxillary teeth in Toluca and their absence in Conopsis. Bogert and Oliver (1945:378) suggested syn- onymizing Toluca with Conopsis. Smith and Laufe (1945:12) de- fined the generic position of Toluca. Actually, in deciding the generic position of these snakes, five genera (Ficimia, Gyalopion, Pseudo- ficimia, Conopsis, and Toluca) must be considered. Of these AMPHIBIANS AND REPTILES OF MICHOACAN lll Ficimia and Gyalopion are closely related; they have been placed in one genus by some workers. Pseudoficimia is intermediate between Ficimia-Gyalopion and Toluca-Conopsis. A workable definition of the supraspecific classification of these snakes must await a thorough review of the species. Trimorphodon biscutatus biscutatus (Duméril, Bibron, and Dumeéril) Dipsas biscutata Duméril, Bibron, and Duméril, Erpétologie générale, 7 (pt. 2):1153, 1854.—México. Type locality restricted to Tehuantepec, Oaxaca, México, by Smith and Taylor (1950a:340). Trimorphodon biscutatus biscutatus, Smith, Proc. U.S. Natl. Mus., 91:159, November 10, 1941. Apatzing4n (11); Cofradia; Cuatro Caminos; El Sabino (2); La Placita; La Playa (2); Lombardia (2); Nueva Italia (2); Rio Tepalcatepec, 27 km. S of Apatzingan; Tafetan. In the arid lowlands of the Tepalcatepec Valley and presumably also in the scrub forest of the coastal lowlands, this is an abundant snake, which is active only at night. Usually snakes of this species are found on the ground, but one large individual was observed at night in a low tree. That individual defied capture by widely open- ing its mouth and striking repeatedly at the collector. The excreta of one specimen contained feathers of an unidentified species of bird. Trimorphodon latifascia Peters Trimorphodon biscutata latifascia Peters, Monats. Akad. Wiss. Berlin, p. 877, 1869.—Puebla, México. Type locality restricted to Izicar de Matamoros, Puebla, México, by Smith and Taylor (1950a:341). peter phedce latifascia, Taylor, Univ. Kansas Sci. Bull., 25:364, July 10, 39. Apatzingan (5); Casada Tzararacua; Coalcoman (2); Lombardia; 14 km. S of Lombardia; Nueva Italia; San Salvador. In Michoacan this species has been collected in semi-arid habitats at elevations from 300 to 1480 meters in the Tepalcatepec Valley and lower slopes of the Cordillera Volcdnica. In this area it occurs sym- patrically with Trimorphodon biscutatus biscutatus. In life, adults have a pale tan dorsal ground color and rich chocolate brown cross-bands; the eye is pale grayish tan. A ju- venile from Coalcoman has black cross-bands on a pale grayish tan ground color. As stated by Schmidt and Shannon (1947:83) and Peters (1954:32), the type specimen of Trimorphodon fascio- lata Smith from Cascada Tzararacua is indistinguishable from specimens of Trimorphodon latifascia. Seven males have 209 to 223 (216.5) ventrals; one female has 227 ventrals. The number of dark cross-bands on the body varies from 12 to 16 (18.5). The relationships of this species are with ye UNIVERSITY OF KANSAS PuBts., Mus. Nat. Hist. Trimorphodon tau on the Mexican Plateau. In fact, additional specimens from the headwaters of the Tepalcatepec Valley and the lower slopes of the Mexican Plateau in eastern Michoacan and adjacent Jalisco may show that the two are conspecific. Tri- morphodon latifascia differs from tau in having fewer dark cross- bands on the body and in lacking an interocular bar. Trimorphodon tau Cope Trimorphodon tau Cope, Proc. Amer. Philos. Soc., 11:151, 1869.—Quuote- pec, Oaxaca, México. Emiliano Zapata (2); between Morelia and Ciudad Hidalgo; Tacicuaro; Tangamandapio. Two of the specimens from Michoacan (UMMZ 118948 from Tangamandapio and UIMNH 191388 from Tacicuaro) have cream- colored, Y-shaped marks on the head. These markings supposedly are characteristic of Trimorphodon upsilon. One specimen from Emiliano Zapata (UMMZ 118950) and one from between Morelia and Ciudad Hidalgo (EHT-HMS 21402) have a cream-colored line on the parietal suture; in another specimen from Emiliano Zapata (UMMZ 118949) the anterior end of this line is expanded, giving the appearance of an incipient “Y”. Thus, the nature of the mark- ings on the head in specimens from Michoacan is intermediate between the typical condition in Trimorphodon tau and the usual condition in T. upsilon. Smith and Taylor (1945:148) gave the range of Trimorphodon tau as: “Central Guerrero, in the Sierra Madre del Sur; central Oaxaca; and the edge of the plateau in central Michoacan.” They gave the range of Trimorphodon up- silon as: “Southern Chihuahua south to central Michoacan, east to central Hidalgo.” Specimens referable to T. taw have been found at La Joya de Salas, near Ciudad Victoria, and near Llera, Tamau- lipas (see Smith and Darling, 1952:85, and Martin, 1958:74). Some of these specimens also show combinations of characteristics of T. tau and T. upsilon. Smith and Darling (loc. cit.) suggested that T. tau and T. upsilon be considered as subspecies. However, if T. tau and T. upsilon are subspecies, intergrades would be ex- pected between the ranges of the two populations and not on the northeastern and southwestern periphery of their combined ranges. Instead, the limited evidence now available suggests that T. tau and T. upsilon are names based on a highly variable character of color pattern of the head, and that only one species is involved. In Michoacan this species inhabits the mesquite grassland on the Mexican Plateau. AMPHIBIANS AND REPTILES OF MICHOACAN 113 Tropidodipsas occidentala Oliver Tropidodipsas occidentala Oliver, Occ. Pap. Mus. Zool. Univ. Michigan, 860:20, November 20, 1937.—Comala, Colima, México. Coalcoman. This specimen was reported by Peters (1954:34), who found it beneath a rock at the mouth of a heavily wooded ravine near Coal- coman at an elevation of 950 meters. The only other known speci- men is from Comala, Colima, a village, like Coalcoman, that is lo- cated near the upper limits of the arid scrub forest. Natrix valida isabelleae Conant Natrix valida isabelleae Conant, Nat. Hist. Misc., 126:7, September 15, 1953. —Pie de la Cuesta, Laguna Coyuca, Guerrero, México. Coahuayana; Playa Azul (2); Punto San Juan de Lima. Three females and one male have, respectively, 183, 135, 135, and 131 ventrals, and 68, 68, 73, and 75 caudals. The grayish stippling on the posterior ventral surfaces mentioned by Conant (1953:9) is not visible on these specimens. In the small individuals from Punto San Juan de Lima and from Coahuayana there are four longitudinal rows of dark spots on the dorsum; in two large females from Playa Azul the spots are barely discernible. All of the specimens from Michoacén were found in the coastal lowlands; those from Playa Azul were collected from a small brack- ish, mangrove-lined lagoon. Storeria storerioides (Cope) Tropidoclonium storerioides Cope, Proc. Acad. Nat. Sci. Philadelphia, 17:190, December 26, 1865.—Mexican Plateau. Type locality restricted to Tres Cumbres, Morelos, México, by Smith and Taylor (1950a:336). Storeria storerioides, Garman, Mem. Mus. Comp. Zool., 8(3):29, June, 1888. Dos Aguas (11); Puerto de Garnica; Tancitaro (11); Tzitzio; Uruapan; 16 km. NW of Zacapu. Three males and six females from the Sierra de Coalcoman have, respectively, 122-128 (125.8) and 126-136 (180.0) ventrals, and 46-47 (46.7) and 38-42 (39.1) caudals. Four males and eleven fe- males from the Cordillera Volcdnica have, respectively, 124-182 (128.5) and 127-139 (136.4) ventrals, and 43-48 (44.7) and 38-44 (40.2) caudals. These data show that, although there is little dif- ference in the number of caudals, specimens from the Sierra de Coalcoman have fewer ventrals than do specimens from the Cordil- lera Volcanica. Of eleven specimens from the Sierra de Coalcoman, two have black bellies. Five others from the Sierra de Coalcomaén and one from Puerto de Garnica in the Cordillera Volcdnica have 114 UNIVERSITY OF Kansas PuBLs., Mus. Nat. Hist. the bellies heavily stippled with black, giving a gray appearance. Melanistic tendencies in this species have been discussed by Ander- son (1960:64), who examined the specimen from Tzitzio. In life, one specimen from Dos Aguas (UMMZ 119451) had a cream- colored belly; the edges of the ventrals were dark brick-red. In Michoacan this snake inhabits pine-oak, pine, and fir forests at elevations between 1550 and 2800 meters in the Cordillera Vol- canica and the Sierra de Coalcoman. Most specimens were found beneath rocks; the one from Tzitzio was removed from the stomach of a Mexican Motmot (Anderson, 1960:66). Thamnophis dorsalis cyclides Cope Thamnophis cyrtopsis cyclides Cope, Proc. Acad. Nat. Sci. Philadelphia, 13:299, December 28, 1861.—Cape San Lucas, Baja California (in error). Type locality restricted to Guanajuato, Guanajuato, México, by Smith and Taylor (1950a:330). Smith, Copeia, no. 2:140, June 8, 1951. Milstead, Texas Jour. Sci., 5:368, September, 1953. Thamnophis eques eques (nec. Reuss), Smith, Zoologica, 27:106, October 23, 1942. Bogert and Oliver, Bull. Amer. Mus. Nat. Hist., 83:356, March 30, 1945. Thamnophis vicinus Smith, Zoologica, 27:104, October 23, 1942.—Temaz- cal, Michoacan, México. Thamnophis dorsalis cyclides, Fitch and Milstead, Copeia, no. 1:112, March 961. Barolosa; Coalcoman; Dos Aguas (3); Los Reyes; Morelia (16); Opopeo; Pino Gordo; Tacicuaro (16); Tancitaro (14); Tangamandapio (2); Temaz- cal (2); Tzintzuntzan; Uruapan. The snakes comprising the former Thamnophis eques-group have undergone extensive taxonomic and nomenclatural shuffling by Smith (1942 and 1951), Bogert and Oliver (1945), Milstead (1953), and Fitch and Milstead (1961). Smith recognized in Michoacan three members of the T. eques (= dorsalis) complex: eques eques, eques postremus, and vicinus. Later, Smith (1951) showed that the specific name eques had been misapplied, so that T. eques eques became T. cyrtopsis cyclides, and T. eques postre- mus became T. cyrtopsis postremus; under this arrangement T. vi- cinus stood unchanged. In the meantime, Bogert and Oliver (1945:359) presented a reinterpretation of Smith’s data and sug- gested that T. vicinus, which differs from T. dorsalis cyclides only in lacking a middorsal stripe, “ is not a species, but only a pattern phase, possibly a simple mutant of T. e. eques” (=T. dorsalis cyclides, by present arrangement). Milstead (1953) agreed with Bogert and Oliver on the status of T. vicinus; further- more, on the basis of only a few specimens, Milstead concluded AMPHIBIANS AND REPTILES OF MICHOACAN 115 that T. cyrtopsis postremus was not subspecifically distinct from T. cyrtopsis cyclides. Recently, Fitch and Milstead (1961) showed that Thamnophis dorsalis Baird and Girard (1853) was the correct name for the snakes that had been recognized as Thamnophis cyrtopsis Kennicott (1860). Consequently, the snakes referred to T. eques eques by Smith (1942) and to T. cyrtopsis cyclides by Smith (1951) and Milstead (1958) are now T. dorsalis cyclides. Aside from one specimen from Temazcal and nine from Morelia (paratypes of T. vicinus), only two other specimens completely lacking the middorsal stripe have been seen; one is a male (UMMZ 102510) having 161 ventrals and an incomplete tail from Pino Gordo, and the other is a male (CNHM 39060) from Tancitaro having 158 ventrals and an incomplete tail. A female from Tan- ie 7 pos 275-3) WO Ee ae SPAS iy ae = ri fe) A Fy ie C8 arr, ae - et ‘ ae Bone eae HOGI ty Fic. 10. Dorsal color pattern of Thamnophis dorsalis cyclides (A) and Thamnophis dorsalis postremus (B). citaro (CNHM 39061) having 153 ventrals and 77 caudals has no lateral stripes and only a narrow middorsal stripe on the anterior part of the body. Throughout the region where T. vicinus-like snakes have been found, typical T. dorsalis cyclides occurs in much greater numbers. I concur with Bogert and Oliver in placing T. vicinus as a synonym of T. dorsalis cyclides. Milstead (1953) had available few specimens of Thamnophis dorsalis from the Tepalcatepec Valley. The large series now in existence shows that the population in the Tepalcatepec Valley differs distinctly from that inhabiting the Mexican Plateau, Cordil- lera Volcinica, and Sierra de Coalcomain. Therefore the name T. dorsalis postremus Smith (1942) is resurrected for the population 116 University OF Kansas Pusts., Mus. Nat. Hist. in the Tepalcatepec Valley. T. dorsalis cyclides and T. dorsalis postremus differ in color pattern (Fig. 10) and in scutellation (Table 6). Specimens from the Mexican Plateau and mountain ranges have a distinct light stripe on the second and third scale rows, a dark brown dorsum having squarish black spots, and a row of dark spots on the first row of dorsal scales. Specimens from the Tepalcatepec Valley have a grayish brown dorsum having smaller and less distinct dark spots and no light stripe on the second and third scale rows; the first, second, and third rows of scales are colored like the venter. In some specimens there are small dark flecks on the first row of dorsal scales. TABLE 6.—VARIATION IN SCUTELLATION IN THAMNOPHIS DORSALIS. Mexican Sierra de Tepalcatepec CHARACTER Plateau Coalcoman Valley Ventrals..... ol aN 31 2 32 Mean 164.0 156.5 144.6 Range 153-171 154-159 138-151 o N 19 2 32 Mean 153.5 154.7 138.3 Range 149-159 149-159 131-141 Caudals...... ao N 28 2 29 Mean 83.8 81.0 73.4 Range 80-100 79-83 70-79 2 ON 14 2 28 Mean 78.0 72.0 68.5 Range 71-87 de, 63-73 One specimen from Uruapan (1550 meters) and one from Coal- comin (950 meters) are intermediate in color pattern between T. dorsalis cyclides and T. dorsalis postremus. Both have indistinct lateral stripes and only small dark spots below the stripes. In scutellation these specimens are like T. dorsalis cyclides. In Michoacdin Thamnophis dorsalis cyclides has been collected in a variety of habitats on the Mexican Plateau: pine-oak forest, fir forest, marshes, and cleared land from 1550 to 2800 meters. In the Sierra de Coalcoman one was taken in broad-leafed forest at 950 meters, three in pine-oak forest at 2100 meters, and one in pine forest at 2300 meters. AMPHIBIANS AND REPTILES OF MICHOACAN Py Thamnophis dorsalis postremus Smith Thamnophis eques postremus Smith, Zoologica, 27:109, October 23, 1942.— El Sabino, Michoacan, México. Thamnophis cyrtopsis postremus Smith, Copeia, no. 2:140, June 8, 1951. Thamnophis cyrtopsis cyclides (part), Milstead, Texas Jour. Sci., 5:368, Sep- tember, 1953. Thamnophis dorsalis postremus, Fitch and Milstead, Copeia, no. 1:112, March 17, 1961. Apatzingin (31); Capirio (2); Charapendo; Cuatro Caminos (22); El Sa- bino; Lombardia (9); Nueva Italia (8); Uruapan (8). The reasons for recognizing the population of Thamnophis dorsalis in the Tepalcatepec Valley as distinct from that on the surrounding highlands are presented in the discussion of Thamnophis dorsalis cyclides. In certain features of coloration and in the low numbers of ventrals and caudals, T. dorsalis postremus shows more re- semblance to T. dorsalis sumichrasti than to T. dorsalis cyclides. According to Milstead (1953:367), T. dorsalis cyclides ranges south- ward from the Rio Balsas in southwestern México. If specimens could be obtained from the upper Balsas Basin they might show that T. dorsalis postremus inhabits that extensive basin. In the Tepalcatepec Valley T. dorsalis postremus is most fre- quently found at night in the rainy season, at which time the snakes are abundant near temporary pools where frogs are breeding. The absence of specimens from the coastal lowlands of Guerrero, Michoa- can, and Colima indicate that, although the species inhabits the low- lands of the Tepalcatepec Valley, its range does not include the coastal lowlands. A female (UMMZ 119402 from Cuatro Caminos) having 189 ventrals and a body length of 576 mm., on June 20, 1958, gave birth to 25 young, of which 18 (9 males and 9 females) were preserved. In body length the males varied from 182 to 141 (137.3) mm.; the females, 125 to 137 (133.1) mm. In tail length the males varied from 38 to 44 (42.4) mm.; females, 35 to 42 (39.7) mm. The males have 138 to 147 (142.2) ventrals and 70 to 75 (72.9) caudals; females have 181 to 140 (185.8) ventrals and 68 to 71 (67.0) caudals. Thamnophis eques eques (Reuss) Coluber eques Reuss, Zool. Misc., p. 152, 1834.—México. Type locality re- stricted to Guadalajara, Jalisco, México, by Smith and Taylor (1950a: Thamnophis macrostemma macrostemma, Smith and Taylor, Bull. U.S. Natl. Mus., 187:163, October 5, 1945. Thauncples subcarinata subcarinata, Smith, Herpetologica, 5:63, May 31, Thamnophis eques eques, Smith, Copeia, no. 2:139, June 8, 1951. Jiquilpan; Lago de Cuitzeo; Lago de P&atzcuaro (17); Patzcuaro (5); Tan- gancicuaro; Tupataro (2); Undameo; Zacapu. 118 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. Although this snake has been collected in open pine-oak forest and in oak-bunch grass associations, it seems to reach its greatest abun- dance in marshes on the Mexican Plateau at elevations of 1550 to 2300 meters. Thamnophis melanogaster canescens Smith Thamnophis melanogaster canescens Smith, Zoologica, 27:117, October 23, 1942.—Chapala, Jalisco, México. Lago de Cuitzeo (5); Lago de Patzcuaro; Patzcuaro; Tacicuaro; Tanga- mandapio (2). This species of garter snake seems to be most abundant in the marshes adjacent to the lakes on the Mexican Plateau in Michoacdn and Jalisco. At these elevations (1550 to 2200 meters) it often is found in association with Thamnophis eques eques and sometimes with Thamnophis dorsalis cyclides. On June 11, 1958, individuals of this species were found in a hyacinth-choked marsh at Tanga- mandapio at night. One specimen from Tangamandapio (UMMZ 119414) had, in life, a dark chocolate brown dorsum, reddish brown sides, and cream-colored belly, chin, and labials. There were no longitudinal dorsal stripes. Thamnophis scalaris scaliger (Jan) Tropidonotus scaliger Jan, Elenco sistematico degli Ofidi, p. 70, 1863.—No type locality designated. Type locality restricted to Mexico City, Dis- trito Federal, by Smith and Taylor (1950a:329). Thamnophis scalaris scaliger, Smith, Zoologica, 27:103, October 23, 1942. Cerro Tancitaro (2); Nahuatzen; Opopeo; 26 km. S of PAtzcuaro. The few specimens of this species from Michoacan have been col- lected at elevations from 1800 to 3400 meters in pine or fir forest in the Cordillera Volcanica. Micrurus distans michoacanensis (Dugés) Elaps diastema michoacanensis Dugés, La Naturaleza, ser. 2, 1:487, 1891. —RMichoacaén. Type locality restricted to Apatzingin, Michoacan, Mé- xico, by Smith and Taylor (1950a:885). Micrurus distans michoacanensis, Zweifel, Amer. Mus. Novitates, 1953:11, June 26, 1959. Apatzingan (6). All specimens were collected in the arid scrub forest of the Tepalcatepec Valley. The number of black rings on the body varies from six to eleven. In this respect they agree with the diagnosis of this subspecies presented by Zweifel (1959b:9). AMPHIBIANS AND REPTILES OF MICHOACAN 119 Micrurus laticollaris (Peters) Elaps marcgravii laticollaris Peters, Monats. Akad. Wiss. Berlin, p. 877, 1869.—Izucar de Matamoros, Puebla, México. Micrurus laticollaris, Schmidt, Publ. Field Mus, Nat. Hist., zool. ser., 20:39, December 11, 1933, El Sabino (2); Lombardia. This species ranges throughout the Balsas-Tepalcatepec Basin westward into Colima; specimens from Michoacan were collected in arid scrub forest at elevations from 500 to 1050 meters. The limited observations on Micrurus distans michoacanensis and M. laticollaris indicate that, at least in the Tepalcatepec Valley, M laticollaris seems to inhabit slightly more mesic areas than does M. distans michoacanensis. Pelamis platurus (Linnaeus) Anguis platura Linnaeus, Systema naturae, ed. 12, 1:391, 1766.—Pine Is- land, Pacific Ocean. Pelamis platurus, Gray, Ann. Philos., p. 15, 1825. Boca de Apiza. In November, 1955, Alfonzo Gonzales, a geographer from the University of Texas, observed sea snakes on the beaches of Michoa- can, In May, 1956, Donald D. Brand of the University of Texas gave me one specimen of Pelamis platurus that he obtained on March 2, 1956, at Boca de Apiza. Furthermore, he supplied me with the following observations based on his field work along the coast of Michoacan from the Rio Coahuayana to Maruata from March 1, to April 15, 1956. At that time many sea snakes were observed; in some places living and dead individuals were seen on the beaches; innumerable snakes were seen in the surf. When live individuals were taken from the beach and thrown into the ocean, they usually swam to shore. Many partially eaten indi- viduals were seen protruding from crab holes. Inquiries among the natives resulted in the following information: Sea snakes are fre- quently seen between November and April, but most commonly in March and April, at which time the water is cold. The natives referred to the sea snakes as “culebra del mar.” Most natives said that the snakes were not poisonous; others did not know of any venomous properties. In May, 1956, I worked the coastal region from the Rio Coahuayana to La Placita and saw no sea snakes. In the summer of 1950 James A. Peters, and in the summer of 1951 I worked nearly the entire coastal region of Michoacdn; during 120 University oF Kansas Pusts., Mus. Nat. Hist. that time no Pelamis were seen. Insofar as I know, this is the first report of such seasonal activity in Pelamis platurus in the Americas. Agkistrodon bilineatus bilineatus Giinther Ancisdrodon bilineatus Giinther, Ann. Mag. Nat. Hist., ser. 3, 12:364, 1863. —Pacific coast of Guatemala. Agkistrodon bilineatus bilineatus, Burger and Robertson, Univ. Kansas Sci. Bull, 34 (1):218, October 1, 1951. Apatzinga4n; El Sabino; La Playa; Los Reyes. All specimens from Michoacan are from inland localities between 300 and 1500 meters. The one from Los Reyes (USNM 46416) was collected by Nelson and Goldman on February 13, 1903. The elevation of Los Reyes (1500 meters) seems unusually high for this species, but otherwise there is no reason to doubt the authenticity of the record. Goldman (1951:192) in his description of Los Reyes stated: “Los Reyes is near the boundary between the Lower Austral and Arid Upper Tropical Zones but is preponderantly trop- ical in zonal character. The regular crops are mainly sugar cane, rice, and corn.” ‘Thus the biotic features of the area are not noticeably different from those at El Sabino and La Playa at lower elevations. The development of extensive agriculture through ir- rigation in the Tepalcatepec Valley and planting of rice and sugar- cane in that area may produce a more widespread habitat for this snake. The absence of specimens from the coastal lowlands is due solely to inadequate collecting; the natives there know the snake and re- port that it is not uncommon in certain areas, Crotalus basiliscus basiliscus (Cope) Caudisonia basilisca Cope, Proc. Acad. Nat. Sci. Philadelphia, September 30, 1864.—Colima. Type locality restricted to Colima, Colima, México, by Smith and Taylor (1950a:828). Crotalus basiliscus basiliscus, Gloyd, Nat. Hist. Misc., 17:1, April 28, 1948. Apatzingan (4); Camachines; Coalcoman; El Ticuiz. Specimens from southern Michoacén have fewer ventrals and caudals than do those from the northern part of the range; three males and three females have, respectively, 178, 182, 182, 185, 186, and 188 ventrals, and 27, 28, 29, 22, 29, and 29 caudals. Klauber (1952:81) gave the following data for Crotalus basiliscus (based on specimens from the entire range, except Oaxaca): ventrals in males, 179-201 (191.4), in females, 185-206 (197.6); caudals in males, 26-86 (30.7), in females, 21-29 (24.4). Klauber (1952:84) re- marked that the one specimen that he had seen from Apatzingan AMPHIBIANS AND REPTILES OF MICHOACAN 1A had fewer ventrals and caudals than most other specimens. The low numbers of ventrals and caudals in specimens from Michoacan, as compared with more northern populations, may be indicative of a trend in the reduction of the numbers of these scutes from north to south. The southernmost examples of Crotalus basiliscus (Cro- talus basiliscus oaxacus from Oaxaca) have 172-175 ventrals and 21 caudals (Gloyd, 1948). In Michoacan Crotalus basiliscus basiliscus has been found in arid habitats on the coast, in the Tepalcatepec Valley, and in the lower parts of the Sierra de Coalcoman. All specimens are from localities below 1070 meters in elevation. Crotalus durissus culminatus Klauber Crotalus durissus culminatus Klauber, Bull. Zool. Soc. San Diego, 26:65, August 8, 1952.—E] Sabino, Michoacan, México. El Sabino (18). These specimens are part of the type series and were collected by Hobart M. Smith near the upper limits of the arid scrub forest at an elevation of about 1050 meters on the lower slopes of the Cordillera Volcanica at the northern edge of the Tepalcatepec Val- ley. They were discussed in detail by Klauber (1952:66-70). Crotalus intermedius intermedius Troschel Crotalus intermedius Troschel, in von Miiller, Reisen in Vereiningten Staaten, Canada und Mexico, vol. 8, p. 618, 1865.—Type locality unknown. Crotalus intermedius intermedius, Klauber, Bull. Zool. Soc. San Diego, 26:9, August 8, 1952. Cerro Tancitaro. The one specimen is from the pine forests on the Cordillera Vol- canica. At the present time this species is known from scattered localities in west-central Veracruz, Oaxaca, Michoacan, and as Cro- talus intermedius omiltemanus in Central Guerrero. Apparently it is restricted to montane environments. Crotalus molossus nigrescens Gloyd Crotalus molossus nigrescens Gloyd, Occ. Pap. Mus. Zool. Univ. Michigan, aie January 28, 1936.—Four miles west of La Colorada, Zacatecas, xiCO, Carapan; Los Conejos; Patzcuaro; Tacicuaro (5). In Michoacan this species has been found in pine forests between 1550 and 2300 meters in the Cordillera Volcdnica. I expected to find it in the Sierra de Coalcoman, but inquiries among the natives living in the pine forests of that mountain range revealed that the people there have no knowledge of a large species of rattlesnake. 122 UNIVERSITY OF KANSAS PuBts., Mus. Nar. Hist. Crotalus polystictus (Cope) Caudisonia polysticta Cope, Proc. Acad. Nat. Sci. Philadelphia, 17:191, December 26, 1865.—Tableland of México. Type locality restricted to Tupataro, Guanajuata, México, by Smith and Taylor (1950a:330). Crotalus polystictus Cope, in Yarrow, Wheeler’s Rept. Geog. Geol. Expl. Surv. W. 100th. Mer., vol. 5, p. 533, 1875. Tacicuaro (4); Tupataro (2). Formerly this species was abundant in the marshes around Lago de Chapala. The draining of these marshes probably resulted in reducing the numbers of these rattlesnakes. The species is known only from the Mexican Plateau at elevations of 1450 to 2400 meters. Crotalus pusillus Klauber Crotalus pusillus Klauber, Bull. Zool. Soc. San Diego, 26:34, August 8, 1952.—Tancitaro, Michoacan, México. ies de las Lleguas (2); Carapan; Cerro Tancitaro (16); Dos Aguas Aside from the type series of Crotalus pusillus from Cerro Tan- citaro and one specimen from Carapan referred to the species by Klauber (1952:38), there are fourteen specimens from the Sierra de Coalcoman. These specimens (UMMZ 112566-7, 118591-9, 118601, 121512-3) are like Crotalus pusillus from Cerro Tancitaro in having the prefrontals paired, a black proximal rattle, and the underside of the tail black. The prefrontals are bordered posteri- orly by one scale in two specimens, by two scales in three speci- mens, and by three scales in the other nine. The snakes from the Sierra de Coalcoman have 40 to 46 (42) dorsal body blotches. Ten males have 150-158 (154.4) ventrals and 29-33 (31.0) caudals; two females have 157 and 160 ventrals, and 25 and 27 caudals. The largest specimen is a male having a body length of 545 mm. and a tail length of 63 mm. The only noticeable difference be- tween the specimens from the Sierra de Coalcoman and the topo- typic series is that the latter have fewer dorsal blotches; the range of variation is 83 to 46 (39.8). Most specimens of this species have a grayish brown dorsum and dark brown dorsal blotches. Two specimens from Dos Aguas (UMMZ 118596 and 118599) are pale brown above and have in- distinct blotches. One specimen from Dos Aguas regurgitated a large Gerrhonotus imbricatus imbricatus; of two others from the same locality, one regurgitated a Sceloporus bulleri and an Eptesicus fuscus. The latter specimen was collected at the entrance of a small cave, where it probably had captured the bat. In the Cordillera Volcdnica Crotalus pusillus has been obtained AMPHIBIANS AND REPTILES OF MICHOACAN 123 in pine-oak forest at elevations between 1550 and 1800 meters. In the Sierra de Coalcoman two specimens were taken in pine forest at an elevation of 2300 meters; ten other were found beneath rocks and logs in pine-oak forest at an elevation of 2100 meters. Crotalus triseriatus aquilus Klauber Crotalus triseriatus aquilus Klauber, Bull. Zool. Soc. San Diego, 26:24, August 8, 1952.—Alvarez, San Luis Potosi, México. Morelia (10); Tacicuaro (2). I am following Klauber (1952) in assigning some of the specimens of this species from Michoacan to the subspecies aquilus and others to C. t. triseriatus. The distinguishing characters of these subspecies are given by Klauber (1952:28). On the basis of the few localities from which the species is known in Michoacan it seems as though C. t. aquilus inhabits the open grassy areas on the Mexican Plateau and the associated open pine-oak or oak-bunch grass habitats to the north and east of the Cordillera Volcdnica. Crotalus triseriatus aquilus has been collected at elevations from 1600 to 2000 meters in Michoacan. Crotalus triseriatus triseriatus (Wagler) Uropsophus triseriatus Wagler, Natiirliches System der Amphibien, p. 176, 1830.—México. (Probably Mexico City. ) Crotalus triseriatus triseriatus, Klauber, Bull. Zool. Soc. San Diego, 26:19, August 8, 1952. Cerro Tancitaro (36); Opopeo; Patzcuaro. This small rattlesnake inhabits rocky areas in pine and pine-oak forests above 1600 meters in the Cordillera Volcdnica; it has been collected at 3270 meters on Cerro Tancitaro. The series reported by Schmidt and Shannon (1947:84) is a mixture of specimens of Crotalus triseriatus and Crotalus pusillus. The two species are found together on Cerro Tancitaro, but only Crotalus pusillus inhabits the coniferous forests of the Sierra de Coalcomin. Klauber (1952:30) stated that despite the proximity of Crotalus triseriatus triseriatus and Crotalus triseriatus aquilus in Michoacan, there is no evidence of intergradation. He went on to suggest that additional material might show that the two named populations actually are distinct species. The specimens that have been studied since Klauber’s in- vestigations also show no evidence of intergradation, but there still is no known sympatry of the populations. The small montane rattlesnakes belonging to the species C. pricei, C. pusillus, and C. triseriatus present a problem in systematics and distribution worthy of intensive investigation. A knowledge of the 9—781i7 124 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. distribution and relationships of the various populations of these snakes, together with other species also living in isolated populations on the higher mountains in México, probably will be of great signifi- cance in understanding dispersal and differentiation of animals dur- ing the Pleistocene. SPECIES OF QUESTIONABLE OCCURRENCE Some species for which there are no authentic records from Michoacan can be expected there on zoogeographic probability. Other species have been recorded from Michoacan, but these records are doubtful for any one of several reasons. Fifteen species of such questionable occurrence are discussed below: Syrrhophus modestus modestus Taylor Syrrhophus modestus Taylor, Univ. Kansas Sci. Bull., 28:304, May 15, 1942. —Hacienda Paso del Rio, Colima, México. Syrrhophus modestus modestus, Duellman, Occ. Pap. Mus. Zool. Univ. Michi- gan, 594:5, June 6, 1958. This small terrestrial frog is not uncommon on the coastal lowlands and foothills in Nayarit and in Colima, where it has been collected within a few kilometers of the Michoacan border. At Tecolapa, Colima, on August 9, 1956, Syrrhophus modestus modestus was found with Tomodactylus nitidus orarius, Bufo marinus, Bufo mar- moreus, Hyla baudini, Hyla smithi, and Phyllomedusa dacnicolor, all of which occur on the coastal lowlands of Michoacdn. Because of its solitary and secretive habits, Syrrhophus modestus modestus is not common in collections. Additional field work on the coast of Michoacan should reveal the presence of the species there. Hyla microcephala sartori Smith Hyla microcephala sartori Smith, Herpetologica, 7:186, December 31, 1951.—1 mi. N of Organos, S of El Triente, Guerrero, México. On August 28, 1960, J. R. Dixon obtained a series of this species from a temporary pond 6 kilometers northeast of La Resolana, Jalisco. Previously, Hyla microcephala sartori had been known only from the lowlands of Guerrero and Oaxaca. The existence of the species in Jalisco provides evidence that this frog also occurs in Michoacan and Colima. Gastrophryne usta usta (Cope) Engystoma ustum Cope, Proc. Acad. Nat. Sci. Philadephia, 18:131, 1866. —Guadalajara, Jalisco, México. Gastrophryne usta usta, Carvalho, Occ. Pap. Mus. Zool. Univ. Michigan, 555:18, July 16, 1954. Smith and Taylor (1948:93-4) listed specimens of this species from Organos and E] Treinta, Guerrero, and frem Paso del Rio, AMPHIBIANS AND REPTILES OF MICHOACAN 125 Queseria, Santiago, and Tecoman, Colima. The species occurs from Sinaloa and central Veracruz southward at low elevations to the Isthmus of Tehuantepec and thence along the Pacific lowlands into Central America. Almost certainly it occurs on the coastal lowlands in Michoacan. Since the amphibian fauna of the Tepal- catepec Valley has been better sampled than that of the coast, I suspect that if Gastrophryne occurred in the Tepalcatepec Valley, I would have found it there. Lepidochelys olivacea (Eschscholtz) Chelonia olivacea Eschscholtz, Zool. Atlas, pt. 1, p. 2, 1829.—Manila Bay, Philippine Islands. Lepidochelys olivacea, Girard, United States Exploring Expedition . . ., vol. 20, Herpetology, p. 435, 1858. According to Smith and Taylor (1950b:15), this sea turtle is known from the entire Pacific coast of México; these authors re- ported the species from Chiapas, Oaxaca, Guerrero, Colima, and Sonora. Although the only sea turtle that I observed in Michoa- can is Chelonia mydas, others probably do use the sheltered beaches for nesting. The scanty records of sea turtles along the Pacific coast of México indicate that Chelonia mydas and Lepi- dochelys olivacea are the most abundant species in that region. There are scattered records of Dermochelys coriacea, Caretta ca- retta, and Eretmochelys imbricata along the Pacific coast. The occurrence of any of these along the coast of Michoacan is prob- able. Geoemyda pulcherrima pulcherrima (Gray) Emys pulcherrima Gray, Catalogue of the Shield Reptiles in British Mu- seum, vol. 1, p. 25.—México. Type locality restricted to Presidio de Mazatlan, Sinaloa, México, by Smith and Taylor (1950b:30). Geoemyda pulcherrima pulcherrima, Wettstein, Sitzb. Akad. Wiss. Wien, 143:18, 1934. Smith and Taylor (1950b:80) recorded this species from Sonora, Sinaloa, Nayarit, Colima, and Guerrero; these records indicate that the species probably is distributed along the Pacific coast of México southward from southern Sonora. It unquestionably oc- curs on the coast of Michoacan. Natives of the coastal lowlands tell of another “tortuga de la tierra” besides Geoemyda rubida. In the collections of the Museum of Natural History of the Uni- versity of Illinois is a specimen of Geoemyda pulcherrima from Mexcala in the Balsas Basin in northern Guerrero. On the basis of this specimen it is highly probable that the species also inhabits the Balsas-Tepalcatepec Basin in Michoacan. 126 Universiry OF Kansas Pusts., Mus. Nat. Hist. Pseudemys scripta ornata (Gray) Emys ornata Gray, Synopsis reptilium, p. 80, 1831.—Mazatlan, Sinaloa, México. Pseudemys scripta ornata, Carr, Herpetologica, 1:135, December 30, 1938. The systematics and distribution of Pseudemys scripta in México and Central America are poorly understood. Smith and Taylor (1950b:32) recorded this turtle from the Pacific lowlands of Sinaloa, Jalisco, Oaxaca, and Chiapas. This species is represented by vi- carious populations throughout the Atlantic lowlands of México, northwestern México, over much of the United States, and also in Baja California. Along the Pacific coast of México the species seems to be extremely rare, or, at least, only locally abundant. Since the species has such a wide distribution, and since it occurs on the Pacific lowlands both to the north and to the south of Michoacan, it is reasonable to expect its presence on the coast of Michoacan. Inquiries among the natives living in the Balsas-Tepalcatepec Basin produced only negative evidence about the occurrence of Pseu- demys in the Rio Tepalcatepec and Rio Balsas. I suspect that the best place to search for these turtles on the coast of Michoacan is in the numerous fresh-water lagoons on the coastal plain. Caiman crocodilus fuscus (Cope) Perosuchus fuscus Cope, Proc. Acad. Nat. Sci. Philadelphia, 20:203, No- vember 9, 1868.—Rio Magdalena, Columbia. Caiman crocodilus fuscus, Mertens, Senckenbergiana, 26:275, December 22, Gadow (1930:50) reported that Caiman sclerops (= Caiman crocodilus fuscus) inhabited the “tierra caliente” in Michoacan. Smith and Taylor (1950b:212) accepted Gadow’s record for the State, although otherwise the species is unknown north of Oaxaca. Peters (1954:10) refuted Gadow’s record on the basis that Gadow’s collections contained no specimens of Caiman. The local name “caiman” refers to both Crocodylus and to Caiman, for, in general, the natives do not distinguish between the two. “Caimanes” are reported from along the coast of Michoacdn, where the name presumably refers to Crocodylus acutus acutus, and in the Balsas- Tepalcatepec Basin (Gadow, 1930:50; Webber, 1946:267). I have seen no specimens of either Crocodylus or Caiman from the Balsas Basin. If crocodilians do occur in the basin, they probably are Crocodylus acutus acutus. There is no basis, whatsoever, for in- cluding Michoacan in the range of Caiman crocodilus fuscus. AMPHIBIANS AND REPTILES OF MICHOACAN 127 Bipes canaliculatus Bonnaterre Bipes canaliculatus Bonnaterre, Encyclopédie méthodique, Erpétologie, p. 68, 1789.—Meéxico. Type locality restricted to Mexcala, Guerrero, México, by Smith and Taylor (1950b:39). Duges (1896:480) reported this species from Morelia, Michoacan. Smith and Taylor (1950b:39), who recorded the species from three localities in the Balsas Basin in Guerrero, rejected Dugés’ record. I, too, am unwilling to accept Dugés’ record. Nevertheless, the species probably occurs throughout much of the Balsas Basin. This idea is strengthened by comments made by Storm (1989:342): “The last hard drop, that afternoon, was down the great Cerro de los Cajones [southwest of Tacambaro], and here in the upper forest we came upon . . .. a lizard with front legs and none behind . . . the animal with hands and no feet that sefor Smith [Hobart M. Smith] was seeking! . . . Theyre named Bipes caniculatus (sic.).” Coleonyx elegans nemoralis Klauber Coleonyx elegans nemoralis Klauber, Trans. San Diego Soc. Nat. Hist., 10: 195, March 9, 1949.—Paso del Rio, Colima, México. Klauber (1945:199) and Smith and Taylor (1950b:43) reported this lizard from the coastal lowlands of Colima and Guerrero. Davis and Smith (1953:101) reported it from 8 kilometers northeast of Temilpa, Morelos, in the upper Balsas Basin. Specimens of this lizard have been collected infrequently; the few locality records and limited ecological data indicate that it inhabits dense scrub forest and tropical semi-deciduous forest. Coleonyx elegans nemo- ralis is to be expected on the coastal lowlands, the seaward foothills of the Sierra de Coalcoman, and on the lower slopes of the Cordil- lera Volcanica along the northern edge of the Tepalcatepec Valley. Phrynosoma orbiculare orbiculare (Linnaeus) Lacerta obricularis Linnaeus, Systema naturae, ed. 12, 1:1062, 1789.— México (by inference). Type locality restricted to México, Districto Federal, by Smith and Taylor (1950b:97). ft ie orbiculare orbiculare, Smith, Trans. Kansas Acad. Sci., 37:290, Gadow (1905:213) inferred that Phrynosoma orbiculare occurred at elevations of more than 3000 feet in Michoacan. There are no specimens of this species known from Gadow’s collections made in Michoacan. Smith and Taylor (1950b:98) apparently accepted Gadow’s statement and recorded the species from Michoacdn: “above 3000 feet (Jorullo?).” Reeve (1952:940) somehow miscon- strued this statement to read “Jorullo, above Zumpango (Smith and 128 UnIversIry OF Kansas Pusts., Mus. Nat. Hist. Taylor, 1950b).” Reeve did not indicate on his map (1952:939) that the species occurred in Michoacan. In the most recent review of the species (Horowitz, 1955), no localities are given in Micho- acan. Since Phrynosoma orbiculare is known from central Jalisco, Guanajuato, Queretaro, and México, its presence at least in north- eastern Michoacan is to be expected, although at the present time there are no specimens known from the state. Eumeces brevirostris (Giinther) Mabouia brevirostris Giinther, Proc. Zool. Soc. London, p. 316, August, 1860.—Oaxaca. Type locality restricted to Oaxaca, Oaxaca, México, by Smith and Taylor (1950b: 168). Eumeces brevirostris, Bocourt, Mission scientifique au Mexique et dans YAmerique Centrale. Reptiles, livr. 6, p. 489, 1879. Smith and Taylor (1950b:168) listed this species: “Michoacdn: No specific record.” I am unaware of any specimen of this skink from the state. As presently recognized, this species contains two subspecies. One of these occurs in the mountains of Oaxaca north- ward into central Veracruz; the other, Eumeces brevirostris biline- atus, occurs in Durango southward to Jalisco, where it inhabits the Sierra Madre Occidental. Possibly the species occurs in the Sierra de los Tarascos in Michoacan. Eumeces callicephalus Bocourt Eumeces callacephalus Bocourt, Mission scientifique au Mexique et dans YAmerique Centrale. Reptiles, livr. 6, p. 431, 1879.—Guanajuato, Guanajuato, México. Dugés (1896) in a paper in which he listed several species of Eumeces in México, reported Eumeces callicephalus from Michoa- cin, but he gave no specific locality within the state. Michoacan was included in the range of the species by Taylor (1936:298) and by Smith and Taylor (1950b:164). The species definitely is known from southeastern Arizona southward to Guanajuato. It may occur in Michoacan, but, since there are three rather widespread species of Eumeces inhabiting the Mexican Plateau and associated moun- tain ranges in the northern and northeastern part of Michoacan, interspecific competition might be a reason for the absence of Eumeces callicephalus there. Leptodeira septentrionalis polysticta Giinther Leptodeira polysticta Giinther, Biologia Centrali-Americana, Reptilia, p. 172, May, 1895.—-Belice, British Honduras. Leptodeira septentrionalis polysticta, Duellman, Bull. Amer. Mus. Nat. Hist., 114:72, February 24, 1958. Although this species occurs from sea level to elevations of about 2000 meters from Nayarit southward into Central America, no AMPHIBIANS AND REPTILES OF MICHOACAN 129 specimens are known from Michoacén. Smith and Taylor (1945: 87) listed the species as occurring in Michoacan, but they had no record on which to base this report. Probably, the species occurs on the coastal lowlands and seaward slopes of the Sierra de Coalco- man, Tropidodipsas fasciata guerreroensis Taylor Tropidodipsas guerreroensis Taylor, Univ. Kansas Sci. Bull., 26:470; Novem- ber 27, 1940.—Buena Vista, Guerrero, México. Tropidodipsas fasciata guerreroensis, Alvarez del Toro and Smith, Herpe- tologica, 12:16, March 6, 1956. Dugés (1896:480) reported a snake, questionably of this species, from Uruapan, Michoacin. Taylor (1940c) suggested that on geographic grounds Dugés’ record might refer to T. f. guerreroensis, which is known definitely only from the type locality. Tropidodip- sas occidentala is known from Comala, Colima, and Coalcoman, Michoac4én. On zoogeopraphic grounds that species might be found at Uruapan. Since the specimen apparently no longer is extant, the identification cannot be ascertained. Micrurus fitzingeri fitzingeri (Jan) Elaps fitzingeri Jan, Rev. Mag. Zool., p. 521, 1858.—México. Type locality restricted to Guanajuato, Guanajuato, México, by Smith and Taylor (1950a:330). Micrurus fitzingeri fitzingeri, Brown and Smith, Proc. Biol. Soc. Washington, 55:68, June 25, 1942. Smith and Taylor (1945:174) recorded the species from Zamora, Michoacan. Hobart M. Smith (in litt.) stated that this record was based on a report of Elaps fulvius from Zamora by Dugeés (1896: 482). Smith guessed that the report was based on a specimen of Micrurus fitzingeri. The specimen has not been seen. Although the species is known from Guanajuato and México, until a specimen is available from Michoacan, the species should not be considered part of the herpetofauna of Michoacan, GAZETTEER The localities in Michoacan here listed are those from which specimens were examined as well as other localities mentioned in the text. The localities are arranged alphabetically according to the most definitive word or words in the total name. For example, Lago de Chapala is listed as “Chapala (Lago de)” and Cerro de Tancitaro is listed as “Tancitaro (Cerro de).” Insofar as has been possible, the following information is given for each locality: geo- graphical co-ordinates to the nearest minute of north latitude and west longitude, elevation in meters above mean sea level, a descrip- 130 UNIvERSITY OF Kansas Pusts., Mus. Nat. Hist. © Sahuayo i 20° } + a 2 Oviguilpan © Zamora i~= =") TangamandapioO Ee a ae -—= Te Tangancicucro o ™ ‘ ‘ = aS 4 A C. Patamba Los ReyesO © Corupu AngahucnO om aa _ . OS. Juan V. Paricutin A C. de Cs r f 7 Tancitaro TancitaroO Lombardia O Buenavista ° ° pele Apatzingano acres O/Cofradia Tepalcatepec C.Barolosa Tepalcatepec Li® @ Dos Aguas Capino ‘ rN OCoahuayona Aguililla El TicuzN© > O 0 I Solitre San Juan de Lima Aqui y, Cue Estopilas A ¢ 7¢| Cantar 6 / on i, ; Lo Placita\© Ostulao® Las Tecatas 7 cee Nee El Espinalo Coire San José de la Montana ‘San Salvodorg S Pomarog (s of GS an Pedro Naranjestila Arteaga Maruata 2 ie & ie & (~) 8 QHuahuo = 3 N Ze) ‘9 Sur AS & ey A oa ore ° . 9 - Lo vd 3 : dt - - = ots Mira / Cc ayo Azu % Oo e Qn | 104° WE Duellman- 1958 Fic. 11. Map of Michoac4n showing important localities mentioned in text. AMPHIBIANS AND REPTILES OF MICHOACAN 181 ee Co ¢ 2 \ Y ~— ® ' 3 N \ OQ Huetamo \ iS<= o Rio gals es i STATE OF MICHOACAN, MEXICO ‘ 18°} SCALE km '0,2,2—!0_ 20 30 40 50 60 70 20 30 40 102° Localities not on this map can be located by directions given in the gazetteer. 132 UNIversiITy OF Kansas Pusts., Mus. Nat. Hist. tion of its geographical location, type of dominant vegetation, and in some cases comments concerning collecting sites in the vicinity. Distances are in kilometers; all are map (air line) distances, unless otherwise indicated. Many localities visited on mule trips are given as being a certain number of “mule hours” in a general di- rection from another town or village. In order to reach most of these localities today, one would have to go by mule, and this is the way the muleteers determine their distances. Some of the elevations are taken from maps, but most of them were obtained from one or more readings of altimeters that we carried in the field. The terms used for describing the vegetation are those defined in the section of the natural landscape. My primary cartographic sources have been: the provisional edition of maps published by the American Geographic Society (Colima, Guadalajara, México, and San Luis Potosi sheets pub- lished between 1933 and 1940), scale 1:1,000,000; the preliminary sheets (Colima, Guadalajara, Guanajuato, and México) published in 1949 with a scale of 1:500,000 of the Carta Geografica de la Re- publica Méxicana (Direccidn de Geografia y Meterologia, Secre- taria de Agricultura y Ganaderia); and the Carta de Cuenca Tepal- catepec (Scale 1:250,000) prepared in 1958 by the Comisién del Tepalcatepec, Secretaria de Recursos Hidraulicos. I have visited most of the 181 localities and have gathered data pertaining to vegetation, altitude, and location. I think, nevertheless, that the accuracy of some of the locations and elevations as given in the gazetteer is questionable. This situation can be rectified only by detailed geographic studies. Most of the important towns, villages, rivers, and high moun- tains are shown on the accompanying map (Fig. 11). Places not shown on this map can be located from directions given in the gazetteer. Acahuata.—Lat. 19° 10’, long. 102° 21’, elev. 1040 m. A village north of Apatzingdn and on the southern slope of Cerro de Tancitaro; transition be- tween arid tropical scrub forest and pine-oak forest; tropical semi-decidu- ous forest in barrancas. Agua Cerca.—Lat. 19° 06’, long. 101° 45’, elev. 1550 m. A ranch south- southwest of Ario de Rosales on the road to La Huacana; pine-oak forest. Aguililla—Lat. 18° 45’, long. 102° 47’, elev. 860 m.; a town in a low valley in the Sierra de Coalcoman; arid tropical scrub forest. Alamo (El).—Lat. 19° 42’, long. 100° 55’, elev. 2300 m. A ranch 5 kilometers by road east of El Temazcal; pine-oak forest. Angahuan.—Lat. 19° 88’, long. 102° 14’, elev. 2440 m. A Tarascan village about 27 kilometers northwest of Uruapan; pine forest. Much of the land is still covered by a deep layer of ashes from the nearby Volcan Paricutin. AMPHIBIANS AND REPTILES OF MICHOACAN 133 Apatzingan.—Lat. 19° 06’, long. 102° 22’, elev. 885 m. The largest town in the Tepalcatepec Valley; arid tropical scrub forest. Apiza (Boca de).—Lat. 18° 42’, long. 108° 44’, sea level. The name of the mouth of the Rio Coahuayana; sandy beach and coco palms. Apo.—Lat. 19° 25’, long. 102° 25’, elev. 2160 m. A village on the western slope of Cerro de Tancitaro; pine-oak forest. Aquila.—Lat. 18° 82’, long. 108° 30’, elev. 150 m. A small village on the Rio Aquila in the seaward foothills of the Sierra de Coalcoman; tropical semi-deciduous forest. Araparicuaro.—Lat. 19° 22’, long. 102° 12’, elev. 1525 m. A village 19 pone. west-southwest of Uruapan on the trail to Tancitaro; pine-oak orest. Araro.—Lat. 19° 54’, long. 100° 50’, elev. 1830 m. A small village at the eastern end of the Lago de Cuitzeo lakebed; mesquite-grassland. Ario de Rosales.—Lat. 19° 12’, long. 101° 42’, elev. 1980 m. A town in the Cordillera Volcdnica on the road from Patzcuaro to La Huacana; mixed hard- woods and pine forest. Arteaga (formerly Carrizal).—Lat. 18° 28’, long. 102° 25’, elev. 850 m. A town in the eastern part of the Sierra de Coalcoman; transition between arid tropical scrub forest and oak forest. Atzimba.—Lat. 19° 89’, long. 100° 47’, elev. 2900 m. A national park in the Cordillera Volcdnica, located between Ciudad Hidalgo and Morelia, 2 Tle by road west-southwest of Ciudad Hidalgo; mixed pine and orest. Axolotl (Rancho).—Lat. 19° 47’, long. 100° 88’, elev. 2900 m. A settlement on the western slopes of Cerro San Andrés; pine, oak, and fir forest. Balsas (Rio).—A large river having its headwaters in Tlaxcala, Puebla, and northwestern Oaxaca, flowing westward through an arid valley to the Pacific Ocean, and in its lower part forming the boundary between Michoacan and Guerrero. Barolosa (Cerro de).—Lat. 18° 52’, long. 102° 57’, elev. 2900-3050 m. Pre- sumably the highest mountain in the Sierra de Coalcoman and located about 13 hours by mule east-northeast of Coalcom4n; open pine-oak-fir forest and alder thickets. Barolosa (Rancho).—Lat. 18° 50’, long. 103° 00’, elev. 2820 m. A small ranch on the west-northwestern slope of Cerro de Barolosa, about 11 hours by mule east-northeast of Coalcoman; open pine-oak forest. Barranca Seca.—Lat. 19° 32’, long. 102° 15’, elev. 2100 m. A small village about 7 kilometers northwest of San Juan de Parangaricutiro; pine forest. Bejuco (Barranca de).—Lat. 18° 07’, long. 102° 48’, elev. 90 m. A barranca in the lower slopes of the Sierra de Coalcom4n just west of the lower reaches of the Rio Nexpa; tropical semi-deciduous forest. Buenavista (Tomatlan).—Lat. 19° 17’, long. 102° 36’, elev. 425 m. A village on the Rio Masiaco in the Tepalcatepec Valley, 33 kilometers by road west-northwest of Apatzing4n; open arid tropical scrub forest. Buena Vista.—Lat. 18° 40’, long. 102° 09’, elev. 600 m. A ranch on the north- eastern slopes of the Sierra de Coalcoman; arid tropical scrub forest. Cachan (Rio).—Lat. 18° 14’, long. 103° 14’. A river formed by the affluence of the Rio Coalcoman and the Rio San José and flowing into the Pacific Ocean at a point indicated by the co-ordinates given above. Sometimes the name is applied to the lower part of the river as used here; other times the name is used for the entire length of the Rio Coalcoman. Camichines.—Lat. 18° 47’, long. 103° 05’, elev. 1070 m. A ranch about 5 kilometers east-northeast of Coalcom4n; transition between arid tropical scrub forest and oak forest. 134 UNIVERSITY OF KAnsAs Pusts., Mus. Nat. Hist. Camécuaro (Lago de).—Lat. 19° 55’, long. 102° 13’, elev. 1615 m. A small lake (depth to about 10 m.) drained by the Rio Duero and located one kilometer north-northwest of Tangancicuaro; mesquite-grassland and some cypress and oak around the lake. Cancita (Rio).—A tributary of the Rio Tepalcatepec flowing southward from the southeastern slope of Cerro de Tancitaro. Cantiles (Los).—Lat. 19° 48’, long. 100° 55’, elev. 2160 m. A ranch 838 kilometers by road east of Morelia; pine forest. Capacuaro.—Lat. 19° 33’, long. 102° 02’, elev. 2070 m. A Tarascan village 18 kilometers by road north of Uruapan; pine forest. Capirio.—Lat. 18° 52’, long. 102° 08’, elev. 180 m. A village on the Rio Tepalcatepec, 22 kilometers by road south of Nueva Italia; open arid tropical scrub forest and some gallery forest along the river. Carapan.—Lat. 19° 52’, long. 102° 02’, elev. 2070 m. A village on the north- ern edge of the Sierra de los Tarascos, 32 kilometers by road west of Zacapu; pine-oak forest at village and to the south; mesquite-grassland immediately to the north. Cerrito (El).—Lat. 18° 45’, long. 103° 40’, elev. 15 m. A ranch about 8 kilometers northeast of Coahuayana; tropical semi-deciduous forest. Chapala (Lago de).—A large lake on the Mexican Plateau at an elevation of 1525 m., partly in the state of Jalisco. It is drained by the Rio Grande de Santiago, which flows northward and then westward into the Pacific Ocean. Immediately to the east of the lake are remnants of once extensive marshes. Charapendo.—Lat. 19° 15’, long. 102° 04’, elev. 975 m. A village 24 kilometers by road south of Uruapan near the upper limit of the arid tropical scrub forest in the Tepalcatepec Valley. Cheran.—Lat. 19° 42’, long. 101° 57’, elev. 28350 m. A Tarascan village 27 kilometers by road south-southeast of Carapan; pine forest. Chichihuas.—Lat. 18° 47’, long. 103° 12’, elev. 1200 m. A ranch about 6 kilometers west-southwest of Coalcoman; scrub oak forest. Chinapa.—Lat. 19° 22’, long. 100° 51’, elev. 930 m. A small village on the Rio Chinapa, 43 kilometers south of El Temzcal on the road to Huetamo; arid tropical scrub forest. Chupio.—Lat. 19° 10’, long. 101° 27’, elev. 1080 m. A village 12 kilometers by road south of Tacdmbaro; transition between arid tropical scrub forest and oak forest. Churumuco.—Lat. 18° 37’, long. 101° 38’, elev. 210 m. A small town in the Balsas Valley; arid tropical scrub forest. Ciénega (La).—Lat. 18° 28’, long. 103° 18’, elev. 900 m. A ranch about 3 hours by mule north of Coire; tropical semi-deciduous forest. Coahuayana.—Lat. 18° 44’, long. 103° 31’, elev. 15 m. A village on the coastal plain near the mouth of the Rio Coahuayana; arid tropical scrub forest and tropical semi-deciduous forest. Coalcoman.—Lat. 18° 47’, long. 103° 08’, elev. 945 m. The largest town in the Sierra de Coalcomén and situated in a valley about 12 by 6 kilometers; arid tropical scrub forest on valley floor; oaks and some tropical semi- deciduous forest on surrounding slopes. Coaleoman (Rio).—A river having its headwaters northeast of the town of Coalcomén and flowing southward to join with the Rio San José to form the Rio Cachan. Coalcoman (Sierra de).—A highland mass outlined by the Rio Coahuayana and its tributaries on the west, the Rio Tepalcatepec on the north, and the Rio Balsas on the east, and the Pacific Ocean on the south. The axis of the sierra extends for about 200 kilometers in a west-northwest to east- southeast direction; the mountains are nearly 80 kilometers in breadth; the highest parts of the range are about 3000 meters above sea level. AMPHIBIANS AND REPTILES OF MICHOACAN 135 Cofradia.—Lat. 18° 56’, long. 102° 17’, elev. 215 m. A ranch about 17 kilometers southeast of Apatzingan; arid tropical scrub forest. Coire.—Lat. 18° 26’, long. 103° 22’, elev. 300 m. A village on the seaward foothills of the Sierra de Coalcomdn on the Rio Coire; tropical semi-de- ciduous forest. Cojumatlan.—Lat. 20° 07’, long. 102° 51’, elev. 1530 m. A village on the southeastern shore of Lago de Chapala; mesquite-grassland. Colola (Rio).—A small river emptying into the Pacific Ocean between Maru- ata and Punto San Telmo. Conejos (Los).—Lat. 19° 22’, long. 102° 07’, elev. 1850 m. A village 6 kilometers west-southwest of Uruapan, and sometimes known as Nuevo San Juan; pine-cak forest. Copandaro.—Lat. 19° 54’, long. 101° 12’, elev. 1800 m. A village on the south edge of the Lago de Cuitzeo lakebed; mesquite-grassland. Copuyo (Capuyo or Copullo).—Lat. 18° 28’, long. 100° 56’, elev. 1200 m. A small village about 5 kilometers by road west of Paso Ancho; transition between arid tropical scrub forest and oak forest. Cordillera Volc4nica.—A mountain range along the southern edge of the Mexi- can Plateau, roughly along the nineteenth parallel, and made up of many volcanos; the range extends from Volc4n de Colima on the west to Cofre de Perote and Orizaba in Veracruz; several of the volcanos reach elevations of more than 4000 meters. Corralito (El).—Lat. 18° 52’, long. 102° 38’, elev. 270 m. A small village in the Tepalcatepec Valley, about 30 kilometers southwest of Apatzingan; arid tropical scrub forest. Corupu (Corupo).—Lat. 19° 28’, long. 102° 19’, elev. 2450 m. A village 29 kilometers northwest of Uruapan; pine forest. Cuatro Caminos.—Lat. 19° 00’, long. 102° 05’, elev. 335 m. A village 4 kilometers south of Nueva Italia; arid tropical scrub forest. Cuilala (Playa).—Lat. 18° 10’, long. 103° 06’, sea level. A sandy beach on the Pacific Ocean just east of La Higuerita. Cuitzeo.—Lat. 19° 58’, long. 101° 09’, 1800 m. A village on the north shore of the Lago de Cuitzeo lakebed; mesquite-grassland. Cuitzeo (Lago de).—A large lakebed on the Mexican Plateau at an elevation of 1800 m. In dry years there is little water in the lake, and most of the lakebed is dry; in very wet years the entire lakebed is flooded. The Rio de Morelia flows into the lake, which has no outlet; surrounding vegetation is mesquite-grassland. Cuseno Station.—Lat. 19° 30’, long. 102° 16’, elev. 2200 m. A field station of the American Geological Society established in 1945 and demolished in 1953; located at the northem edge of the lava flow at Volcan Paricutin; remnants of pine forest. Diezmo (El).—Lat. 18° 26’, long. 103° 19’ elev. 850 m. A ranch about 8 kilo- meters north of Coire; tropical semi-deciduous forest. Dos Aguas.—Lat. 18° 45’, long. 102° 55’, elev. 2100 m. A lumber camp on the eastern slope of Cerro de Barolosa, located about 22 kilometers west- northwest of Aguililla; pine-oak forest and some fir forest in sheltered ravines. Duero (Rio).—A small river having its headwaters near Tangancicuaro and flowing northwestward into the Rio Lerma; source of irrigation water for surrounding agricultural area. Emiliano Zapata.—Lat, 18° 59’, long. 102° 39’, elev. 1600 m. A town 10 kilometers east of Jiquilpan; mesquite-grassland and irrigated fields. Erongaricuaro.—Lat. 19° 35’, long. 101° 48’, elev. 2150 m. A Tarascan village on the western shore of Lago de Patzcuaro; pine forest. Espinal (El).—Lat. 18° 27’, long. 102° 07’, elev. 500 m. A ranch in the northern foothills of the Sierra de Coalcoman, 9 kilometers by road north- northeast of San Salvador; arid tropical scrub forest. 136 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. Estopilas (Salitre de).—Lat. 18° 30’, long. 103° 23’, elev. 130 m. A small village about 10 kilometers east of Ostula; tropical semi-deciduous forest and arid tropical scrub forest. Garica (Cerro).—Lat, 19° 48’, long. 100° 48’, elev. 8000 m. A mountain about 8 kilometers north of Pino Gordo; pine-oak-fir forest. Garnica (Puerto de).—Lat. 19° 42’, long. 100° 51’, elev. 2840 m. A moun- tain pass 46 kilometers by road west of Ciudad Hidalgo; pine and fir forest. Gregorio (San).—Lat. 19° 25’, long. 101° 24’, elev. 2200 m. A ranch about 16 kilometers southeast of Patzcuaro; pine forest. Guayabo.—Lat. 18° 45’, long. 102° 15’, elev. 760 m. A village in the Sierra de Coalcoman about 32 kilometers north-northeast of Arteaga; upper limits of arid tropical scrub forest. Herradero (Barranca de).—Lat. 18° 17’, long. 103° 08’, elev. 200-250 m. A barranca south of San Pedro Naranjestila in the Sierra de Coalcoman; tropical semi-deciduous forest. Hidalgo (Ciudad).—Lat. 19° 32’, long. 100° 34’, elev. 2100 m. A town in the valley of the Rio Tuxpan; mesquite-grassland and pine-oak forest. Higuerita (La).—Lat. 18° 12’, long. 103° 06’, sea level. A place name on the Pacific coast; sandy beach and arid tropical scrub forest. Higuertas (Las).—Lat. 18° 89’, long. 103° 17’, elev. 1600 m. A ranch about 7 hours by mule southwest of Coalcoman; pine-oak forest. Hondo (Puerto).—Lat. 19° 25’, long. 100° 13’, elev. 2750 m. A pass in the mountains, 14 kilometers by road east of Zitacuaro (just west of Macho de Agua); pine, oak, and fir forest. Huancana (La).—Lat. 18° 58’, long. 101° 50’, elev. 550 m. A village in the Balsas Basin; arid tropical scrub forest. Huahua (La).—Lat. 18° 12’, long. 103° 00’, sea level. A small village on the Pacific coast; arid tropical scrub forest and gallery forest along the Arroyo de Huahua. Huetamo.—Lat. 18° 88’, long. 100° 53’, elev. 800 m. A town in the Balsas Valley; arid tropical scrub forest. Huingo.—Lat. 19° 55’, long. 100° 50’, elev. 1800 m. A village on the eastern edge of the Lago de Cuitzeo lakebed; mesquite-grassland. Jacona.—Lat. 19° 57’, long. 102° 18’, elev. 1600 m. A small town, 4.3 kilometers by road southwest of Zamora; mesquite-grassland. Jaramillo.—Lat. 19° 20’, long. 102° 02’, elev. 1500 m. A ranch 9 kilometers by road south of Uruapan; pine-oak forest. Jazmin.—Lat. 18° 52’, long. 101° 58’, elev. 275 m. A village in the Tepalcate- pec Valley, 32 kilometers by road southeast of Cuatro Caminos; open arid tropical scrub forest. Jerahuaro.—Lat. 19° 52’, long. 100° 35’, elev. 2600 m. A town in the north- ern part of the state and located east of Lago de Cuitzeo; pine-oak forest. Jiquilpan.—Lat. 19° 59’, long. 102° 43’, elev. 1570 m. A town just southeast of Lago de Chapala; mesquite-grassland. Jorullo (Volcan).—Lat 19° 00’, long. 101° 45’, elev. 1300 m. (crest). A cinder and lava cone rising from the foothills of the Cordillera Volcanica; arid tropical scrub forest on lower slopes and pine-oak forest on top. Jungapeo.—Lat. 19° 26’, long. 100° 29’, elev. 1480 m. A village in the valley of the Rio Tuxpan, about 13 kilometers south of Tuxpan on the southern plenes of the Mexican Plateau; tropical semi-deciduous forest and pine-oak orest. Lengua de Vaca (Puerto de).—Lat. 19° 26’, long. 100° 13’, elev. 2900 m. A pass in the mountains at the Michoacén-Mexico border through which passes the Mexico City-Morelia highway; pine and fir forest. AMPHIBIANS AND REPTILES OF MiICHOACAN 13% Lerma (Rio).—A river originating in the state of México and flowing westward, and forming the northern boundary of the state of Michoacan, to Lago de Chapala. Lima (San Juan de).—Lat. 18° 29’, long. 102° 42’, sea level. A ranch on the Pacific coast; arid tropical scrub forest and tropical semi-deciduous forest. Lima (Punta San Juan de).—Lat. 18° 38’, long. 102° 43’, sea level. A rocky promontory jutting into the Pacific Ocean just southwest of San Juan de Lima; arid tropical scrub forest. Limoncito.—Lat. 18° 45’, long. 102° 43’, elev. 730 m. A ranch 10 kilometers north of Aguililla; arid tropical scrub forest; tropical semi-deciduous gallery forest along the nearby Rio Tepecuate. Lombardia.—Lat. 19° 08’, long. 102° 02’, elev. 640 m. A town in the Tepalca- epee Valley, 38 kilometers by road south of Uruapan; arid tropical scrub orest. Lleguas (Acuaro de las).—Lat. 18° 48’, long. 102° 52’, elev. 28320 m. A place name for a stream and meadow (Llano de la Llegua) surrounded by pine- oak forest, located about 10 hours by mule east of Coalcoman. Macho de Agua.—Lat. 19° 25’, long. 100° 15’, elev. 2850 m. A ranch just west of Puerto de Lengua de Vaca and 16 kilometers by road east of Zita- cuaro; mixed oak, pine, and fir forest. Maquili—Lat. 18° 36’, long. 103° 32’, elev. 120 m. A village on the Rio Aquila about 3 kilometers south-southwest of Aguila; tropical semi-deciduous forest. Maravatio.—Lat. 19° 53’, long. 100° 27’, elev. 2010 m. A town in the Rio Lerma Valley; irrigated fields on flats and pine-oak forest on slopes. Marquez (Rio).—A tributary to the Rio Tepalcatepec, flowing through a deep gorge (Barranca del Marquez) between Lombardia and Nueva Italia. The stream originates from springs near Uruapan, where the stream is known as the Rio Cupatitzio. Maruata.—Lat. 18° 17’, long. 103° 20’, sea level. Place name for a Nineteenth Century port of little importance near the mouth of the Rio Coire; sandy beach, fresh-water lagoon, and arid tropical scrub forest. Mexcala (Laguna).—Lat. 18° 29’, long. 103° 41’, sea level. A brackish la- goon surrounded by mangroves, located just southwest of El Ticuiz. Mil Cumbres.—Lat. 19° 39’, long. 100° 47’, elev. 2800 m. A name for a look- out on the México-Morelia highway in Atzimba National Park, about 382 kilometers by road west-southwest of Ciudad Hidalgo; pine and fir forest. Mira (La).—Lat. 18° 05’, long. 102° 20’, elev. 20 m. A small village about 5 kilometers north-northeast of Playa Azul; arid tropical scrub forest. Morelia.—Lat. 19° 48’, long. 101° 10’, elev. 1900 m. Capital of and largest city in Michoac4n; mesquite-grassland on flats and pine-oak forest on sur- rounding hills. Morelia (Rio de).—A small, intermittent stream originating in the mountains south of Morelia and emptying into Lago de Cuitzeo. Motin del Oro.—Lat. 18° 14’, long. 108° 48’, sea level. A ranch on the Pa- cific coast; arid tropical scrub forest. Motin (Rio).—Lat. 18° 138’, long. 108° 48’ (mouth). A small river flowing from the Sierra de Coalcom4n into the Pacific Ocean. Nahuatzen (Nauhuatzin).—Lat. 19° 42’, long. 101° 50’, elev. 2450 m. A Tarascan village in the mountains west of Lago de Patzcuaro; pine forest. Nexpa (Rio).—Lat. 18° 05’, long. 102° 47’ (mouth). A large river draining the central part of the Sierra de Coalcom4n, originating near Aguililla, and flowing into the Pacific Ocean. Nogueleras.—Lat. 18° 84’, long. 103° 17’, elev. 1600 m. A ranch about 10 hours by mule south-southwest of Coalcom4n; oak forest. Nueva Italia.—Lat. 19° 02’, long. 102° 07’, elev. 880 m. A town in the Tepal- eines Valley, 59 kilometers by road south of Uruapan; arid tropical scrub orest. 138 Unrversiry OF Kansas Pusts., Mus. Nat. Hist. Nuevo (Rancho).—Lat. 18° 26’, long. 102° 07’, elev. 520 m. A ranch 7 kilometers by road north-northeast of San Salvador in the northern foothills of the Sierra de Coalcoman; arid tropical scrub forest. Ocorla.—Lat. 18° 38’, long. 103° 07’, elev. 885 m. A ranch about 6 hours by mule south-southeast of Coalcoman; scrubby oak forest. Opopeo.—Lat. 19° 24’, long. 101° 37’, elev. 2800 m. A village 16 kilometers south of Patzcuaro; pine and fir forest. Orilla (La).—Lat. 18° 00’, long. 102° 12’, elev. 10 m. The site of a former hacienda of the same name near the mouth of the Rio Balsas; arid tropical scrub forest. Oropeo.—Lat. 18° 52’, long. 101° 48’, elev. 300 m. A village in the Tepal- Gatepec Valley about 13 kilometers south of La Huacana; arid tropical scrub orest. Ostula.—Lat. 18° 30’, long. 103° 28’, elev. 120 m. A village in the seaward foothills of the Sierra de Coalcoman, located on the Rio Ostula about 16 kilometers east-southeast of La Placita; arid tropical scrub forest and scat- tered tropical semi-deciduous forest. Ozumatlan (Sierra de).—A range in the Cordillera Volc4nica extending east- northeast from a point south of Morelia to Queréndaro and reaching eleva- tions in excess of 2600 m. Palma (La).—Lat. 20° 09’, long. 102° 46’, elev. 1525 m. A village on the southeastern shore of Lago de Chapala; lake-shore marshes and mesquite- grassland. Paracho.—Lat. 19° 39’, long. 102° 02’, elev. 2375 m. A Tarascan village in the Cordillera Volcanica, located 35 kilometers by road north of Uruapan; pine forest. Parangaricutiro (San Juan de).—Lat. 19° 30’, long. 102° 15’, elev. 2200 m. A former Tarascan village that was destroyed by the eruption of Volcan Pari- cutin; lava and volcanic ash amidst open pine forest. Paricutin (Volc4n).—Lat. 19° 30’, long. 102° 16’, elev. 2200 m. at base and 2700 m. at summit. A volcano born in February, 1948; it ceased to be active in December, 1951, and is located at the north-northeastern base of Cerro de Tancitaro; volcanic ash and lava amidst open pine forest. Paso Ancho.—Lat. 19° 28’, long. 100° 52’, elev. 1100 m. A small village 30 plometers south of El] Temazcal on the road to Huetamo; arid tropical scrub orest. Patamba (Sierra).—Lat. 19° 45’, long. 102° 21’, elev. 3700 m. at summit. A mountain, the summit of which is about 22 kilometers southwest of Tan- gancicuaro; pine forest from 2000 to 2600 m.; fir forest above 2600 m. Patzcuaro.—Lat. 19° 80’, long. 101° 36’, elev. 2200 m. A town near the south- eastern shore of Lago de Patzcuaro; pine forest. Patzcuaro (Lago de).—A large lake on the southwestern part of the Mexican Plateau at an elevation of 2165 m. It has no outlet. The lake is surrounded by mountains supporting pine and pine-oak forest. Along the southern and eastern shores of the lake are small marshes. Peas (Las).—Lat. 18° 08’, long. 102° 38’, sea level. A small village on the Pacific coast; arid tropical scrub forest. Pichi (Estero).—Lat. 18° 01’, long. 102° 24’, sea level. A brackish lagoon surrounded by mangroves and coconut groves, located just east of Playa Azul. Pino Gordo.—Lat. 19° 42’, long. 100° 45’, elev. 2600 m. A ranch 37 kilometers by road west of Ciudad Hidalgo; pine-oak forest. Placita (La).—Lat. 18° 32’, long. 103° 87’, elev. 20 m. A village on the coastal lowlands, located on the Rio Aquila; arid tropical scrub forest; tropical semi- deciduous forest along the river. Playa (La).—Lat. 18° 57’, long. 102° 33’, elev. 800 m. A small village on the western edge of the lava flow of Volcan Jorullo; arid tropical scrub forest and some tropical semi-deciduous forest in ravines. AMPHIBIANS AND REPTILES OF MICHOACAN 139 Playa Azul.—Lat. 18° 01’, long. 102° 25’, sea level. A village on the Pacific coast near the mouth of the Rio Carrizal; arid tropical scrub forest; coconut plantations; mangrove-lined lagoons. Pémaro.—Lat. 18° 18’, long. 103° 17’, elev. 300 m. An Indian village in the southern foothills of the Sierra de Coalcoman, located about 8 hours by mule north-northeast of Maruata; tropical semi-deciduous forest. Pozos (Los).—Lat. 18° 30’, long. 103° 17’, elev. 300 m. A ranch located about 5 hours by mule north of Coire; tropical semi-deciduous forest. Queréndaro.—Lat. 19° 48’, long. 100° 53’, elev. 1900 m. A town on the Mexi- can Plateau south of Lago de Cuitzeo; mesquite-grassland. Quiroga.—Lat. 19° 42’, long. 101° 30’, elev. 2200 m. A Tarascan town on the north edge of Lago de Patzcuaro; mesquite-grassland and pine-oak forest. Reyes (Los).—Lat. 19° 35’, long. 102° 28’, elev. 1500 m. A town in western Michoacan, 50 kilometers south-southwest of Zamora; mesquite-grassland, oak and pine forest. Sabino (El).—Lat. 19° 14’, long. 102° 03’, elev. 1050 m. A hacienda about 24 kilometers south of Uruapan; arid tropical scrub forest, many streams, rice fields. Sahuayo.—Lat. 20° 05’, long. 102° 48’, elev. 1550 m. A town just south of the eastern end of Lago de Chapala; mesquite-grassland. Salada (La).—Lat. 19° 07’, long. 102° 00’, elev. 580 m. A ranch southwest of Lombardia; arid tropical scrub forest. Salto (Arroyo El).—Lat. 18° 45’, long. 103° 04’, elev. 1870 m. A valley of ae Rio Flores about 3 hours by mule east-southeast of Coalcom4n; pine-oak orest. San Andrés (Cerro).—Lat. 19° 48’, long. 100° 35’, elev. 3950 m. at summit. A mountain, the summit of which is about 16 kilometers north-northwest of Ciudad Hidalgo; oak forest to 2500 m. and pine and fir forest above 2500 m. San José (de la Cumbre).—Lat. 19° 41’, long. 100° 50’, elev. 2750 m. A ranch 51 kilometers by road east of Morelia; pine and fir forest. San José (de la Montafia).—Lat. 18° 25’, long. 103° 06’, elev. 750 m. A village sometimes called La Guitarra, located 14 hours by mule south-south- east of Coalcomén; tropical semi-deciduous forest. San Pedro Naranjestila.—Lat. 18° 17’, long. 103° 06’, elev. 500 m. An Indian village in the southern foothills of the Sierra de Coalcoman; tropical semi- deciduous forest. San Salvador.—Lat. 18° 25’, long. 102° 08’, elev. 700 m. A small village in the Sierra de Coalcoman, 37 kilometers by road northeast of Arteaga; arid tropical scrub forest. San Telmo (Ojos de Agua de).—Lat. 18° 37’, long. 103° 42’, sea level. A small settlement at the base of Punto San Juan de Lima; tropical semi- deciduous forest and groves of oil palms. San Telmo (Punta).—Lat. 18° 18’, long. 103° 29’, sea level. A rocky promon- tory jutting into the Pacific Ocean, on which there is a lighthouse (El Faro); arid tropical scrub forest. Santa Ana.—Lat. 18° 27’, long. 102° 06’, elev. 600 m. A ranch about 4 kilo- meters by road northeast of San Salvador; arid tropical scrub forest. Tacambaro.—Lat. 19° 05’, long. 101° 22’, elev. 1820 m. A town in the Cordillera Volcanica; pine forest. Tacicuaro.—Lat. 19° 38’, long. 101° 18’, elev. 2000 m. A village 21 kilo- meters east-southeast of Quiroga; mesquite-grassland and scrubby oak forest. Tafétan.—Lat. 19° 43’, long. 100° 52’, elev. 1000 m._ A village 40 kilometers by road south of El Temazcal; arid tropical scrub forest. Tancitaro.—Lat. 19° 20’, long. 102° 22’, elev. 1850 m. A small town on the southern slope of Cerro de Tancitaro; pine-oak forest. 10—7817 140 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Tancitaro (Cerro de).—Lat. 19° 25’, long. 102° 18’, elev. 3870 m. at summit. An old volcano in the Cordillera Volcanica; the southerm slope drops into the Tepalcatepec Valley; the summit is about 30 kilometers west of Uruapan; pine and oak forest on lower slopes replaced by pine or fir forest above. Tangamandapio.—Lat. 19° 56’, long. 102° 25’, elev. 1700 m. A small town on the Mexican Plateau between Jiquilpan and Zamora; mesquite-grassland and irrigated fields, Tangancicuaro.—Lat. 19° 52,’ long. 102° 13’, elev. 1770 m. A town 12 kilo- meters by road southeast of Zamora; mesquite-grassland and irrigated fields. Tarascos (Sierra de los).—A name applied to that part of the Cordillera Vol- canica extending eastward from Cerro de Tancitaro and Sierra Patamba to Patzcuaro. Tarécuaro.—Lat. 19° 58’, long. 102° 29’, elev. 1700 m. A village on the Mexi- can Plateau, 26 kilometers southwest of Zamora; mesquite-grassland and pine-oak forest. Tecatas (Las).—Lat. 18° 86’, long. 103° 17’, elev. 1950 m. A ranch located about 10 hours by mule south-southwest of Coalcoman; oak forest. Temazcal (El).—Lat. 19° 40’, long. 100° 56’, elev. 2200 m. A road junction, 29 kilometers east of Morelia; here the road to Huetamo leads south from the Mexico City-Morelia highway; pine forest. Tepalcatepec.—Lat. 19° 10’, long. 102° 50’, elev. 570 m. A village in the up- per Tepalcatepec Valley; arid tropical scrub forest. Tepalcatepec (Rio).—A large river having its headwaters in southeastern Ja- lisco and flowing through a broad valley, which separates the Cordillera Volcanica from the Sierra de Coalcoman, to the Rio Balsas. Ticuiz (El).—Lat. 18° 40’, long. 103° 40’, elev. 10 m. A village on the coastal plain about 11 kilometers south of Coahuayana; arid tropical scrub forest and tropical semi-deciduous forest. Tinguidin.—Lat. 19° 45’, long. 102° 28’, elev. 1800 m. A small town, 17 kilometers north of Los Reyes; pine-oak forest. Tizupan (Rio).—Lat. 18° 09’, long. 102° 55’ (mouth). A small river flowing southward from the Sierra de Coalcoman to the Pacific Ocean. Tlalpujahua.—Lat. 19° 48’, long. 100° 10’, elev. 2600 m. A mining town in the northeastern part of the state; pine and fir forest. Tumbiscatio.—Lat. 18° 32’, long. 102° 20’, elev. 900 m. A town in the Sierra de Coalcoman; arid tropical scrub forest. Tupataro.—Lat. 19° 58’, long. 100° 15’, elev. 2050 m. A village in the north- poten corner of the state, 18 kilometers northwest of Tlalpujahua; oak orest. Tuxpan.—Lat. 19° 35’, long. 100° 27’, elev. 1850 m. A town in a basin nearly surrounded by mountains and near the headwaters of the Rio Tuxpan, 19 kilometers by road east-southeast of Ciudad Hidalgo; arid mesquite-grassland and irrigated fields. Tuxpan (Rio).—A river draining the mountains in the eastern part of the state and flowing southward into the Rio Balsas. Tzararacua (Cascada).—Lat. 19° 18’, long. 102° 02’, 1480 m. A waterfalls of the Rio Cupatitzio, 10.5 kilometers by road south of Uruapan; oak forest with scattered pines. Tzintzuntzan.—Lat. 19° 38’, long. 101° 35’, elev. 2170 m. A village at the site of the seat of the ancient Tarascan empire on the eastern shore of Lago de Patzcuaro; grasslands and marshes. Tzitzio—Lat. 19° 35’, long. 100° 55’, elev. 1680 m. A village 16 kilometers by road south of El] Temazcal; pine-oak and arid tropical scrub forest. Ucareo (Serrania de).—A part of the Cordillera Volcdnica, including Cerro San Andrés. AMPHIBIANS AND REPTILES OF MICHOACAN 141 Undameo.—Lat. 19° 34’, long. 101° 17’, elev. 2000 m. A village 20 kilometers west-southwest of Morelia; mesquite-grassland. Uruapan.—Lat. 19° 25’, long. 102° 02’, elev. 1630 m. A large town on the southern slopes of the Cordillera Volcanica; pine-oak forest. Zacapu.—Lat. 19° 48’, long. 101° 47’, elev. 2000 m. A town on the Mexican Plateau; mesquite-grassland. Zamora.—Lat. 19° 59’, long. 102° 17’, elev. 1570 m. A large town on the Mexican Plateau; mesquite-grassland. Zicuiran.—Lat. 18° 53’, long. 101° 55’, elev. 190 m. A small village 28 kilo- meters east-southeast of Cuatro Caminos; arid tropical scrub forest. Zinapécuaro.—Lat. 19° 52’, long. 100° 49’, elev. 1900 m. A town near the southeastern end of Lago de Cuitzeo; mesquite-grassland and pine-oak forest. Ziracuaretiro.—Lat. 19° 25’, long. 101° 52’, elev. 12830 m. A village 19 kilo- meters by road east of Uruapan; transition between pine-oak forest and arid tropical scrub forest. Zirimicuaro,—Lat. 19° 24’, long. 101° 56’, elev. 1800 m. A hacienda 13 kilo- meters by road east of Uruapan; pine-oak forest and fields of sugar cane. Zitacuaro.—Lat. 19° 25’, long. 100° 21’, elev. 2100 m. A town in the high- lands of eastern Michoacan; pine-oak forest. Zurumbeneo.—Lat. 19° 43’, long. 101° 02’, elev. 2100 m. A ranch 19 kilo- meters by road east of Morelia; scrubby oak forest. SUMMARY The preceding analysis of the amphibians and reptiles of the state of Michoacan shows that the herpetofauna is composed of 176 spe- cies and subspecies definitely recorded from the state, plus ten others that probably occur there. Ten species are reported for the first time from Michoacan: Pseudoeurycea robertsi, Leptodactylus occi- dentalis, Microbatrachylus pygmaeus, Pternohyla fodiens, Hypo- pachus caprimimus, Phyllodactylus homolepidurus, Anolis dunni, Sceloporus bulleri, Sceloporus heterolepis, and Geagras redimitus. Five species that have been reported previously from Michoacan are based on specimens having unreliable locality data or on misidenti- fications; therefore, the following species are not considered to be a part of the herpetofauna of Michoacan: Caiman crocodilus fuscus, Urosaurus irregularis, Geophis nasalis, Tropidodipsas fasciata gue- rreroensis, and Micrurus fitzingeri fitzingeri. Systematic studies based at least in part on specimens from Micho- acan have resulted in a redefinition of nine species and subspecies: Bufo marmoreus, Bufo perplexus, Anolis nebulosus, Anolis nebu- loides, Sceloporus bulleri, Sceloporus heterolepis, Sceloporus me- lanorhinus calligaster, Hypsiglena torquata torquata, and Hypsiglena torquata ochrorhyncha. Nine species that previously have been recognized as valid have been placed in synonymy. These are: Bufo horribilis Wiegmann, 1833, and Bufo angustipes Smith and Taylor, 1945, as synonyms of 142 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Bufo marinus (Linnaeus), 1758. Microbatrachylus albolabris Taylor, 1940, Microbatrachylus minimus Taylor, 1940, and Microbatrachylus imitator Taylor, 1942, as synonyms of Microbatrachylus pygmaeus (Taylor), 1987. Phrynohyas corasterias Shannon and Humphrey, 1957, as a synonym of Phrynohyas inflata (Taylor), 1944. Hyla microeximia Maslin, 1957, as a synonym of Hyla eximia Baird, 1854. Hylella azteca Taylor, 1943, as a synonym of Hyla smaragdina Tay- lor, 1940. Loxocemus sumichrasti Bocourt, 1876, as a synonym of Loxocemus bicolor Cope, 1861. Eleutherodactylus vocalis Taylor, 1940, is considered to be a subspecies of Eleutherodactylus rugulo- sus. The populations of Thamnophis dorsalis in the Tepalcatepec Valley are shown to be distinct from those inhabiting the highlands of the state; Thamnophis dorsalis postremus Smith, 1942, is revived for the population in the Tepalcatepec Valley. Descriptions are given of the tadpoles of Bufo occidentalis and Hyla bistincta. LITERATURE CITED ANDERSON, J. D. 1960. Storeria storerioides in westem Mexico. Herpetologica, 16:63-6, March 81. BocErtT, C. M., and Oxrver, J. A. 1945. A preliminary analysis of the herpetofauna of Sonora. Bull. Amer. Mus. Nat. Hist., 83:297-426, March 80. BouLENGER, G. A 1885. ‘Catalogue of the lizards in the British Museum, vol. II, London, xiii ++ 497 pp., November 15. 1894. 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The tadpole of Hyla robertsorum with comments on the affinities of the spees. Occ. Pap. Mus. Zool. Univ. Michigan, 563:1-9, arch 29. REEVE, W. L. 1952. Taxonomy and distribution of the horned lizards genus Phrynosoma. Univ. Kansas Sci. Bull., 34 (pt. 2) :817-960, February 15. Scumor, K. P., and SHANNON, F. A. 1947, Notes on amphibians and reptiles of Michoacan, Mexico. Fieldiana, zool., 31:63-85, February 20. 146 UNIvERSITY OF Kansas Pusts., Mus. Nar. Hist. SHANNON, F. A., and Humpurey, F. L. 1957. A new species of Phrynohyas from Nayarit. Herpetologica, 13: 15-18, March 30. 1958. A discussion of the polytypic species, Hypopachus oxyrrhinus, with a description of a new subspecies. Herpetologica, 14:85-95, July 23. Situ, H. M. 1938. The lizards of the torquatus group of the genus Sceloporus Wieg- mann, 1828. Univ. Kansas Sci. Bull., 24:539-698, February 16. 1939. The Mexican and Central American lizards of the genus Sceloporus. Zool. ser., Field Mus. Nat. 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The status of the Mexican lizards of the genus Mabuya. Univ. Kansas Sci. Bull., 88:1303-1813, March 2. WELLMAN, J. 1959. Notes on the variation in and distribution of the Mexican colubrid Boe Coniophanes lateritius. Herpetologica, 15:127-128, Septem- er 10. Woopsury, A. M., and Woopsury, D. M. 1944. Notes on Mexican snakes from Oaxaca. Jour. Washington Acad. Sci., 34:360-373, November 15. ZWEIFEL, R. G. 1957. A new frog of the genus Rana from Michoacan, Mexico. Copeia, 1957 (2):78-83, July 15. 1959a. Variation in and distribution of lizards of western Mexico related to Cnemidophorus sacki. Bull. Amer. Mus. Nat. Hist., 117:57-116, April 27. 1959b. Additions to the herpetofauna of Nayarit, Mexico. Amer. Mus. Novitates, 1953:1-18, June 26. 1959c. Snakes of the genus Imantodes in westem Mexico. Amer. Mus. Novitates, 1961:1-18, September 16. 1960. Results of the Puritan-American Museum of Natural History Ex- pedition to western Mexico. 9. Herpetology of the Tres Marias Islands. Bull. Amer. Mus. Nat. Hist., 119:77-128, February 29. 1961. Relationship of two whiptail lizards (genus Cnemidophorus) in western Mexico. Copeia, no. 1:98-103, March 17. Transmitted April 21, 1961. Vol. 8. Vol. 9. Vol. 10. (Continued from inside of front /cover) Nos. 1-10 and index. Pp. 1-675, 1954-1956. 1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 fig- ures in text. December 10, 1955. Additional records ead extensions of ranges of mammals from Utah. By Stephen D, Durrant, M. Raymond Lee, and Richard M, Hansen. Pp. 69- 80. December 10; 1955. 8. A new long-eared myotis ee vibage from northeastern Mexico. By Rol- lin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955. 4. Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming. By Sydney Anderson, Pp. 85-104, 2 figures in text. May 10, 1956. 5. The condylarth genus Ellipsodon. |By Robert W. Wilson. Pp. 105-116, 6 figures in text. May 19, 1956. 6. Additional remains of the multituberculate genus Eucosmodon. By Robert W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956. 7. Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures in text. June 15, 1956. 8. Comments on the taxonomic status of Aporeanes peninstlae, with description of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 387-346, 1 figure in text, 1 table. August 15, 1956. 9. Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp. 847-3851. August 15, 1956. 10. A> new bat (Genus Leptonycteris) from Coahuila. By Howard J. Stains. Pp. 3538-356. January 21, 1957. 11. A new species of pocket sephes (Genus sn Sia se from Jalisco, Mexico. By Robert J. Russell. Pp. 357-861. January 21, 12. Geographic variation in the pocket gopher, Ep Area pathads in Colorado. gar M. Youngman. Pp. 368-387, 7 figures in text. February 21], 18. New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones, Jr. Pp. 385-888. May 12, 1958. 14. Pleistocene bats from San Josecito Cave, Nuevo Ledén, /México. By J. Knox Jones, Jr. Pp. 389-396. / December 19, - 15. New Subspecies of the rodent Baiomys from Central America. By Robert L. Packard. Pp. 397-404. December 19, 1958. 16 « Mammals of the Grand Mesa, ae ia By Sydney Anderson. Pp. 405- 414, 1 figure in text. May 20 Distribution, variation, and relationships of the montane vole, Peon a ed mon- tanus. By Emil K. Urban. Pp. 415-511. 12 figures in text, 2 tables. August 1, 1959. Conspecificity of two pocket mice, Perognathus goldmani and P. By E. Raymond Hall and Marilyn Bailey Ogilvie. Pp. 518-518, ais “map. January 14, 1960. - Records of harvest mice, Reithrodontomys, from Central America, with description of a new subspecies from age corr By Sydney Anderson and J. Knox Jones, Jr. Pp. 519-529. January 14, Small carnivores from San Josecito Cave Piao), Nuevo ee papatee By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, . Pleistocene pocket gophers from San Josecito Cave, Nuevo ia orale 0 By Robert J. Russell. Pp. 539-548, 1 figure in text. January 14, . Review of the insectivores of Korea. . By J. i Jones, Jr., and David H. Johnson. Pp. 549-578. February 23, 1960 Speciation and evolution of the pygmy mice, ae Baiomys. By Robert L. Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960. Index Pp. 671-690. 1. 2. 8. Studies of birds killed in nocturnal migration. By Harrison B. Tordoff and Robert M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 1956. Comparative breeding behavior of Ammospiza caudacuta and A. maritima. By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956. The forest habitat of the University of Kansas Natural History Reservation. By Henry S. Fitch and Ronald R. McGregor.. Pp. 77-127, 2 plates, 7 figures in text, 4 tables. December 31, 1956. Aspects of reproduction and development in the prairie vole (Microtus ochro- helices Py Tey S. Fitch. Pp. 129-161, 8 figures in text, 4 tables. Decem- er Birds found on the Arctic slope of northern Alaska. By James W. Bee. Pp. 163-211,' pls. 9-10, 1 figure in text. March 12, 1958. 6. The wood rats of Colorado: distribution and ecology. By Robert B. Finley, Jr. Pp. 218-552, 34 plates, 8 figures in text, 85 tables. November 7, 1958. Home ranges and movements of the eastern cottontail in Kansas.. By Donald W. Janes. Pp. 553-572, 4 plates, 8 figures in text. May 4, (Continued on outside of back cover) 8, (Continued from inside of back cover) Natural history of the salamander, Aneides hardyi. By Richard F. Johnston and Schad Gerhard. Pp. 578-585. October 8, 1959. 9. x new subspecies of lizard, Cnemidophorus sacki, from Michoac&n, México. 10. By William E. Duellman. Pp. 587-598, 2 figures in text. May 2, 1960. A taxonomic study of the Middle American Snake, Pituophis deppei. By William E. Duellman. Pp. 599-612, 1 plate, 1 figure in text. May 2, 1960. . Index Pp. 611-626. Vol. 11. Nos. 1-10 and index. Pp, 1-703, 1958-1960. Vol. 12, 1, 2. 8. 4, Vol. 13. 1. 2 Vol. 14. 1. 2: Vol. 15. 1. Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry A. Vaughan. Pp. 1-158, 4 plates, 24 figures in text, July 8, 1959. The ancestry of modern Amphibia: a review of the Sa es By Theodore H. Eaton, Jr. Pp. 155-180, P10 figures in text. July 1 959. The baculum in microtine rodents. By Sydney Westies Pp. 181-216, 49 figures in text. February 19, 1960. A new order of fishlike Seb from the Pennsylvanian of Kansas. By Theodore H. ere Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in text. _May 2, More numbers abe I in volume 12. Five natural hybrid combinations in minnows (Cyprinidae). By Frank B. Cross and W. L. Minckley. Pp. 1-18. June 1, 1960. . A distributional study of the emplibians of the isthmus of Tehuaneae Misc By William E. Duellman. Pp. 19-72, pls. 1-8, 8 figs. August 1 A new subspecies of the slider turtle (Pseudemys scripta) from Coahuila, Merce Nos John M. Legler. Pp. 73-84, pls. 9-12, 8 figures in text. August Autecology of the Copperhead. By Henry S. Fitch. Pp. 85-288, pls, 18-20, 26 figures in text. November 30, 1960. Occurrence of the See Snake, Thamnophis sirtalis, in the Great Plains and Rocky Mountains. By Henry S. Fitch and T. Paul 'Meslin Pp. 289-308, 4 figures in text. February 10, 1961. Fishes of the Wakarusa River in Kansas. By James E. Deacon and Artie L, Metcalf. Pp. 809-322, 1 figure in text. February 10, 1961. Geographic variation in the North American Cyprinid Fish, Hybopsis gracilis. By Leonard J. Olund and Frank B. Cross. Pp. 323-348, pls. 21-24, 2 figures in text. February 10, 1961. Descriptions of two species of frogs, Genus Ptychohyla—studies of American hylid frogs, V. By William E. Duellman. Pp. 349-357, pl. 25, 2 figures in text. April 27, 1961. Fish populations, following a drought, in the Neosho and Marais des Cygnes rivers of Kansas. By James Everett Deacon. Pp. 359-427, pls. 26-30, figs. in text. August 11, i. More numbers will appear in volume 18. Neotropical bats from western: México. By Sydney Anderson. Pp, 1-8. October 24, 0. Geographic Oe ena in the harvest mouse Reithrodontomys megalotis on the central Great Plains and in adjacent regions. San at Snot Jones, Jr. eng B. Mursaloglu. Pp. 9-27, 1 figure in text. July 2 Mammals of Mesa vows National Park, Cae rs ‘Sydney ‘Atleeon Pp. 29-67, pls. 1-2, 8 figures in text. July 24, 1961. More numbers will appear in volume 14, The amphibians and reptiles of Michoacin, México. . By William E. Duell- man. Pp. 1-148, pls. 1-6, 11 figs. December 20, 1961. = “ BIN CIAS SLING pe) a NC ART UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Volume 15, No. 2, pp. 149-173 January 31, 1962 Some Reptiles and Amphibians from Korea BY ROBERT G. WEBB, J. KNOX JONES, JR., AND GEORGE W. BYERS UNIVERSITY OF KANSAS LAWRENCE ~ 1962 UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HIsTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 15, No. 2, pp. 149-173 Published January 31, 1962 UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY JEAN M. NEIBARGER, STATE PRINTER TOPEKA, KANSAS 1962 Some Reptiles and Amphibians from Korea’ 0 BY : ROBERT G. WEBB, J. KNOX JONES, JR., AND GEORGE W. BYERS In 1954, two of us (Jones and Byers) collected reptiles and amphibians in Korea incidental to field studies relating to hemor- rhagic fever. The 382 specimens thus obtained were deposited either in the Museum of Natural History of The University of Kansas (KU), or in the Museum of Zoology of the University of Michigan (UMMZ), and are the basis for the present report. Con- tinuous American military operations of one sort or another in Korea since 1945 have afforded opportunities for interested persons to obtain there collections of amphibians and reptiles, the study of which has resulted in several recent publications (Babb, 1955; Dixon, 1956; Hahn, 1959 and 1960; Shannon, 1956 and 1957; Stewart, 1958 and 1954; Tanner, 1953; Walley, 1958a and 1958b). This paper, which contains comments on the natural history and taxonomy of 22 species, all previously reported from Korea, supple- ments earlier studies, especially Shannon’s (1956) annotated list of the herpetofauna of the country. Shannon (loc. cit.) recorded 86 kinds of reptiles and amphibians from Korea. Subsequently, Bufo stejnegeri (previously omitted) was added by Shannon (1957), Takydromus takydromoides oldi was described by Walley (1958a), and Takydromus kwangakuen- sis was relegated to synonymy under T. amurensis by Walley (1958b). Presently, then, 87 kinds are on record from the Korean Peninsula. In the accounts beyond, Jones and Byers are mostly responsible for the remarks on natural history, whereas Webb is mostly re- sponsible for the taxonomic comments. The synonymies include (1) the original description, which is followed by (2) the first use of the name-combination here employed if it differs from the name as originally proposed, and (3) any synonyms having type localities in Korea. All measurements are in millimeters and all dates refer to the year 1954 unless otherwise indicated. A gazetteer of lo- calities mentioned in the text and a list of literature cited follow the accounts of species. We are grateful to the officers, enlisted men and civilians asso- ciated in 1954 with the Field Unit of the Commission on Hemor- (151) 152 UNIVERSITY OF KANnsAs PusLs., Mus. Nar. Hist. rhagic Fever, Armed Forces Epidemiological Board, who aided our efforts in Korea; we are especially mindful of the contributions of Dr. Albert A. Barber, Dr. Marshall Hertig, Mr. Louis J. Lipovsky and Dr. Warren D. Thomas. We are grateful also to Mr. Yoshinori Imaizumi, National Science Museum of Japan, for his translations of several papers in Japanese, and to Dr. Edward H. Taylor for making certain pertinent references available to us. Hynobius leechii Boulenger Hynobius Leechit Boulenger, Ann. Mag. Nat. Hist., ser. 5, 19:67, January, 1887 (type locality, Gensan [= Wonsan], Korea). Hynobius leechii quelpaertensis Mori, Jour. Choser Nat. Hist. Soc., 6:47 (Japanese) and 53 (English), March 25, 1928 (type locality, Quelpart Island [= Cheju Do], Korea). Specimens examined (3).—1 mi. SW Inje, 1 (KU); 4 mi. NNE Sogwi-ri, Cheju Do, 2 (KU). Description (KU 38774 from 1 mi. SW Inje).—Total length, 86 (head 18, body 40, tail 38); costal grooves (including axillary and inguinal), 18; two costal grooves between adpressed toes; length of inner branch of series of vomerine teeth less than distance between outer border of naris and peak of opposite series (tooth-rows V-shaped, approximately as long as broad); dorsal surface yellowish brown or buff (yellowish in life), having numerous blackish marks; venter yellowish cream, having an indistinct grayish mottling. Remarks.—The salamander described above was found in a fox- hole with another desiccated individual (not saved) on a military compound on April 24. The only other occurrence of H. leechii on the mainland to come to our attention was the report of several larvae that were seen in a small pool on a hillside near Chip’o-ri in the summer of 1953. Each of the two specimens from Cheju Do (KU 38775-76) differs from KU 88774 in having (1) the length of inner branch of the vomerine series slightly greater than the distance between outer border of naris and peak of opposite series (tooth-rows V-shaped, longer than broad), (2) a dorsal and ventral keel on the tail, (3) one costal groove (rather than two costal grooves) between adpressed toes, and (4) in being darker both dorsally and ventrally. The area of buff on the dorsal surface of each specimen is reduced by a fine, blackish mottling and stippling, and the venter of each is grayish. Respective total lengths of KU 38775 and 38776 are 84 (head 12, body 86, tail 36) and 89 (12, 35, 42), and the number of costal grooves 13 and 14. Although the two specimens are of approximately the same size, the tail of KU 38776 is noticeably the longer; the tail of KU 38775 is thicker and deeper than that of KU 38776. REPTILES AND AMPHIBIANS FROM KOREA 153 The two specimens from 4 mi. NNE Sogwi-ri were taken on Sep- tember 9 in damp substrate under volcanic rocks along a little-used road; although many rocks were overturned, only these two indi- viduals were found. A South Korean soldier informed us that salamanders were fairly common on Cheju Do. Mori (1928a:16) first mentioned in Japanese text the alleged distinctiveness of the salamander occurring on Cheju Do. Later, Mori (1928b:47 in Japanese, and 1928c:53 in English) provided valid descriptions of the subspecies, Hynobius leechii quelpaer- tensis. Okada (1934:17) questioned the validity of H. I. quelpaer- tensis and Sato (1943) regarded the salamanders of Cheju Do as inseparable from the Korean H. leechii. The English description of quelpaertensis is briefer than the preceding one in Japanese and lacks comparisons with related forms. KU 88775-76 seemingly differ appreciably from the description of quelpaertensis only in having the series of vomerine teeth narrowly V-shaped and longer than broad. We tentatively follow Sato in regarding quelpaertensis as a synonym of leechii. Bombina orientalis (Boulenger) Bombinator orientalis Boulenger, Ann. Mag. Nat. Hist., ser. 6, 5:148, pl. IX, fig. 2, February, 1890 (type locality restricted to Chefoo, China, by Pope, Bull. Amer. Mus. Nat. Hist., 61:435, August 29, 1981). Bombina orientalis, Stejneger, Bull. U. S. Nat. Mus., 58:51, figs. 30-43, pl. VII (reproduction from Boulenger, supra), July 22, 1907. Specimens examined (87).—2 mi. N Chip’o-ri, 8 (KU), 5 (UMMZ); 1 mi. SW Inje, 1 (KU); 15 mi. NE Mosulp’o, Cheju Do, 6 (KU); Sangdaehwa, 2 (KU); Taehoesan-ni, 1 (KU); 1 mi. W Tangjonggok, 32 (KU), 17 (UMMZ); Tangnim-ni, 2 (KU); 3 mi. SW Yanggu, 1 (KU); 2 mi. N Yongdae-ri, 8 (KU), 4 (UMMZ). Remarks.—Most of our specimens were taken from breeding con- gresses after heavy rains in rice fields and other shallow temporary waters. Thirteen individuals from Chip’o-ri were collected from foxholes around the edge of a military compound (two pairs in amplexus; no egg masses seen), and represent a small sample of frogs that were everywhere following a heavy rain on the night of May 18-14. On April 23, 32 Bombina orientalis and three Rana temporaria dybowskii were trapped in the water-filled bottom of an unused grease pit near Tangjonggok. Many tadpoles and two kinds of egg masses (small clusters and beadlike strings) were present; the small clusters of Bombina were commonest. On June 13 at the same locality, thousands of these toads were observed (hundreds in axillary amplexus) in foxholes, temporary rain pools, and backwashes along the Puk-ch’on [river]. On June 12 near 154 UNIVERSITY OF KANSAs Pusts., Mus. Nat. Hist. Yongdae-ri many individuals were seen (several pairs in amplexus), along Route 24 paralleling the Puk-ch’on, in rain pools and in ditches and backwashes from the river; almost all available water contained small (approximately 10 x 10 mm.) egg masses. Num- bers of eggs per mass, selected at random, were 5, 2, 2, 5, 2, 8, 8, 2 and 5. Some that were saved subsequently hatched on June 15-17. The call is a quiet low trill or series of staccato whistles rising slightly at the beginning; a short peeplike note also was heard. The specimens from Cheju Do, which are generally smaller than those collected on the mainland in spring, were taken on September 6 in a small stream that had large volcanic rocks in many places and that was flanked by thick brush and small trees. The earliest and latest dates on which B. orientalis was collected were April 21 and September 6, respectively. In the breeding season, males are distinguished from females by the large blackish (probably brownish earlier in season) areas on the anteroventral surface of the antebrachium, the metacarpal tubercle, and the inner surface of the first finger (sometimes also the second and third). Also, males have conspicuous black-tipped tubercles on the back (usually absent in females) that extend onto the limbs (usually smooth in females, at least laterally). Field observations by one of us (Byers) suggested that the dorsal pat- tern of males had greater contrast than that of females and that the venter was brighter reddish. Eight females from Tangjonggok averaged 47.9 (43-51) in snout-vent length, whereas 24 males from there averaged 50.0 (46-55), indicating little, if any, size difference between the sexes. Okada (1931:29) recorded variation in color of live Korean individuals (green or brown dorsally and pale yellow or red ven- trally) and variation in extent of black markings on the belly (op. cit.:fig. 12). The specimens from Cheju Do (28, 32, 32, 32, 37 and 46 in snout-vent length) have less black ventrally than speci- mens from the mainland. Bufo bufo gargarizans Cantor Bufo gargarizans Cantor, Ann. Mag. Nat. Hist., ser. 1, 9:483, August, 1842 (type locality, island of Chusan, China). Bufo bufo gargarizans, Stejneger, Bull. U. S. Nat. Mus., 58:59, July 22, 1907. Specimens examined (10).—Central National Forest, near Pup’yong-ni, 1 (KU); 5 mi. NW Choksong, near Imjin River, 1 (KU); 5 mi. E Seoul, 1 (KU); 6 mi. E Seoul, 4 (KU), 2 (UMMZ); 1 mi. S Yami-ri, 1 (KU). Description (nine females)—Dorsal surface brownish, having indistinct pale areas, some of which tend to form longitudinal stripes that extend pos- REPTILES AND AMPHIBIANS FROM KOREA 155 teriorly from parotoid glands; blackish mark, usually on lateral part of parotoid, having short spurs directed posteriorly and ventrally; edge of upper jaw and warts on dorsal surface becoming blackish with increasing size; small, con- spicuous group of warts near angle of jaw below parotoid; middorsal warts tending, at level of posterior edge of parotoids, to form a V that has its apex between the parotoids; ventral surface pale yellowish, sometimes having well- defined blackish marks; granular underparts of large specimens having small blackish tubercles. Male (KU 40118 from 5 mi. E Seoul).—Snout-vent length, 65; no vocal sacs or slits; dorsal and inner surfaces of first and second fingers, and inner surface of third finger black; canthus rostralis indistinct (a well-defined ridge on right side); nostrils closer to tip of snout than to eye, their distance from each other slightly less than interorbital width; interorbital width (6.2) greater than width of eyelid (4.7); tympanum distinct, circular, its diameter (8.0) less than length of eye (6.5), and approximately twice distance (1.6) of tympanum from eye; no cranial crests; parotoid gland elongate, approxi- mately twice as long as broad (12.5 x 5.0), narrowly separated from posterior edge of eyelid; head elongate (width at posterior edge of tympanum, 28.6); length from posterior edge of tympanum to tip of upper jaw, 18.9; first finger slightly longer than second, fourth finger about two-thirds as long as third; most subarticular tubercles divided; outer palmar tubercle larger than inner; heels not touching when folded legs placed at right angles to longitudinal axis of body; tibiotarsal articulation just reaching eye when leg laid forward; tarsometatarsal articulation not reaching beyond snout; foot large (tibiotarsal articulation to tip of fourth toe approximately 46.0); fourth toe approximately half webbed, other toes more than half webbed; edges of webs somewhat crenulate; some subarticular tubercles divided; length of inner metatarsal tubercle (4.5) more than half length of first toe (7.0); inner metatarsal tubercle larger than outer, both darkened; tarsal fold extending from inner metatarsal tubercle for approximately two-thirds length of tarsus; tips of toes (not fingers) darkened; dorsal surface of back and proximal part of hind legs coarsely granular, of rounded, pavement-type tubercles lacking sharp tips; small group of warts near angle of jaw below parotoids; dorsal pattern con- trasting and irregular (especially on limbs), of dark brown and pale gray; conspicuous black mark (interrupted) on lateral surface of parotoid having two, well-defined spurs that project posteroventrally; undersurface granular, lacking markings except for two indistinctly-margined dark spots on chest, and black spot on left leg. Remarks.—This nocturnal, introduced species (Okada, 1931:47) is presumably widespread in Korea and seemingly prefers lowland habitats. Individuals were taken in sparse vegetation on a sand flat near the Han River, at the edge of a rice field in a light rain, along a road at night, and in millet fields adjacent to the Han River, which was flooding at that time (July 9). As is obvious from the foregoing descriptions, the male (KU 40118), which was obtained on March 19, differs considerably from the nine females; neither does it agree with Stejneger’s (1907:66) 156 UNIVERSITY OF KANsAs PuBLs., Mus. Nat. Hist. or Okada’s (op. cit.:45-46, fig. 18) description of males of Bufo bufo asiaticus [= gargarizans] from Wonsan and Seoul. Upon cursory examination, KU 40118 is notable for having a contrasting dorsal pattern and elongate, ranidlike proportions. Some of the characteristics resemble those of Bufo raddei Strauch as given by Stejneger (op. cit.:70-72, figs. 53-57), Okada (1935:9, figs. 2 and 32-34, pls. II-III), and Liu (1950:203-205, fig. 43). Stejneger (op. cit.:59-68) recognized B. b. gargarizans as occur- ring in southern China, and Bufo bufo asiaticus as the subspecies occurring in northern China. Subsequently, asiaticus was relegated to synonymy under the earlier-named gargarizans—see discussions by Pope and Boring (1940:33) and Liu (op. cit.:220). Kaloula borealis (Barbour) Cacopoides borealis Barbour, Bull. Mus. Comp. Zool., 51(12):231, April, 1908 (type locality, Antung, Manchuria). Kaloula borealis, Noble, Amer. Mus. Novit., 165:6, April 16, 1925. Specimens examined (8).—5 mi. ESE Seoul, 1 (KU); 6 mi. E Seoul, 8 (UMMZ); 7 mi. ESE Seoul, 4 (KU). Remarks.—On April 5, one male and three females were un- covered by a bulldozer from between one and two feet below the surface of the ground in an old Korean burial mound; one individual was completely surrounded by compact soil. All quickly became active when placed in water. Two of the females (48 and 44 in snout-vent length) contained masses of immature eggs. A male obtained on June 4 was found during a rain; the three UMMZ specimens were obtained on July 8 on banks above the Han River. Breeding of this species seems to coincide with the rainy season in late spring and early summer when males were noted calling around flooded ditches and swales in deep grass. The local Korean name of the species, which sounds something like “maeng-kongi,” is said to come from the call, which is best described as a monoto- nous, snoring sound that rarely is heard in two parts as suggested by the name. Hyla arborea japonica Giinther [Hyla arborea] Var. japonica Giinther, Catalogue of the Batrachia Salientia in the ‘ British Museum, p. 109, 1858 (type locality, Japan). Hyla pire var. savignyi Boulenger, Ann. Mag. Nat. Hist., ser. 5, 19:67, January, 1887 (type locality, Gensan [= Wonsan], Korea}, Hyla stepheni Boulenger, Proc. Zool. Soc. London, p. 579, pl. 51, fig. 1 (for 1887), April, 1888 (type locality, Port Hamilton, a small island between Korea and Japan). Specimens examined (44).—Central National Forest, near Pup’yong-ni, 2 (KU); 8 mi. NW Chip’o-ri, 1 (KU); 1 mi. N Mosulp’o, Cheju Do, 1 (KU); REPTILES AND AMPHIBIANS FROM KOREA 157 1 mi. NW Oho-ri, 13 (KU), 3 (UMMZ); Sangbonch’on-ni, 2 (UMMZ); 5 mi. ESE Seoul, 8 (KU); 6 mi. E Seoul, 10 (KU), 3 (UMMZ); 7 mi. NNE Sogwi-ri, Cheju Do, 1 (UMMZ). Remarks.—Hylids from 5 mi. ESE Seoul were collected from a cement-walled pit at the Seoul City Water Works; a specimen of Elaphe rufodorsata taken in the pit had eaten one hylid. Two individuals were taken in the morning of May 29 on leaf litter in a wooded valley in the Central National Forest where a number were calling in a light rain, but the species was rarely found in woods. Two frogs were found along a rocky stream at Sang- bonch’on-ni. Most individuals were taken while calling, on grasses and reeds or on the ground, along the edges of rice fields. Sixteen hylids collected 1 mi. NW Oho-ri were calling in shallow water of a rice field on a hillside, but none was heard in a large lake nearby or in adjacent fields. On May 15, 4 mi. ESE Ch’orwon, approxi- mately one hundred tadpoles, thought to be of this species, congre- gated near a drain (into a lower field) of an unused rice field; the tadpoles were well-developed, some having hind legs. The earliest and latest dates of collection represented in our material are May 8 and October 29. The call is best described as a raspy “waak,” “week,” or “wiick” in the middle register. The listing of “Hylae arboreae var japonicae descript pars Schleg. in Fauna Japon. p. 112 . . .” by Giinther (1858:81) in syn- onymy under the account of Polypedates schlegelii, implies that Schlegel was the first author to use the name-combination Hyla arborea japonica. Boulenger (1882:86, 381) went so far as to credit Schlegel as the author of the name japonica. The reason for this action is not known because Schlegel (in von Siebold, 1838:112) referred to this hylid only under the name “Hyla arborea.” Rana rugosa Schlegel Rana rugosa Schlegel, Reptilia [Saurii et Batrachii], in von Siebold, Fauna Japonica, p. 110, pl. 3, figs. 3-4, 1838 (type locality designated as Japan, probably near Nagasaki, by Stejneger, Bull. U. S. Nat. Mus., 58:128, July 22, 1907). Specimens examined (26).—Central National Forest, near Pup’yong-ni, 13 (KU), 1 (UMMZ); 2 mi. S Ch’orwon, 4 (KU); 2 mi. E Hoengsong, 8 (KU); 1 mi. NW Oho-ri, 1 (KU); 5 mi. ESE Seoul, 1 (KU); 6 mi. E Seoul, 3 (UMMZ). Remarks.—Rana rugosa was associated with Rana nigromaculata and Rana amurensis coreana at all localities where the species was taken save at 2 mi. E Hoengsong, where R. a. coreana was not observed. Three specimens of R. rugosa were collected among 158 UNIVERSITY OF KANSAS PusLs., Mus. Nat. Hist. grasses and reeds in water along the edge of Ch’orwon Reservoir, 2 mi. S Ch’orwon, where they were difficult to find in the thick vegetation even though their low, soft calls were heard; the speci- men from 1 mi. NW Oho-ri was found in a rice field. Otherwise, habitats recorded indicate a preference for small, fast-flowing streams, especially in wooded valleys. On one occasion, indi- viduals were found trapped in cement-walled pits about old ruins on a wooded hillside in the Central National Forest. The earliest and latest dates of capture among our specimens are May 15 and November 6. In addition to the localities listed above, the species was observed 4 mi. W Ch’ungju. R. rugosa may have an extensive breeding season as suggested by the variation in size of frogs collected or observed in 1954. Of nine frogs obtained on May 29, five ranged in snout-vent length from 26 to 28, and four from 42 to 54. Three specimens collected on October 9 measured 89, 41, and 55, and two obtained on No- vember 6 measured 25 and 37. Rana nigromaculata Hallowell Rana nigromaculata Hallowell, Proc. Acad. Nat. Sci. Philadelphia, [12]:500 (tor 1860), 1861 (type locality, Simoda, Japan). Specimens examined (47).—Central National Forest, near Pup’yong-ni, 6 (KU); 2 mi. S Ch’orwon, 5 (KU); 4 mi. W Ch’ungju, 2 (KU); 7 mi. W Ch’ungju, 1 (KU); 2 mi. E Hoengsong, 1 (KU); 8 mi. SW Kunsan, 1 (KU); 1 mi. NW Oho-ri, 5 (KU); 5 mi. ENE Pusan, 2 (KU); 5 mi. ESE Seoul, 9 (KU); 6 mi. E Seoul, 3 (KU), 10 (UMMZ); 6 mi. NNE Sogwi-ri, Cheju Do, DK) Description——Back brownish or grayish (greenish in life), having few, indistinct or well-defined, dark blotches, or extensive blackish areas (repre- senting fusion of markings); pale, middorsal stripe (pale green, buff or yellowish in life) from snout to anus, sometimes as wide as interorbital width, but indistinct or lacking when pattern on back absent; blackish bar often present behind tympanum; posterior surface of thigh mottled; underparts pale yellow to whitish, sometimes having a few dusky marks on throat; longitudinal ridges between dorsolateral folds indistinct in some small frogs; largest female and male having respective snout-vent lengths of 100 and 70. Remarks.—Rana nigromaculata is the most abundant ranid in central Korea and, in a general way, the ecological equivalent of Rana pipiens in temperate North America. The species is associated with most aquatic habitats, from rocky streams to rice fields and large impoundments. In the vicinity of Seoul the din of large breeding congresses was heard more or less continuously from mid-April to mid-May. Large numbers of juveniles (approximately one inch long) were noticed first on July 8 and were present REPTILES AND AMPHIBIANS FROM KOREA 159 thereafter for about three weeks, being commonest in standing water after heavy rains or during prolonged showers. These data and the different sizes of individuals collected at the same time suggest either variable growth or, more probably, an extensive breeding season. Our earliest and latest dates of collection are April 16 and October 7. The Korean name for “frog,” most often applied to R. nigromaculata, sounds something like “keg-oh-ree.” The call is a prolonged, raspy, staccato croak, sometimes with a rising inflection at the end. In addition to the localities listed above, the species was observed 5 mi. W Kwangju and 3 mi. S Osan. Despite a high degree of individual variation, Rana nigromaculata seemingly varies geographically as well; some subspecies probably should be recognized, but the species as a whole has never been thoroughly studied systematically. The division of R. nigromacu- lata into three subspecies by Schmidt (1927:563-567) was con- sidered untenable by Fang and Chang (1931:95-98), and it has been regarded by most recent authors as a variable, monotypic species. The named subspecies R. n. chosenica (Okada, 1931:89, with type locality at Seoul, and geographically restricted to Korea) was considered a subspecies of Rana plancyi by Shannon (1956:36). The most trenchant characters of planyci seem to be the wide dorso- lateral folds, the uniform greenish dorsum, the presence of dermal pustules on the back between the dorsolateral folds, and the lack of a mottled pattern on the posterior surface of the thigh. Among our specimens of R. nigromaculata, the width of the dorsolateral folds is variable, a uniform greenish dorsum is found only in large males, dermal pustules are mixed with ridges in only one male (KU 38733), and all have a spotted or mottled pattern on the posterior surface of the thigh. All of our specimens having an indistinct pattern on the back, or lacking a pattern, are males and resemble the photographs of males published by Moriya (1954: pl. I, fig. 5) and Liu (1936: pl. IV, figs. 1-2); juveniles of both sexes and large females have contrasting patterns. None of our frogs seems, therefore, clearly referable to the species plancyi, although some characters are suggestive of plancyi. Moriya (op. cit.:19), who studied variation of R. nigromaculata in Japan, noted that one of the most distinct populations there (R. n. brevipoda) resembled Rana plancyi. Ting (1939) discovered that nigromaculata and plancyi were cross-fertile and raised hybrid 160 UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist. larvae through metamorphosis. Pope and Boring (1940) suggested hybridization between the two species in eastern China, and the above mentioned facts suggest to us the possibility of hybridization in other regions. Rana amurensis coreana Okada Rana temporaria coreana Okada, Annot. Zool. Japon., 11:140 (footnote), July 25, 1927, nomen nudum. Rana temporaria coreana Okada, Jour. Chosen Nat. Hist. Soc., 6:19, pl. 1, fig. 7, 1928 (type locality, Keijo [= Seoul], Korea). Rana amurensis coreana, Shannon, Herpetologica, 12:38, March 6, 1956. Specimens examined (9).—Central National Forest, near Pup’yong-ni, 1 (KU); 2 mi. S Ch’orwon, 1 (KU); 4 mi. W Ch’ungju, 1 (KU); 1 mi. N Oho-ri, 1 (KU); 5 mi. ESE Seoul, 8 (KU); Yongp’yong, 2 (KU). Remarks.—One individual of R. a. coreana (KU 38698) and one of Rana temporaria dybowskii (KU 38715) were collected on May 29 along a stream in the Central] National Forest in association with R. rugosa and R. nigromaculata. Specimens of coreana from the vicinity of Seoul were found in rice fields. The earliest date of collection was April 13 at Yongp’yong. Our largest specimen of coreana measured 47 in snout-vent length. Because R. a. coreana and Rana temporaria dybowskii are sym- patric in central Korea and closely resemble one another, the two species were not distinguished in the field and the following obser- vations may pertain to either (or both) species. Wood frogs were observed 2 mi. E Songdong-ni on July 12 in paddies (rice fields) along with individuals of Hyla arborea, Rana rugosa, and Rana nigromaculata. At Chip’o-ri on April 6, individuals (probably R. t. dybowskii) were seen in a seepage pool from an abandoned rice field; R. nigromaculata also was seen there. Six or seven egg masses (some having small tadpoles) were observed in the shallow water, but it was not certain to which species the eggs belonged. Completely metamorphosed young (probably R. a. coreana) were first seen 1 mi. N Oho-ri on June 9. At Taehoesan-ni on November 12, several sluggish frogs were seen in a small pool that was cov- ered by a thin layer of ice. On September 26 in the Central National Forest, many wood frogs of various sizes were observed. R. amurensis and R. tempo- raria probably have extended breeding seasons that correspond to those of R. rugosa and R. nigromaculata. Judging from our obser- vations, amurensis prefers the proximity of water, whereas tempo- raria may occur some distance from permanent water. In our specimens, R. amurensis coreana differs from R. temporaria REPTILES AND AMPHIBIANS FROM KOREA 161 dybowskii in having (1) smaller maximal size, (2) more slender body, (8) shorter legs, (4) incompletely webbed toes, (5) no mottling or barring on lips, (6) no contrasting barred pattern on hind legs, (7) dark brown stripes (usually) between dorsolateral folds, (8) a dark brown, linear mark below canthus, and (9) an immaculate ventral surface. Two additional distinguishing char- acters, which we found difficult to evaluate, are the nearly straight, dorsolateral folds, and lack of vocal sacs or ostia in males of R. a. coreana (Shannon, 1956:38). Some of the differences between the two species were illustrated by Okada, 1931:107, fig. 48, R. temporaria temporaria [= R. t. dybowskii] and 128, fig. 54, R. temporaria coreana [= R. amurensis coreana]. Rana temporaria dybowskii Giinther Rana Dybowskii Giinther, Ann. Mag. Nat. Hist., ser. 4, 17:387, May, 1876 (type locality, Abrek Bay, near Vladivostok, Siberia). Rana temporaria dybowski, Shannon, Herpetologica, 12:38, March 6, 1956. Specimens examined (20).—Central National Forest, near Pup’yong-ni, 7 (KU Y Chip’o-ri, 2 (KU); 1 mi. SW Inje, 6 (KU); 8 mi. SW Kangnung, 1 (KU); Taegwang-ni, 1 (KU); 1 mi. SW Tangjonggok, 3 (KU). Remarks.—On October 9 in the Central National Forest, five individuals were found in a concrete-walled pit in old ruins on a wooded hillside; no specimens of Rana amurensis coreana were taken there. R. ¢. dybowskii was most often taken on high, moist slopes, and seemed to be especially common in forests. The speci- men from 8 mi. SW Kangnung was obtained in a wooded area along a mountain stream. The earliest date of collection of a specimen of dybowskii was March 7 at Taegwang-ni. See also the remarks under the preceding account of Rana amurensis coreana. The largest male among our specimens measured 65 in snout- vent length and the largest female, 79. Five gravid females had snout-vent lengths of 64, 68, 69, 69 and 70. Trionyx sinensis Wiegmann Trionyx (Aspidonectes) sinensis Wiegmann, Nova Acta Acad. Leopold.- Carol., 17:189, 1835 (type locality, near Macao, China). Specimen examined.—Han River, 5 mi. ESE Seoul, 1 (KU). Remarks.—Our only specimen was purchased from a man who had captured it by hand in the Han River; it was the only turtle seen during our stay in Korea. Koreans eat turtles, and the elaborate (and relatively permanent) fish-traps that they construct across streams and small rivers probably reduce the size of populations of T. sinensis and other species. 162 Untversiry OF Kansas Pusts., Mus. Nar. Hist. Eremias argus Peters Eremias argus Peters, Monatsber. preuss. Akad. Wiss., Berlin, p. 61, fig. 3 (for 1869), 1870 (type locality, Chefoo, China). Specimens examined (23).—Chip’o-ri, 1 (KU); 5 mi. E Seoul, 3 (KU), 3 (UMMZ); 5 mi. ESE Seoul, 4 (KU); 6 mi. E Seoul, 10 (UMMZ); 7 mi. ESE Seoul, 2 (KU). Remarks.—Individuals of E. argus were most often seen on dry hillsides having a relatively sparse cover of vegetation. The first lizard of this species was taken on April 2. In 1954 the last part of March and early part of April were generally warm, with tem- peratures above 70° F. on several occasions; probably some E. argus were active in late March. KU 38773 (snout-vent length, 51) laid three eggs between June 4 and 14; KU 38768 (snout-vent length, 58) obtained on May 8 was gravid, containing four eggs. Testes of lizards in the breeding season measure approximately 4.0 x 2.5 (KU 88772, obtained on June 16). The snout-vent length of our largest female is 61, that of the largest male, 57. The snout-vent length of 11 specimens averaged 77 (67-96) per cent of length of tail. Tachydromus amurensis Peters Tachydromus amurensis Peters, Sitzungsber. Gesell. naturf. Freunde Berlin, p. 71, 1881 (type locality, Kossakewitcha, Amurland). Specimens examined (8).—Central National Forest, near Pup’yong-ni, 1 (KU), 1 (UMMZ); Majon-ni, 1 (KU). Remarks.—One of our specimens was found among grasses along a small stream in the Central National Forest. The other two were obtained by other persons and we lack knowledge of conditions of their capture. A juvenile (KU 39416, snout-vent length, 25) that was obtained on September 9 is tentatively referred to this species. There seem to be three femoral pores on the left leg but the number is indistinct on the right. The specimen is dark and lacks a pattern. Its condi- tion precludes counts of ventral scales (not keeled), but scalation is otherwise the same as a male (KU 40120, snout-vent length ap- proximately 47, length of tail, 124). The third specimen, a male (UMMZ 113442, snout-vent length, 51, length of tail, 115), agrees with KU 40120, except in having 82 instead of 29 dorsal scales at midbody, 4-4 instead of 3-3 femoral pores, and in lacking a pale stripe from eye through ear to shoulder. REPTILES AND AMPHIBIANS FROM KOREA 163 Tachydromus wolteri Fischer Tachydromus Wolteri Fischer, Jahrb. Wiss. Anst. Hamburg, 2:82 (for 1884), 1885 (type locality, Chemulp’ o, Korea). Specimens examined (2).—Yongp’yong, 2 (KU). Remarks.—On April 14, two females (57 and 45 in snout-vent length, the tail of the latter measuring 103) were easily captured by hand on a burned-over rice field. Lygosoma reevesii (Gray) Tiliqua Reevesti Gray, Ann. [Mag.] Nat. Hist., ser. 1, 2:292, December, 1838 (type locality, China). [Lygosoma (Liolepisma) laterale] var. reevesi, Boettger, Katalog der Ba- trachier-Sammlung . . ., p. 104, 3. Specimens examined (6).—Central National Forest, near Pup’yong-ni, 3 (KU); 4 mi. NNE Sogwi-ri, Cheju Do, 1 (KU); 7 mi. NNE Sogwi-ri, Cheju Do, 1 (UMMZ); 16 mi. NE Mosulp’o, Cheju Do, 1 (UMMZ). Remarks.—On October 23 an individual was captured while sunning on a stump on a wooded hillside in the Central National Forest; two others at this locality were collected on damp ground- cover on the same hillside. A juvenile from Cheju Do was found among moss-covered rocks in a stream bed; the other specimens from Cheju Do were found among moss-covered rocks on the western slope of Halla San. Each ovary of a female obtained on October 23 contained five enlarged follicles, about 1 mm. in diameter. The left testis of a male obtained on August 10 seemed enlarged, indicating possible sexual activity, and measured approximately 6 x 2mm. The snout- vent length of our largest male is 41, that of our largest female, 48. The prefrontals are in contact in all of our specimens save one (UMMZ 113446). There is disagreement among herpetologists concerning the generic name of the small lygosome skink in the United States and its ecological equivalent in China and Korea. We tentatively use Lygosoma (Conant, 1951:207-208), although Mittleman (1950) pointed out reasons for using Scincella. Shannon (1956:41) dis- cussed the debated issue whether or not the lygosome skinks of the New and Old worlds are conspecific. 164 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. Rhabdophis tigrina lateralis (Berthold) Tropidonotus lateralis Berthold, Nachrichten Gesell. Wiss. Gottingen, p. 180, 1859 (type locality, China). Specimens examined (26).—Central National Forest, near Pup’yong-ni, 1 (KU); 2 mi. N Chip’o-ri, 2 (KU); 8 mi. NW Chip’o-ri, 4 (KU); 4 mi. N Ch’onan, 1 (KU); 3 mi. S Kumhwa, 1 (KU); 1 mi. SW Naegong-ni, 1 (KU); 4 mi. E Seoul, 1 (KU); 5 mi. E Seoul, 2 (UMMZ); 5 mi. ESE Seoul, 4 (KU); 6 mi. E Seoul, 1 (KU), 4 (UMMZ); 7 mi. ESE Seoul, 1 (KU); 6 mi. NNE Sogwi-ri, Cheju Do, 2 (KU); 5 mi. NE Taejon, 1 (KU). Remarks.—This common, vagrant species was found on brushy hillsides, near buildings on hills above rice fields, in tall grasses near streams, in rice fields, and along drainage and irrigation ditches. The earliest and latest dates of collection were April 5 and Novem- ber 7. On the first date mentioned an individual was found in hibernation with five Elaphe rufodorsata and one Agkistrodon halys in an earthen Korean burial mound. The specimen was uncovered by a bulldozer at a depth of about one foot below the surface. We were told that 18 snakes of this species were found in the same place (7 mi. ESE Seoul) the previous winter. The stomach of each of four individuals contained one Rana nigromaculata. The stomach of another individual contained a R. nigromaculata and remains of a carabid beetle, whereas another contained three small, partially-digested frogs that appeared to be Hyla arborea. P. M. Youngman reported to us that he found a snake of this species that was attempting to swallow a toad, Bufo bufo gargarizans. One of the small individuals from Cheju Do was being eaten by a Zamenis spinalis when found. One specimen was parasitized by three nematodes, Kalicephalus natricis (see Olsen, 1957:208). Two females of this oviparous species (lengths of body, 680 and 700) collected on May 14 contained nine eggs (18 mm. long), and 13 eggs (15 mm.) respectively; a third (length of body, 610) ob- tained on June 26 contained 10 eggs that were approximately 18 mm. long. A female (UMMZ 113458, length of body, 710), which was captured on July 10 and kept alive in captivity, laid 11 eggs on August 12 between 9 and 10 in the morning. The weight of nine of these eggs averaged 3.82 (8.0-8.6) grams; the last two eggs deposited were small and weighed only 1.3 and 1.4 grams. The eggs were incubated unsuccessfully. One that was opened on September 14 and another opened on September 26 contained young easily recognized as of this species. In captivity the parent REPTILES AND AMPHIBIANS FROM KOREA 165 snake underwent ecdysis on about July 20 and again on August 26. Our largest female and largest male have respective total lengths of 1013 (840 + 173) and 740 (575 + 165). Our smallest specimens, captured on September 9, measured 215 and 230 mm. in length of body, and probably represent young of the year. The snake found in hibernation on April 5 measured 275 in length of body. The ventrals of 11 males averaged 161.3 (158-171) and those of 14 females, 165.1 (160-170); subcaudals of eight males averaged 69.6 (66-74) and those of 14 females, 61.5 (52-73). Males seem to have small scales in the anal region that are more strongly keeled than scales elsewhere on the body (the scales catch on finger tips when rubbed in a posteroanterior direction), but males lack small tubercles on the upper and lateral parts of the head as mentioned by Maslin (1950:433). The comments of the same author (op. cit.:434) concerning integumental poison glands in the nuchal region of this species are of interest in view of several reports that we received of swollen extremities resulting from handling snakes of this species. In using the generic names Rhabdophis and Amphiesma for species formerly placed in the genus Natrix, we follow Malnate (1960), who divided Natrix (auct.) into five distinct genera. Amphiesma vibakari ruthveni (Van Denburgh) Natrix vibakari ruthveni Van Denburgh, Proc. California Acad. Sci., ser. 4, 13(2):8, July 26, 1923 (type locality, Pusan, Korea). Specimens examined (5).—Central National Forest, near Pup’yong-ni, 1 (KU); 4 mi. SW Ch’ongyang-ni, 1 (KU); 10 mi. NE Mosulp’o, Cheju Do, 1 (UMMZ); 6-7 mi. NNE Sogwi-ri, Cheju Do, 1 (KU), 1 (UMMZ). Remarks.—The specimen from the Central National Forest was captured on August 18 near a stream on a damp ground-cover of leaves. The specimens from Cheju Do were taken in early Septem- ber, one in a grassy area, and the other two on earthen banks of road-cuts on the slopes of Halla San. The stomach of one individual from Cheju Do contained an earthworm. Our largest specimen, a male having 154 ventrals and 68 subcaudals, measured 508 (380 + 128). The subcaudal counts of 68 (KU 38861) and 69 (UMMZ 113461) on two males from Cheju Do are higher than the maximal count known for the subspecies ruthveni in Korea, and resemble those of Amphiesma vibakari vibakari of the Japanese islands. The sub- caudals average 61 (55-65) in ruthveni and 71 (63-83) in vibakari 166 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. according to Van Denburgh (1923:3-4). A juvenile from the Central National Forest (KU 38862), lacking the tip of the tail, has 64 subcaudals. Dinodon rufozonatum (Cantor) Lycodon rufo-zonatus Cantor, Ann. Mag. Nat. Hist., ser. 1, 9:488, August, 1842 (type locality, island of Chusan, China). Dinodon rufozonatus, Peters, Sitzungsber. Gesell. naturf, Freunde Berlin, p. 89, 1881. Specimens examined (4).—Central National Forest, near Pup’yong-ni, 3 (KU); Yongsan (Seoul), 1 (UMMZ). Remarks.—The three specimens from the Central National Forest were taken in the period August 12-26. Two were caught in live- traps set for small mammals in deep forest among granite outcrops. The specimen from Yongsan was obtained on October 27 in a partly wooded area. Ventrals and subcaudals of our four specimens (all males) numbered, respectively, 198, 200, 198, 205, and 74, 75, 75, —. Total length of the largest specimen was 960 (790 + 170). We follow Chang (1932:54) and most subsequent authors in regarding D. rufozonatum as a monotypic species. Zamenis spinalis (Peters) Masticophis spinalis Peters, Monatsber. preuss. Akad. Wiss., Berlin, p. 91 (for 1866), 1867 (type locality, unknown—“Mexico” erroneously listed). Zamenis spinalis, Giinther, Ann. Mag. Nat. Hist., ser. 4, 9:22, January, 1872. Specimens examined (2).—5 mi. ESE Seoul, 1 (KU); 6 mi. NNE Sogwi-ri, Cheju Do, 1 (KU). Remarks.—The specimen from Cheju Do was captured on Sep- tember 9 in tall grass near a small stream and was eating a small Rhabdophis tigrina. The female from near Seoul was obtained from a Korean on June 10, and was gravid (six eggs, each approxi- mately 35 mm. in length). The length of body measured approxi- mately 550 and the length of incomplete tail 168 in one specimen (KU 38777, female from 5 mi. ESE Seoul), 540 and 183 in the other (KU 38778, female from Cheju Do). Respective ventral and subcaudal counts of the two females are 204, 194, and 74 ++, 86. There is some disagreement in the literature as to the proper generic name of this snake. Differences in dentition between Old World species (referable to Zamenis) and the American species (referable to Coluber) are discussed by Bogert and Oliver (1945: 365). The species spinalis has been referred to Coluber by several authors (see Pope, 1935:226). REPTILES AND AMPHIBIANS FROM KOREA 167 Elaphe dione (Pallas) Coluber dione Pallas, Reise durch verschiedene Provinzen des Russischen Reichs, 2:717, 1773 (type locality, “Salt steppes toward the Caspian Sea according to Stejneger, Bull. U. S. Nat. Mus., 58:315, July 22, 1907). Elaphis dione, Duméril and Bibron, Erpétologie générale . . ., 7:248, 1854. Specimens examined (10).—Choksong, 1 (KU); 4 mi. N Ch’onan, 1 (KU); Seoul, 1 (KU); 5 mi. E Seoul, 1 (KU), 2 (UMMZ); 5 mi. ESE Seoul, 1 (KU); 6 mi. E Seoul, 1 (UMMZ); Taegwang-ni, 1 (KU); 2 mi. WSW Tongjonggok, CK). Remarks.—This species seemingly occurs in upland habitats. Specimens were taken on rocky hillsides, on sparsely wooded hill- sides, and in cultivated fields. November 21 was the latest date of capture of an active individual (UMMZ 113451), the head of which was seen many times prior to capture protruding from a hole be- neath the concrete floor of a building. A female (KU 38855), measuring 915 (775 + 140) in total length, and obtained on June 13, contained nine eggs (32 mm. long). One juvenile had eaten a half-grown house mouse, Mus musculus; the stomach of a male con- tained three mice, one a striped field mouse, Apodemus agrarius, the other two probably also of that species but too far digested for certain identification. Eggs probably hatch in late summer. A young of the year (length of body, 340) was captured on September 30; another juvenile (length of body, 285) was obtained in May. Our largest male (KU 40123) measured 904 (719 + 185) in total length. Ventrals and subcaudals of six females averaged 205.8 (198-211) and 62.2 (55-69), respectively, whereas corresponding counts of four males averaged 196.8 (190-214), and 71.0 (69-74). Each of nine specimens had dorsal scales in 23-25-19 rows except one (UMMZ 118451), which had 23-25-23 rows. Elaphe rufodorsata (Cantor) Tropidonotus rufodorsatus Cantor, Ann. Mag. Nat. Hist., ser. 1, 9:483, August, 1842 (type locality, island of Chusan, China). Elaphe rufodorsata, Stejneger, Bull. U. S. Nat. Mus., 58:310, figs. 269-271, July 22, 1907. Specimens examined (27).—7 mi. NW Changhowan-ni, 1 (KU); 3 mi. NW Chip’o-ri, 8 (KU); 7 mi. W Ch’ungju, 2 (KU), 1 (UMMZ); 8 mi. S Kumhwa, 2 (KU); 1 mi. NW Oho-ri, 1 (KU); 4 mi. E Seoul, 1 (KU); 5 mi. E Seoul, 2 (KU); 5 mi. ESE Seoul, 2 (KU); 6 mi. E Seoul, 3 (KU), 2 (UMMZ); 7 mi. ESE Seoul, 5 (KU); 4 mi. N Uijongbu, 1 (KU); 5 mi. NE Uijongbu, 1 (UMMZ). Remarks.—E. rufodorsata was commonly observed and collected on barren hillsides, on country roads, in rice fields, and along drain- 168 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. age ditches and small streams. One was found sunning outstretched on aroad. Two individuals were trapped in cement-walled pits at the Seoul City Water Works. On April 5, five snakes of this species with one Rhabdophis tigrina and one Agkistrodon halys, all partly caked with earth, were found sunning in a shallow depression on the side of a Korean burial mound, which was presumably a hiber- naculum. Aside from one juvenile, four of the E. rufodorsata were of approximately the same size, having bodies ranging in length from 385 to 455. Copulation was observed on April 25 (male, KU 38811, length of body, 400, and female, KU 38812, length of body, 565), and on May 4 (female, KU 38816, length of body, 620). Eggs doubtless hatch at various times in summer. One of five snakes obtained on April 5 (see above) measured 310 (250-+ 60) in total length. Another juvenile (KU 38828), obtained on October 18, was 478 (385 +93) long, and our smallest specimen of this species (KU 38821), captured on June 26, measured 275 (230 + 45). The stomachs of two snakes each contained a Rana nigromacu- lata; another individual had eaten a Hyla arborea, and a fourth specimen had eaten a small fish. One specimen was parasitized by a cestode. The largest female from our series (KU 38816) measured 740 (620 + 120), and the largest male (KU 38813), 595 (475 + 120). Respective ventral and subcaudal counts of 13 males averaged 170.5 (167-174) and 60.0 (56-68), ventrals of 12 females averaged 178.3 (169-182), and subcaudals of 11 averaged 51.0 (46-56). Elaphe schrencki anomala (Boulenger) Coluber anomalus Boulenger, Ann. Mag. Nat. Hist., ser. 8, 17:243, March, 1916 (type locality, Chihfeng, China). Elaphe schrencki anomala, Pope, The reptiles of China, p. 266, fig. 57, May 11, 1985. Specimens examined (7).—Central National Forest, near Pup’yong-ni, 2 (KU), 1 (UMMZ); 4% mi. W Chip’o-ri, 1 (KU); 5 mi. N P’yong-taek, 1 (KU); 5 mi. E Seoul, 1 (KU); 5 mi. ESE Seoul, 1 (KU). Remarks.—Individuals were observed or taken on dry, scrubby hillsides and in grassy upland areas. One of the three snakes from the Central National Forest was captured on a steep, forested hill- side among granite outcroppings; another was obtained there along a stream bank and had eaten three bats, Murina aurata (see Jones, 1960:265), and one mouse, Apodemus sp. (tail only found). P. M. Youngman reported (personal communication ) finding a rat (Rattus sp.) in the stomach of one individual. A female (KU 38830, length REPTILES AND AMPHIBIANS FROM KOREA 169 of body, 1180) that was obtained on June 2 contained 17 eggs, each approximately 82 mm. long. The ventrals of two females numbered 223 and 229, and the subcaudals of the latter 70. Ventrals and subcaudals of five males were, respectively, 211, 213, 214, 215, 216, and 71,775,075; 6937/5. The coloration and pattern of our seven specimens are of interest in view of the probable intergradation between E. s. anomala and E. s. schrencki in northern Korea (see comments by Shannon, 1956: 46). The smallest specimen (KU 38831), having a total length of 335 (280 + 55), was obtained 4 mi. N P’yong-taek on September 24, It is nearly uniform pale brown (lacks a dorsal pattern) and additionally is characterized as follows: incomplete pattern on the head; no black postocular band (pale brown with black posterior border); ventrolateral extensions of the head pattern that form longitudinal stripes of white on the third row of scales; a pale whitish stripe on the sixth and seventh scale rows that extends posteriorly to the level of the fortieth ventral and that has a narrow black border (sometimes interrupted ); small and indistinct blackish markings and pale stripes on sides (no higher than sixth row of scales); underside of the head whitish; and venter grayish, having blackish margins on the ends of ventrals posteriorly. KU 388831 is unusual and perhaps anomalous in having a pattern that does not conform to the juvenal pattern of either subspecies. A female (KU 38830), having a total length of 1390 (1180 + 210), from 5 mi. ESE Seoul conforms to descriptions of anomala in being uniformly pale brown above and in having indistinct dark smudges on the sides; the ventral surface is whitish having indistinct dark smudges, brown spots at the ends of each ventral, and the posterior edge of each ventral brown. A male (KU 40125), measuring 1090 (890 + 200) in total length, from 5 mi. E Seoul, is pale brown above and lacks markings on the anterior part of the body. Indistinct dark markings occur at midbody, whereas the posterior quarter of the body and tail have well-defined black bands on a buff back- ground. The black bands posteriorly are arranged in pairs; each pair of bands is separated by two and a half to three scales, whereas the bands of each pair are separated by only one and a half scales. The ventral surface has an obscure marbled pattern. Our largest specimen, a male (UMMZ 113454) having a total length of 1488 (1230 + 258), from the Central National Forest, resembles KU 40125, except that pale brown blotches (29 on body, one blackish on neck) and dark lateral spots occur anteriorly on the body. 170 UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist. Another female (KU 88860, body length, 970) from 4% mi. W Chip’o-ri, our northernmost locality of record, has a fairly distinct pattern dorsally. The 30 dark brown, black-edged blotches that are separated by a buff background are not arranged in pairs (as in KU 40125); the dorsal blotches sometimes alternate with small lateral blotches. The ventral surface is marbled throughout. Two males from the Central National Forest, having total lengths of 1105 (920 + 185) and 8380 (690 + 140), generally resemble one another in having the head and neck dark brown or blackish and the anterior part of body dark brown, but discernibly blotched. The posterior part of the body and tail of each bears well-defined blotches (dark brown or black) with buffy interspaces; the dorsal blotches are sometimes arranged in pairs. The ventral surface of each is marbled throughout. These two males are noteworthy in that the pattern anteriorly is obscure, but the ground color is dark, not pale as in the two specimens from the vicinity of Seoul. Agkistrodon halys brevicaudus Stejneger Agkistrodon blomhoffii brevicaudus Stejneger, Bull. U. S. Nat. Mus., 58:463, July 22, 1907 (type locality, Pusan, Korea). Agkistrodon halys brevicaudus, Okada, A catalogue of vertebrates of Japan, p. 108, 1938. Specimens examined (12).—Central National Forest, near Pup’yong-ni, 4 (KU); 3 mi. NW Chip’o-ri, 2 (KU); 16 mi. NE Mosulp’o, Cheju Do, 1 (UMMZ); 5 mi. E Seoul, 1 (UMMZ); 6 mi. E Seoul, 2 (KU); 7 mi. ESE Seoul, 1 (KU); 7 mi. NNE Sogwi-ri, Cheju Do, 1 (UMMZ). Remarks.—Individuals of Agkistrodon were collected on brushy or wooded hillsides, along rock walls or in piles of rocks, and in damp, rocky, wooded ravines near streams. Many were docile when captured. One specimen was infested with nematodes, another with cestodes. One specimen had eaten a striped field mouse, Apodemus agravius, and another had eaten a gray hamster, Cricetulus triton. One female, obtained on May 22, 6 mi. E Seoul, contained 14 embryos. Another female, obtained on August 25 in the Central National Forest, contained three well-developed em- bryos. We have not included descriptive or taxonomic remarks concern- ing A. halys because Dr. Howard K. Gloyd, University of Arizona, who currently is studying the systematics of the genus Agkistrodon, has our specimens on loan. REPTILES AND AMPHIBIANS FROM KOREA 171 Gazetteer Listed below are all localities mentioned in the accounts of species; the latitude (north) and longitude (east) are given for each. All place-names can be found in “Gazetteer to maps of Korea,” 8 vols., AMS 2, U. S. Army Map Service, September, 1950, and, except for the two marked by an asterisk, can be located on AMS map series L552 (Korea, 1:250,000). The McCune-Reischauer system of romanization of Korean names is used. Changhowan-ni. 87°07’, 127°38’ Central National Forest. A small mixed forest 15-18 mi. NE Seoul and immediately west of the vil- lage of Pup’yong-ni; most of our collecting there was done approxi- mately at 37°45’, 127°10’ Cheju Do (Quelpart Island). A large island in the East China Sea off the southwestern tip of the Korean mainland (see Mosulp’o and Sogwi-ri) Chip’o-ri. 38°08’, 127°19’ Choksong. 87°58’, 126°57’ Ch’onan. 36°48’, 127°09’ *Ch’ongyang-ni. 38°15’, 127°23' Ch’orwon. 38°15’, 127°13' Ch’ungju. 36°58’, 127°57’ Halla San. A central, volcanic moun- tain on Cheju Do (see above) Hoengsong. 87°29’, 127°59’ Inje. 38°04’, 128°11’ Kangnung. 37°45’, 128°54’ Kumhwa. 38°17’, 127°28’ Kunsan. 85°59’, 126°43’ Kwangju. 35°09’, 126°55’ Majon-ni. 87°52’, 126°46’ Mosulp’o. 83°13’, 126°15’ Naegong-ni. 87°41’, 127°10’ Oho-ri. 88°20’, 128°32’ Osan. 87°09’, 127°04’ Pup’yong-ni. 37°44’, 127°12’ Pusan. 85°08’, 129°04’ P’yong-taek [= P’yongt’aeng-ni]. 36° 59's 127205: Sangbonch’on-ni. 87°27’, 127°16' Sangdaehwa. 87°30’, 128°26’ Seoul. 37°82’, 127°00’ Sogwi-ri. 33°15’, 126°34’ Songdong-ni. 38°01’, 127°16’ Taegwang-ni. 38°11’, 127°06’ *Taehoesan-ni. 38°04’, 127°14’ Taejon. 36°20’, 127°26’ Tangjonggok. 38°11’, 128°19’ Tangnim-ni. 37°50’, 127°37’ Uijongbu. 37°44’, 127°03’ Wonsan. 39°09’, 127°27’ Yami-ri. 88°08’, 127°16’ Yanggu. 38°06’, 128°00’ Yongdae-ri. 38°18’, 128°23’ Yongp’yong. 88°01’, 127°13’ Literature Cited Bass, G. H., Jr. 1955. An annotated checklist of the amphibians and reptiles of Korea. Bull. Philadelphia Herp. Soc., 1:17-28. Bocenrt, C. M., AND OLIVER, J. A. 1945. A preliminary analysis of the herpetofauna of Sonora. Bull. Amer. Mus. Nat. Hist., 83:297-426, 8 pls., 13 figs., 2 maps, March 80. BouLENcER, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum. Taylor and Francis, London, xvi + 503 pp., 30 pls. Cuance, M. L. Y. 1932. Notes on a collection of reptiles from Szechuan. Contrib. Biol. Lab., Sci. Soc. China, Zool. Ser., 8:9-95, 28 figs., February. 72, UNIVERSITY OF KAnsAsS Pusts., Mus. Nat. Hist. Conant, R. 1951. The reptiles of Ohio. Second edition (with revisionary addenda). Amer. Midland Nat., Univ. Notre Dame Press, 284 pp., 27 pls. Drxon, J. R. 1956. A collection of amphibians and reptiles from west central Korea. Herpetologica, 12:50-56, 1 fig., March 6. Fane, P. W., AND CHANG, M. L. Y. 1931. Amphibia of Nanking. Contrib. Biol. Lab., Sci. Soc. China, Zool. Ser., 7:65-114, 18 figs., May. GUNTHER, A. 1858. Catalogue of the Batrachia Salientia in the collection of the British Museum. Taylor and Francis, London, xvi + 160 pp., 12 pls. Haun, D. E. 1959. Observations and collecting notes on Rana n. nigromaculata in central Korea. Jour. Ohio Herp. Soc., 2:7-8, September 20. 1960. Collecting notes on central Korean reptiles and amphibians. Jour. Ohio Herp. Soc., 2:16-24, 4 figs., September 20. Jongs, J. K., Jr. 1960. The least tube-nosed bat in Korea. Jour. Mamm., 41:265, May 20. Liv, C. 1936. Secondary sex characters of Chinese frogs and toads. Zool. Ser., Field Mus. Nat. Hist., 22:115-156, 12 pls., October 81. 1950. Amphibians of western China. Fieldiana: Zool. Memoirs, Chicago Nat. Hist. Mus., 2:1-400, 10 pls., 100 figs., June 15. MALNATE, E. V. 1960. Systematic division and evolution of the colubrid snake genus Natrix, with comments on the subfamily Natricinae. Proc. Acad. Nat. Sci. Philadelphia, 112:41-71, 2 figs., 4 maps, September 23. MasLin, T. P. 1950. Snakes of the Kiukiang-Lushan area, Kiangsi, China. Proc. Cali- fornia Acad. Sci., ser. 4, 26:419-466, 10 figs., April 28. MITTLEMAN, M. B. 1950. The generic status of Scincus lateralis Say, 1823. Herpetologica, 6:17-20, June 5. Morr, T. 1928a. A presumption of the age in which the separation of Japan and Korea had occurred and the general condition at that age estimated from the distribution of animals in Saishuto and Tsushima. Chosen, pp. 14-25, January, 1928 (in Japanese). 1928b. On amphibians and reptiles of Quelpaert Island. Jour. Chosen Nat. Hist. Soc., 6:47-52, March 25 (in Japanese). 1928c. On a new Hynobius from Quelpaert Is]. Jour. Chosen Nat. Hist. Soc., 6:58, March 25. Mokrrya, K. 1954. Studies on the five races of the Japanese pond frog, Rana nigro- maculata Hallowell. I. Differences in the morphological char- acters. Jour. Sci. Hiroshima Univ. (ser. B, div. 1), 15:1-21, 2 pls., 1 fig., December. Oxapa, Y. 1931. The tailless batrachians of the Japanese Empire. Imp. Agric. Exp. Sta., Tokyo, 215 pp., 29 pls., 97 figs., March 30. 1934. A contribution toward a check list of the urodeles of Japan. Copeia, 1934 (1):16-19, April 24. 1935. Amphibia of Jehol. Rept. First Sci. Exped. Manchoukuo, sect. V, div. II, pt. II, art. 1, pp. 1-24 (in Japanese) and 25-47 (in English), 7 pls., 9 figs., November 30. REPTILES AND AMPHIBIANS FROM KOREA 178 OLsENn, L. S. 1957. A new species of Neoascaris (Nematode) from a Korean wood mouse, ‘Trans. Amer. Micros. Soc., 76:205-208, 9 figs., April. Pope, C. H. 1985. The reptiles of China (Natural History of Central Asia, Vol. X). reece Nat. Hist., New York, lii + 604 pp., 27 pls., 78 figs., ay 11. Pope, C. H., AND Borine, A. M. 1940. A survey of Chinese Amphibia. Peking Nat. Hist. Bull., 15:18-86, 1 map, September. ATO, I. 1943. The tailed batrachians of Japan. Tokyo, 520 pp., illustrated (in Japanese). SCHLEGEL, H. 1888. Reptilia (in von Siebold, Fauna Japonica), pp. 1-144, illustrated. Scum mvt, K, P. 1927. Notes on Chinese amphibians. Bull. Amer. Mus, Nat. Hist., 54: 553-575, 2 pls., October 14. SHANNON, F. A. 1956. The reptiles and amphibians of Korea. Herpetologica, 12:22-49, 1 pl., 1 fig., March 6. 1957. Addition to the herpetofauna of Korea, Herpetologica, 13:52, March 80. STEJNEGER, L. 1907. Herpetology of Japan and adjacent territory. Bull. U. S. Nat. Mus., 58:xx + 577, 85 pls., 409 figs., July 22. StEwakt, G. D. 1953. Notes on a collection of amphibians from central Korea. Herpeto- logica, 9:146-148, October 80. 1954. A small collection of reptiles from central Korea. Copeia, 1954 (1):65-67, February 19. TANNER, V. M. 1953. Pacific Islands herpetology No. VIII, Korea. Great Basin Nat., 13:67-78. Tine, H. 1989. A study of the reciprocal hybrids of two species of frogs, Rana nigromaculata and Rana plancyi. Peking Nat. Hist. Bull., 18: 181-200, 3 pls., March. VAN DENBURGH, J. 1923. A new subspecies of watersnake (Natrix vibakari ruthveni) from eastern Asia. Proc. California Acad. Sci., ser. 4, 13:8-4, July 26, WALLEY, H. D. 1958a. A new lacertid lizard from Korea. Herpetologica, 14:208-205, 1 fig., December 1. 1958b. “The status of Takydromus kwangakuensis Doi.” Copeia, 1958 (4):838, December 22. Transmitted June 30, 1961. 28-8517 2! see i Ras pi ae via fs iS Yah wha iP ie ; hi Ps iy | oe Hagel agohoielt btie pera Rone AN ol iby mi dpe vy aE eR KLE hes, see en an ae 2 . aa ik at P rie Wee y tare oo nl. a re y Ba ey, ae hy a, shir Ppt Tah y ‘ 4 ry ith a/5, “o> | , Reet hunt tt mea ab ks has as ai pW" aegypti ae ne mph Wien ibs pueiebably « ae eh Ae’ emt ipa ty ae aw Pie W yt i} po? hy ly my Os appeal al ani? eo: 4 | dg? nat ot her 4 og ‘haat ci, we. q CR Res nee eee A itt tio uaa penta i on gee A eee aff they Wa at wi hse Ni,” ie ae rave q yr a aad ‘a ab es eee a fie ex ci ca my Ae Be Fab » cea! Bonne . Lo 5 aia a baled 1 Sceptre ‘a, fit wet Watton a ve: 5 Wei p oils at he, f ied a 47 RS ame We eos gL , i ofa is Wi? le Se Datew oy tO ali WP Be an vl a 7 soya 8? 7 4 ‘ae g a) ” ‘ a se ‘ ‘ : ; o : Z y) a Miia ht Mela 2 bas i ese agae a8 tata MUS COMP, rAtit i Banani UNIVERSITY OF KANSAS PUBLICATIO Museum or Natura History Volume 15, No. 3, pp. 175-181, 1 £&. March 7, 1962 A New Species of Frog (Genus Tomodactylus) from Western Mexico BY ROBERT G. WEBB UNIVERSITY OF KANSAS _LAWRENCE 1962 UNIVERSITY OF KANSAS PuBLICATIONS, MusEUM OF NATURAL HIsTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 15, No. 3, pp. 175-181, 1 fig. Published March 7, 1962 UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY JEAN M. NEIBARGER, STATE PRINTER TOPEKA, KANSAS f re i MUS. COMP. Z9n1 LIBRARY A New Species of Frog (Genus Tomoda from Western México 5 5 § BY [buen iF ROBERT G. WEBB Thirteen specimens of frogs collected in the summers of 1960 and 1961 in the Mexican states of Durango and Sinaloa represent a heretofore unnamed species. The specimens have been deposited in the Museum of Natural History of the University of Kansas (KU) and in the Museum of Michigan State University (MSU). The species may be named and described as follows: Tomodactylus saxatilis new species Holotype—KU 63326 (Fig. 1); obtained eight miles west of El Palmito, Sinaloa, approximately 6100 feet, on 23 June 1961; original field number, 2354 of Robert G. Webb. Paratypes.—A total of 12 specimens: KU 63327-33, same data as holotype, 23-25 June 1961; MSU 4085-88, two miles north of Pueblo Nuevo, Durango, approximately 6000 feet elevation, 24 July 1960; MSU 4089, one half mile west of Revolcaderos, Durango, approximately 6600 feet, 29 July 1961. Diagnosis—A species of Tomodactylus possessing the following combina- tion of characters: (1) tips of two outer fingers truncate, about twice width of narrowest part of digit; (2) tympanum small, less than one half diameter of eye; (3) ventral surfaces smooth; (4) contrasting marbled pattern on back and top of head, and (5) venter whitish, lacking dark marks. Description of holotype—Adult male; snout-vent length, 31.5 (measure- ments are in millimeters and were taken by means of dial calipers reading to one tenth of a millimeter); width of head, 11.2; length of head, 10.3; horizontal diameter of eye, 3.1, and of tympanum, 1.2; distance from eye to nostril, 3.8; internarial width, 2.9; interorbital width, 4.1; width of eyelid, 2.5; lumbar gland (left side), 7.0x 2.6; distance from axilla to groin, 15.2; tibial length, 12.7; length of foot, 13.1. Head slightly wider than body; tip of snout rounded, slightly truncate; canthus rounded; tympanum small, less than one half diameter of eye; tym- panum having posterior margin ill-defined, separated from eye by distance about equal to diameter of tympanum; diameter of eye slightly less than dis- tance from eye to nostril; width of eyelid about two thirds interorbital width; paratoid gland indistinct; lumbar glands high, separated from insertion of leg by about one millimeter; back and sides of body having low, scarcely elevated pustules; top of head, limbs and venter smooth; few low, whitish pustules below and behind tympanum, and low on sides of body; posterior surface of thighs and anal region pustulate; one pair of whitish postanal spots; ventral disc attached near insertion of legs, lacking conspicuous transverse fold; skin loose on throat, chest and abdomen. C177) 178 UNIVERSITY OF KANSAS PuUBLS., Mus. Nat. Hist. Digits not webbed; tips of two outer fingers truncate, having terminal transverse grooves, about twice width of narrowest part of digit; digits of first and second fingers slightly expanded; fingers from shortest to longest, ]-2-4-8, first only slightly shorter than second; three palmar tubercles; inner palmar tubercle about one third size of large median tubercle; outer tubercle about one tenth size of large median tubercle; four supernumerary palmar tubercles; tips of toes slightly wider than narrowest part of digits; toes from shortest to longest, 1-2-5-3-4, second only slightly shorter than fifth; inner metatarsal tubercle about four times size of small outer metatarsal tubercle; supernumerary tubercles on foot small; no tarsal fold; heels touching when tibiae adpressed to thighs; tibiotarsal articulation reaching eye when leg adpressed to side of body. Contrasting marbled pattern on back and top of head; contrasting, mostly barred, pattern on limbs; ventral surfaces whitish, lacking dark marks, but having minute dark peppering; marbling of dorsal surfaces blackish and whitish in preservative. Vomerine teeth lacking; internal choanae lateral, partly concealed by maxillaries; tongue smooth, elongate, shallowly notched distally, free for about half its length; vocal sac median; internal vocal slits large and near angle of jaw. Variation—Twelve males closely resemble the holotype. Two specimens from Pueblo Nuevo are soft and not well preserved. The ranges of variation (means in parentheses) for the 18 males comprising the type series are: snout- vent length, 25.5-31.5 (27.9); width of head, 9.7-11.2 (10.5); diameter of eye, 2.9-3.6 (3.2); horizontal diameter of tympanum, 1.1-1.8 (1.4); length of tibia, 11.1-13.1 (12.1); length of foot, 11.0-138.1 (12.1). The pale ground color of the marbled pattern in most specimens is least extensive on the back and arms, but most extensive on the legs. The lumbar glands are slightly elevated and conspicuous, and in KU 68328 are extremely protuberant, or (KU 68330) evident on left side but flattened and indistinct on right side. The back is rough having low, scarcely elevated pustules, but becomes less rough anteriorly and most of the top of head is smooth. The three specimens from Pueblo Nuevo, Durango, differ slightly from the other specimens examined in lacking pairs of postanal white spots, and in having smooth backs (slightly pustulate in MSU 4088). The tibiotarsal articulation fails to reach the eye in KU 63330. The small inner palmar tubercle is continuous with the large median tubercle on the right hand of KU 63330, and lacking on both hands of KU 68329 and on the left hand of KU 63328. The tip of the tongue is entire in some specimens and in others has an irregular margin. Coloration of living specimens.—Marbled pattern on back and top of head of dark brown to blackish on yellowish-gold; pattern slightly less contrasting on limbs than on back, consisting of brown to grayish on pale yellow; side of head and body grayish sometimes having pale yellow to whitish spots; iris blackish having fine reticulation of yellowish to greenish-gold; venter dirty white. Habitat—The three records of occurrence for Tomodactylus saxatilis are in a mixed boreal-tropical habitat, which is transitional between a pine-oak forest at higher elevations and a tropical decidu- A New SpEcrEs oF Froc ( TOMODACTYLUs ) 179 ous forest at lower elevations. The mixed boreal-tropical habitat is most conspicuous at elevations between approximately 7800 and 5500 feet on southerly exposed slopes of barrancas and arroyos of the dissected plateau of the Sierra Madre Occidental. The mixed boreal-tropical habitat occurs for approximately 80 miles along the paved highway (Mexican Highway 40) between Cd. Durango, Durango, and Mazatlan, Sinaloa. The records of occurrence in those states that are along this highway are separated by 14.5 miles (via road). Fic. 1. Tomodactylus saxatilis new species, adult male, KU 63326, holotype (x 2), dorsal view. The terrain consists of occasional level areas, but is mostly of steep hillsides. Dominant trees are large oaks and pines; a char- acteristic pine is the sad or drooping-needle pine, locally called “pino triste.” The vegetational cover is usually open, including grasses, small oaks and pines, broad-leaved shrubs and herbs, prickly pears, magueys, thorny acacias, bracken fern, and epiphytes in trees. Ferns occur in moist protected places, and orchids are occasional, sometimes in trees. 180 UNIVERSITY OF KANSAS Pusu3s., Mus. Nat. Hist. Outcrops of rock, boulder-strewn areas, and occasional rock slides (talus) also characterize the terrain. Tomodactylus saxatilis seems to be restricted to rocky habitats. The individuals collected were detected when they called at night from within crevices of rocks or from exposed perches on rocks and boulders; some calling frogs were out of reach on steep rock walls. The call is a single, loud, high peep. Comparisons.—Dixon (1957) recognized six species of Tomodactylus (nitidus, dilatus, albolabris, angustidigitorum, fuscus and grandis) in his revision of the genus. Another species (rufescens) was subsequently described by Duellman and Dixon (1959). Tomodactylus saxatilis differs from all the species named immediately above by the combination of characters given in the diagnosis. Tomodactylus saxatilis differs from nitidus, angustidigitorum and grandis in having the tips of the two outer fingers widened and truncate; saxatilis differs from dilatus, albolabris, fuscus and rufescens in having a smooth venter (not pustulate), a contrasting pale and dark marbled pattern on the back, and a lack of “flash” colors on the femora. Tomodactylus saxatilis, having lumbar glands, also resembles three species referred to the genus Syrrhophus. Tomodactylus macrotympanum was de- scribed by Taylor (1940:496, 497) as having a large, moderately distinct lumbar gland; the species was referred to the genus Syrrhophus by Dixon (op. cit.:384). According to Firschein (1954:55), Syrrhophus smithi and S. petrophilus have elongate lumbar glands shaped like those in Tomodactylus. Tomodactylus saxatilus resembles macrotympanum, smithi and petrophilus more than it does other species; all four attain large maximal sizes, and have lumbar glands, mostly smooth ventral surfaces, three palmar tubercles (some- times absent in saxatilis), and usually contrasting dorsal patterns (reduced to flecks and spots in all species except saxatilis). Tomodactylus saxatilis differs from macrotympanum in having an extensive marbled dorsal pattern and a small tympanum, and differs from smithi and petrophilus in having a marbled dorsal pattern and the tips of the outer two fingers widened and truncate. Tomodactylus saxatilis differs from all other named species of Syrrhophus in having conspicuous lumbar glands and in lacking inguinal glands. Remarks.—The characteristics delimiting the genera Tomodac- tylus and Syrrhophus are not agreed upon by all workers (see dis- cussions by Firschein, 1954:50; Langebartel and Shannon, 1956:164; and Dixon, 1957:383). I have referred saxatilis to the genus Tomo- dactylus on the basis of a lumbar gland, which was considered a distinguishing character for the genus by Smith and Taylor (1948:46) and Langebartel and Shannon (1956:165). Lumbar glands are longer than broad, at least one third the distance from axilla to groin, lateral and usually high, and often conspicuous and protuberant. The elevation or flatness of the lumbar glands seems to be due to individual variation; living specimens in the field had conspicuous and protuberant, or non-elevated, indistinct lumbar glands. Lumbar glands are not to be confused with inguinal glands, A New Species oF Froc (ToMopACTYLUS ) 181 which are roundish, often yellowish, sometimes diffuse, lateral but low, often inconspicuous, and usually not protuberant. Inguinal glands occur in the genus Microbatrachylus and in some species of Eleutherodactylus, and have been described as flat, or low, or small, or indistinct for most species of Syrrhophus. For financial assistance with field work I am grateful to Rollin H. Baker, and those individuals who administer the Michigan State University Development Fund and the Bache Fund of the National Academy of Sciences (Grant No. 463). I am grateful also to J. Keever Greer, Donald F. Switzenberg, and Rudolph A. Scheibner for aid in the field, to Edward H. Taylor, James R. Dixon, and William E. Duellman for profitable discussions, and to Thomas Sweringen for figure 1. The specific name alludes to the habitat (Latin, saxatilis = found among rocks). LITERATURE CITED Drxon, J. R. 1957. Geographic variation and distribution of the genus Tomodactylus in Mexico. Texas Journ. Sci., 9(4):379-409, 5 figs., 1 map, De- cember. DvELLMAN, W. E., and Drxon, J. R. 1959. A new frog of the genus Tomodactylus from Michoacan, Mexico. Texas Journ. Sci., 11(1):78-82, 1 fig., 1 table, March. FirscuHeIn, I. L. 1954. Definition of some little-understood members of the leptodactylid genus Syrrhophus, with a description of a new species. Copeia, 1954, 1:48-58, February 19. LANGEBARTEL, D. A., and SHANNON, F. A. 1956. A new frog (Syrrhophus) from the Sinaloan lowlands of Mexico. Herpetologica, 12(3):163-165, 2 figs., September 1. Smitu, H. M., and Tay.or, E. H. 1948. An annotated checklist and key to the Amphibia of Mexico. Bull. U. S. Nat. Mus., No. 194, pp. iv + 118. TayLor, E. H. 1940. Herpetological miscellany. Univ. Kansas Sci. Bull., 26(15):489- 571, 10 pls., 7 figs., November 15. O 29-1923 UNIVERSITY OF KANSAS PUBLICATIONS |...” MuSEUM OF NATURAL HISTORY Volume 15, No. 4, pp. 183-204 October 26, 1962 —————— Type Specimens of Amphibians and Reptiles in the Museum of Natural History, The University of Kansas BY WILLIAM E. DUELLMAN AND BARBARA BERG UNIVERSITY OF KANSAS LAWRENCE 1962 UNIVERSITY OF KANSAS PUBLICATIONS, MusSEUM OF NATURAL HISTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 15, No. 4, pp. 183-204 Published October 26, 1962 UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY JEAN M. NEIBARGER, STATE PRINTER TOPEKA, KANSAS Type Specimens of Amphibians and Reptiles 7) in the Museum of Natural History, = = The University of Kansas BY WILLIAM E. DUELLMAN AND BARBARA BERG In recent years various museums have issued lists of the type specimens in their collections. These lists are valuable to systema- tists by providing these workers with information that otherwise sometimes is difficult to obtain, Although by comparison with the large museums in the eastern part of the United States, such as the American Museum of Natural History, the United States National Museum, and the Museum of Comparative Zoology at Harvard, the herpetological collections in the Museum of Natural History at The University of Kansas are relatively small (70,000 specimens), a large number of types are contained in the collection. The largest number of types are of species described from Costa Rica by Ed- ward H. Taylor between 1949 and 1958. Beginning in 1932 and continuing until World War II, Taylor described many species from México; types of these species were contained in the Edward H. Taylor—Hobart M. Smith collection. Inquiries from herpetolo- gists reveal that many workers suppose that the EHT-HMS collec- tion is in the Museum of Natural History. Such is not the case, for the EHT-HMS collection has been divided between the Chicago Natural History Museum and the University of Illinois Museum of Natural History. The list of type specimens presented here includes all holotypes and paratypes that are known in the museum as of April 15, 1962. Only specimens actually designated as paratypes by the original describer are considered as such. Each species or subspecies is listed in the combination as it was originally proposed. If the name has been subjected to subsequent nomenclatural change or has been placed in synonymy, the name in current use is given after an equals sign, together with a citation to the authority for the change. The arrangement in the following list is alphabetical by genus and species within orders (suborders in the Squamata). Current combinations of names that have been changed subsequent to their original proposal are listed in the correct alphabetical place with a cross-reference to the original combination. (185) 186 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. All original citations have been checked, and actual dates of publication are given for each. The checking of original citations, museum catalogues, labels, and sometimes collectors’ field cata- logues has revealed numerous errors. In some cases the catalogue numbers of the holotypes and paratypes given by Taylor actually were his field numbers. In the following list the museum numbers that were subsequently assigned are given. Certain localities have been given incorrectly; for example, Taylor described several species from Moravia de Chirrip6, which he gave as being in Limon Province, Costa Rica. Actually Moravia de Chirrip6 is in Cartago Province. We have attempted to correct all such errors. The holotype and two paratypes of Rana taylori Smith are stated to be in the Museum of Natural History. Smith (Herpetologica, 15:214, December 8, 1959) proposed the name Rana taylori for three specimens discussed by Taylor (Univ. Kansas Sci. Bull., 35:906, July 1, 1952), who gave the numbers 2976-2978. Number 2978 was designated the holotype of Rana taylori by Smith, who never saw the frogs. The numbers 2976-2978 are E. H. Taylor’s field numbers; the specimens never have been catalogued in the Museum of Natural History, and their present location is unknown. The 125 species and subspecies represented by holotypes and paratypes in the collection are distributed among the orders of amphibians and reptiles as follows: AMPHIBIA REPTILIA Gymnophiona (Caecilians) ... 3 Testudines (Turtles) ...... 5 Caudata (Salamanders) ..... 20 Sauria (Lizards) «2.1.08 Ja. 81 Salientia (Frogs and Toads) .. 89 Serpentes (Snakes) ......... 27 GYMNOPHIONA Dermophis costaricense Taylor, Univ. Kansas Sci. Bull., 37:506, fig. 2, October 15, 1955. Ho.otypr.—No. 36343, Cinchona, Alajuela Province, Costa Rica; Edward H. Taylor; July 14, 1954. PARATYPES.—Nos. 36337-36340, 36342, 36344-36347, data same as for holotype. Nos. 86447 and 86448, Moravia de Chirripé, Cartago Province, Costa Rica; Edward H. Taylor; August 15-16, 1954. Dermophis glandulosus Taylor, Univ. Kansas Sci. Bull., 37:509, fig. 3, October 15, 1955. Ho.totypE.—No. 29979, San Isidro del General, San José Province, Costa Rica; Virgil Cave; 1947. Dermophis occidentalis Taylor, Univ. Kansas Sci. Bull., 37:503, fig. 1, October 15, 1955. AMPHIBIAN AND REPTILE SPECIMENS 187 Hotoryre.—No. 36298, 15 miles west-southwest of San Isidro del General, San José Province, Costa Rica; Edward H. Taylor; August 3, 1954. PaRATYPES.—Nos. 36296 and 36297, same locality; Edward H. Taylor; July 19, 1952. CAUDATA Bolitoglossa alvaradoi Taylor, Univ. Kansas Sci. Bull., 36:604, fig. 2, June 1, 1954. Hototyre.—No. 30484, Moravia de Chirrip6, Cartago Province, Costa Rica; Edward H. Taylor; August 16, 1951. Paratypres.—Nos. 34903-34905, same locality; Edward H. Taylor; June 25-26, 1952. Bolitoglossa arborescandens Taylor, Univ. Kansas Sci. Bull., 36:600, fig. 2, June 1, 1954. Ho.totyre.—No. 34925, Moravia de Chirripé, Cartago Province, Costa Rica; Edward H. Taylor; August 16, 1951. Bolitoglossa palustris Taylor, Univ. Kansas Sci. Bull., 33:283, April 20, 1949. Hototyre.—No. 23817, San Isidro del General, San José Province, Costa Rica; Edward H. Taylor; August 23, 1947. Bolitoglossa veracrucis Taylor, Univ. Kansas Sci. Bull., 34:189, fig. 1, Oc- tober 1, 1951. Ho.otype.—No. 26941, 35 kilometers southeast of Jestis Carranza, Vera- cruz, México; Walter W. Dalquest; April 6, 1949. ParaTyPEs.—Nos. 26933-26940, 26942-26953, data same as on holotype. Chiropterotriton abscondens Taylor, Proc. Biol. Soc. Washington, 61:177, November 12, 1948. ParatyPE.—No. 29971, Isla Bonita, Heredia Province, Costa Rica; David Allee and Sr. Machado; August 1, 1947. Notre.—This specimen was listed by Taylor (1948:177) as Richard C. Taylor No. 1416; subsequently the specimen was cataloged as KU 29971. Magnadigita cerroensis Taylor, Univ. Kansas Sci. Bull., 34:724, pl. 80, February 15, 1952. Ho.totyre.—No. 29961, Cerro de la Muerte, 2 kilometers below Millville, San José Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; August 15, 1947. Magnadigita nigrescens Taylor, Univ. Kansas Sci. Bull., 33:282, April 20, 1949, Ho.LotyPe.—No. 23816, Boquete Camp, Cerro de la Muerte (on highway between Millville and San Isidro del General), San José Province, Costa Rica; Edward H. Taylor; August 17, 1947. Magnadigita pesrubra Taylor, Univ. Kansas Sci. Bull., 34:707, pl. 76, text fig. 2, February 15, 1952. HoLotype.—No. 25093, Summit of Cerro de la Muerte, Cartago Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; August 14-15, 1947. 188 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Paratypes.—(All collected by Edward H. Taylor and Richard C. Taylor in August, 1947) Nos. 25083-25086, 25142-25156, 25157 (166 specimens), Millville, Cerro de la Muerto, San José Province, Costa Rica: Nos. 25082, 25087-25092, 25094-25100, same locality as holotype. Magnadigita torresi Taylor, Univ. Kansas Sci. Bull., 34:718, pl. 77, Febru- ary 15, 1952. Ho.otype.—No, 25104, Summit of Cerro de la Muerte, Cartago Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; August 17, 1947. ParatypPes.—Nos. 25101-25103, 25105-25116, data same as holotype. Norte.—Taylor (1952:718) listed 25116a (30 specimens); these are not present in the collection. Oedipina allent Taylor, Univ. Kansas Sci. Bull., 36:607, fig. 8, June 1, 1954, Ho.oryrPe.—No. 34926, Palmar, Puntarenas Province, Costa Rica; Edward H. Taylor; September 2, 1952. Oedipina bonitaensis Taylor, Univ. Kansas Sci. Bull., 84:762, pl. 86, fig. 8, text fig. 8, February 15, 1952. Ho.otyre.—No, 29972, Volcan Pods, 6 miles west of Isla Bonita, Alajuela Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; August 3, 1947, ParaTyPes.—Nos. 29973 and 29974, data same as on holotype. Oedipina cyclocauda Taylor, Univ. Kansas Sci. Bull., 34:764, pl. 87, fig. 4, text fig. 9, February 15, 1952. Horotype.—No. 25066, Los Diamantes, 1 kilometer east of Guadpiles, Limén Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; Sep- tember 1-8, 1947. PARATYPES.—Nos. 25038-25065, data same as on holotype. Oedipina gracilis Taylor, Univ. Kansas Sci. Bull., 34:773, February 15, 1952. Hotoryre.—No. 29969, Los Diamantes, 1 kilometer east of Gudpiles, Limén Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; September 8, 1947. Paratypes.—Nos, 29968 and 29970, data same as on holotype. Oedipina inusitata Taylor, Univ. Kansas Sci. Bull., 34:277, pl. 87, fig. 8, text fig. 18, February 15, 1952. Ho.totyre.—No. 29964, 3 miles northeast of Santa Cruz, south slope of Volcan Turrialba, Cartago Province, Costa Rica; Edward H. Taylor; August 9, 1947. Oedipina longissimus Taylor, Univ. Kansas Sci. Bull., 34:771, pl. 87, fig. 5, text fig. 11, February 15, 1952. Hototypre.—No. 29963, Cervantes, Cartago Province, Costa Rica; Edward H. Taylor; July 9, 1947. Oedipina pacificensis Taylor, Univ. Kansas Sci. Bull., 34:775, pl. 87, fig. 2, text fig. 12, February 15, 1952. Ho.totyPe.—No. 29965, 2 miles east of San Isidro del General, San José Province, Costa Rica; Edward H. Taylor; August 23, 1947. AMPHIBIAN AND REPTILE SPECIMENS 189 Oedipina serpens Taylor, Univ. Kansas Sci. Bull., 83:286, April 20, 1949. Hototyre.—No. 23815, Morehouse Finca, 5 miles southwest of Turrialba, Cartago Province, Costa Rica; Edward H. Taylor; July 2, 1947. Oedipina syndactyla Taylor, Proc. Biol. Soc. Washington, 61:179, No- vember 12, 1948. Hototype.—No. 29962, Volcdn Pods, 4 miles west of Isla Bonita, Alajuela Province, Costa Rica; Edward H. Taylor; August 5, 1947. Notre.—This specimen was listed by Taylor (1948:179) as No. 843. This is the E. H. Taylor field number; subsequently the specimen was catalogued as KU 29962. Thorius minutissimus Taylor, Amer. Mus. Novitates, 14387:5, figs. 1-2, December 7, 1949. ParaTyPE.—No. 28080, Santo Tomas Tecpdn, Oaxaca, México; Thomas C. MacDougall, 1948. Norre.—Received in exchange; formerly this specimen was American Mu- seum of Natural History No. A 53930. Typhlotriton nereus Bishop, Copeia, 1944(1):1, figs. 1-3, April 21, 1944. Paratypes.—Nos. 16036-16088, 16043-16045, near Galena, Cherokee County, Kansas; Wallace Lane and William McNown; March 26, 19382. No. 16199, York Springs, Imboden, Lawrence County, Arkansas; Byron C, Mar- shall; February, 1930: Nos. 16350, 16352-16354, 16357-16360, near Riverton, Cherokee County, Kansas; Edward H. Taylor; March 26, 1932. Note.—Nos. 16350, 16352-16354, 16357-16360 were listed erroneously by Bishop (1944:1-2) as being from Galena, Cherokee County, Kansas. SALIENTIA Agalychnis alcorni Taylor [= Phyllomedusa dacnicolor Cope, according to Duellman (Herpetologica, 18:29, 1957)], Copeia, 1952(1):381, figs. 1-2, June 2,. 1952. Hototyre.—No. 29763, South bank of Rio Tepalcatepec, 17 miles south of Apatzingan, Michoacan, México; J. R. Alcor; November 30, 1950. PARATYPE.—No. 28100, 0.5 miles east of San Blas, Nayarit, México; J. R. Alcorn; January 6, 1949. Agalychnis saltator Taylor [= Phyllomedusa saltator (Taylor), according to Funkhouser (Occas. Papers Nat. Hist. Mus. Stanford Univ., 5:36, 1957)], Univ. Kansas Sci. Bull., 87:527, fig. 10, October 15, 1955. Ho.otyre.—No. 385615, Finca Bosco, 5 kilometers north-northeast of Tilaran, Guanacaste Province, Costa Rica; Edward H. Taylor; August 14, 1954. ParaTyPes.—Nos. 385609-10, 35612-35614, 85616-35620, 35622-35626, data same as for holotype. Nore.—Taylor (1955:527) erroneously gave the type locality as 4 kilometers north-northeast of Tilaran; in Taylor’s field catalogue and in the museum cata- logue the locality is listed as 5 kilometers north-northeast of Tilaran. Paratypes 35611 and 35621, listed by Taylor, have been exchanged with the Stanford Natural History Museum. Atelopus loomisi Taylor—See Atelopus varius loomis Taylor. 190 UNIvERSITY OF Kansas Pusts., Mus. Nat. Hist. Atelopus varius loomisi Taylor [= Atelopus loomisi Taylor, according to Taylor (Univ. Kansas Sci. Bull., 37:517, 1955)], Univ. Kansas Sci. Bull. 35:625, fig. 7, July 1, 1952. Hototype.—No. 24746, Isla Bonita, Heredia Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; June 26, 1947. Paratypres.—Nos. 24742-24745, same locality as on holotype, August 1-5, 1947. Bufo holdridgei Taylor, Univ. Kansas Sci. Bull., 35:607, fig. 3, July 1, 1952. Ho.otyre.—No. 30885, Volcan Barba, Heredia Province, Costa Rica; Ed- ward H. Taylor; July 15, 1951. Centrolenella spinosa Taylor [= Teratohyla spinosa (Taylor), according to Taylor (Proc. Biol. Soc. Washington, 64:35, 1951)], Univ. Kansas Sci. Bull., 33:259, April 20, 1949. Ho.iotyre.—No. 23809, Los Diamantes, 1 kilometer east of Guapiles, Limén Province, Costa Rica; Edward H. Taylor and Richard C, Taylor; Sep- tember 7, 1947. Paratypes.—Nos, 23810-23811, data same as on holotype. Cochranella albomaculata Taylor, Univ. Kansas Sci. Bull., 33:267, April 20, 1949. Hototryrpe.—No. 23814, Los Diamantes, 1 kilometer east of Guapiles, Limén Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; Sep- tember 7, 1947. Cochranella chirripoi Taylor, Univ. Kansas Sci. Bull., 39:59, fig. 10, No- vember 18, 1958. Ho.iotyre.—No. 36865, Cocales Creek, Suretka, Limén Province, Costa Rica; Edward H. Taylor; June 29-80, 1954. ParatyPes.—Nos. 36862-36864, 36866-36870, data same as on holotype. Cochranella colymbiphyllum Taylor, Univ. Kansas Sci. Bull., 33:262, April 20, 1949. Hototyre.—No. 23812, Isla Bonita, Heredia Province, Costa Rica; Ed- ward H. Taylor and Richard C. Taylor; July 29, 1947. Cochranella decorata Taylor, Univ. Kansas Sci. Bull., 39:50, fig. 6, Novem- ber 18, 1958. Hototyre.—No. 36896, Hacienda La Florencia, about 8 miles west of Turrialba, Cartago Province, Costa Rica; Jack Reark; November, 1952. ParATYPEs.—Nos. 36883-36884, San José, San José Province, Costa Rica; Marco Tulio Pacheco; summer, 1954. Note.—The locality of the paratypes is not definite; Taylor presumed that they were from San José. See note under the following species. Cochranella millepunctata Taylor, Univ. Kansas Sci. Bull., 39:53, fig. 7, November 18, 1958. Ho.otrype.—No. 36887, La Palma, San José Province, Costa Rica; Edward H. Taylor, June 20, 1954. AMPHIBIAN AND REPTILE SPECIMENS 191 Paratypes.—No. 32948, Moravia de Chirripé6, Cartago Province, Costa Rica; Edward H. Taylor; June 26, 1952. Nos 36885-36886, 36888-36894, data same as holotype. No. 36897, Sarchi, Alajuela Province, Costa Rica; Robert Hunter; summer, 1954. Note.—Taylor (1958:53) erroneously listed 36883 and 36884 as paratypes of this species; on page 50 of the same publication he correctly designated these specimens as paratypes of Cochranella decorata. Also, Taylor (1958:53) erroneously listed 32943 as 23948. Cochranella reticulata Taylor, Univ. Kansas Sci. Bull., 39:61, fig. 11, No- vember 18, 1958. Ho.Lotyrpe.—No. 32922, Bridge across Rio Reventazén, Inter-American In- stitute of Agriculture, Turrialba, Cartago Province, Costa Rica; Edward H. Taylor; August 1, 1952. PaRATYPES.—No. 32916, Moravia de Chirripé, Cartago Province, Costa Rica; Edward H. Taylor; June 26, 1952. Nos. 32917-32920, Cariblanco, Heredia Province, Costa Rica; Edward H. Taylor; July 20, 1952. No. 32921, data same as holotype. Nos. 32923 and 32924, Palmar, Puntarenas Province, Costa Rica; Edward H. Taylor; September 3, 1952. No. 82925, Golfito, Pun- tarenas Province, Costa Rica; Edward H. Taylor; September 7, 1952. Nos. 36878-36882, Cocales Creek, Suretka, Limén Province, Costa Rica; Edward H. Taylor; June 30, 1954. Cochranella talamancae Taylor, Univ. Kansas Sci. Bull., 35:781, 783, July 1, 1952. Ho.Lotype.—No. 30887, Moravia de Chirripéd, Cartago Province, Costa Rica; Edward H. Taylor; August 16, 1951. NotEe.—Taylor (1952:781) erroneously listed this specimen as KU 4143; actually that is Taylor’s field number. Subsequently the specimen was cata- logued as KU 30887. Dendrobates granuliferus Taylor, Univ. Kansas Sci. Bull., 39:10, fig. 5, November 18, 1958. Ho.otyre.—No. 43874, Low mountains north of Rio Diquis, about 3 miles north of Palmar, Puntarenas Province, Costa Rica; Edward H. Taylor; Sep- tember 3, 1952. ParaTyPes.—No, 43875, San Isidro del General, San José Province, Costa Rica; Edward H. Taylor; summer, 1952. Nos. 48876-43880, data same as holotype. Note.—Taylor (1958:10) erroneously listed 48875 as being from the type locality. Eleutherodactylus crassidigitatus Taylor, Univ. Kansas Sci. Bull., 35:740, fig. 31, July 1, 1952. Hototyre.—No. 28396, Isla Bonita, Heredia Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; July 22-August 3, 1947. ParATyPEs.—Nos, 28360, 28361, 28440, Morehouse Finca, 5 miles south- west of Turrialba, Cartago Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; July 11-21, 1947. Nos. 28362-28368, 28370-28374, data same as on holotype. 192 UnIverSITY OF Kansas Pusts., Mus. Nat. Hist. Note.—Taylor (1952:740) listed 28360, 28361, and 28440 as being from “Turrialba and surroundings.” Eleutherodactylus dubitus Taylor, Univ. Kansas Sci. Bull., 35:707, fig. 22, July 1, 1952. Hotorypre.—No. 24942, Isla Bonita, Heredia Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; July 22-August 3, 1947. ParaTypes.—Nos. 24910 and 24911, Morehouse Finca, 5 miles southwest of Turrialba, Cartago Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; July 11-16, 1947. Nos. 24912-24914, Santa Cruz Volcan, Cartago Province, Costa Rica; Edward H. Taylor; July 17, 1947. Nos. 24915-24941, 24943-24948, data same as on holotype. Notre.—Taylor (1952:707) erroneously listed 24910-24914 as being from the type locality. Eleutherodactylus minimus Taylor, Univ, Kansas Sci. Bull., 37:517, fig. 7, October 15, 1955. Hototryre.—No. 37128, 5 kilometers north-northeast of Tilaran, Guanacaste Province, Costa Rica; Edward H. Taylor; August 13, 1954. ParatypEs.—Nos. 37126, 37127, 37129-87132, data same as on holotype. Eleutherodactylus persimilis Barbour [= Microbatrachylus persimilis (Bar- bour), according to Taylor (Univ. Kansas Sci. Bull., 35:670, 1952)], Occas. Papers Boston Soc. Nat. Hist., 5:193, pl. 14, January 30, 1926. ParaTyPpE.—No. 47166, Suretka, Limén Province, Costa Rica; Emmett R. Dunn and Chester B. Duryea; 1923. Notre.—Received in exchange; formerly this specimen was Museum of Comparative Zoology No. 11609. Eleutherodactylus taurus Taylor, Univ. Kansas Sci. Bull., 39:17, fig. 7, November 18, 1958. Hoxrotyre.—No. 43866, Golfito, Puntarenas Province, Costa Rica; Edward H. Taylor; September 6, 1952. ParaAtyPes.—Nos. 43867, 43869-43871, 43873, same locality as holotype; John Baker and Edward H. Taylor; September 2-6, 1952. Nos. 48868, 43872, Palmar, Puntarenas Province, Costa Rica; Edward H. Taylor; September 2, 1952. Eleutherodactylus vocator Taylor, Univ. Kansas Sci. Bull., 37:522, fig. 8, October 15, 1955. Hototyre.—No. 87001, Agua Buena, Puntarenas Province, Costa Rica; Edward H. Taylor; July 25, 1954. ParatyPpEs.—Nos. 36999 and 37000, data same as on holotype. Hyla alvaradoi Taylor, Univ. Kansas Sci. Bull., 35:882, fig. 63, July 1, 1952. Ho.totypeE.—No. 30886, Moravia de Chirripd, Cartago Province, Costa Rica; Edward H. Taylor; August 16, 1951. Hyla bivocata Duellman and Hoyt, Copeia, 1961(4):414, figs. 1-3, De- cember 19, 1961. Ho.totype.—No. 58446, 6.2 kilometers by road south of Rayén Mescalapa, Chiapas, México; William E. Duellman and Dale L. Hoyt; August 5, 1960. AMPHIBIAN AND REPTILE SPECIMENS 193 ParATYPEs.—Nos. 58447-58450, data same as on holotype. Hyla chaneque Duellman, Herpetologica, 17:1, figs. 1-2, April 15, 1961. Hototyre.—No. 58439, 6.2 kilometers by road south of Rayén Mescalapa, Chiapas, México; William E. Duellman; August 5, 1960. ParatTyPes.—Nos. 58440-58442, same as holotype. No. 58444, 5.6 kilometers by road south of Rayén Mescalapa, Chiapas, México; William E. Duellman; June 15, 1960. Note.—No. 58443, listed by Duellman (1961:1), has been exchanged with the Museum of Zoology at the University of Michigan. Hyla cyclomaculata Taylor, Copeia, 1949( 4) :272, pl. 1, December 15, 1949. Ho.otyPe.—No, 26954, Huatusco, Veracruz, México; Walter W. Dalquest; December 2, 1948. Hyla dalquesti Taylor—See Hyla proboscidea Taylor. Hyla debilis Taylor, Univ. Kansas Sci. Bull., 35:880, fig. 62, July 1, 1952. Ho.otyre.—No. 28184, Isla Bonita, Heredia Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; August 1, 1947. Hyla dulcensis Taylor, Univ. Kansas Sci. Bull., 39:87, fig. 17, November 18, 1958. Ho.totype.—No. 32168, Golfito, Puntarenas Province, Costa Rica; Edward H. Taylor; September 7, 1952. ParaTyPEs.—Nos. 32166 and 32167, same locality as holotype; John Baker and Edward H. Taylor; September 7 and 8, 1952. Hyla immensa Taylor, Univ. Kansas Sci. Bull., 35:815,, figs. 42-43, July 1, 1952. Hototyre.—No. 30404, Inter-American Institute of Agricultural Sciences, Turrialba, Cartago Province, Costa Rica; Edward H. Taylor; June 25, 1951. Hyla legleri Taylor, Univ. Kansas Sci. Bull., 39:33, figs. 15-16, November 18, 1958. Hototrypre.—No. 32982, 15 kilometers west-southwest of San Isidro General, San José Province, Costa Rica; Edward H. Taylor; July 10, 1952. ParATyPES.—Nos. 32981, 32983-32988, 35171, same locality as holotype; John Baker and Edward H. Taylor; July 10, 1952. Note.—Taylor (1958:33) erroneously listed 32989-32992 as paratypes of this species; specimens bearing those numbers are Smilisca gabbi. Hyla manisorum Taylor, Univ. Kansas Sci. Bull., 36:630, fig. 11, June 1, 1954. Ho.otyre.—No. 34927, Bataan, Limén Province, Costa Rica; Edward H. Taylor; July 22, 1951. Hyla mixomaculata Taylor, Copeia, 1950(4):274, pl. 1, December 22, 1950. Ho.otyre.—No, 26777, Coscomatepec, Veracruz, México; Walter W. Dal- quest; December 1, 1948. PaRATYPES.—Nos. 26776, 26778-26780, 26782-26785, data same as on holo- type. No. 26781, Huatusco, Veracruz, México; Walter W. Dalquest; Decem- ber 2, 1948. 194 UNIversITY OF Kansas Pusts., Mus. Nar. Hist. Note.—Taylor (1950:274) erroneously listed 26786 as a paratype of this species; the specimen bearing that number is a Hyla staufferi. Hyla moraviensis Taylor, Univ. Kansas Sci. Bull., 35:865, fig. 57, July 1, 1952. Hototypr.—No. 30284, Moravia de Chirripé, Cartago Province, Costa Rica; Edward H. Taylor; August 16, 1951. ParatyPes.—Nos. 30283, 30285 and 30286, data same as on holotype. Hyla proboscidea Taylor [= Hyla dalquesti Taylor, according to Taylor (Copeia, 1949(1):74, April 15, 1949—substitute name for Hyla proboscidea Taylor, preoccupied by Hyla proboscidea Brongersma, 1933)], Univ. Kansas Publ. Mus. Nat. Hist., 1:259, figs. a-c, August 16, 1948. Hotorype.—No. 23626, 2 kilometers west of Jico (= Xico), Veracruz, México; Walter W. Dalquest; October 28, 1946. PaRATYPES.—Nos. 23623-23625, 23627, data same as on holotype. Nore.—Taylor (1948:259) erroneously listed 23628 as a paratype of this species; the specimen bearing that number is a Heloderma suspectum. How- ever, 23623 not listed by Taylor is a Hyla proboscidea; this specimen is noted as a paratype in the museum catalogue. Hyla rivularis Taylor, Univ. Kansas Sci. Bull., 35:847, fig. 53, July 1, 1952. Hototyrr.—No. 28197, Isla Bonita, Heredia Province, Costa Rica; Ed- ward H. Taylor and Richard C. Taylor; August 1, 1947. ParaTyPEes.—Nos. 28196 and 28198, data same as on holotype. Hyla rufioculis Taylor, Univ. Kansas Sci. Bull., 35:827, fig. 47, July 1, 1952. Ho.otypPr.—No. 28216, Isla Bonita, Heredia Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; July 28, 1947. ParatypPrs.—Nos. 28204-28210, Morehouse Finca, 5 miles southwest of Turrialba, Cartago Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; July 14, 1947. Nos. 28211-28215, 28217-28222 same locality as holo- type; Edward H. Taylor and Richard C. Taylor; July 22-August 3, 1947. Nos. 31072-31078, Moravia de Chirripé, Cartago Province, Costa Rica; Edward H. Taylor; August 16, 1951. Hyla wellmanorum Taylor [= Smilisca wellmanorum (Taylor), according to Starrett (Copeia, 1960(4):303, 1960)], Univ. Kansas Sci. Bull., 35:843, fig. 52, July 1, 1952. Hototyre.—No. 30302, Bataan, Limén Province, Costa Rica; Edward H. Taylor; July 20, 1951. ParatyPres.—Nos. 30300 and 30301, data same as on holotype. Hyla weyerae Taylor, Univ. Kansas Sci. Bull., 36:633, June 1, 1954, Hotorype.—No. 34850, Esquinas Forest Preserve, Las Esquinas, between Golfito and Palmar, Puntarenas Province, Costa Rica; Mrs. Albert E. Weyer; October, 1952. Microbatrachylus persimilis (Barbour )—See Eleutherodactylus persimilis Barbour. Microbatrachylus rearki Taylor, Univ. Kansas Sci. Bull., 85:678, fig. 16, July 1, 1952. AMPHIBIAN AND REPTILE SPECIMENS 195 Hototyre.—No. 31533, Bataan, Limén Province, Costa Rica; Edward H. Taylor; July 25, 1951. ParatTyPeEs.—Nos. 30909-30911, 30913, 31539, 31540, 31543-31546, Tur- rialba, Cartago Province, Costa Rica; Edward H. Taylor; June 26-July 3, 1951. Nos. 30912, 31534-31538, data same as on holotype. Nos. 31541 and 31542, Los Diamantes, 1 kilometer east of Gudpiles, Limén Province, Costa Rica; Edward H. Taylor; July 26, 1951. Plectrohyla brachycephala Taylor [= Plectrohyla matudai brachycephala Taylor, according to Bumzahem and Smith (Herpetologica, 10:62, 1954)], Amer. Mus. Novitates, 1487:16, fig. 6, December 7, 1949. ParaTyPpE.—No. 28079, Tributary of Rio Ostuta, foot of Sierra Madre, be- tween Sierra Madre and Cerro Atravesado, Oaxaca, México; Thomas C. Mac- Dougall; 1948. Nortre.—Received in exchange; formerly American Museum of Natural History No. A 53758. Phyllomedusa saltator (Taylor)—See Agalychnis saltator Taylor. Ptychohyla chamulae Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 13:354, pl. 25, fig. 2, April 27, 1961. Hototyre.—No. 58063, 6.2 kilometers by road south of Rayén Mescalapa, Chiapas, México; William E. Duellman, Dale L. Hoyt, and John Wellman; August 5, 1960. ParaTyPEs.—Nos. 58064-58073, data same as on holotype. Smilisca wellmanorum (Taylor)—See Hyla wellmanorum Taylor. Teratohyla spinosa (Taylor)—See Centrolenella spinosa Taylor. Tomodactylus saxatilis Webb, Univ. Kansas Publ. Mus. Nat. Hist., 15:177, fig. 1, March 7, 1962. Ho.otyPe.—No. 63326, 8 miles west of El Palmito, Sinaloa, México; Rob- ert G. Webb; June 23, 1961. ParatyPes.—Nos. 63327-63333, data same as on holotype. TESTUDINES Gopherus flavomarginatus Legler, Univ. Kansas Publ. Mus. Nat. Hist., 11: 837, fig. 1, pls. 7-8, April 24, 1959. ParATYPE.—No. 39415 (carapace only), 8 miles east and 2 miles South of Americanos, Coahuila, México; Robert W. Dickerman; May 19, 1954. Pseudemys scripta taylori Legler, Univ. Kansas Publ. Mus. Nat. Hist., 13:75, figs. 1-3, pls. 9-12; August 16, 1960. Ho.totyre.—No. 46952, 16 kilometers south of Cuatro Ciénegas, Coahuila, México; John M. Legler, September 6, 1958. ParRATYPES.—Nos, 46932-46951, 46953-46969, data same as on holotype. Nos. 46970-46974, 6 miles west of Cuatro Ciénegas, Coahuila, México; John M. Legler; September 3-6, 1958. No. 46976, Rio Chiquito, 10 kilometers south of Cuatro Ciénegas; John M. Legler; September 9, 1958. Trionyx ater Webb and Legler, Univ. Kansas Sci. Bull., 40:21, pls. 1-2, April 20, 1960. 196 Universiry oF Kansas Pusts., Mus. Nat. Hist. Ho.toryPe.—No. 46903, 16 kilometers south of Cuatro Ciénegas, Coahuila; John M. Legler, Wendell L. Minckley, and Robert B. Wimmer; September 6, 1958. ParaTyPEes.—Nos. 46904-46906, 46908-46912, data same as on holotype. Trionyx muticus calvatus Webb, Univ. Kansas Publ. Mus. Nat. Hist., 11: 519, pls. 13-14, fig. 1, August 14, 1959. ParaTyPEes.—Nos. 47117-47119, Pearl River, 1 mile south to 4 miles north of Monticello, Lawrence County, Mississippi; Robert G. Webb; April 10, 1957. Trionyx spinifera pallidus Webb, Univ. Kansas Publ. Mus. Nat. Hist., 18: 522, pls. 39-40, February 16, 1962. PaRATYPES.—Nos. 2934, 2947, 2966, 2967, 2973, Lewisville, Lafayette County, Arkansas; R. E. McEntyre; May 1, 1926. Nos. 40175 and 40176, Lake Texoma, 2 miles east of Willis, Marshall County, Oklahoma; John M. Legler; summer, 1955. No. 50832, 4 miles southwest of Kingston, Mouth of Caney Creek, Marshall County, Oklahoma; Virgil Dowell; summer, 1954. SAURIA Alopoglossus plicatus Taylor [= Ptychoglossus plicatus (Taylor), according to Ruibal, Bull. Mus. Comp. Zool., 106:501, 1952)], Univ. Kansas Sci. Bull., 83:272, fig. 1, August 20, 1949. Ho.otrypre.—No. 23800, Morehouse Finca, 5 miles southwest of Turrialba, Cartago Province, Costa Rica; Edward H. Taylor; July 21, 1947. Ameiva festiva occidentalis Taylor, Univ. Kansas Sci. Bull., 38:264, fig. 66, December 20, 1956. Paratypes.—Nos. 40545-40548, 40553-40558, Agua Buena, Cafias Gordas, Puntarenas Province, Costa Rica; Edward H. Taylor; July 25-26, 1954. Nos. 40549 and 40550, 15 kilometers west-southwest of San Isidro del General, San José Province, Costa Rica; Edward H. Taylor; August 2, 1954. Anadia metallica arborea Taylor, Univ. Kansas Sci. Bull., 37:542, fig. 13, October 15, 1955. Hototype.—No. 34225, Las Flores, Tenoria de Las Cafias, Guanacaste Province, Costa Rica; John Baker and Edward H. Taylor; August 22, 1952. Anadia metallica attenuata Taylor, Univ. Kansas Sci. Bull., 37:537, fig. 12, October 15, 1955. Ho.otyre.—No. 84223, Pacuare, Rio Pacuare, Cartago Province, Costa Rica; Edward H. Taylor; August 17, 1951. ParatypPes.—No, 34224, Moravia de Chirripéd, Cartago Province, Costa Rica; Edward H. Taylor; June 26, 1952. Anolis achilles Taylor, Univ. Kansas Sci. Bull., 38:153, fig. 43, December 20, 1956. Ho.otyre.—No. 40634, La Palma, San José Province, Costa Rica; Edward H. Taylor; June 20, 1954. Anolis allisoni Barbour, Proc. New England Zool. Club, 10:58, July 26, 1928. AMPHIBIAN AND REPTILE SPECIMENS 197 ParaTyPe.—No. 47167, Coxen Hole, Ruatan Island, Honduras; Thomas Barbour; 1928. Norre.—Received in exchange; formerly this specimen was Museum of Comparative Zoology No. 26737. Anolis aquaticus Taylor, Univ. Kansas Sci. Bull., 38:141, fig. 39, December 20, 1956. Hototyre.—No. 34276, Palmar, Puntarenas Province, Costa Rica; John Baker; September 3, 1952. ParaTyPes.—Nos. 34277 and 384278, Golfito, Puntarenas Province, Costa Rica; Edward H. Taylor; September 3, 1952. Anolis biscutiger Taylor, Univ. Kansas Sci. Bull., 38:81, fig. 19, December 20, 1956. Ho.otyre.—No. 40771, Golfito, Puntarenas Province, Costa Rica; Edward H. Taylor, September 4-8, 1952. ParatyPes.—No. 40764, 10-15 kilometers west-southwest of San Isidro del General, San José Province, Costa Rica; Edward H. Taylor; July 10, 1952. Nos. 40766 and 40768, Palmar, Puntarenas Province, Costa Rica; John Baker and Edward H. Taylor; September 2, 1952. Nos. 40769, 40770, 40772- 40774, data same as holotype. Notes.—Taylor (1956:81) erroneously listed 40769, 40770, 40772-40774 as being from Palmar, instead of Golfito; paratypes 40765 and 40767 have been exchanged with the Museum of Comparative Zoology. Anolis humilis marsupialis Taylor, Univ. Kansas Sci. Bull., 38:97, fig. 28, December 20, 1956. Hototyre.—No. 40893, 15 kilometers west-southwest of San Isidro del General, San José Province, Costa Rica; Edward H. Taylor; July 14, 1952. ParATyPEs.—Nos. 40889, 40890, 40892, 40895 and 40896, data same as on holotype. Nore.—Paratypes 40891 and 40894 listed by Taylor (1956:97) have been exchanged with the Museum of Comparative Zoology. Anolis nelsoni Barbour, Mem. Mus. Comp. Zool., Harvard, 44:287, March, 1914. ParaTyPE.—No, 47168, Swan Island, Honduras; George Nelson; March, 1912. Nore.—Received in exchange from the Museum of Comparative Zoology. Anolis woodi attenuatus Taylor, Univ. Kansas Sci. Bull., 38:118, fig. 32, December 20, 1956. Hototype.—No. 34257, Isla Bonita, Heredia Province, Costa Rica; Ed- ward H. Taylor; July 29, 1947. PaRATYPE.—No. 34256, data same as holotype. Aristelliger nelsoni Barbour, Mem. Mus. Comp. Zool., Harvard, 44:258, March, 1914. 198 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Paratype-—No. 47169, Swan Island, Honduras; George Nelson; March, 1912. Nore.—Received in exchange; this specimen formerly was Museum of Comparative Zoology No. 9602. Crotaphytus collaris auriceps Fitch and Tanner, Trans. Kansas Acad. Sci., 54:553, December, 1951. Hototyrr.—No. 29934, 3.5 miles north-northeast of Dewey, west side of Colorado River, Grand County, Utah; Henry S. Fitch; June 28, 1950. PARATYPES.—Nos. 29935-29942, data same as on holotype. Eumeces brevirostris bilineatus Tanner, Great Basin Nat., 18:59, fig. 2, November 15, 1958. Hotorype.—No. 44732, 10 miles southwest of El Salto, Durango, México; Charles M. Fugler; June 28, 1955. Paratyprs.—Nos. 44728, 44730, 44731, 9 miles southwest of El Salto, Durango, México; Charles M. Fugler and Robert G. Webb; June 26-28, 1955. Nos. 44726, 44729, same locality as holotype; Rollin H. Baker and Robert G. Webb; June 25, 1955. No. 44727, 15 miles southwest of E] Salto, Durango, México; Charles M. Fugler; June 25, 1955. Eumeces dicei pineus Axtell, Copeia, 1960(1):19, figs. 1-2, March 25, 1960. HotoryreE.—No. 39539, Cafion de Carolina, 13 miles east of San Antonio de las Alazanas, Coahuila, México; Robert G. Webb; July 6, 1955. Eumeces gaigei Taylor [= Eumeces multivirgatus epipleurotis Cope, ac- cording to Axtell (Texas Jour. Sci., 13:345, 1961)], Univ. Kansas Sci. Bull., 22:219, fig. 1, April 15, 1985. Hototryre.—No. 7300, near Taos, Taos County, New Mexico; Edward H. Taylor; June 18, 1929. PaRATYPE.—No. 7301, data same as on holotype. Eumeces inexpectatus Taylor, Univ. Kansas Sci. Bull., 20:251, pls. 17-18, October 1, 1932. Hotoryrpe.—No. 8232, Citrus County, Florida; Charles E. Burt; March 7, 1980. Eumeces lynxe durangoensis Tanner, [= Eumeces lynxe lynxe (Wiegmann), according to Legler and Webb (Southwest. Nat., 5:16, 1960)], Great Basin Nat., 18:57, fig. 1, November 15, 1958. Hototyee.—No. 44737, 30 miles east of El Salto, Durango, México; Charles M. Fugler; June 30, 1955. ParRATYPES.—Nos. 44734 and 44735, same as holotype. No. 44736, 10 miles east of El Salto, Durango, México; Charles M. Fugler; June 30, 1955. Eumeces multivirgatus taylori Smith—See Eumeces taylori Smith. Eumeces taylori Smith [= Eumeces multivirgatus taylori Smith, according to Lowe (Herpetologica, 11:2338, 1955)], Proc. New England Zool. Club, 21:93, November 14, 1942. Paratype.—No. 13161, Black River, 2 miles east of entrance to Carlsbad Caverns, Eddy County, New Mexico; Edward H. Taylor; September 11, 1931. AMPHIBIAN AND REPTILE SPECIMENS 199 Holbrookia bunkeri Smith [= Holbrookia maculata bunkeri Smith, accord- ing to Williams and Smith (Herpetologica, 13:265, 1958)], Univ. Kansas Sci. Bull., 22:185, April 15, 1935. Hoxioryre.—No. 19236, 15 miles south of Juarez, Chihuahua, México; David H. Dunkle and Hobart M. Smith; June 19, 1934. Nore.—Formerly this specimen was H. M. Smith No. 80, as given in the original description. Holbrookia maculata bunkeri Smith—See Holbrookia bunkeri Smith. Leiolopisma cherriei lampropholis Taylor [= Scincella cherriei lampropholis (Taylor), according to generic arrangement by Mittleman (Smithsonian Misc. Coll., 117(No. 17):23; 1952)], Univ. Kansas Sci. Bull., 88:287, figs. 71-72, December 20, 1956. Hototyre.—No. 344386, Bataan, Limén Province, Costa Rica; Edward H. Taylor; July 28-25, 1951. ParatyPEs.—(all from Costa Rica) No. 25169, 5 kilometers southeast of Tilaran, Guanacaste Province; E. Raymond Hall and Marco Tulio Pacheo; March 10, 1947. Nos. 25524-25535, 25538-25542, Interamerican Institute of Agriculture, Turrialba, Cartago Province; Edward H. Taylor and others; June and July, 1947. Nos. 25536 and 25537, Morehouse Finca, 5 miles southwest of Turrialba, Cartago Province; Edward H. Taylor and Richard C. Taylor; July 11-12, 1947. Nos. 34409-34414, Los Diamantes, 1 kilometer east of Guapiles, Limén Province; John Baker and Edward H. Taylor; July and August, 1951. Nos. 34415-34485, 34437 and 34439, data same as on holotype. Nos. 34456-34469, Interamerican Institute of Agriculture, Turrialba, Cartago Province; Edward H. Taylor; July and August, 1951. Nos. 34470-34483, La Lola, Limén Province; John Baker and Edward H. Taylor; August 7-11, 1952. No. 84484, Puerto Limén, Limén Province; Edward H. Taylor; August 3, 1952. No. 34485, Mountain Cow Creek, near Limén, Limén Province; Ed- ward H. Taylor; August 6, 1952. Nos. 34502-34510, Interamerican Institute of Agriculture, Turrialba, Cartago Province; Edward H. Taylor; June, 1952. Nos. 34511-34539, 34864, Interamerican Institute of Agriculture; John Baker and Edward H. Taylor; September, 1952. Nos. 40514-40516, Suretka, Limén Province; Edward H. Taylor; June and July, 1954. No. 40521, Puerto Viejo, Heredia Province; Edward H. Taylor; August 7, 1954. Lepidophyma anomalum Taylor [= Lepidophyma obscurum Barbour, ac- cording to Taylor (Univ. Kansas Sci. Bull., 38:220, 1956)], Univ. Kansas Sci. Bull., 37:554, fig. 14, October 15, 1955. Hototrype.—No. 34207, Los Diamantes, 1 kilometer east of Guapiles, Limén Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; Sep- tember 8, 1947. PaRaTyPEs.—Nos. 34205 and 34206, data same as holotype. No. 34208, Morehouse Finca, 5 miles southwest of Turrialba, Cartago Province, Costa Rica; Edward H. Taylor; July 2, 1951. No. 34209, no specific locality in Costa Rica. No. 34210, Bataan, Limén Province, Costa Rica; Edward H. Taylor; July 24, 1951. Nos. 34211, 34212, 34214-34216, La Lola, Limon Province, Costa Rica; John Baker and Edward H. Taylor; August 8-12, 1952. No. 34213, Los Diamantes, 1 kilometer east of Gudpiles, Limén Province, Costa Rica; John Baker; August 12, 1952. 200 UNIVERSITY OF KANsas Pusis., Mus. Nat. Hist. Lepidophyma ophiophthalmum Taylor, Univ. Kansas Sci. Bull., 37:558, fig. 15, October 15, 1955. Ho.otypre.—No. 36250, 5 kilometers north-northeast of Tilaran, Guanacaste Province, Costa Rica; Edward H. Taylor; August 16, 1954. ParaTyPes.—Nos. 34217-84219, Las Flores, Tenorio de Las Cafias, Guana- caste Province, Costa Rica; Edward H. Taylor and John Baker; August 22, 1954. Lepidophyma reticulatum Taylor, Univ. Kansas Sci. Bull., 37:551, fig. 14, October 15, 1955. Ho.totyre.—No. 86245, Agua Buena, Puntarenas Province, Costa Rica; Edward H. Taylor; July 25, 1954. PaRrATyPEs.—Nos. 34189-34192, 3 kilometers east of San Isidro del Gen- eral, San José Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; September 1-8, 1947. No. 34193, San Isidro del General, San José Province, Costa Rica; John Baker; July 7, 1952. Nos. 34194-34202, 34204, 15-17 kilometers west-southwest of San Isidro del General, San José Province, Costa Rica; John Baker and Edward H. Taylor; July 10, 1952. Nos. 36230-36237, 4-5 kilometers northwest of Tilaran, Guanacaste Province, Costa Rica; Edward H. Taylor; August 10-15, 1954. Nos. 36238-36244, 36246-36249, data same as on holotype. Leposoma bisecta Taylor [= Leposoma southi southi Ruthven and Gaige, according to Ruibal (Bull. Mus. Comp. Zool., 106:482, 1952)], Univ. Kansas Sci. Bull., 33:275, fig. 2, April 20, 1949. Hototyre.—No. 23801, 0.5 mile west of San Isidro del General, San José Province, Costa Rica; Edward H. Taylor; August 21, 1947. Leposoma southi orientalis Taylor, Univ. Kansas Sci. Bull., 37:546, Oc- tober 15, 1955. HototyPe.—No. 36124, Volio, Limén Province, Costa Rica; Edward H. Taylor; June 24, 1954. ParatyPeE.—No. 36125, same as on holotype. Mabuya brachypoda Taylor, Univ. Kansas Sci. Bull., 38:308, fig. 76, De- cember 20, 1956. Hototypre.—No. 36258, 4 kilometers east-southeast of Tilaran, Guanacaste Province, Costa Rica; Edward H. Taylor; August 15, 1954. ParatyPEs.—Nos. 34810-34318, mouth of Rio Barranca, Puntarenas Province, Costa Rica; John Baker and Edward H. Taylor; July 26, 1952. Nos. 84314-84817, Tenorio de Las Camas, Guanacaste Province, Costa Rica; John Baker and Edward H. Taylor; August 21-24, 1952. No. 34826, Palmar, Puntarenas Province, Costa Rica; Edward H. Taylor; September 2, 1952. Nos. 36312 and 36313, Barranca, Puntarenas Province, Costa Rica; Edward H. Tay- lor; August 21, 1954, Mabuya macularia postnasalis Taylor and Elbel, Univ. Kansas Sci. Bull., 88:1105, March 20, 1958. Ho.totrype.—No. 40110, Phu Lom Lo (mountain), Kok Sathon (village), Dan Sai (district), Loei (province), Thailand; Robert E. Elbel; March 80, 1954. Paratypes.—Nos. 40109, 40111, 40112, data same as on holotype. AMPHIBIAN AND REPTILE SPECIMENS 201 Sceloporus poinsetti macrolepis Smith and Chrapliwy, Herpetologica, 13: 268, January 24, 1958. Paratypes.—No. 33855, 4 miles southwest of Coyotes, Durango, México; William M. Lynn; July 14, 1953. Nos. 38097 and 38098, 2 miles west of Sain Alto, Zacatecas, México; Robert W. Dickerman; June 7, 1954. Scincella cherriei lampropholis (Taylor)—See Leiolopisma cherriei lam- propholis Taylor. Urosaurus ornatus caeruleus (Smith)—See Uta Caerulea Smith. Uta caerulea Smith [= Urosaurus ornatus caeruleus (Smith), according to Mittleman (Bull. Mus. Comp. Zool., 91:139, 1942)], Univ. Kansas Sci. Bull., 22:172, April 15, 1935. Hotoryre.—No. 19237, about 380 miles north of Ciudad Chihuahua, Chihuahua, México; David H. Dunkle and Hobart M. Smith; June 21, 1934. ParatypPrs.—No. 19238, data same as on holotype. Nos. 19239 and 19240, 20 miles south of Ciudad Chihuahua, Chihuahua, México; David H. Dunkle and Hobart M. Smith; June 21, 1934. NotE.—Formerly these specimens were respectively H. M. Smith Nos. 132, 128, 141 and 148, as given in the original description. Xenosaurus nemanorum Taylor, Univ. Kansas Sci. Bull., 33:183, April 20, 1949. PaRATypEs.—Nos. 24043-24047, Xilitla region, San Luis Potosi, México; Marcella Newman; April 27-June 1, 1947. Note.—Formerly these specimens were respectively Louisiana State Uni- versity Nos. 502, 496, 492, 503, and 489. SERPENTES Bothrops schlegeli supraciliaris Taylor, Univ. Kansas Sci. Bull., 36:791, fig. 89, July 15, 1954. Hototyre.—No. 31997, mountains near San Isidro del General, San José Province, Costa Rica; collector and date unknown. Coniophanes fissidens obsoletus Minton and Smith, Herpetologica, 16:108, fig. 1, June 17, 1960. ParatyPes.—Nos. 30976-30979, 34834, Turrialba, Cartago Province, Costa Rica; John Baker and Edward H. Taylor; June-August, 1951. Dipsas costaricensis Taylor [= Sibon longifrenis (Stejneger), according to Peters (Misc. Publ. Mus. Zool. Univ. Michigan, 114:192, 1960)], Univ. Kansas Sci. Bull., 34:63, pl. 7, fig. 2, October 1, 1951. Hototyre.—No. 25703, Morehouse Finca, 5 miles southwest of Turrialba, Cartago Province, Costa Rica; Edward H. Taylor; July 16, 1947. Dipsas tenuissima Taylor, Univ. Kansas Sci. Bull., 36:771, figs. 28-29, July 15, 1954. Horotyre.—No. 31961, 15 kilometers west-southwest of San Isidro del General, San José Province, Costa Rica; Edward H. Taylor; July 10, 1952. Dryadophis sanguiventris Taylor, Univ. Kansas Sci. Bull., 36:722, July 15, 1954, 202 University OF Kansas Pusts., Mus. Nar. Hist. Hototyre.—No. 31978, Esquinas Forest Reserve, Las Esquinas, between Golfito and Palmar, Puntarenas Province, Costa Rica; Edward H. Taylor; September 3, 1952. Ficimia streckeri Taylor, Copeia, 1931(1):5, March 31, 1931. Ho.otyre.—No. 9140, 3 miles east of Rio Grande City, Starr County, Texas; Edward H. Taylor; July 18, 1930. Note.—Taylor (1931:5) erroneously listed the number of the holotype as 4140. Geophis acutirostris Taylor, Univ. Kansas Sci. Bull., 36:691, fig. 3, July 15, 1954. Hototyre.—No. 34670, Cot, south slope of Volcan Irazi, Cartago Province, Costa Rica; Edward H. Taylor; July 2, 1952. Geophis aquilonaris Legler, Univ. Kansas Publ. Mus. Nat. Hist., 11:329, figs. 1-2, January 28, 1959. Hototyre.—No. 44265, 23 miles south and 1.5 miles east of Creel, Chihuahua, México; Ronald H. Pine; July 23, 1957. ParaAtyPEe.—No. 44266, data same as on holotype. Geophis bakeri Taylor, Univ. Kansas Sci. Bull., 36:689, July 15, 1954. Hototype.—No. 31983, Isla Bonita, Heredia Province, Costa Rica; Ed- ward H. Taylor; July 19, 1952. ParatypEs.—No. 30940, Pacayas, Cartago Province, Costa Rica; Edward H. Taylor; August 6, 1951. Nos. 31982, 31984, data same as on holotype. Geophis zeledoni Taylor, Univ. Kansas Sci. Bull., 36:693, fig. 4, July 15, 1954. Hototype.—No. 31992, Finca Zeledon, between Volcan Barba and Volcan Pods, Heredia Province (?), Costa Rica; Edward H. Taylor; July 24, 1952. ParatyPe.—No. 31951, data same as on holotype. Lampropeltis doliata taylori Tanner and Loomis, Trans. Kansas Acad. Sci., 60:31, pls. 3-4, April 23, 1957. ParaTyPE.—No, 38952, Vernal, Uintah County, Utah; Lawrence Atwood; 1955. Lycodon osmanhilli Taylor, Univ. Kansas Sci. Bull., 33:562, pl. 20, March 20, 1950. Horotype.—No, 24141, Colombo, Ceylon; W. C. Osman Hill; date un- known. Micrurus alleni richardi Taylor [= Micrurus richardi Taylor, according to Taylor (Univ. Kansas Sci. Bull., 36:777, 1954)], Univ. Kansas Sci. Bull., 34: 169, pl. 23, fig. 7, October 1, 1951. Ho.otyre.—No, 25189, Los Diamantes, 1 kilometer east of Guapiles, Limdén Province, Costa Rica; Edward H. Taylor and Richard C. Taylor; September 8, 1947. Micrurus pachecoi Taylor, Univ. Kansas Sci. Bull., 34:165, pl. 22, fig. 6, October 1, 1951. Ho.otyre.—No. 25188, Guanacaste Province, Costa Rica; collector and date unknown. AMPHIBIAN AND REPTILE SPECIMENS 203 Micrurus richardi Taylor—See Micrurus alleni richardi Taylor. Natrix harteri paucimaculata Tinkle and Conant, Southwestern Naturalist, 6:34 figs. 1-2, June 10, 1961. PARATYPES.—Nos. 59884 and 59885, Colorado River south of city limits of Robert Lee, Coke County, Texas; Donald W. Tinkle and Don L. McGregor; October 8, 1960. Natrix rigida sinicola Huheey, Copeia, 1959(4):305, fig. 1, December 30, 1959. PARATYPE.—No. 24500, Steep Hill Creek, Natchitoches Parish, Louisiana; Henry S. Fitch; August 15, 1947. Ninia cerroensis Taylor, Univ. Kansas Sci. Bull., 36:699, figs. 5-6, July 15, 1954. Hotoryre.—No. 31935, Pacific slope of Cerro de la Muerte, 7500 feet, San José Province, Costa Rica; Edward H. Taylor; July 8, 1952. Nothopsis torresi Taylor, Univ. Kansas Sci. Bull., 34:31, pl. 1, October 1, 1951. Ho.otyPE.—No. 25719, Morehouse Finca, 5 miles southwest of Turrialba, Cartago Province, Costa Rica; Edward H. Taylor; July 16, 1947. Note.—Taylor (1951:31) erroneously listed the number of the holotype as 28719. Pliocercus annellatus Taylor, Univ. Kansas Sci. Bull., 34:107, pl. 10, fig. 4, October 1, 1951. Hototypre.—No. 25730, Morehouse Finca, 5 miles southwest of Turrialba, Cartago Province, Costa Rica; Edward H. Taylor; July 11, 1947. Notre.—Taylor (1951:107) erroneously listed the number of the holotype as 25370. Rhadinaea altamontanus Taylor, Univ. Kansas Sci. Bull., 36:740, fig. 16, July 15, 1954. Ho.otyre.—No. 30962, National Forest Reserve, Pan-American Highway, Talamanca Range, Cartago Province, Costa Rica; Edward H. Taylor and Jack Reark; August, 1951. Rhadinaea decipiens rubricollis Taylor, Univ. Kansas Sci. Bull., 36:739, July 15, 1954. Ho.otypr.—No. 31956, Cinchona, Alajuela Province, Costa Rica; Edward H. Taylor; June 23, 1952. Sibynophis collaris triangularis Taylor and Elbel, Univ. Kansas Sci. Bull., 38:1130, fig. 23, March 20, 1958. Ho.otyre.—No. 33520, Nongko (village), Siracha (district), Chon Buri (province), Thailand; Boonsong Lekagul; August 18, 1953. Tantilla costaricensis Taylor [= Tantilla schistosa costaricensis Taylor, ac- cording to Smith (Herpetologica, 18:16, 1962)], Univ. Kansas Sci. Bull., 36:766, fig. 25a, July 15, 1954. Hotoryre.—No. 30995, Cervantes, Cartago Province, Costa Rica; Edward H. Taylor; August 30, 1951. Tantilla schistosa costaricensis Taylor—See Tantilla costaricensis. 204 UNIVERSITY OF KANSAS PuBts., Mus. Nat. Hisv. Tantilla schistosa taylori Smith, Herpetologica, 18:17, April 9, 1962. Ho.iotyre.—No. 35627, Suretka, Lim6én Province, Costa Rica; Edward H. Taylor; August 1, 1954. Thamnophis melanogaster chihuahuaensis Tanner, Herpetologica, 15:170, figs. 1-2, December 8, 1959. Paratypes.—Nos. 44275-44278, 23 miles south and 1.5 miles east of Creel, Chihuahua, México; Sydney Anderson and Ronald H. Pine; July 19-21, 1957. Notre.—Tanner (1959:170) erroneously listed these specimens as being from 1.5 miles east of Creel. Tropidoclonion lineatum annectens Ramsey, Herpetologica, 9:12, June 1, 1953. Hototyre.—No. 30005, Tulsa, Tulsa County, Oklahoma; Harold Dundee; May 3, 1951. PARATYPES.—Nos. 30006-30011, data same as holotype. Typhlops silus Legler [= Typhlops lumbricalis (Linnaeus), according to Richmond (Copeia, 1961(2):222, 1961)], Herpetologica, 15:105, June 2, 1959. Hototyre.—No. 47469, Banes, Oriente Province, Cuba; Chapman Grant; April, 1956. ParatyPes.—Nos. 47470 and 47471, data same as on holotype. Transmitted June 28, 1962. O 28-7817 UNIVERSITY OF KANSAS PUBLICATIONS | j_ Wiel gs MusEuM OF NATURAL HISTORY Volume 15, No. 5, pp. 205-249, pls. 7-10, 6 figs. October 4, 1963 Amphibians and Reptiles of the Rainforests of Southern El Peten, Guatemala BY WILLIAM E. DUELLMAN UNIVERSITY OF KANSAS LAWRENCE 1963 UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HisTory Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Vol. 15, No. 5, pp. 205-249, pls. 7-10, 6 figs. Published October 4, 1963 UNIVERSITY OF KANSAS Lawrence, Kansas Li 1963 j ; UNIVERSITY | AI One EE 8 I sO PRINTED BY JEAN M. NEIBARGER, STATE PRINTER TOPEKA, KANSAS Amphibians and Reptiles of the Rainforests of Southern El Petén, Guatemala BY WILLIAM E. DUELLMAN CONTENTS PAGE INTRODUCTION) lhe. exe Sisco te nee ee es oe eee eran 207 Acknowledgments’)... ..20. 0... 000. toes ie oie te 208 IDESCRIPTION OF AREA... ccc nade de eee ee eee 208 Physiograply eo. ste. s,| e- «ke She visas tes Pee eee 209 Glimate:- % 500 Oe es CHA eS Soles eee OO ee eee 209 Veretationsdt 2 2. atone be Spode ohana et er ea eee 209 GAGETTERR Cee ORIe Vo oh M5 Se ERR Ree ee re 210 Tar HeRPETOFAUNA OF THE RAINFOREST: . 720222 .-- 052 oe sae 214 Composition of the: Fauna, 1023. .52 ty 3> «oe 212 Ecology. of the Herpetofauna sie s):2 5. ae so ee 212 Relationships,of the fauna .. 5.45720. 0-4) eee ee 217 ACCOUNTS OF SPECIES 55, cpus. «sie. sickens Bee ee 218 ELYPOTHETICAL, LIST OF SPECIES: as 04 en eee ee eee 246 SUMMARY seit dk tt. eldlon.0s ae ee ee ee 247 erreRATORENCrrEeD (ee. es PS ee A ee 247 INTRODUCTION Early in 1960 an unusual opportunity arose to carry on biological field work in the midst of virgin rainforest in southern El Petén, Guatemala. At that time the Ohio Oil Company of Guatemala had an air strip and camp at Chinaja, from which place the company was constructing a road northward through the forest. In mid- February, 1960, J. Knox Jones, Jr. and I flew into El Petén to collect and study mammals, reptiles, and amphibians. While en- joying the comforts of the fine field camp at Chinaja, we worked in the surrounding forest and availed ourselves of the opportunity to be on hand when the road crews were cutting the tall trees in the forest, thereby bringing to the ground many interesting speci- mens of the arboreal fauna. We stayed at Chinajé until late March, with the exception of a week spent at Toocog, another camp of the Ohio Oil Company located 15 kilometers southeast of La Libertad and on the edge of the savanna. Thus, at Toocog we were able (207) 208 UNIvERSITY OF Kansas Pusts., Mus. Nat. Hist. to work both in the forest and on the savanna. In the summer of 1960, John Wellman accompanied me to E] Petén for two weeks in June and July. Most of our time was spent at Chinaja, but a few days were spent at Toocog and other localities in south-central E] Petén. Many areas in Guatemala have been studied intensively by L. C. Stuart, who has published on the herpetofauna of the forested area of northeastern El Petén (1958), the savannas of central El Petén (1935), and the humid mountainous region to the south of El Petén in Alta Verapaz (1948 and 1950). The area studied by me and my companions is covered with rainforest and lies to the north of the highlands of Alta Verapaz and to the south of the savannas of central El] Petén. A few specimens of amphibians and reptiles were obtained in this area in 1935 by C, L. Hubbs and Henry van der Schalie; this collection, reported on by Stuart (1937), contained only one species, Cochranella fleishmanni, not present in our collection of 77 species and 617 specimens. Acknowledgments I am grateful to L. C. Stuart of the University of Michigan, who made the initial arrangements for our work in El] Petén, aided me in the identification of certain specimens, and helped in the preparation of this report, J. Knox Jones, Jr. and John Wellman were able field companions, who added greatly to the number of specimens in the collection. In Guatemala, Clark M. Shimeall and Harold Hoopman of the Ohio Oil Company of Guatemala made available to us the facilities of the company’s camps at Chinaji and Toocog. Alberto Alcain and Luis Escaler welcomed us at Chinaji and gave us every possible assistance. Juan Monteras and Antonio Aldafia made our stay at Toocog enjoyable and profitable. During our visits to southem E] Petén, Julio Bolén C, worked for us as a collector, and between March and June he col- lected and saved many valuable specimens; his knowledge of the forest and its inhabitants was a great asset to our work. Jorge A. Ibarra, Director of the Museo Nacional de Historia Natural in Guatemala assisted us in obtaining necessary permits and extended other kindnesses. To all of these people I am indebted for the essential parts that they played in the completion of this study, Field work in the winter of 1960 was made possible by funds from the American Heart Association for the purposes of collecting mammalian hearts. My field work in the summer of 1960 was supported by a grant from the Graduate Research Fund of the University of Kansas. DESCRIPTION OF THE AREA A vast lowland region stretches northward for approximately 700 kilometers from the highlands of Guatemala to the Gulf of Mexico, The northern two-thirds of this low plain is bordered on three sides by seas and forms the Yucatan Peninsula. The lowlands AMPHIBIANS AND REPTILES OF EL PETEN 209 at the base of the Yucatan Peninsula make up the Departamento El Petén of Guatemala. The area with which this report is con- cerned consists of the south-central part of El Petén. Physiography Immediately south of Chinaja is a range of hills, the Serrania de Chinaja, having an almost due east-west axis and a crest of about 600 meters above sea level. South of the Serrania de Chinaj4 are succeedingly higher ridges building up to the Meseta de Coban and Sierra de Pocolha and eventually to the main Guatemalan highlands. The northern face of the Serrania de Chinajaé is a fault scarp dropping abruptly from about 650 meters at the crest to about 140 meters at the base. From the base of the Serrania de Chinajé northward to the Rio de la Pasién at Sayaxché the terrain is gently rolling and has a total relief of about 50 meters, North of the Rio de la Pasidn is a low dome reaching an elevation of 170 meters at La Libertad; see Stuart (1935:12) for further discussion of the physiography of central El Petén. The rocks in southern El Petén are predominately Miocene marine limestones; there are occasional pockets of Pliocene deposits. There is little evidence of subterranean solution at Chinajé, but northward in central El Petén karsting is common, The upper few inches of soil is humus rich in organic matter; below this is clay. Climate The climate of El Petén is tropical with equable temperatures throughout the year. Temperatures at Chinaji varied between a night-time low of 65° F. and a daytime high of 91° F. during the time of our visits. In the Képpen system of classification the climate at Chinaja and Toocog is Af. Rain falls throughout the year, but there is a noticeable dry season. To anyone who has traveled from south to north in E] Petén and the Yucatan Peninsula, it is obvious from the changes in vegetation that there is a decrease in rainfall from south to north. There is a noticeable difference between Chinaj& and Toocog. Although rainfall data are not available for Chinaj4 and Toocog, there are records for nearby stations (Sapper, 1932). At Paso Caballos on the Rio San Pedro about 40 kilometers northwest of Toocog the average annual rainfall amounts to 1620 mm.; the driest month is March (21 mm.), and the wettest months are June (269 mm.) and September (265 mm.). At Cubiquitz, Alta Verapez, about 35 kilometers south-southwest of ChinajA and at an elevation of 300 meters, the average annual rainfall is 4006 mm.; the driest month is March (128 mm.), and the wettest months are July (488 mm.) and October (634 mm.). During the 18 days in February and March, 1960, that we kept records on the weather at ChinajA4 moderate to heavy showers occurred on seven days. During our stay there in June and July rain fell every day, as it did in Toocog. However, during the week spent at Toocog in March no rain fell. Vegetation The vegetation of northem and central El Petén has been studied by Lundell (1937), who made only passing remarks concerning the plants of the southern part of El Petén. No floristic studies have been made there. The 210 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. following remarks are necessarily brief and are intended only to give the reader a general picture of the forest. I have included names of a few of the commoner trees that I recognized. Chinaja is located in a vast expanse of unbroken rainforest. In this forest there is a noticeable stratification of the vegetation. Three strata are apparent; in the uppermost layer the tops of the trees are from 40 to 50 meters above the ground. The spreading crowns of the trees and the interlacing vines form a nearly continuous canopy over the lower layers. Among the common trees in the upper stratum are Calophyllum brasiliense, Castilla elastica, Cedrela mexicana, Ceiba pentandra, Didalium guianense, Ficus sp., Sideroxylon lundelli, Swietenia macrophylla, and Vitex sp. (Pl. 1, fig. 1). The middle layer of trees have crowns about 25 meters above the ground; these trees in some places where the upper canopy is missing form the tallest trees in the forest. This is especially true on steep hillsides. Common trees in the middle layer include Achras zapote, Bombax ellipticum, Cecropia mexicana, Orbignya cohune, and Sabal sp. The lowermost layer reaches a height of about 10 meters; in many places in the forest this layer is absent. Common trees in the lower stratum include Crysophila argentea, Cymbopetalum penduliflorum, Casearia sp., and Hasseltia dioica. The ground cover is sparce; apparently only a few small herbs and ferns live on the heavily shaded forest floor. Important herpetological habitats include the leaf litter, rotting stumps, and rotting tree trunks on the forest floor and the buttresses of many of the gigantic trees, especially Ceiba pen- tandra (Pl. 2). Epiphytes, especially various kinds of bromeliads, are common. Most frequently these are in the trees in the upper and middle strata. At Toocog there is sharp break between savanna and forest (Pl. 7, fig. 2). The forest is noticeably drier and more open than at Chinaja (Pl. 9). The crowns of the trees are lower, and there is no nearly continuous canopy between 40 and 50 meters above the ground. Although Swietenia macrophylla and other large trees occur, they are Jess common than at Chinaji. Especially common at Toocog are Achras zapote, Brosimum alicastrum, and various species of Ficus, GAZETTEER The localities from which specimens were obtained are cited below and shown on the accompanying map (Fig. 1). Chinaji.—Lat. 16° 02’, long. 90° 13’, elev. 140 m. Camp of the Ohio Oil Company of Guatemala and formerly a small settlement. On some maps Chinaja is located just to the north of the Alta Verapaz—E]l Petén boundary; recent surveys place the location just to the south of the imaginary line through the rainforest. Field work was conducted in the immediate vicinity of the camp, on the lower slopes of the Serrania de Chinaj4, and at several sites to the northwest and north-northwest of Chinaj4, where the forest was being cleared. The entire area supports rainforest. La Libertad—Lat. 16° 47’, long. 90° 07’, elev., 170 m. A town on the savannas in central El Petén; although we collected there in the rainy season, the specimens obtained on the savannas are not included in this report. Paso Subin.—Lat. 16° 38’, long. 90° 12’, elev. 90 m. A small settlement on the Rio Subin, a tributary of the Rio de la Pasién. Specimens were ob- tained in rainforest in the immediate vicinity of the settlement. Rio de la Pasién.—A large river flowing northward through southern El Petén and thence westward into the Rio Usumacinta. Specimens were obtained along the river between the Rio Subin and Sayaxché. AMPHIBIANS AND REPTILES OF EL PETEN 211 MEXICO oie Seer as Fs BU dat fet She Mel SE Se weit c ' i t { i i i Sent oUaxactun we oTikal Bs A \ - o Piedras Negras GX fio Son Pedro BI PETEN Logo El Petén Bie Frio Usumacinta eines oLa Libertad BRITISH HONDURAS Kilometers S ALTA VERAPAS Fic. 1. Map of El Petén, Guatemala, showing localities mentioned in text. Rio San Rom4n.—A river flowing northward in south-central E] Petén to the Rio Salinas (Usumacinta). We collected along the river at a place about 16 kilometers north-northwest of Chinaja, approximately at Lat. 16° 10’, long. 90° 17’, elev. 110 m. In the dry reason the river was clear; it is surrounded by rainforest. Sayaxché.—Lat. 16° 31’, long. 90° 09’, elev. 80 m. A town on the southem bank of the Rio de la Pasion. Specimens were obtained in the rainforest and in cleared areas in the immediate vicinity of the town. Toocog (formerly Sojio).—Lat. 16° 41’, long. 90° 02’, elev. 140 m. A camp of the Ohio Oil Company of Guatemala located at the rainforest-savanna edge, 15 kilometers southeast of La Libertad. Although we collected on the savannas as well as in the forest, especially to the east of the camp, only species obtained in the forest are considered in this report. THE HERPETOFAUNA OF THE RAINFOREST In presenting an account of the herpetofauna of southern El Petén three items need to be considered: (1) The composition of the fauna; (2) the ecology of the fauna; (3) the relationships of the fauna. Each of these topics is discussed briefly below. Logically a discussion of the origin of the fauna should follow, but this is being withheld for inclusion in a report on the herpetofauna of the entire El Petén by L. C. Stuart and the author; at that time the above topics will be expanded to cover the herpetofauna of the whole region. 212 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Composition of the Fauna A total of 78 species of amphibians and reptiles has been found in the rainforests in southem El] Petén; a break down into families and genera is given in table 1. Another 13 species probably occur in southern El] Petén (see TABLE 1.—COMPOSITION OF THE HERPETOFAUNA IN SOUTHERN EL PETEN, GUATEMALA. Group Families Genera Species Gymn ophiona:. sig jase see eee (1)* (1) (1) Caudata ar . orocu Shc tcer tea etb aioe 1 1 2 DAL EMoLa Sites hatte ace ere 6 10 (1) 19 (1) Crocodiligz een see er: 1 1 1 Mestudines a ik.cco case etes Meee teases ae 4 a 8 Sauriayie: 24 tei tcicheeo ne ama 6 13 (1) 19 (1) Derpembes 32 ./o'S5 ss vais wus, chee amie 4 PAI) 29 (10) BOG a Pe Te RR ee 22 (1) 53 (10) 78 (13) * Numbers in parenthesis indicate the number of additional taxa that probably occur. Hypothetical List of Species). The fauna primarily is composed of typical humid lowland forest inhabitants, such as: Hyla ebraccata Eumeces sumichrasti Hyla loquax Ameiva festiva edwardsi Phyllomedusa callidryas taylori Imantodes cenchoa leucomelas Smilisca phaeota cyanosticta Leptophis ahuetulla praestans Anolis biporcatus Xenodon rabdocephalus mexicanus Anolis capito Bothrops nasutus Anolis humilis uniformis Bothrops schlegeli schlegeli Nevertheless, the region also provides at least a limited amount of habitat suitable for some species that are more frequently found in open forest of a drier nature; such species include: Hyla microcephala martini Anolis sericeus sericeus Hyla staufferi Eumeces schwartzei Hupenocie cuneus nigroreticu- Oxybelis aeneus aeneus atus Because of the absence of sufficiently open habitat or owing to the presence of competitors, some conspicuous members of sub-humid forests are not present in southern El] Petén. Conspicuous absentees are the following: Rhinophrynus dorsalis Ameiva undulata Phrynohyas spilomma Cnemidophorus angusticeps Triprion petasatus Conophis lineatus Anolis tropidonotus Masticophis mentovarius Ctenosaura similis mentovarius Ecology of the Herpetofauna Our two visits to Chinaj4 and Toocog afforded the opportunity to gather data on the ecology of the rainforests of southern El Petén and to study the relationships between the environment and mem- bers of the herpetofauna. Tropical rainforests present the opti- PEAGES7 ee, te along Fic. 1. Edge of rainforest lS ieee i MMA cee ere S a be ih Ca RPS A Rainforest at edge of savanna at Toocog, El] Petén, Guatemala. 4 tev Bic. 2: Interior of rainforest at Chinaja. Notice size of buttresses on large tree (Ceiba pentandra). PLATE 9 Interior of rainforest at Toocog. Notice less dense vegetation as compared with Pl. 8. PLATE 10 \ eT : a) Tai A Fic. 1. Rainforest along Rio San Roman, 16 kilometers north-northwest of Chinaja. Fic. 2. Rain pond in forest at Toocog. This was a breeding site for six species of frogs. AMPHIBIANS AND REPTILES OF EL PETEN 213 mum conditions for life, and it is in this environment that life reaches its greatest diversity. Here, too, biological inter-relation- ships are most complex. This complexity is illustrated by the presence of many species of some genera, all of which are found together in the same geographic region. In the rainforests of southern E] Petén there are six species of Anolis, five of Hyla, four of Bothrops, and three of Coniophanes. Obviously, the di- versity of ecological niches in the rainforest is sufficient to support a variety of related species. Of the examples mentioned above, fairly adequate ecological data were obtained for most of the species of Anolis, which will be used to show the ecological diver- sity and vertical stratification of sympatric species in the rainforests. Of the six species of Anolis, all except A. sericeus are typically found in humid forests. Anolis sericeus sericeus is poorly repre- sented in the collections from southern El Petén, where it may be in competition with Anolis limifrons rodriguezi that resembles Anolis s. s€riceus in size, coloration, and habits. Therefore, Anolis sericeus sericeus is excluded from the following discussion. The com- mon terrestrial species is Anolis humilis uniformis; sometimes this small species perches or suns on the bases of small trees or buttresses of some large trees. When disturbed it takes to the ground and seeks cover in the leaf litter or beneath logs or palm fronds. Anolis lemurinus bourgeaei is about twice the size of Anolis humilis uniformis and is usually observed on buttresses of large trees or on the lower two meters of tree trunks. Individuals were seen foraging on the ground along with Anolis humilis uniformis. At no time were Anolis lemurinus bourgeaei observed to ascend the trunks of large trees; they always took refuge near the bases of trees. Anolis limifrons rodriguezi is found on the stems and branches of bushes. It is a small species that sometimes is observed on the ground but was never seen ascending large trees. Anolis capito is about the same size as Anolis lemurinus bourgeaci and lives on the trunks of large trees. In the tops of the trees lives a large green species, Anolis biporcatus. Similar segregation habitatwise can be demonstrated for other members of the herpetofauna. The avoidance of interspecific competition in feeding is well illustrated by three species of snakes that probably are the primary ophidian predators on frogs. Drymobius margaritiferus margaritiferus is diurnal and terrestrial; it feeds on frogs at the edges of breeding ponds by day. Also during the day Leptophis mexicanus mexicanus feeds on frogs in 214 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. bushes and trees. At night the activities of both of these species is replaced by those of Leptodeira septentrionalis polysticta, which not only feeds on the frogs in the trees and bushes, but descends to the ground and even enters the water in search of food. From the examples discussed above, the importance of the three dimensional aspect of the rainforest is apparent. The presence of a large and diverse habitat above the ground is of great significance in the rainforest, for of the non-aquatic com- ponents of the herpetofauna in the rainforests of southern El Petén, 42, per cent of the species spend at least part of their lives in the bushes and trees. Another important part of the forest is the subterranean level—the rich mulch, underground tunnels, and rotting subterranean vegetation. Of the 78 species of amphibians and reptiles in southern E] Petén, seven are primarily fossorial, and half-a-dozen others are secondarily fossorial. Probably the fos- sorial members of the fauna are the least well represented in the collection, for such widespread species as Dermophis mexicanus mexicanus, Rhadinaea decorata decorata and Tantilla schistosa schistosa were expected, but not found. In the following discussion of the ecological distribution of amphibians and reptiles in the rainforest I have depended chiefly on my observations made in southern El Petén, but have taken into consideration observations made on the same species in other regions, together with reports from other workers. The reader should keep in mind that the evidence varies from species to spe- cies. Of some species I have observed only one animal in the field; of others, I have seen scores and sometimes hundreds of individuals. For species on which I have few observations or rather inconclusive evidence, the circumstance of inadequate data is mentioned. In analyzing the ecological distribution within the forest, it is convenient to recognize five subdivisions (habitats); each is treated below as a unit. 1. Aguatic.—This habitat includes permanent streams and rivers (Pl. 10, fig. 1), some of which are clear and others muddy. In the rainy season temporary ponds form in depressions on the forest floor (Pl. 10, fig. 2); these are important as breeding sites for many species of amphibians. Aquatic members of the herpeto- fauna are here considered to be those species that either spend the greatest part of their lives in the water or usually retreat to water for shelter. Seven species of turtles and one crocodilian are AMPHIBIANS AND REPTILES OF EL PETEN 915 aquatic. Of these, Dermatemys mawi, Staurotypus triporcatus, and Pseudemys scripta ornata inhabit clear water, whereas Chelydra rossignoni, Claudius angustatus, Kinosternon acutum, and K. leu- costomum inhabit muddy water. Crocodylus moreleti apparently inhabits both clear and muddy water, for in the dry season it lives along the clear rivers, but in the rainy season inhabits flooded areas in the forest as well. 2. Aguatic Marcin.—Extensive marshes were lacking in the part of southern El Petén that I visited; consequently, the aquatic margin habitat is there limited to the edges of rivers and borders of temporary ponds. Bufo marinus, Rana palmipes, and Rana pipiens are characteristic inhabitants of the aquatic margin, although in the rainy reason Bufo marinus often is found away from water. Observations indicate that Tretanorhinus nigroluteus lateralis inhabits the margins of ponds and streams and actually spends considerable time in the water. Although Iguana iguana rhinolopha is arboreal, it lives in trees along rivers, into which it plunges upon being disturbed. Species included in this category are those that customarily spend most of their lives at the edge of permanent water. Frogs and toads that migrate to the water for breeding and the snakes that prey on the frogs at that time are not assigned to the aquatic-margin habitat. 3. FossorraL.—Characteristic inhabitants of the mulch on the forest floor are Bolitoglossa moreleti mulleri, Lepidophyma flavi- maculatum flavimaculatum, Scincella cherriei cherriei, Ninia sebae sebae, Pliocercus euryzonus aequalis, and Micrurus affinis apiatus. Other species of snakes that spend most of their lives above ground often forage in the mulch layer; among these are Coniophanes bipunctatus biserialis, Coniophanes fissidens fissidens, Coniophanes imperialis clavatus, Lampropeltis doliata polyzona, and Stenorrhina degenhardti. Among the amphibians, at least Hypopachus cuneus nigroreticulatus, Eleutherodactylus rostralis, and Syrrhophus leprus are known to seek shelter in the mulch. 4, TERRESTRIAL.—One turtle, Geoemyda areolata, is primarly terrestrial. Among the lizards, conspicuous terrestrial species are Anolis humilis uniformis and Ameiva festiva edwardsi; Anolis lemurinus bourgeaei and Basiliscus vittatus spend part of their lives on the ground, but also live on trees and in bushes. Eumeces schwartzei and E, sumichrasti apparently are terrestrial. The only terrestrial lizard that is nocturnal is Coleonyx elegans elegans, which 216 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist. by day hides in the leaf litter or below ground. Nocturnal am- phibians that are terrestrial include Bufo marinus, Bufo valliceps valliceps, Eleutherodactylus rugulosus rugulosus, Syrrhophus leprus, and Hypopachus cuneus nigroreticulatus. A large number of active diurnal snakes are terrestrial; these include Boa constrictor im- perator, Clelia clelia clelia, Dryadophis melanolomus laevis, Dry- marchon corais melanurus, Drymobius margaritiferus margaritiferus, Pseustes poecilonotus poecilonotus, and Spilotes pullatus mexicanus. Nocturnal terrestrial snakes include three kinds of Bothrops (B. atrox asper, B. nasutus, and B. nummifer nummifer), all of which seem to be equally active by day. 5. ARBOREAL.—In this habitat the third dimension (height) of the rainforest probably is the most complex insofar as the inter- relationships of species and ecological niches are concerned. I have attempted to categorize species as to microhabitats within the arboreal habitat; in so doing, I recognize four subdivisions— bushes, tree trunks, tree tops, and epiphytes. Bush inhabitants include several species of lizards and snakes, all of which have rather elongate, slender bodies, and long tails. Common bush-inhabitants in southern El Petén are Anolis limifrons rodriguezi, Basiliscus vittatus, Laemanctus deborrei, Leptophis mexicanus mexicanus, and Oxybelis aeneus aeneus. All of these are diurnal, and all but Laemanctus have been observed sleeping on bushes at night. Tree-trunk inhabitants include five species of lizards. Theca- dactylus rapicaudus lives on the trunks of large trees; Sphaero- dactylus lineolatus lives beneath the bark on dead trees and on corozo palms. Anolis lemurinus bourgeaei lives on the bases and butresses of large trees, from which it often descends to the ground, Corythophanes cristatus and Anolis capito were found only on tree trunks and large vines. The least information is available for the species living in the tree tops. The following species were obtained from tops of trees when they were felled, or have been observed living in the tree tops: Anolis biporcatus, Iguana iguana rhinolopha, Celestus rozel- lae, Leptodeira septentrionalis polysticta, Leptophis ahaetulla praestans, Sibon dimidiata dimidiata, and Sibon nebulata nebulata. Epiphytes, especially the bromeliads, provide refuge for a variety of tree frogs and small snakes. Of the tree frogs, Hyla picta, Hyla staufferi, Phyllomedusa callidryas taylori, Similisca baudini, and Similisca phaeota cyanosticta have been found in bromeliads; other AMPHIBIANS AND REPTILES OF EL PETEN 217 species probably occur there. Among the snakes, Imantodes cenchoa leucomelas, Leptodeira frenata malleisi, Leptodeira septentrionalis polysticta, Sibon dimidiata dimidiata, and Sibon nebulata nebulata are frequent inhabitants of bromeliads; all of these snakes are nocturnal. Relationships of the Fauna Most of the 78 species of amphibians and reptiles definitely known from the rainforest in southern El Petén have extensive ranges in the Atlantic lowlands of southern México and Central America; many extend into South America. Sixty-two (80%) of the species belong to this group having extensive ranges in Middle America. Three species (Syrrhophus leprus, Leptodeira frenata, and Kinosternon acutum) are at the southern limits of their dis- tributions in southern El Petén and northern Alta Verapaz, whereas Eleutherodactylus rostralis and Thecadactylus rapicaudus are at the northern and western limits of their distributions in El Petén. Nine (11%) species have the center of their distributions in El Petén and the Yucatan Peninsula; representatives of this group include Claudius angustatus, Dermatemys mawi, Laemanctus deborrei, and Eumeces schwartzei. In determining a measure of faunal resemblance, I have departed from the formulae discussed by Simpson (1960) and have analyzed the degree of resemblance by the following formula used to cal- culate an index of faunal relationships: C (2) Ni + Ne C=species common to both faunas. Ni = number of species in the first fauna. Neo = number of species in the second fauna, R = degree of relationships (when R = 1.00, the faunas are identical; when R= 0, the faunas are completely different). = R, where The herpetofauna of southern El] Petén has been compared with that in the Tikal-Uaxacttin area (Stuart, 1958), that in the humid lowlands of Alta Verapaz (Stuart, 1950, plus additional data), and that in the Mexican state of Yucatin (Smith and Taylor, 1945, 1948, and 1950). The herpetofaunas of lowland Alta Verapaz and Yucatan are the largest, having respectively 94 and 91 species, where as there are 78 species known from southern E] Petén and 64 from the Tikal-Uaxactiin area. An analysis of faunal relation- ships (Table 2) shows that the faunas of the rainforests of southern El Petén and lowland Alta Verapaz are closely related. The re- lationships between these two areas and the Tikal-Uaxactin area 218 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. TABLE 2.—INDEX OF FAUNAL RELATIONSHIPS BETWEEN SOUTHERN EL PETEN AND OTHER REGIONS. Lowland | Southern Tikal- setae Petén ager an es Lowland Alta: Verapaz. ie 5 .j.. ‘2 li" 2 os A ay At ! 1 pie Ve roe! a f "1 b } a J i ¥ I iy i ny # " nt ) veo i f a - cn i r 5, 7 4 hy hy ae nm a T18) 7 Vol. 10. 28. (Continued from inside of front cover) Conspecificity of two Sooke mice, Perognathus goldmani and P; ‘artus.. By . Raymond Hal! and Marilyn Bailey Ogilvie. Pp. 518-518, L map. Janu- ary 14, 1960. Records of harvest mice, Reithrodontomys, from Central America, with de- scription of a new subspecies from Nicaragua. By Sydney Anderson and J. Knox Jones, Jr. Pp. 519-529. January 14, 1960. : Small carnivores from San Josecito Cave (Pleistocene), Nuevo: Leén, ears By E. Raymond Hall. Pp. 581-538, 1 figure in text. January 14, 1960 Pleistocene pocket gophers from San Josecito Cave, Nuevo Leon, México. By Robert J. Russell. Pp. 539-548, 1 if age in text. January 14, 1960. Review of the insectivores of Korea. y J. Knox Jones, Jr., and David H. Johnson. Pp. 549-578. February 23, 1860. Speciation and_ evolution of the pygmy mice, genus Baimoys.. By Robert L. Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960. Index. Pp. 671-690 1. 2. 8. 10. Studies of birds killed in nocturnal migration. By Harrison. B. Tordoff and Robert M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 1956. Comparative breeding behavior-of Ammospiza caudacuta and A. maritima. By Glen E. Woolfenden, Pp. 45-75, 6 plates, 1 figure. December 20, 1956. The forest habitat of the University of Kansas Natural History Reservation, By Henry S. Fitch and Ronald R. McGregor: Pp. 77-127, 2 plates, 7 figures in text, 4 tables. December 31, 1956. Aspects of reproduction and development in the prairie vole (Microtus ochro- gaster). By Henry S. Fitch. Pp.)129+161, 8 figures in text, 4 tables. Decem- ber 19, 1957. Birds found on the Arctic slope of northern Alaska. By James W. Bee. Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958. The wood rats of Colorado: distribution and ecology. By Robert B. Finley, Jr. Pp. 218-552, 34 plates, 8 figures in text, 35 tables. November 7, 8. ioe ranges and movements of the eastern cottontail in Kansas. By Donald Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, 1959. weit history of the salamander, Aneides hardyi. By Richard F, Johnston and Gerhard A. Schad. Pp. 573-585. October 8, 1959. A new’ subspecies of lizard, Cnemidophorus sacki, from Michoacan, México. By William E, Duellman. Pp. 587-598; 2 figures in text. May 2, 1960. A taxonomic study of the middle American snake, Pituophis-deppei. By William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960. Index. Pp. 611-626. Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960. Vol. 12, 3. Functional morpnorery of three bats: Sumops, Myotis, ag ieee By Terry Vol. 13. ee 3. *4, S | A. Vaugha 153, 4 plates, 24 figures in text. July 8 The Sigrales of er “Amphibia: a review of the evidence, By Theodore H. Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, The baculum in microtine rodents. By Sydney Anderson. oe 181-216, 49 figures in text. “February 19, 1960. A new order of fishlike Amphibia from _ the Pennsylvanian of Kansas. -;By Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in text. May 2, 1960. : Natural histcry of the bell vireo. By Jon C.-Barlow. Pp. 241-296, 6 figures in text. March 7, 1962 Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C, Fox. Pp. 297-307, 6 figures-in text. May 21, Vertebrates from the barrier island of Tamaulipas, México. By Robert K. Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 309- 845, pls. 5-8, June 18, 1962. Teeth of Edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 fig- ures in text. October 1, 1962. More numbers will appear in volume 12. Five natural hype combinations in minnows. (Cyprinidae). By Frank B. Cross and W. L. Minckley. Pp. 1-18. June 1, 1960, A eee aal "study of the amphibians of the Isthmus of Tehuantepec, México. By William E, Duellman. Pp. 19-72; pls: 1- 8, 3 figures in text. August 16, 1960. A. new subspecies of the slider turtle (Pseudemys scripta) from Coahulia, Seren By John M. Legler. Pp. 73-84, pls. 9-12, 3 figures in text. August Autecology of the copperhead. By mene S. Fitch. Pp. 85-288, pls. 13-20, 26 f&eures in text. November 80, 1960. Occurrence of the garter snake, Thonaopkis sirtalis, in the Great Plains and Rocky Mountains. By Henry S. Fitch and T. Paul Maslin, Pp. 289-308, 4 figures in text. February 10, 1961. — Fishes of the Wakarusa river in Kansas. By James. E. Deacon and Artie L. Metcalf. Pp. 309-322, 1 figure in text. February 10, 1961. Geographic variation in the North American cyprinid fish, Hybopsis gracilis. By Leonard J. Olund and Frank B. Cross. Pp. 823-348, pls. 21-24, 2 figures in text. February 10, 1961. (Continued on outside of back cover) 9, 10. (Continued from inside of back cover) Decriptions of two species of frogs, genus Ptychohyla; studies of Ameri- can hylid frogs; V. By William E. Duellman. Pp. 349-357, pl. 25, 2 figures in text. April 27, 1961. Fish populations, following a_drought, in the Neosho and Marais des Cygnes rivers of Kansas. By James Everett Deacon. Pp. 359-427, pls. 26-30, 8 figs. August 11, 1961. Recent soft-shelled turtles of North America (family Trionychidae). By / Robert G. Webb. Pp, 429-611, pls. 31-54, 24 figures in text. February 16, 1962. { Index. Pp. 613-624. Volk. 14. 1. 2. 8. Neotropical bats from western México. By Sydney Anderson. Pp. 1-8. October 24, 1960. Geographic variation in the harvest mouse. Reithrodontomys megalotis, on the central Great Plains and in adjacent regions. By J. Knox Jones, Jr., and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961. Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson. Pp. 29-67, pls. 1 and 2, 3 figures in text. July 24, 1961. A new subspecies of the black myotis (bat) from eastern Mexico. By E. A Raymond Hall anad Ticul Alvarez. Pp, 69-72, 1 figure in text. December Vol. 15,)/ 1. 2. 3. 4, 29, 1961. North American yellow bats, ‘‘Dasypterus,” and a list of the named kinds of the genus Lasiurus Gray. ‘By E. Raymond Hall and J. Knox Jones, Jr. Pp. 73-98, 4 figures in text. _December 29, 1961. Natural history of the brush mouse (Peromyscus boylii) in Kansas with description of a new subspecies. By Charles A. Long. Pp. 99-111, 1 figure in text. December 29, 1961. i Taxonomic status of some mice of the Peromyscus boylii group in eastern Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 113- 120, 1 figure in text. December 29, 1961. A new subspecies of ground’ squirrel (Spermophilus spilosoma) from Ta- -maulipas, Mexico. By, Ticul Alvarez. Pp, 121-124. March 7, 1962. Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J. reer Jones, Jr., and Ticul Alvarez. Pp. 125-183, 1 figure in text. March 7, A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene, of Texas. By E. Raymond Hall and Walter W. Dalquest.. Pp. 135-138, 2 figures in text. April 30, 1962. . , A new subspecies of wood rat (Neotoma) from northeastern Mexico. By Ticul Alvarez. Pp, 139-1438. April 30, 1962. Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul ee and M. Raymond Lee. Pp. 145-159, 1 figure in text. May 18, A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, 1 figure in text. May 21, 1962. The mammals of Veracruz. By E. Raymond Hall anad Walter W. Dalquest. Pp. 165-362, 2 figures. May 20, 1963. The recent mammals of Tamaulipas, Méxicd, By Ticul Alvarez. Pp. 363- 478, 5 figures in text. May 20, 1963. More numbers will appear in volume 14. : The amphibians and reptiles of Michoacdn, México. By William E. Duell- man. Pp. 1-148, pls. 1-6, 11 figures in text. December 20, 1961. Some reptiles and amphibians from Korea. By Robert G, Webb, J. Knox Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962. A new species of frog (Genus Tomodactylus) from western México, By Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962. ; Type specimens of amphibians and reptiles in the Museum of Natural His- tory, the University of Kansas. By William E. Duellman and Barbara Berg. Pp. 183-204. October 26, 1962. Amphibians and Reptiles of the Rainforests of Southern El Petén, Guatemala. By William E. Duellman, Pp. 205-249, pls. 7-10, 6 figures in text. October 4, 1968. More numbers will appear in volume 15. UNIVERSITY OF KANSAS PUBLICATIONS | Ae MusEuM oF NaTuRAL History STB a oe Volume 15, No. 6, pp. 251-295, 9 figs. October 4, 1963 A Revisionof Snakes ofthe Genus Conophis (Family Colubridae, from Middle America) BY JOHN WELLMAN UNIVERSITY OF KANSAS LAWRENCE 1963 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Institutional libraries interested in publications exchange may obtain this series by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas. Copies for individuals, persons working in, a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of Kansas, Lawrence, Kansas. There is no provision for sale of this series by the University Library, which meets institutional requests, or by the Museum of Natural History, which meets the requests of individuals. However, when individuals request copies from the Museum, 25 cents should be included, for each separate number that is 100 pages or more in length, for the purpose of defraying the costs of wrapping and mailing. * An asterisk designates those numbers of which the Museum’s supply (not the Library’s supply ) is exhausted. Numbers published to date, in this series, are as follows: Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950. *Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text, April 9, 1948. Vol. 8. *1,..The avifauna of Micronesia, its origin, evolution, and distribution. By Rol- lin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951. *2, A quantitative study of the nocturnal migration of birds. By George H. Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951. 8. Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473- 580, 49 figures in text, 13 tables.. October 10, 1951. *4. Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and wares W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, Index. Pp. 651-681. £Vol. 4. (Complete) American weasels. By E. Raymond Hall, Pp. 1-466, 41 plates, 31 figures in text. December 27, 1951. { Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953. *Vol: 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. Durrant. Pp. 1-549, 91 figures in text, 80 tables. August 10, 1952. Vol. 7. Nos. I-15 and index. Pp. 1-651, 1952-1955. Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956. Vol. 9. *1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 figures in text. December 10, 1955. 2. Additional records and extension of ranges of mammals from Utah. By Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80. December 10, 1955. 8. A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rol- lin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955. 4. Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956. 5. The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6 figures in text. May 19, 1956. 6. Additional remains of the multituberculate genus Eucosmodon. By Robert W. Wilson. Pp, 117-123, 10 figures in text. May 19, 1956. 7. Mammals of Coahulia, Mexico. By Rollin H. Baker. Pp. 125-8385, 75 figures in text. June 15, 1956. c 8. Comments on the taxonomic status of Apodemus peninsulae, with. description of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 337-346, 1 figure in text, 1 table. August 15, 1956. 9. Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp. 847-351. August-15, 1956. ; 10. A new bat (Genus Leptonycteris) from Coahulia. By Howard J. Stains. Pp. 353-356. January 21, 1957. : : 11. A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico. By Robert J. Russell. Pp. 357-361. January 21, 1957. : 12. Geographic variation in the pocket_ gopher, Thomomys bottae, in Colorado. By Phillip M. Youngman... Pp. 363-387, 7 figures in text. February 21, 1958. 13. New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones, Jr.. Pp. 885-388. May 12, 1958. é re , 14. Pleistocene bats from San Josecito Cave, Nuevo Leén, México. By J. Knox Jones, Jr. Pp. 389-396. December 19, 1958. : 15. New subspecies of the rodent Baiomys from Central America. By Robert L. Packard. , Pp. 397-404. December 19, 1958. 16. Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405- 414, 1 figure in text, May 20, 1959. Z 17. Distribution, variation, and relationships of the montane vole, Microtus mon- tanus. By Sydney Anderson. © Pp. 415-511, 12 figures in text, 2 tables. August 1, 1959. (Continued on inside of back cover) < UNIVERSITY OF KANSAS PUBLICATIONS MuSsEUM OF NATURAL HISTORY Volume 15, No. 6, pp. 251-295, 9 figs. October 4, 1963 A Revision of Snakes of the Genus Conophis (Family Colubridae, from Middle America) JOHN WELLMAN UNIVERSITY OF KANSAS LAWRENCE 1963 Universtry oF KANnsAs PusBLicaTIONS, MusEUM OF NaTuRAL HIsToRY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 15, No. 6, pp. 251-295, 9 figs. Published October 4, 1963 UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY JEAN M. NEIBARGER, STATE PRINTER TOPEKA, KANSAS 1963 A Revision of Snakes of the Genus Conophis (Family Colubridae, from Middle America ) BY JOHN WELLMAN CONTENTS PAGE ENTER ODUGCTIO Ne 24 gop cece ve oes es dae Sea eae ee ene 253 ACKNOWLEDGMENTS 12. 3f05ishs chee cin ds aes feorer sh cree Te ee 254 MATERTAL STAND UMETHODS!1, crisluisicntss de eieichtle CAC Eon CR ena 254 GENUS) Conophis> Reters), boc... oS 26 50s G2) shoe tle ee eer eI 255 Key to the Species and Subspecies Analysis of Characters Scutellationy 305.255 aegis, Hehe a sh sett od oe Toso OA EOS ee 258 Sizeviand' !Proportions) gas, ach). oes siekee oe, Soe Ae ee 258 Golors Battermig sage siers cess tesus: fisces ths aaa tle Mr ee Ree te 260 SexualOimorp hiss 0... Asset ade de ws ein teres RUE Oe eee 260: GRETEN CAE S Eis Ne Rg fo NTS RS OO OT NT 262 GER EGHUS GUNNS Fes hee ee EO ETE eT ice 262 Giilineatusnlineatusiee 2 lcs caches See Oe ee 267 GaLTiNneatusnCOncOlOn Nee ec sos Scie Oe eee 270 GONE VETMANN) oxo. ee ee ee es eee 202 (Cire (iii lel Vo, Bal Bae ace dee tenes eMC ADC RNA Nr Pak omit, wit wid ode g 274 CACTUS Oe PE A te ee eee a ee eae 277 HIST be peas ered ean Ar MEMES YE MMORIUON UMA DOM BE os Ao UE ED 282 WDemtitiOn os ee cree ogee eee ae h tees tens ei negca ye CLES Rae 288 WETtODYA Gl acs ee ee tee te ee ao a ae ee 288 Hemipenes) «2 ct acdsion,: lo simi sacl ak dae. eee ka ee ee eee 289 Foodyiand., Feeding. 2.25..4.: «c7)-'+. ss, steaeie enh pce ee reer 289 Bifect of Poison. sere. eda oie sD niee e Oe OOO Ce ees 290 “LAXONOMIC) RELATIONSHIPS AND. LE VOLULION) 5 2). ee eee eee 291 SUNNGAR YW) SH. sid. 6 cube se agbeoneus tl axel oxchis) oho .iosle maw eWabere era ersie Rey see Ore NeR omen come 292 LITERATURE - CLrEn ©) ijcccciee fo.0 oi onethcwer eer o etens ogee Roe ererer ONO coher eerone eee 293 INTRODUCTION Need for a comprehensive systematic review of the snakes of the genus Conophis was pointed out by Stuart (1954a, b). Since these snakes appeared to be of zoogeographic importance in the Central American region, I undertook the review as set forth on the follow- ing pages. (253) 254 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist. ACKNOWLEDGMENTS For permission to examine specimens, and for information concerning speci- mens in their care, J am grateful to Mr. L. C. Battersby and Miss Alice G. C. Grandison, British Museum (Natural History); Mr. Charles M. Bogert and Dr. Richard G, Zweifel, American Museum of Natural History; Dr. Doris M. Cochran, United States National Museum; Prof. William B. Davis, Agricultural and Mechanical College of Texas; Dr. Josef Eiselt, Naturhistorisches Museums, Vienna; Prof. Norman Hartweg and Prof. Laurence C. Stuart, Museum of Zoology, University of Michigan; Dr. Robert F. Inger, Chicago Natural His- tory Museum; Dr. Alan E. Leviton, Califomia Academy of Sciences; Mr. Edmond V. Malnate, Academy of Natural Sciences, Philadelphia; Prof. George S. Myers, Stanford University Natural History Museum; Mr. Wilfred T. Neill, Ross Allen’s Reptile Institute; Mr. Neil D. Richmond, Camegie Museum; Dr. William J. Riemer, University of Florida Collections; Prof. Robert C. Stebbins, Museum of Vertebrate Zoology, University of California; Prof. Hobart M. Smith, University of Illinois Natural History Museum; and Dr. Emest E. Williams, Museum of Comparative Zoology, Harvard. Prof. William E. Duellman supplied invaluable information and guidance in my study. I am grateful to Prof. E. Raymond Hall for use of facilities of the Museum of Natural History and editorial assistance. I thank Prof, Laurence C. Stuart and Prof. Edward H. Taylor for information and sugges- tions. My own field experience in Middle America came as a result of assist- ing Professor Duellman in his own researches supported by a grant from the National Science Foundation (NSF-G 9827). For these things I am deeply grateful. Specimens that I have seen alive were collected by field companions Dale L. Hoyt and Jerome B. Tulecke. Finally, I am grateful to my wife, Margaret L. Wellman, for much help including typing much of the manu- script, MATERIALS AND METHODS Of the 325 specimens of the genus Conophis available to me, representing most of those in museum collections, scale counts were made in the usual manner on 309. Ventrals were counted following the system proposed by Dowling (1951:97-99); the anal plate was not included. The anteroposterior position of the place where reduction occurs in the number of the dorsal rows of scales is designated by citing the number of the ventral scale directly beneath that place. Measurements were taken to the nearest millimeter by means of a milli- meter stick. Body length is the distance from the tip of the snout to the posterior edge of the anal plate; tail length, from the latter point to the tip of the tail; and total length, the sum of the body plus tail. Descriptions of color are based on preserved specimens. Where descrip- tions of the color of living individuals are given, the data were taken from Kodachrome slides made available to me by William E. Duellman. Due to the transient nature of the longitudinal dark stripes in these snakes, no standard terminology has been devised, except that the posterior continuations of the stripes which on the head pass through the eye are termed lateral stripes; the posterior continuations of the median stripe of the head are termed dorso- MmpLe AMERICAN SNAKES (CONOPHIS ) 255 lateral stripes. A paravertebral stripe is one that is present on the scale-row on either side, of, but not including, the mid-dorsal (vertebral) scale-row. In order to reduce confusion in the discussion of variation, the numbers designating the rows of dorsal scales are written as Ist, 2nd, whereas the numbers designating the stripes are written as first, second. Except in three dried skeletons, teeth were counted on dentigerous bones in situ. Since teeth are often missing, the sockets were counted in order to obtain an accurate count. In accounts of the species and subspecies, the observed range of variation is followed by the mean in parentheses; in some instances the mean is fol- lowed by the standard deviation, also in parentheses. An example is 65-79 (70.6+38.93). Each synonymy includes all generic and specific combinations known to me that have been used for the genus, and, in addition, references to catalogues, checklists, and reports of collections. Localities of occurrence that are not plotted on the distribution maps are recorded in italic type under Specimens Examined. In the list of Specimens Examined the localities and specimens are listed in the following order: countries in alphabetical order; states or departments in alphabetical order in each country; localities in alphabetical order in each state or department; museum numbers in numerical order after the abbreviations of names of museums. When more than one specimen bears a single catalogue number, the number of specimens is given in parentheses following the museum cata- logue number. Specimens for which data are given only as to country or to state or department are listed first after the name of that political unit under “no specific locality.” The abbreviations for the museum collections are: AMNH American Museum of Natural History ANSP Academy of Natural Sciences of Philadelphia BMNH British Museum (Natural History) CAS California Academy of Sciences CNHM Chicago Natural History Museum ERA-WTN E. Ross Allen-Wilfred T. Neill, Ross Allen’s Reptile Institute KU University of Kansas Museum of Natural History MCZ Museum of Comparative Zoology, Harvard MVZ Museum of Vertebrate Zoology, University of California NMW Naturhistorisches Museums Wien, Vienna SU Stanford University Natural History Museum TCWC Texas Cooperative Wildlife Collection, Agricultural and Mechanical College of Texas UF University of Florida Collections UIMNH University of Illinois Museum of Natural History UMMZ University of Michigan Museum of Zoology USNM United States National Museum Family CoLUBRIDAE Subfamily Xenodontinae Genus Conophis Peters Tomodon (part) Duméril, Bibron and Duméril, Erpétologie Genérale, 7(pt.2):936, February 25, 1854 (lineatus and vittatus); Salvin, Proc. Zool. Soc. London, 28:455, 1860 (pulcher). 256 UNIVERSITY OF KANSAS Pusts., Mus. Nar. Hist. Psammophis (part), Giinther, Catalogue of Colubrine Snakes in the Col- lection of the British Museum, London, 1858:135 (lineatus). Conophis Peters, Monatsb, Akad. Wiss. Berlin, 1860:519-520, pl., fig. 3 ( vittatus); Cope, Proc. Acad. Nat. Sci. Philadelphia, 13:300, December 28, 1861 (lineatus concolor); Proc. Acad. Nat. Sci. Philadelphia, 18:318-319, February 20, 1867 (lineatus concolor); Proc. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876 (pulcher); Bocourt in Dumeéril, Bocourt and Mocquard, Mission Scientifique au Mexique et dans l’Amerique Centrale, 2:643-644, pl. 38, fig. 5, 1886 (lineatus lineatus); Cope, Proc. Amer. Philos. Soc., 23:489, October 28, 1886; Hoffmann, Klassen und Ordnungen des Thier-Reichs. Reptilien. Bd. 6, 3:1707, 1890; Cope, Trans. Amer. Philos. Soc., 18:207, April 15, 1895; Dunn, Bull. Antivenin Inst. Amer., 2(1):21, 24, April, 1928; Copeia, no. 4:214, December 31, 1937 (nevermanni). Tachymenis (in part), Garman, Bull. Essex Inst., 16:33, January 9, 1884 (vittatus and lineatus). Erythrolamprus (in part), Ditmars, Bull. Antivenin Inst, Amer., 2(2):27-29, June. Coniophanes (in part), Wettstein, Sitz. Akad. Wiss. Wien, mathem-naturw. kl. 143:37-38, 1934 (nevermanni). Historical summary.—In 1854 Duméril, Bibron and Duméril described and figured Tomodon lineatum from America. In 1860 Peters described and figured as a new genus and species, Conophis vittatus, based on a specimen that he had obtained from a dealer in Hamburg. The provenance of this specimen is not known, for it was discovered aboard a ship near the mouth of the Mississippi River. It was not until 1871 that Cope included lineatus in the genus Conophis. Cope (1861) proposed the name Conophis vittatus (nec Peters, 1860). Later (1900) he changed its name to Conophis lineaticeps. Early uncertainty of the relationships of the species lineatus caused Giinther (1858) to place it in the genus Psammophis. With the exception of Garman (1884a and 1884b) who placed lineatus in the genus Tachymenis, and Wett- stein (1934) who reported five specimens of Conophis nevermanni as Conio- phanes i. imperialis, all specimens reported after 1876 were placed in the genus Conophis. The only previous attempt to review the systematics of this genus was made by Smith (1941) who based his study primarily on specimens in the United States National Museum. He examined only 28 specimens, including none of one species (nevermanni). Description —Hemipenis slightly bifurcate having forked sulcus spermaticus, large spines near base, and smaller spines or papillae on flounces nearer apices; prediastemal maxillary teeth 8-12, subequal in length, and followed by short diastema and one enlarged fang or two; fangs grooved, only one functional at any one time, unless snake is in process of shedding teeth; teeth 6-10 on palatine, 15 to 19 on pterygoid, 15 to 21 on dentary; teeth on dentary decreasing in size posteriorly; large parotid (venom) gland on either side of head in temporal region; head shields of basically unmodified colubrid type excepting decurved rostral; rostral concave below and therein modified for burrowing; internasals and prefrontals paired; nasals divided; loreal single; preocular one, rarely two; postoculars, two; supralabials, 7-8, 3rd and 4th or 4th and 5th under eye; infralabials, 8-11, usually 9 or 10; temporals, normally 1 plus 2 plus 3; chin-shields subequal in length; ventrals, 149-183, rounded and overlapping; caudals, 55-89, paired and imbricate; anal divided; dorsal Mpie AMERICAN SNAKES (CONOPHIS ) 257 scales smooth and in 19 rows at mid-body with no apical pits or keels; scale reduction normally involving fusion of 3rd and 4th rows, resulting in 17 scale-rows near tail; tail length more than 20 per cent of body length; maximum total length exceeding 1.1 meters; dorsal color pattern consisting of dark stripes, or no darkening, on paler ground-color; ventral surfaces im- maculate pale yellowish or white, except on specimens having single lateral dark spots on some or all ventrals; pupil round; diurnal or crepuscular; feeding primarily on small lizards, sometimes on small mammals or other snakes. Distribution—Semiarid regions of southern México and Central America as far south as Costa Rica. Key TO THE SPECIES AND SUBSPECIES Although many juveniles differ greatly in general coloration from the adults, both the juveniles and the adults of any species or subspecies can be identified from the following key; juveniles differ from adults in extent and intensity of dark pigmentation but not in rows of scales involved. 1. Seven supralabials (38rd and 4th below orbit); 3 to 8 dark stripes along? body ben Be) SEER OS OS ee ein eee ee eee 2 Eight supralabials (4th and 5th below orbit); unstriped or with more than 4 dark stripes along body, or dark with 2 or 4 pale stripes... .. 3 2. Dark stripes involving no more than one longitudinal SCALETOW? Hato re srende ek kt C. lineatus lineatus (part), p. 267 Dark stripes involving at least two adjacent scale-rows .. .C. vittatus, p. 277 8. Supralabials having black borders above; head and body generally black with 2 or 4 white lines running length of BAYA ke sk nly sete eos Wen ak Ne oer eee C. nevermanni, p. 272 Supralabials immaculate or having dark borders below; head and body usually pale with dark stripes, or without stripes .................. 4 4, Lateral dark stripe through eye involving upper half of second scale- row; dark stripe on paravertebral row, at least DOSECTIOELY, oo ea see ik rege Sane nee eed C. pulcher, p. 274 Lateral dark stripe becoming indistinct on body, or restricted to 4th or 8rd and 4th rows anteriorly, not involving 2nd scale-row on anterior 1% of body (an auxiliary lateral stripe sometimes present involving 2nd row:); no’ paravertebral’ stripes. 4. <... 2-05 se ie ae eee 5 5. Stripes disappearing posteriorly (except for small ee of pigment on scale-row 4 or 7); lst scale-row unpigmented .. .C. lineatus concolor, p. 270 Stripes present posteriorly; lst scale-row pigmented............... 6 6. Lateral stripes narrow on nape, restricted to 4th scale-row on |Srove tes ne tt ee ena, OE ORO, 2 C. lineatus lineatus (part), p. 267 Lateral stripes involving 38rd and 4th rows, at least on WAPO de ey hee tetas Mee atsis he ue Deena C. lineatus dunni, p. 262 Analysis of Characters Characters showing inter-specific and intra-specific variation and that have a wide range of variation were analyzed statistically, when possible, in order to determine extent of variation. One character (see table 3) was analyzed for sexual dimorphism, and for it the coefficient of difference is also given. The statistical terms and formulae have been adopted from Mayr, Linsley and Usinger (1953). Dorsal head shields varied individually and were of no taxonomic importance. Osteological and hemipeneal characters did not show enough variation to be considered here. 258 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist. Scutellation Labials, dorsals, ventrals, and subcaudals were the most useful scales. Labials—All species usually have eight supralabials except C. vittatus, which has seven. The only other population having a relatively high frequency of occurrence of seven supralabials is C. 1. lineatus. In specimens having eight supralabials, the fourth and fifth enter the orbit; in specimens having seven supralabials, the third and fourth enter the orbit (the second and third are fused). Usually there are ten infralabials, sometimes nine or eleven; speci- mens having seven supralabials usually have nine infralabials, sometimes eight, rarely ten. Dorsals—Although there is no variation in the number of rows of dorsal scales, there is some in the method of scale reduction. There are 19 rows of dorsal scales from close behind the head to about midway on the body where two rows are lost, leaving 17 rows from there to near the base of the tail. This reduction is accomplished by fusion of the scales of the 3rd and 4th rows or sometimes by the dropping out of the 3rd row. The place at which reduction occurs in number of dorsal scales in relation to the ventral (scale) directly below is highly variable and of little taxonomic importance (table 1). TABLE 1.—VARIATION IN THE PLACE OF DorsAL SCALE REDUCTION IN CoNoOPHIS. Z 8 « § g Pie Taxon ne Ee Ps 58 — 5. SH f=) 3 a 3 5 SF co En = s aA em es 3 & ° as a = A RD 6) LxCOnCOLOM a Acide rok: 45 89-114 102.5 yee 0.83 5.43 VE QUINN oe sok noes 36 91-111 102.1 4.59 0.77 4.50 lslaneatusee a..c o5: 26 91-107 100.2 3.59 O72 3.58 nevermannt........ 6 84-— 97 93.2 4.71 1.92 5.05 DUICKEnaeenere Hae 26 94-119 104.6 4.90 0.96 4.68 0.16 6.45 VULOLUS Ee Te ee Sc 170 84-118 102.3 6.60 Ventrals—The number of ventral scutes varies from 149-183, and shows no significant variation in the means (table 2). Subcaudals—The number of subcaudal scutes varies from 55 to 89. In some populations there is no overlap in the range of variation of males and females, The total variation and sexual dimorphism are analyzed in table 3. Size and Proportions Although considerable variation in size is observable, little taxonomic use is made of size since sufficient series are not available to determine age classes. The subspecies attaining the largest size is C. lineatus concolor; all others are smaller and of about the same size and proportions. The longest specimen, a male of C. l. concolor, has a body length of 893 mm., a tail length of 274 mm., and a total length of 1167 mm. Miwp.e AMERICAN SNAKES (CONOPHIS ) 259 TABLE 2.—VARIATION IN THE NUMBER OF VENTRALS IN CONOPHIS. 8 nm — ws 8 s re Taxon oe go Ps a 5 38 g SE er Ew = = aQ a eS = 3 v gs Ss 5 Z om = oD) ™ Oo PE COnCOLOr or) NETS TE: 45 158-170 163.7 1.56 0.23 0.95 Lidunntn.8haet8 «. 36 | 159-178 | 167.2 4.56 0.76 2:72 Lstaneatus’.- 55. es: 26 | 157-169 | 163.5 3.59 0.72 2.20 nevermannt........ 6 173-183 176.5 4.00 1.63 2.27 ALIC LER Mea eT ene 26 149-180 169.5 5.31 1.04 3.13 UUULALUSAE A. Hee ral 149-180 163.7 6.33 0.15 3.87 TABLE 3.—SEXUAL DIMORPHISM AS INDICATED BY VARIATION IN THE NUMBER OF SUBCAUDALS IN CONOPHIS. S r= g a] & 3/ 8 ae o — & =] o Taxon cE 78 a) z a ge US) Ihe a5 | = | ‘Ss | 5A gu | a | sQO| & | &s| €s te i ton bilo (ies anal besa Sa n| Ss eo} = a wm 1 'e) lineatus concolor...... fot 22 68-74 | 70.3 | 2.14 | 0.46 | 3.04 sy ibe °) 16 56-65 | 61.8 | 2.18 | 0.55 | 3.53 lineatus dunni....... fof 14 67-80 | 74.5 | 3.86 | 1.03 | 5.18 ae 9 | 16 | 60-72 | 67.1 | 3.91 | 0.97 | 5.82 | lineatus lineatus...... fot ial 67-73 | 69.8 | 6.17 | 1.85 | 8.84 aveo Q 9 60-66 | 62.4 | 6.17 | 2.06 | 9.89 NeverMaNNt.......... fot 3 82=89 | 85,430 seveceercllices eters | etereeacke Q 2 71276 73. Be lae eae ae A Pie DULChET Ee See eee fot 7 10=79" V4.3 eel er ea et9) s 0.9 Q 11 65-71 | 68.2 | 3.42 | 1.08 | 5.01 ULLLOLUSH, eet ee o 95 59-76 | 67.8 | 3.33 | 0.34 | 4.91 a ihe ? 58 | 55-66 | 60.0 | 2.75 | 0.36 | 4.58 260 UNnIversiTy OF Kansas Pusts., Mus. Nat. Hist. Color Pattern This is the primary feature used to separate species and subspecies in this genus. The color pattern consists of three black or deep brown stripes on the dorsal part of the head, one mid-dorsally, and one on each side of the head passing through the eye. On the body, there are usually dark longitudinal stripes on a pale tan or white background. There may be as few as three in vittatus, and as many as 13 in I. dunni; except that there is none in C. l. concolor. There are two pairs of primary dark stripes. The first is the body stripe that is the posterior extension of the stripe which on the head passes through the eye and is termed the lateral stripe. The other primary stripe is the posterior continuation of the mid-dorsal head stripe. Usually it is split into two dorsolateral stripes on the body. Stripes may be present on the scale-row to either side of the primary stripe. These stripes are usually dark brown or black and are the secondary stripes. Finally, additional stripes may be present that are paler brown and bear no direct relationship to the primary stripes. These are auxiliary stripes. Every stripe originates either as broad continuous stripe or as a row of spots or dashes, forming a discontinuous stripe, which in some specimens becomes continuous posteriorly. The stripes are usually black or deep brown, although auxiliary stripes are sometimes paler. The dorsal ground color is pale brown, tan, olive, or white; usually the ground color is palest ventrally and darkest dorsally. In some specimens of Conophis the lateral tips of the ventrals are spotted, one spot on each end of each ventral. Otherwise, the ventrals are immaculate white. In some species there is considerable ontogenetic change in color pattern, although the juveniles bear the basic color characteristics of the adults. For example, juveniles of the sympatric species C. lineatus dunni and C. pulcher can be separated on the basis of which scale-rows are darkly pigmented. C. 1. dunni has eight stripes in juveniles and as many as 13 in adults. Juveniles show a greater contrast between the black stripes and the pale ground color than do adults. With increased age (size) the stripes in some populations become paler and are split; simultaneously the ground color becomes darker. Sexual Dimorphism Sexual dimorphism is evident in all species and subspecies of Conophis. Differences always exist in the number of subcaudals and in the tail/body ratio; males have more subcaudals and rela- tively longer tails than do females (table 3). Otherwise, there is little sexual dimorphism in these snakes. Males and females cannot be differentiated by any feature of coloration. Formulation of a biological concept of the species as defined by Mayr (1942) is difficult when most of the data primarily relied upon are from preserved specimens. Nevertheless, a total view of variation was attempted so that differences within and between populations could be recognized. Differences, between popula- 261 oo ESOS SC Y 2 Soe ais I 3 Se H MippLE AMERICAN SNAKES (CONOPHIS ) tions, that seem to be part of a continuous or internal cline (Huxley, G 1942) are not used for characterizing subspecies. sie Ome SPOS eS Ge ee) ne SS 8a cad ae Qe eel lies =pno) — (ome, Go 298 Pea? BO8 8087 Gag A-~ VVMEALHDE acd S552 2 Fad gs & mee setae. OX 3 09 2 p> ee g0 8 eG ees ~ 338 S5ORSAS no} Re Q f8ga~5eSon 5 Gt on Oe ty fe) o 8 ae) 7 Oe => 6H el HO) a, Bad 479 = s Ps egedgnchb< ayers Ori Le RS ore Bs pS s S88 a0 7535 6885 oO ‘ oe eeTs. SSH CSRS gop RUSS cg eS 8h 8255 ¢ o iS 3 Beds OMe ete S2AZ, Seer ¥ Zi epi Sons ro) S288 os* FEL Tamora om lm} = OvonN =O. . << og p S Baud a "BD Be Aa ea HO aie CBR ENTAN OSES BERS? ON Ze os g OA ga Bo Se — Gy = oO ag "risa sgee BUNS 8a 3 oa. SBaSCoR~ gi 6 =} SH OMss eens eS - HS Neioa er See] Ses rhe eer eek ey aoe ee bee oe asa nN bd o ES > get aso OOS Z eSeoOuon Bs SPP SHS Bagel enato 262 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist. Conophis lineatus (Duméril, Bibron and Duméril ) Tomodon lineatum (in part) Duméril, Bibron and Duméril, Erpétologie Genérale, 7( pt. 2):936-938, February 25, 1854. Diagnosis—No dark pigmentation posterior to nape; lateral dark stripe anteriorly passing through eye and posteriorly involving 4th or 8rd and 4th scale-rows only; first scale-row darkly pigmented; no paravertebral dark stripe; six to thirteen (or no) dark stripes at mid-body; usually eight (sometimes seven) supralabials immaculate white or having dark ventral margins. Variation.—The variation in this species is discussed more completely in the descriptions of the subspecies. One hundred and seven specimens have 157 to 178 (164.8) ventrals. Eighty-eight of these snakes having complete tails have 56 to 80 (68.0) subcaudals; the number of ventrals plus subcaudals varies from 222 to 247 (238.5) in 87 of these. On 107 specimens the reduc- tion from 19 to 17 dorsal scale-rows takes place between ventrals 89 and 114 (101.8). Sexual dimorphism is evident in the number of subcaudals; there are, on the average, fewer subcaudals in females than in males of each sub- species. The largest specimen is a male C. I. concolor (USNM 46345) from Chichén Itz4, Yucatan, México, having a body length of 893 mm., a tail length of 274 mm. and a total length of 1167 mm. The smallest is a juvenile C. 1. dunni (MCZ 49749) from Tegucigalpa, Honduras, having a body length of 162 mm., a tail length of 51 mm. and a total length of 213 mm. The greatest variation is in coloration. Dark color, or lack thereof has been used to separate the subspecies of C. lineatus. The ground-color is pale brown, pale olive or white, either with no stripes on the body or with eight to thirteen dark stripes at mid-body. Specimens having dark stripes on the body always have black or dark brown pigmentation on the first, 4th and 7th dorsal scale-rows. In some there is dark pigmentation on the 2nd, 8rd, 8th and 10th rows of scales. The stripes appear on the nape or farther pos- teriorly, usually on the anterior third of the body, either as a series of spots or dashes that form a continuous stripe farther posteriorly or as a continuous stripe. The ventrals usually have more or less conspicuous dark spots laterally on those specimens having dark stripes present on the dorsum; spots are absent on all specimens having no dorsal stripes and on some specimens having dorsal stripes. Except for the dark lateral spots (when present) the ventrals are immaculate white. Usually the dorsal ground-color is pale tan, especially on the striped forms. The ground-color is usually palest on the lower dorsal scale rows and darkest dorsally. Three populations are separable as subspecies; one has no stripes on the body and occurs in the Yucatan Peninsula. The other two have stripes on the dorsum and vary clinally in coloration from the north (Veracruz, México) to south (Costa Rica) (Fig. 2). Reasons for separating these widespread, vari- able snakes into two subspecies are that they are discontinuous in distribution (the population in Veracruz is disjunct from the one that extends from Guate- mala to Costa Rica), and that these populations have distinctly different color patterns. Conophis lineatus dunni Smith Psammophis lineatus, Giinther, Catalogue of Colubrine Snakes in the Col- lection of the British Museum, p. 185, 1858. Muppie AMERICAN SNAKES (CONOPHIS ) 263 A- C.1. concolor @ — ©.1. dunni He —C.1 lineatus SCALE OF MILES Fic. 2. Selected locality records for the subspecies of Conophis lineatus. Conophis lineatus, Cope, 3rd Ann. Rept. Peabody Acad. Sci., p. 82, 1871; Proc. Acad. Nat. Sci. Philadelphia, 23:204, October 24, 1871; Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; Bull. U. S. Natl. Mus., 32:77, 1887; Giinther, Biologia Centrali-Americana, p. 165, March, 1895; Boulenger, Catalogue of the Snakes in the British Museum (Natural History ), 3:122-123, 1896; Werner, Arch. Naturges., 90, abt. A, 12:143, 1925; Schmidt, Zool. Ser. Field Mus. Nat Hist., 12:199-200, November 21, 1928; Amaral, Mem. Inst. Butantan, 4:212, 1929; Werner, Zool. Jahrb., 57:184, 1929; Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 292:5, June 29, 1934; Dunn, Copeia, no. 4:214, December 31, 1937. Conophis lineatus similis Smith, Journ. Washington Acad. Sci., 31:123-124, March 15, 1941 (Type.—United States National Museum, No. 79963; type locality—Managua, Nicaragua; nec Bocourt in Duméril, Bibron and Mocquard, Mission Scientifique au Mexique et dans lAmerique Centrale, 2:647-648, 1886); Cochran, Bull. U. S. Natl. Mus., 220:167, 1961. Conophis lineatus dunni Smith, Proc. U. S. Natl. Mus. 92:394-395, No- vember 5, 1942; Savage, Trans. Kansas Acad. Sci., 50:483-486, Decem- ber 31, 1949; Taylor, Univ. Kansas Sci. Bull., 34(pt. 1):145, October 1, 1951; Neill and Allen, Publ. Res. Div. Ross Allen’s Rept. Inst., 2:56, November 10, 1959; Herpetologica, 16:146-148, fig. 2, September 23, 1960. Conophis pulcher pulcher, Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, 69:79, June 12, 1948; Contr. Lab. Vert. Biol. Univ. Michigan, 45:24, 264 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. May, 1950; Contr. Lab. Vert. Biol. Univ. Michigan, 49:14, August, 1951; Contr. Lab. Vert. Biol. Univ. Michigan, 65:19-20 (part), March, 1954. Conophis pulcher plagosus, Mertens, Zool. Anz., 148:98, February, 1952; Abhand. Senken. Naturw. Gesell., 487:61-62, December 1, 1952. Conophis lineatus nevermanni, Taylor, Univ. Kansas Sci. Bull., 37(pt. 1): 563-565, fig. 16, October 15, 1955. Type.—United States National Museum, no. 79963, obtained by Lt. H. C. Kellers. Type locality: Managua, Nicaragua. There are also three para-~ types; one a topotype (USNM 79964), one from “Nicaragua” (USNM 25237), and one from Esparta, Costa Rica (USNM 87758). Diagnosis—Lateral dark stripe anteriorly passing through eye and poste- riorly involving 3rd and 4th scale-rows; Ist scale-row darkly pigmented; no paravertebral dark stripe, although vertebral row sometimes darkly pigmented; six to thirteen stripes at mid-body; eight supralabials immaculaate or having dark ventral margins. Variation.—Thirty-six specimens have 159 to 178 (167.2+4.56) ventrals. Thirty of these snakes having complete tails have 60 to 80 (70.55.36) subcaudals; the number of ventrals plus subcaudals varies from 224 to 247 (237.6). In 36 specimens the reduction from 19 to 17 dorsal scales takes place between ventrals 91 and 111 (102.1+ 4.59). Sexual dimorphism is evident in the number of subcaudals; 16 females have 60 to 72 (67.1), and 14 males have 67 to 80 (74.5) subcaudals. The largest specimen (ERA-WTN BH-300) is a female from Augustine, British Honduras, having a body length of 732 mm., a tail length of 183 mm. and a total length of 915 mm. A juvenile (MCZ 49794) from Tegucigalpa, Honduras, has a body length of 162 mm., a tail length of 51 mm. and a total length of 213 mm. The greatest variation is in coloration. The ground-color is pale brown or white with dark stripes of black or deep brown present dorsally and laterally. Some specimens from Costa Rica have as many as 13 dark stripes at mid- body (fig. 1, C). In these snakes the first row of dorsal scales bears a series of large, slightly elongated, dark spots; on the 2nd row a narrow dark brown stripe on the middle of the scales; on the 3rd a black stripe on the dorsal one- third to one-half of the scales; on the 4th and the 7th rows black stripes on the medial half of the scales of each row; on the 8th and 10th (vertebral) rows dark brown stripes on the medial third of the scales of each row. A specimen from Guatemala (UMMZ 107339) shows the greatest reduction of stripes and dark pigmentation (fig. 1, A); it has only eight stripes at mid- body: on the first row of dorsal scales a discontinuous stripe is formed by a series of dashes; the 3rd row bears a series of small black spots near the base and tip of each scale; the 4th and 7th rows bear continuous black stripes on the medial third to fourth of the scales of each row; the 8th row has extremely small dark spots near the tips of some scales. The primary stripes, chacteristic of the species lineatus, are those on the Ist, 4th and 7th rows of dorsal scales; these are the most prominent stripes. In some specimens these primary stripes begin as spots or dashes on the nape and become continuous stripes posteriorly; in others they are continuous for. the length of the body. The stripe on the Ist row is most variable; usually it consists of only a discontinuous series of dashes for most of its length. The secondary stripes are those on the 3rd and 8th rows; of these, only the one on the 3rd scale-row is present on the nape. The stripe on the 3rd row in. MippLE AMERICAN SNAKES (CONOPHIS ) 265 combination with the dark stripe on the 4th row is the posterior continuation of the dark stripe that on the head passes through the eye; this stripe is characteristic of C. lineatus dunni. Both secondary stripes usually begin anteriorly as a series of spots or dashes and become continuous stripes posteriorly; occasionally near the base of the tail they fuse with the primary stripes on the 4th and 7th rows. In some specimens in Costa Rica indistinct stripes are present on the 10th (posteriorly the 9th) rows, and in some specimens in Honduras, Nicaragua, and Costa Rica similar indistinct stripes are present on the 2nd row. Usually there are more or less conspicuous dark spots laterally on the ventrals, but in some specimens there are no spots. Except for the dark lateral spots (when present) the ventrals are immaculate white. The dorsal ground-color is a pale brown or brownish white in preserved specimens on the Ist, 2nd, 38rd and 4th rows of scales where dark stripes or spots are not present, The ground-color of the dorsum between the 5th rows on each side is a somewhat darker shade of pale to medium brown. Never is more than the lower one-third of each of the supralabials brown. In many specimens little or no brown is present on the lower margins of these scales. Some of the specimens having brown on the supralabials also have dusky markings of tan or gray on the chin and infralabials. Specimens from the northern part of the range (Guatemala) less frequently have dark chins and supralabials than do specimens from the southern part of the range (Costa Rica). There is, nevertheless, at any one locality considerable varia- tion in the amount of dark pigmentation present on the chin and supralabials, thereby indicating that the slight geographic trend in this character is not significant. Probably the most common pattern of dorsal coloration consists of eight or ten dark stripes (fig. 1, B). In snakes having this pattern the stripes on the Ist, 3rd, 4th and 7th rows are always present and prominent, although those on the lst and 3rd rows sometimes are present as discontinuous rows of dashes, The ground-color from the venter to the 7th row is usually pale brown, and that dorsally between the 7th rows on each side is usually a darker, medium brown. A series of spots or dashes or a continuous stripe is some- times present on the 8th row of scales. Snakes having a larger number of dark stripes and more dark pigmentation occur in the southern part of the range. There seems to be a cline from paler snakes having fewer stripes in the north to darker snakes in the south. AWM a Fic. 8. Patterns of dorsal coloration at mid-body of juveniles of two sym- patric species of Conophis. > A exoccipitat y = ae Lin supraoccipital prefronta! A postfrontal > aa __—~s € SS Ss BR Ny eZ ae e@ ; columaita ZG ar aN °. LE nasal “= cg ea squamosal iy WH vomer frontal j premaxillary KL | RS f i, ip tl = f_- columella ia ~OX\, = = ; pm Co SS. aS FZ )"\ | Za p basioccipital Legh , C Pe SZ exoccipital septomoxillary \/! ¥ porasphenoid G prefrontal henoid prootic postfrontal Fic. 6. The skull, lacking dentigerous bones, of Conophis lineatus concolor (UMMZ S-788) showing (A) dorsal, (B) lateral, and (C) ventral views. 38. 8—5936 284 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. vomer is approximately U-shaped, when viewed from below. It has no posterior process and does not articulate with the parasphenoid; there is a sizeable gap between the two bones. The septo-maxillary has a lateral process that terminally is directed slightly anteriorly. Cranium and associated elements.—The frontal is almost three times as long as it is wide; it is flat above with an emarginate dorsolateral margin that forms the upper limit of the optic capsule. Ventrally the frontal is concave and forms the median limits of the optic cavity. Farther ventrally the frontal joins with the parasphenoid, which at this place forms the ventral extent of the skull, and together with the basisphenoid forms the ventral part of the posterior three-fourths of the skull. In ventral aspect, the parasphenoid is a long, thin bone, slightly expanded anteriorly. It forms the anterior floor of the optic foramen; whereas the frontal forms the anterior roof of the same opening, The frontal and its septo-maxillary process surround the olfactory fenestra. The prefrontal articulates with the anterolateral process of the frontal. The posterior surface of the prefrontal forms the anterior wall of the orbit of the eye. The articulating surface upon which the median process of the maxillary bone rests is situated ventrally. The anterior dorsal surface of the prefrontal, together with the anterolateral edge of the frontal, extends slightly over the nasal cavity, affording some degree of protection for the contained organs and forming the posterior border of the cavity. A small nasal process also extends anteriorly from the ventrolateral surface of the prefrontal. The orbital-nasalis foramen is located in the anterior surface of the prefrontal. The parietals are fused into one large bone that forms the roof and sides of the middle part of the cranial cavity. From its suture with the frontal, the dorsal surface of the parietal is relatively flat in the area bounded laterally by the parietal crests, which extend posteromedially from the anterolateral comers of the bone and converge medially at a point near its posterior margin. A slight posterior extension of the parietal crests forms the supratemporal crest, which is present on the posterior part of the parietal and on the anterior part of the supraoccipital. The postfrontals are attached to the anterolateral processes of the parietal. Together the anterior surfaces of these two bones form the posterior rim of the orbit of the eye. The post- frontal extends laterally and ventrally and has a terminal extension that projects anterolaterally. Im an articulated skull the transpalatine articulates with the ventrolateral articulating surface of the postfrontal. Anteromedially, the parietal forms the roof and posterior margin of the optic foramen. The basisphenoid, which is fused with the parasphenoid, also forms a small part of the posteroventral margin of the optic foramen. The basisphenoid forms the floor of the middle part of the cranial cavity and the ventromedial down- pouching that contains the pituitary body. Posterolateral to the parietal and dorsal to the posterior part of the basisphenoid is the prootic. Laterally this bone is deeply emarginate; posteriorly it forms a large part of the otic notch, through which the columella passes. The columella is a Jong, thin bony rod that terminates posteriorly in cartilage. It is the cartilagenous part of the columella that connects with the external sound detecting mechanism. There are several foramina on the lateral surface of the prootic. On the anterolateral surface of the prootic, branches of the trigeminal nerve pass through three foramina whereas the facial nerve passes through the single posterior foramen MippLE AMERICAN SNAKES (CONOPHIS ) 285 near the otic notch. The squamosal is attached dorsoventrally to the posterior part of the parietal and to the lateral part of the prootic. At this place of attachment there is on the prootic a relatively heavy crest that forms a rather broad articulating base. The squamosal is long, flat, and curves slightly in a dorsal direction throughout its length; it becomes thinner and narrower posteriorly. The posterior third of the squamosal forms a broad base by means of which the squamosal articulates with the quadrate. The columella and the squamosal extend posteriorly beyond the limits of the braincase. Posteriorly the skull consists of four bones: an unpaired median dorsal supraoc- cipital, an unpaired median ventral basioccipital and two lateral exoccipitals. The basioccipital does not have noticeable pterygoid processes, but is rather smooth ventrally and only slightly emarginate on its posterolateral margins. Posteriorly, this bone forms the ventral part of the occipital condyle. The rest of the condyle, on each side, is formed by the exoccipitals. The exoc- cipitals also form part of the base to which the squamosal is attached. The exoccipitals extend around the sides of the foramen magnum and meet dorsally. Each exoccipital also forms the posterior part of the otic notch, which traverses the exoccipital. The exoccipitals bear moderate occipital crests that extend pterygoid palatine . A trans -palatine maxillary trans- palatine maxillary pterygoid palatine Fic. 7. The maxillo-palatal-pterygoid arch of Conophis lineatus concolor (UMMZ S-788) showing (A) dorsal, (B) lateral, and (C) ventral views. 3. Teeth shown by means of broken lines were represented only by their sockets. 286 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. posterolaterally across the supraoccipital as branches from the supratemporal crest. The supraoccipital also has a medial crest that extends a short distance posteriorly from the supratemporal crest onto the exoccipitals at their dorsal line of fusion. Maxillo-palatal-pterygoid arch—In an articulated skull, the anterior edge of the maxillary is immediately posterior to the lateral tip of the premaxillary (fig. 7). The maxillary is curved moderately laterally and is not robust at its tip, but it becomes heavier about one-third of its length posteriorly. A dorsomedian process begins at about one-third of its distance from the anterior end; the prefrontal articulates with this process. The process is broad and almost flat, except that at its medial end, an elongate, rounded knob extends ventrally. The dorsomedian process of the maxillary extends toward, but does not meet, a lateral process from the palatine. The maxillary teeth are set in sockets on the ventral surface of the bone. Just posterior to the level of the last prediastemal tooth is the median trans-palatine process that articulates with the anteromedian part of the trans-palatine. Immediately posterior to this process, the maxillary narrows slightly; then it broadens to form an obliquely oriented knob. The posteroventral surface of the posterior knob of the maxillary bears one or two enlarged maxillary teeth. (These teeth are discussed further in the section on Dentition.) The anterolateral part of the trans-palatine articulates with the dorsal surface of the posterior knob of the maxillary. Toward the middle of its length, the trans-palatine narrows considerably; then it broadens again and articulates with the pterygoid. The palatine is slightly rounded at its anterior end, which extends anteriorly to the posterior margin of the vacuity containing Jacobson’s organ. The palatine extends posteriorly to the trans-palatine process of the maxilla, where the palatine articulates with the pterygoid. A posterior pterygoid process from the palatine projects posteromedially from the end of the palatine and overlaps the anterior end of the pterygoid. Just less than one-half the distance from the anterior end of the palatine, there is a lateral process that curves ventrolaterally forming a blunt tip posteriorly. Slightly more pos- teriorly and on the medial side of the palatine, is a medial sphenoid process, which is thin, rather broad, and curves ventromedially; ultimately it comes to lie near the anterior part of the parasphenoid. The palatine teeth are set in shallow sockets on the ventral edge of the bone. Of the bones of the maxillo-palatal-pterygoid arch, those on the pterygoid extend farthest pos- teriorly. The pterygoid is broad medially and posteriorly, although pointed at its posterior tip. The trans-palatine articulates in a broad line at about one-third of the distance along the lateral margin of the pterygoid. Immedi- ately posterior to this articulation, there is a median ridge on the pterygoid; lateral to the pterygoid ridge is an abrupt hollow, the pterygoid groove. Posteromedially, this groove becomes gradually more shallow and disappears. The dorsal surface of the pterygoid is rounded anteriorly and somewhat flattened posteriorly, whereas the ventral surface is gently rounded along its length, except that there is a high median crest. The pterygoid teeth are situated in shallow sockets along this crest. The teeth diminish in size posteriorly. Mandible.—The dentary (fig. 8) is compressed laterally and rounded be- low. The teeth, which are longest about one-third of the way from the anterior end of the dentary, are set in sockets on the medial side of the bone. MippLE AMERICAN SNAKES (CONOPHIS ) 287 dentary articular B articular Fic. 8. The left mandible and associated quadrate of Conophis lineatus concolor (UMMZ S-788) showing (A) lateral and (B) medial views. <3. Teeth shown by means of broken lines were represented only by their sockets. angular surangular The posterior half of the dentary overlies the fused surangular-prearticular part of the articular. Ventrally, the posterior part of the dentary underlies the splenial, which is set in a median trench within the dentary. Near the com- mon suture of the dentary and the splenial is the large inferior alveolar foramen; completely within the splenial and ventral to the inferior alveolar foramen is the anterior mylohyoid foramen. Posterior to the splenial and also forming a part of the ventral surface of the mandible is the wedge-shaped angular, which lies directly beneath the fused surangular-prearticular. As has been implied, the articular, the surangular, and the prearticular are fused. The prearticular part of this bone forms a part of Meckel’s canal. In the surangular part, immediately posterior to the end of the dentary, is the large surangular foramen. Lying in a longitudinal axis along the medial surface of the articular is a high crest, dorsal to which is a deep hollow. The lateral wall of the articular above this hollow is thin and rounded dorsally; the ven- tral surface is uniformly round and slightly curved dorsally, except that it ends with a short tympanic crest, which projects beyond the articulation with the quadrate. Where the quadrate articulates with the dorsolateral surface of the posterior portion of the squamosal, the former is broad and has a high mid- lateral crest, which extends about one-third of the distance down the quadrate before disappearing. The columellar process (the place of fusion of the 288 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. columella) is about two-thirds of the way down the medial surface of the quadrate. Ventrally the quadrate has a narrow neck dorsal to its articulation with the articular. The articulation is formed by two lateral flanges of the quadrate that fit over a medial ridge formed by the articular. Dentition Teeth on the maxillary and pterygoid decrease in size posteriorly, whereas those of the dentary do likewise except for the first one or two that are usually slightly smaller than those immediately posterior. The palatine teeth are subequal in size. The enlarged, grooved teeth on the maxillary are in shallow sockets on the posteroventral surface of the posterior knob of the maxillary. These teeth point posteriorly. The grooves are deep and are situated anterolaterally. One or two enlarged grooved teeth are present on a given maxillary. There seems to be a correlation between the type of pres- ervation, the age of the snake, and the number of grooved teeth. Old (large) individuals always have only one grooved tooth that is rooted and functional, whereas some of the younger animals have two in place. Usually replacement teeth are present in alcoholic specimens, but these unrooted teeth are lost in the preparation of dried skeletons. Thus, it seems that in Conophis only one pair of grooved teeth is functional at any one time, although usually re- placement teeth are present behind and beside the functional one. Some specimens have one tooth in the medial socket on one side and one in the lateral socket on the other. Replacement teeth on the maxillary and dentary are present in the buccal tissue on the medial side of the bones, whereas on the palatines and pterygoids, the replacement teeth are present laterally. Apparently there are no significant differences in dentition among the mem- bers of the genus Conophis. Vertebrae The fiftieth vertebra of Conophis vittatus (UMMZ 82642) can be described as follows: The neural spine is elongate, thin and low; the posterior edge is sharply emarginate, and the anterior edge is only slightly emarginate. The zygosphene is thin dorsoventrally; in a ventral or dorsal view the zygosphene has a slightly concave anterior edge, the flat surface of which is oriented ventrolaterally, The centrum is elongate and triangular from below; it is widest at the paradiapophyses and narrowest at the short condylar neck. The condylus is directed posteriorly. The centrum, when viewed laterally, is slightly concave and has prominent subcentral ridges that extend from the median side of the paradiapophysial articular surfaces posteriorly to the neck of the condylus. The paradiapophysial articular surfaces are well developed and have two facets. The diapophysial surface is larger and more spherical than the parapophysial one. The parapophysial process projects beyond the parapophysial articular surface and is nearly even with the lip of the cotyle, which is slightly oval. The neural arch is slightly depressed; its width is somewhat less than the width of the cotyle. The articular surfaces of the postzygapophyses are oval and are directed posterolaterally. There is a strongly developed concave interzygapophysial ridge. A well-developed accessory spine extends laterally beyond the oval articular facets of the pre- zygapophysis and forms a slightly flattened, blunt spine. Excellent drawings MippLe AMERICAN SNAKES (CONOPHIS ) 289 of the middle thoracic vertebra of Conophis lineatus dunni from Honduras were published by Auffenberg (1958:6). Hemipenes The hemipenes of Conophis are moderately caliculate, having spines cover- ing the surface from the base to near the apex (fig. 9). These spines are largest near the base and are re- duced to small papillate projections near the apex. The apex terminates in a small disc having three to five laminae in C. vittatus and one lamina in C. lineatus concolor. The sulcus is bifurcate; the fork is near the base and almost gives the appear- ance of two sulci on some _ speci- mens. Distally the apices are widely separated, and the _inter- vening space gives the hemipenis a slightly bilobed appearance in some species (especially C. vittatus) or a deeply bilobed appearance in others (especially C. lineatus con- color). The everted hemipenis reaches posteriorly to the eighth subcaudal scale. The sulcus bifurcates at the third subcaudal scale. The situation is similar in situ (Cope, 1895:pl. 28, fig; 2). There are no apparent hemipenial differences among the species of the genus Conophis. As can be seen in fy¢.9, The everted left hemipenis of the above description, the hemipenis Conophis vittatus (UMMZ 82650). of C. vittatus is less bilobed and has a x 5. more pronounced disc at the apex than the others. The hemipenis of C. lineatus concolor is most bilobed, but has the smallest apical disc. The other species and subspecies vary widely within these extremes. Food and Feeding Conophis eats mostly small lizards, especially Cnemidophorus. In México Conophis occurs in semi-arid habitat where Cnemi- dophorus is common. A specimen each of Conophis vittatus and C. lineatus lineatus were obtained while I was collecting Cnemi- dophorus. The only record of Conophis having fed on a warm- blooded vertebrate was obtained in the course of this study, when I recovered from the stomach of a Conophis lineatus concolor (CNHM 36299) from Chichén Itza, Yucatén, a heteromyid rodent (Heteromys gaumeri). 290 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Ralph Axtell (personal communication) observed Conophis ac- tively searching for food at dusk. His observations were made near Tehuantepec, Oaxaca, and the snakes were seen to chase lizards of the genus Cnemidophorus. Near Alvarado, Veracruz, in the late afternoon, I watched a Conophis lineatus lineatus follow a lizard into a hole. Mittleman (1944:122) presents the only discussion of the mode of feeding of a captive specimen of Conophis lineatus ssp. When presented with a Thamnophis slightly smaller than itself, the Conophis struck, and within eight minutes immobilized the Thamnophis, Within one-half hour the Thamnophis was swal- lowed. Three days later the Conophis ate another Thamnophis, though still distended from its first meal; nine days later it ate a Storeria. In the course of several months, the Conophis ate various toads and hylids and two more Storeria. Apparently members of the genus Conophis do not constrict their prey, but rely upon a combination of loss of blood and action of the venom to completely immobilize their prey. Ditmars (1931:pls. 26-27) showed three photographs of “Cono- phis lineatus” (actually Conophis pulcher) ingesting another snake, identified by him as a young Ophis (= Xenodon) colubrinus. Effect of Poison The rear fangs of these snakes are large for the size of the snake. Various collectors have been bitten, and several reports of the effect of the poison have been published. The snakes are aggressive and bite constantly while being handled. A field companion, Dale L. Hoyt, was bitten on the forefinger by a specimen of C. I. lineatus and immediately felt a burning sensation. The finger swelled, much as it would if stung by a wasp, but it returned to normal size in about twenty-four hours. Ditmars (1931:legend pl. 27) reported immediate burning pain and a localized swelling, an inch in diameter and half an inch high, which lasted for several hours. Mertens (1952b:83) reported merely that the hand of the gardener at the Instituto Tropical in San Salvador bled strongly for a full hour. Edward H. Taylor was bitten by a specimen of Conophis vittatus (Taylor and Smith, 1939:252); pain and swelling lasted for some time. Taylor (personal communication) is still troubled by damage incurred by that bite, which apparently resulted in mechanical damage to the second joint of the middle finger, for the joint swells when the finger is used or exercised. William E. Duellman (personal communication ) was bitten on the hand in July, MippLE AMERICAN SNAKES (CONOPHIS ) 291 1956. There was immediate pain and localized swelling, both of which disappeared several hours later. TAXONOMIC RELATIONSHIPS AND EVOLUTION The genus Conophis is known only from the Recent. Except that Conophis belongs to the subfamily xenodontinae and probably is of New World origin, little is known about the relation- ships of the genus. Auffenberg (1958) described a new genus and species of fossil colubrid snake from the Miocene of Montana as Dryinoides oxyrhachis and compared it with several recent genera. This specimen, of which there is a relatively complete skull and a series of vertebrae, seems most closely to resemble a specimen of Conophis lineatus dunni (UF 7657) from Honduras, with which it was compared in basic osteology. The two genera could be related, for the progenitors of Conophis possibly inhabited much of North America in the Miocene. Another possibility is that the main stock of the xenodontines reached South America in earliest Tertiary times, and that the formation of the Panamanian and Colombian seaways that separated South America and Central America from the Late Paleocene to the middle of the Pliocene left the Conophis stock isolated in Middle America where members of the genus dispersed through semi-arid habitats. Turning our attention now to the species within the genus, instead of the genus as a whole, Conophis vittatus is readily set apart from other members of the genus on the basis of the universal presence of seven supralabials. In basic coloration it also differs, having no stripe on the Ist scale-row, or spots on the venter, and a maximum of four broad stripes on the body. The other species appear to be more closely related; these make up the C. lineatus- group. Conophis nevermanni differs so much from the other species that it might be placed in a separate group. Nevertheless, the basic striped pattern, which is masked by the increased melanism of many specimens, indicates that nevermanni is more closely related to the lineatus-group than to vittatus. The lineatus-group, thus, consists of pulcher, nevermanni and the three subspecies of lineatus. In this group the color pattern is characterized by the high fre- quency of ventral spotting, darkening of part of the supralabials, dark pigmentation on the Ist scale-row, and more than four dark stripes on the body of adults. Conophis lineatus concolor, on which the dark pigmentation on the body apparently is secondarily lost, is an exception. 292 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. If differences in color pattern be used as an indication of the relationships between the species and subspecies of the genus Conophis, I would consider C. vittatus the most divergent unit. The subspecies of lineatus closely resemble one another and, as a unit, resemble pulcher from which they differ primarly in the position of the dorsalmost stripes. Conophis nevermanni is more divergent than is pulcher from the species lineatus, but probably is not so far removed from lineatus as is vittatus. In the light of what has been pointed out immediately above with respect to resemblances of, and differences between, the spe- cies, an hypothesis to account for their formation and for their pres- ence in the areas where they are today is the following: Concur- rent with climatic fluctuations in the Late Pliocene and Pleistocene, the northernmost population differentiated into the species vittatus, and has subsequently spread north and west from the region of Tehuantepec, México. During the same period nevermanni be- came isolated in northern Costa Rica. The species pulcher probably differentiated from the remaining lineatus stock during the Early Pleistocene orogenic upheaval in Guatemala. The pulcher stock was isolated on the Pacific Coastal slopes of Guatemala, while lineatus moved through the subhumid corridor of northern Middle America into México and southward toward Costa Rica (Stuart, 1954a). In the Late Pleistocene and Recent, pulcher moved back across the central Guatemalan high- lands occupying its present range in northern Middle America. Primarily because of the formation of unsuitable habitat (wet for- est) that presently separates the geographic ranges of populations of lineatus, this species differentiated into three subspecies. SUMMARY The genus Conophis Peters, 1860, contains four species. Three are monotypic and the fourth has three subspecies, making a total of six taxa. The genus is characterized by maxillary teeth of equal size fol- lowed by a diastema and two enlarged grooved fangs. The scales are smooth, in 19 rows at mid-body, and 17 nearer the tail. The anal is divided, apical pits are lacking, the head shields are normal for a colubrid, and the hemipenis is bilobed having many large basal spines. The six taxa are separated primarily on the basis of color pattern, but characters of scutellation, including numbers of dorsals, ven- MippLe AMERICAN SNAKES (CONOPHIS ) 293 trals, caudals, and places of reduction of the number of dorsal scale-rows, were analyzed. Snakes of this genus are distributed throughout semi-arid en- vironments from southern México southward into Costa Rica. They feed upon lizards, primarily of the genus Cnemidophorus; in addi- tion they are known to eat small rodents and other snakes. Conophis is a member of the subfamily Xenodontinae and, as presently understood, has no known living close relatives. A single specimen of Dryinoides from the Miocene of Montana has been compared with this genus. The genus Conophis is thought to have evolved in Middle America. The present distribution and differ- entiation probably are primarily the result of climatic fluctuations in Middle America, which produced the areas of subhumid en- vironment where Conophis presently lives. LITERATURE CITED AUFFENBERG, W. 1958. A new genus of colubrid snake from the Upper Miocene of North America. Amer, Mus. Novitates, 1874:1-16. February 27. Corer, E. D. 1861. Contributions to the ophiology of Lower California, México and Central America. Proc. Acad, Nat. Sci. Philadelphia, 13:292-306. December 28. 1867. Fifth contribution to the herpetology of tropical America. Proc. Acad, Nat. Sci. Philadelphia, 18:317-323. February 20. 1871. Ninth contribution to the herpetology of tropical America. Proc. Acad, Nat. Sci. Philadelphia, 23(2):200-224, October 24. 1876. On the batrachia and reptilia of Costa Rica. Journ. Acad, Nat. Sci. Philadelphia, series 2, 8(4):93-154, 6 pls. 1895. The classification of the ophidia. Trans. Amer. Philos, Soc., 18:186-219, 33 pls. April 15. 1900. The crocodilians, lizards, and snakes of North America, Ann, Rept. U. S. Natl. Mus. for 1898, pp. 153-1270, 36 pls. Dirmars, R. L. 1931. Snakes of the World. New York, The MacMillan Company, 1931. xi + 207 pp., 84 pls. Dow inc, H. G. 1951. A proposed standard system of counting ventrals in snakes. British Journ. Herpetology, 1(5):97-99, fig. 1. DvuELLMAN, W. E. 1958. A preliminary analysis of the herpetofauna of Colima, Mexico. Occas. Papers Mus. Zool. Univ. Michigan, 589:1-22, March 21. Dumert, A. M. C., Brsron, G., AND DuMErRML, A. H. A. 1854. Erpétologie genérale, ou histoire naturelle des reptiles. Paris, 7( pt. 2):xii-++ 785. February 25. Atlas, 24 pp., 108 Be Dumerit, A. H. A., Bocourt, M., AnD Mocaguarp, F. 1870-1909. Mission Scientifique au Mexique et dans |’Amerique Centrale Soa Etudes sur les Reptiles. Paris, vol. 2:xiv-+ 1012 pp., pls. GarMan, S. 1884a. The North American reptiles and batrachians. Bull. Essex Inst., 16:1-46. January 9. 294 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. 1884b. The reptiles and batrachians of North America. Mem. Mus. Comp. Zool., 8(3):xxxi + 185 pp., 9 pls. July. GunTuer, A. C. L. G. 1858. Calan of colubrine snakes in the collection of the British Museum. London. xiv+ 281 pp. Huxtey, J. 1942. Evolution. The Modern Synthesis, London. 645 pp. JAN, G. AND SoRDELLI, F. 66. Iconographie Generale des Ophidiens. Milano, livr. 19, pls. 1-6, December. 1881. Iconographie Generale des Ophidiens. Milano. livr. 50, pls. 1-7. November. Mayr, E. 1942. Be Te and the Origin of Species. New York, x+ 884 pp., gs. Mayr, E., Linsey, E. G., AND UsINGER, R. L. 1953. Methods and Principles of Systematic Zoology. New York. ix + 328 pp., 45 figs. MERTENS, R. 1952a. Neues uber die Reptilienfauna von El Salvador. Zool. Anz., 148:87-93. February. 1952b. Die Amphibien und Reptilien von El Salvador auf grund der reisen von R. Mertens und A. Zilch, Abhand. Senken. Naturw. Gesell., 487:83, 1 Kart., 16 taf. December 1. MiTTLEMAN, M. B. 1944. Feeding habits of a Central American opisthoglyph snake. Copeia, no. 2:122. June 80. NEILL, W. T. AND ALLEN, R. 1961. Further studies on the herpetology of British Honduras. Her- petologica, 17(1):387-52. April 15. ParKER, W. K. 1878. On the structure and development of the skull in the common snake (Tropidonotus natrix). Phil. Trans. Roy. Soc. London, pt. 2: 885-417, pp., pls. 27-33. PETERS, W. 1860. Drei neue amerikanisches Schlangen. Monatsb. Akad. Wiss. Ber- lin, 1860:517-521, pl., fig. 3. October. RapDovaAnovic, M. 1937. Osteologie des Schlangenkopfs. Jenaische Zeitschr. Naturw., 71(2):179-812. SAvAGE, J. M. 1949. Notes on the Central American snake, Conophis lineatus dunni Smith, with a record from Honduras. Trans. Kansas Acad. Sci., 50:483-486. December 31. Scumopt, K. P. 1928. Reptiles collected in Salvador for the California Institute of Tech- nology. Zool. Ser. Field Mus. Nat. Hist., 12(16):193-201. No- vember 21. Scum, K. P. and Incrr, R. F. 1957. Living Reptiles of the World. Garden City, New York, Hanover House. 287 pp. SmitH, H. M. 1941. Notes on snakes of the genus Conophis. Journ. Washington Acad. Sci., 31(3):117-124. March 15. Smitu, H. M. and Taytor, E. H. 1950. Type localities of Méxican reptiles and amphibians. Univ. Kansas Sci. Bull., 83:313-380. March 20. MiwpLe AMERICAN SNAKES (CONOPHIS ) 295 Stuart, L. C. 1948. The amphibians and reptiles of Alta Verapaz, Guatemala. Misc. Publ. Mus. Zool. Univ. Michigan, 69:1-109. June 12. 1954a.A description of a subhumid corridor across northern Central America, with comments on its hertetofaunal indicators. Contr. Lab. Vert. Biol. Univ. Michigan, 65:1-26 pp., 6 pls. March. 1954b. Herpetofauna of the southeast highlands of Guatemala. Contr. Lab. Vert. Biol. Univ. Michigan, 68:1-65 pp., 3 pls. November. Szunyocuy, J. 1932. Beitrage zur vergleichenden Formenlehre des Colubridenschadels, nebst einer Kraniologischen Synopsis der fossilen Schlangen Un- garns. Acta Zool., 13:1-56. Taytor, E. H. 1955. Additions to the known herpetological fauna of Costa Rica with comments on other species. No. II. Univ. Kansas Sci. Bull., 87:299-575. October 15. Tay or, E. H. and Smiru, H. M. 1939. Miscellaneous notes on Mexican snakes. Univ. Kansas Sci. Bull., 95:239-258. July 10. WETTSTEIN, O. 1934. Ergibnisse der osterreichischen biologischen Costa Rica—Expedi- tion 1930. Die Amphibia und Reptilien. Stiz. Akad. Wiss. Wien, mathem-naturw. kl., Abt. 1, bd. 1438:1-39. Transmitted November 80, 1962. L 29-5936 | 7 one Wie a ry a a - dia! pitt oy int 7 ai as Lag iad diet : D hes alae, se cen via (SI I Ale ot “iL a wo » «nem Yo ee eee in > OR) ON. i nt Aaa © * 4. by Te ood an Mn ‘a ne ee >, a ay, ay, y | ate.) yo ae ; ie oe Fe Ley j iW ~ A vy | A? ei | » a iy aT 7 j i oh ere Te . Hu ee Ai ied be td \ ’ mc a . i : ove AG 7 ((. A Tr at i Cm is haan ’ , pan) = : payee bd = eee [Pw P 4 ~ ae ee Wits a F wn Pa ; _— i @ = ’ f a 7, Chine oer il) “Hh li : ; %) wa . is > i AAV pa OP pees rie) | ai ee ’ a 4 ee Ate : ; 7 f ; 4 ; 7 1 we ee br Pe “Ke pay Ne hus al whe Pg lee tabi Thy a » tt Py ; ‘9 vi » @ 9 | ty ida php | ry ‘ a, re 7 haga \ oe 3 . % re F : 7 i | et ee id ‘in th ee Aisa spent ciel ey his AM : a0 a he ‘i : ie on, . ron Ken a woh be ‘ *”~ arn 7h iva a in te J Ned - Samii ae ee pues MBAs) af is ter} “S ‘Gi’ ioe, ee ; 1 nee et xi! inpag ae A teak a ey Tp 1 ga an ah ha) uy bi i . . i me al he je _ 7 : + sh a ii a ar t rie 7 i Vol. 10. Vol. 12. Vol. 18. 18. 19. 20. 21, 22. 23. (Continued from inside of front cover) Conspecificity of two pocket mice, Perognathus goldmani and P, artus. By E. Raymond Hall and Marilyn Bailey Ogilvie. Pp. 513-518, 1 map. Janu- ary 14, 1960. Records of harvest mice, Reithrodontomys, from Central America, with de- scription of a new subspecies from Nicaragua. By Sydney Anderson and J. Knox Jones, Jr. Pp. 519-529. January 14, 60. Small carnivores from San Josecito Cave (Pleistocene), Nuevo Leén, México, By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, 1960. Pleistocene pocket gophers from San Josecito Cave, Nuevo Leon, México. By Robert J. Russell. Pp. 539-548, 1 pure in text. January 14, 1960. Review of the insectivores of Korea. y J. Knox Jones, Jr., and David H. Johnson, Pp. 549-578, February 23, 1960. Speciation and evolution of the pygmy mice, genus Baimoys. By Robert L. Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960. Index. Pp. 671-690 ale ° aie 8. 4, *6. Studies of birds killed in nocturnal migration: By Harrison B. Tordoff and Robert M. Mengel. Pp. 1-44,.6 figures in text, 2 tables. September 12, 1956. Comparative breeding behavior. of ‘Ammospiza _ caudacuta and A. maritima, By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956. The forest. habitat of the University of Kansas Natural History Reservation. By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures in text, 4 tables. December 81, 1956. Aspects of reproduction and development in the prairie vole: (Microtus ochro- gaster). By Henry S, Fitch. Pp, 129-161, 8 figures in text, 4 tables. Decem- ber 19, 1957. Birds found on the Arctic slope of northern Alaska. ye James W. Bee. Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958 : The wood rats of Colorado: distribution and ecology. By Robert B. Finley, Jr. Pp. 218-552, 34 plates, 8 figures in text, 85 tables. November 7, 1958. Home ranges and movements of the eastern cottontail in Kansas. By Donald W. Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, 1959. Natural history of the salamander, Aneides hardyi. By Richard F. Johnston and Gerhard A. Schad. Pp. 5738-585. October 8, 1959. 9. A new ee of lizard, Cnemidophorus sacki, from Michoacan, México, 10. By William E, Duellman. Pp. 587-598, 2 figures in text. May 2, 1960. A taxonomic study of the middle American snake, Pituophis deppei. By William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960. Index. Pp. 611-626. Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960. 1. *2. 3. *4, Functional morpgloRy of three bats: Sumops, Myotis, Macrotus._ By Terry A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. ‘July 8, 1959. The ancestry of modern “Amphibia: a review of the evidence. By Theodore H, Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, 1959. The baculum in microtine Rey By Sydney Anderson. Pp. 181-216, 49 figures in text. February 19, 1960. A new order of fishlike eee from the Pennsylvanian of Kansas. By Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp..217-240, 12 figures in text. May 2, 1960. Natural history of the bell vireo. By Jon C, Barlow. Pp. 241-296, 6 figures Ds in text. March 7, 1962. Two new \pelycosaurs from the lower Permian of Oklahoma. By Richard C. Fox. Pp. 297-307, 6 figures in text. May 21, 19 Vertebrates from the barrier island of Tamaulipas, México. By Robert K. Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 309- 845, pls. 5-8. June 18, 1962. Teeth of Edestid sharks. By Theodore H. Eaton, Jr. Pp. 847-362, 10 fig- ures in text. October 1, 1962. More numbers will appear in volume 12. Five natural Any combina eon in minnows (Cyprinidae). By Frank B. Cross and W. L. Minckley. Pp. 1-18. June 1, 1960. A ofS AS eaek "study of the Ate. of the Isthmus of Tehuantepec, México. By William E. Duellman. Pp. 19-72, pls. 1-8; 3 figures in text. August 16, 1960. A new subspecies. of the slider turtle (Pseudemys scripta) from Coahulia, Cee By John M. Legler. Pp. 73-84, pls. 9-12, 8 figures in text. August Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, pls. 13-20, 26 figures in text. November 30, 1960, Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and Rocky Mountains. By Henry S. Fitch and T. Paul. Maslin. Pp. 289-308, 4 figures in text. February 10, 1961. Fishes of the Wakarusa river in Kansas. By Jams E. Deacon and Artie L. Metcalf. Pp. 309-322, 1 figure in text. February 10, aly Geographic variation in the North American cyprinid fish, Hybopsis gracilis. By Leonard J. Olund and Frank B. Cross. Pp. 323-348, pls. 21-24, 2 figures in text. February 10, 1961 (Continued on outside of back cover) 8. 9. 10. (Continued from inside of back cover) Decriptions of two species of frogs, genus Ptychohyla; studies of Ameri- can hylid frogs, V. By William E. Duellman. Pp. 349-357, pl. 25, 2 figures in text. April 27, 1961. Fish populations, following a drought, in the Neosho and Marais des Cygnes rivers of Kansas» By James Everett Deacon. Pp. 359-427, pls, 26-30, 8 figs. August 11, 1961. Recent soft-shelled turtles of North America (family Trionychidae). By Cea Me Webb. Pp. 429-611, pls. 31-54, 24 figures in text. February Index. Pp. 613-624. Vol. 14. 1. 2. 8. Neotropical bats from western México. By Sydney Anderson. Pp. 1-8. October 24, 1960. Geographic variation in the harvest mouse. Reithrodontomys megalotis, on the central’Great Plains and in adjacent regions. By J, Knox Jones, Jr., and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961. Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson. Pp. 29-67, pls. 1 and 2, 8 figures in text. July 24, 1961. A new subspecies of the black myotis (bat) from eastern Mexico. By E. SESS Hall anad Ticul Alvarez. Pp. 69-72, 1 figure in text. December North American yellow bats, “‘Dasypterus,’” and a list of the named kinds of the genus Lasiurus Gray. By E. Raymond net and J. Knox Jones, Jr. Pp. 73-98, 4 figures in text. December 29, 196 Natural history of the brush mouse Tecate boylii) in Kansas with description of a new a tae By Charles A. Long. Pp. 99-111, 1 figure in text. December 29, 19 Taxonomic status of Bite mice of the Peromyscus boylii group in eastern Mexico, with description of a new Te ean By Ticul Alvarez. Pp. 118- 120, 1 figure in text. December 29, A new subspecies of ground bea iScuae ualare spilosoma )- ecg Ta- maulipas, Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962 Taxonomic status of the free-tailed bat, Tadarida yucatanica Mille By J. , Saas Jones, Jr., and Ticul Alvarez. Pp. 125-138, 1 figure in text. March 7, A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene, of Texas. By E. Raymond Hall and Walter W. Dalquest. Pp. 135-138, 2 figures in text. April 80, 1962. A new subspecies of wood rat (Neotoma) from northeastern Mexico. By Ticul Alvarez. Pp. 189-148. April 80, 1962. Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul rehie and M. Raymond Lee. Pp. 145-159, 1 figure in text. May 18, A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, 1 figure in text. May 21, 1962. The mammals of Veracruz. By E. Raymond Hall anad Walter W. Dalquest. Pp. 165-362, 2 figures. May 20, 1963. The recent mammals of Tamaulipas, México, By Ticul Alvarez. Pp. 863- 478, 5 figures in text. May 20, 1963 More numbers will appear in volume 14, The gcteltens and reptiles of Michoacan, México. By William E. Duell- * man. Pp. 1-148, pls. 1-6, 11 figures in text. December 20, 1961. Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox Jones, Jr., and George W. Byers, Pp. 149-173. January 31, 1962. A new species of frog (Genus Tomodactylus ) from western México. By Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962. eee specimens of amphibians *and reptiles in the Museum of Natural His- the University of Kansas. By William E. Duellman\and Barbara Berg. Pe 183-204. October 26, 1962 Amphibians and Reptiles of ae “Rainforests of Southern El Petén, Guatemala. By William E. Duellman.» Pp. 205-249, pls. 7-10, 6 figures in text. October 4, 1963. A revision of snakes of the genus Conophis (Family Colubridae, from Middle America). By John Wellman. Pp. 251-295, 9 figures in text.. October 4, 1963. More numbers will appear in volume 15. Sb tT A = Za ay UNIVERSITY OF KANSAS PUBLICATIONS MusEUM OF NATURAL HISTORY Volume 15, No. 7, pp. 297-349, pls. 11-18, 7 figs. 2 October 18, 1968 _A Review of the Middle American Tree Frogs of the Genus Ptychohyla BY WILLIAM E. DUELLMAN J UNIVERSITY OF KANSAS LAWRENCE 1963 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Institutional libraries interested in publications exchange may obtain this series by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas. Copies for individuals, persons. working in a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of Kansas, Lawrence, Kansas. There is-no provision for sale of this series by the University Library, which meets institutional requests, or by the Museum of Natural History, which meets the requests of individuals, Nevertheless, when individuals request copies from the Museum, 25 cents should be included, for each separate number that is 100 pages or more in length, for the purpose of defraying the costs of wrapping and mailing, - * An asterisk designates those numbers of which the Museum’s supply (not the Library’s supply) is exhausted. Numbers published to date, in this series, are as follows: Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950. *Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text. April 9, 1948. Vol. 8. *1. The avifauna of Micronesia, its origin, evolution, and distribution. By ‘Rol- lin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951. *2. A quantitative study of the nocturnal migration of, birds. By George H. Lowery, Jr. Pp. 861-472, 47 figures in text. June 29, 1951. 8. Phylogeny of the waxwings and allied birds. By M. Dale Arvey. - Pp. 4738- 580, 49 figures in text, 13 tables. October 10, 1951. *4, Birds from the state of Veracruz, Mexico. By George H. Lowery, ‘Jr., and Neer W.,. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, Index. Pp. 651-681. £Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 figures in text. December 27, 1951. Vol. 5, Nos. 1-87 and index. Pp. 1-676, 1951-1958. *Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. Durrant. Pp. 1-549, 91 figures in text, 80 tables. August 10, 1952. Vol..7. Nos. 1-15 and index. Pp. 1-651, 1952-1955. Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956. Vol. 9. *1. Speciation of the wandering shrew. By James S. Findley. -Pp. 1-68, 18 figures in text. December 10, 1955. 2. Additional records and extension of ranges of mammals from Utah. By Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80. December 10, 1955. 8. A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rol- lin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955. 4, Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956. 5. The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6 figures in text. May 19, 1956. 6. Additional remains of the multituberculate genus Eucosmodon. By Robert W. Wilson. Pp, 117-123, 10 figures in text. May 19, 1956. 7. Mammals of Coahulia, Mexico. By Rollin H. Baker. Pp. 125-835, 75 figures in text. June 15, 1956. 8. Comments on the taxonomic status of Apodemus peninsulae, with description of a new subspecies from North China.- By J. Knox Jones, Jr. Pp. 337-346, 1 figure in text, 1 table. August 15, 1956. 9. Extensions ef known ranges of Mexican bats. By Sydney Anderson. Pp. 847-351. August 15, 1956. : 10. A new bat (Genus‘ Leptonycteris) from Coahulia. By Howard J. Stains. Pp. 353-3856. January 21, 1957. : : 11. .A-new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico. By Robert J. Russell. Pp. 3857-861. January 21, 1957. . 12. Geographic variation in the pocket gopher, Thomomys bottae, in Colorado. By Phillip M. Youngman. Pp. 363-387, 7 figures in text. February 21, 1958. 18. New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones, Jr. Pp. 385-388," May 12, 1958. | os, ; 14, Pleistocene bats from San Josecito Cave, Nuevo Leén, México. By J. Knox Jones, Jr. Pp. 889-396. December 19, 1958. } 15. New subspecies of the rodent Baiomys from Central America. By Robert L. Packard. Pp. 397-404. December 19, 1958, 16. Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405- 414, 1 figure in text, May 20, 1959. 17. Distribution, variation, and relationships of the montane vole, Microtus mon- | tanus. By Sydney Anderson. Pp. 415-511, 12 figures in text, 2 tables. August 1, 1959. (Continued on inside of back cover) UNIVERSITY OF KANSAS PUBLICATIONS MusEuUM OF NATURAL HISTORY Volume 15, No. 7, pp. 297-349, pls. 11-18, 7 figs. October 18, 1963 A Review of the Middle American Tree Frogs of the Genus Ptychohyla WILLIAM E. DUELLMAN UNIVERSITY OF KANSAS LAWRENCE 1963 UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 15, No. 7, pp. 297-349, pls. 11-18, 7 figs. Published October 18, 1963 UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY JEAN M. NEIBARGER, STATE PRINTER TOPEKA, KANSAS 1963 A Review of the Middle American Tree Frogs of the Genus Ptychohyla BY WILLIAM E. DUELLMAN CONTENTS PAGE INTRODUCTION 55 8.6 35.24 12 SSSR ge reo ee eee 301 Keknowledements: 0. .oco.cc tau ore ee ee 301 Matenialseands Methods) ..(.cies.c2 4 aces 2 ee Oe 302 NAT YSISUGEP ID ATA eo... ice. 32. < Ses then o> Hee cere ee eae 303 External Morphology. 4:2: 46> i an See eee 303 Coloriandwrattertie 2: 2,4.) 1:0 e seea-y oe een oe ee 307 (ON (200) (0), fk i nee eee ARIA AO n'a idlol oe oe 307 FRAC DOVES feist thers c &:5 ov oaks REAR Geno oe ee eee 310 Breeding Gall 5 ee so 'os, oho le aan Denteah ne aan eee ee 312 SYSTEMATIC WA CCOUNTS jiiip.i:)+ ccs Slee dona aca Lak Leta ee ee 314 Piychohyla Maylors 1944. ook ee 314 Key tovAGultsie. dit: asc tisenatedh Ge eee re eee 315 Key to: Tadpoles %o. 23. «oc es wag cls Stee Sea eee eee 315 Piychohyla- euthysanota,Group —. 22.0). ):25 25 --ecee 315 Piychohyla euthysanota. <.../.<2 i. ee ee 315. Ptychohyla euthysanota euthysanota (Kellogg) ........... 315 Ptychohyla euthysanota macrotympanum (Tanner) ........ 320 Ptichohyla leonhardschultze: (AI) ...- 2° ae 323 Ptychohyla spinipoller (Schmidt) .:.2.25..->-.--- 9) 327 Ptychohula schmidtorum Group: ©... 6 92 20ers ee ee 331 Piychohyla schmidtorum .. 2.2... 02...) 2-)4203e ee eee 331 Ptychohyla schmidtorum schmidtorum Stuart ............. got Ptychohyla schmidtorum chamulae Duellman ............. 334 Piychohyla iznicolor Duellman -.-........5,-7 3522: 337 DISTRIBUTION AND) ECOLOGY =. {2.44 nad. oe Ia eee 340 Geographic Distribution of the Species ................... 340 Habitat Preterence?s:< cc ac: 1h hee ee eee 342: Interspecific: Competition. ....42.5.95. 47 322 224) eee eee 343 Reproduction and Development ..................-.+--.. 344 (299) 300 University OF Kansas Pusts., Mus. Nat. Hist. PHYLOGENY OF PryCHOHYLA Piychohyla asa Natural Assemblage 77 2.....))...022.).% =<: 345 Generic Relationships Interspecific Relationships LITERATURE CITED .... INTRODUCTION Probably no ecological group of hylid frogs (some Hyla plus Plectrohyla and Ptychohyla) in Middle America is so poorly known as those species that live in the cloud forests on steep mountain slopes and breed in cascading mountain streams. During the last half of the nineteenth century most of the species of hylids living in the lowlands of southern México and northern Central America were named and described. Despite the extensive collecting by Salvin and Godman, Nelson and Goldman, and the various expedi- tions of the Mission Scientifique, no members of the genus Ptycho- hyla were obtained until 1927, when in the mountains of El Salvador Ruben A. Stirton found a small tree frog that subsequently was described and named Hyla euthysanota by Kellogg (1928). Until recently frogs of this genus were know from few specimens and in the literature by nearly as many names. Although I first collected Ptychohyla in 1956, it was not until 1960 that special efforts were made to obtain specimens of this genus. The summer of 1960 was spent in southern México and Guatemala, where every accessible stream in the cloud forests was searched for tree frogs, especially Ptychohyla and Plectrohyla. Similar, but less extensive, investigations were carried out in 1961 and 1962. The result of this field work is a rather large collection of Ptychohyla representing all of the known species, plus tape recordings of the breeding calls and tadpoles of all of the species. Previously, I have discussed the nomenclature of one of the species (Duellman, 1960) and have described two new species (Duellman, 1961). In the latter paper I made reference to a future account (this one) that would deal with the systematics and biology of the entire genus. Although I have series of specimens, tadpoles, osteological preparations, and recordings of breeding calls, thereby having a wide array of data at my disposal, much still remains to be learned about these frogs, especially about various aspects of their life histories. Even the validity of the genus is open to question; this problem is discussed at length in the section beyond entitled “Ptychohyla as a Natural Assemblage.” Acknowledgments I am indebted to the following persons for permitting me to examine speci- mens in their care: Miguel Alvarez del Toro, Museo Zoologia de Tuxtla Gutierrez, México (MZTG); Charles M. Bogert and Richard G. Zweifel, Ameri- can Museum of Natural History (AMNH); Doris M. Cochran, United States (301) 302 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. National Museum (USNM); Norman Hartweg and Charles F. Walker, Uni- versity of Michigan Museum of Zoology (UMMZ); Robert F. Inger, Chicago Natural History Museum (CNHM); Hobart M. Smith, University of Illinois Museum of Natural History (UIMNH); Heinz Wermuth, Zoologisches Museum Berlin (ZMB); and Ernest E. Williams, Museum of Comparative Zoolegy (MCZ). The abbreviations following names of institutions will be used throughout the text; the Museum of Natural History at the University of Kan- sas is abbreviated KU. Throughout my work on these frogs I have profited from discussions with L. C. Stuart, who has made many valuable suggestions and with his char- acteristic generosity has placed at my disposal his extensive collections of tad- poles from Guatemala. For his aid I am indeed grateful. I am grateful to Thomas E. Moore for tapes of breeding calls of two species. My own field work was made more enjoyable and profitable through the assistance of Dale L. Hoyt, Craig E. Nelson, Jerome B. Tulecke, and John Wellman, all of whom spent many hours in often unsuccessful attempts to collect specimens and record breeding calls of Ptychohyla. I am indebted to many residents of México, Guatemala, and EI Salvador for permission to work on their land and for providing shelter, food, and guides. I am especially grateful to Mr. and Mrs. Horatio Kelly of “Colegio Linda Vista” at Pueblo Nuevo Solistahuacan, Chiapas, for a pleasant stay at their school; Jordi Julia ‘Z. of the Comisién del Papaloapan, Ciudad Aleman, Veracruz, for arranging for field work in northern Oaxaca in 1959; Walter Hannstein and Lothar Menzel for the use of facilities at Finca La Paz, Guatemala, in 1960; Alan Hempstead for the use of facilities at Finca Los Alpes, Guatemala in 1960 and 1961; and Julio Aguirre C. of the Instituto Tropical de Investigaciones Cientificas, San Salvador, El] Salvador, for providing comfortable working quar- ters and transportation and guides to the mountains in northem El Salvador. Without the cheerful efforts of Jorge A. Ibarra, Director of the Museo Nacional de Historia Natural in Guatemala, my field work would have been greatly re- stricted during politically precarious times in that country. Permits to collect in México were furnished by the late Luis Macias Arellano of the Direccién General de Caza. Each of these individuals has my profound thanks for his indispensable aid. Field work on hylid frogs in Middle America has been supported by the National Science Foundation, Grant NSF-G9827, and this is the 9th publica- tion on the results of study of the material from America. Materials and Methods During the course of this study I have examined 247 frogs that I assign to the genus Ptychohyla, plus 40 lots of tadpoles and 12 skeletal preparations. Furthermore, I have examined all of the type specimens. I have studied each of the species and subspecies in the field and have examined from seven (P. euthysanota macrotympanum) to 33 (P. spinipollex) living individuals of each species. Measurements given in the analysis of data and in the descriptions of the species are those described by Duellman (1956). In the descriptions of living colors the capitalized names are from Ridgway (1912). All interpretations of osteological characters are based on specimens cleared in potassium hydroxide and stained with alizarin red. MippLeE AMERICAN TREE FROGS 803 Recordings of the breeding calls were made with a Magnemite Portable Tape-recorder; audiospectrographs were made on a vibralyzer (Kay Electric Company ) using normal pattern and wide bandwidth. ANALYSIS OF DATA Data that are used to arrive at a systematic arrangement of the species of Ptychohyla are analyzed and discussed below for the values inherent in the analysis. These data are of some value also in the recognition of species and subspecies but if employed for that purpose the data must be used in combination with the keys and the diagnoses of the individual species and subspecies. External Morphology Each of the external morphological characters used in the sys- tematic treatment of Ptychohyla, as well as the nature of the tongue, is discussed below. SIZE AND Proportions.—Comparisons of size and certain propor- tions are given in Table 1. Frogs of this genus are small; the largest specimen examined is a female of P. euthysanota euthysanota hav- ing a snout-vent length of 53.3mm. The species comprising the Ptychohyla schmidtorum group are smaller; the largest specimen examined is a female of P. schmidtorum schmidtorum having a snout-vent length of 38.0 mm. An analysis of the various measure- ments and proportions shows few constant differences. Ptychohyla ignicolor differs from all of the other species in having the head slightly wider than long and the tympanum noticeably less than half the size of eye. Ptychohyla spinipollex has a relatively narrow in- terorbital distance, approximately equal to the width of the eyelid, whereas in all of the other species that distance is much more than the width of the eyelid. SnouTt.—All species have a blunt snout. In P. leonhardschultzei and P. ignicolor the snout is nearly square in lateral profile; in P. schmidtorum the snout is slightly rounded above and below, and in the other species it is rounded above. Ptychohyla leonhardschultzei and P. spinipollex have a vertical fleshy rostral keel on the snout; in these speces, because of this keel, the snout in dorsal profile is pointed. The nostrils are slightly protuberant in all species, and in P. schmidtorum the internarial region is slightly depressed. Hanv.—The species in the Ptychohyla euthysanota group have a vestige of web between the first and second fingers; the other fingers are about one-third webbed. Breeding males have a cluster of horny nuptial spines on the thumb. The spines are largest in P. 304 Unrversirty OF Kansas Pusis., Mus. Nat. Hist. TABLE ],—VARIATION IN CERTAIN CHARACTERS IN THE SPECIES OF PTYCHOHYLA. (MEANs ARE IN PARENTHESES BELOW THE RANGES.) 2) od Sil. Ss a\ ab) &| 8 5 iB Species i= gs me yas S| 2 S8iaag| 8/52 aa z a&i( #2] ala m g Mueiesieul ss wine 2 6 m ie = > patel ed Tle ead toe rod Ne aN Eee Si Ee te ae he oe P. euthysanota euthysanota....... rot 17 | 38.1 | 44.4-55.0 | 48.6-63.8 | 4-6 (48.7) (56.3) (S21) 2 15 | 53.3 | 46.5-56.6 | 42.9-56.4 | 6-18 (51.4) (51.4) (9.6) P. euthysanota macrotympanum...| & 5 | 38.0 | 48.8-52.0 | 50.0-57.1 Q-4 (50.2) (54.1 (2.6) °) 5 44.8 | 46.4-54.1 48 .7-58.9 8-10 (50.2) (53.7) (9.2) P. leonhardschulizei...)| o& 16 | 35.6 | 48.8-56.1 | 48.7-61.9 | 6-9 552 (52.1 (6.5) Q 3 41.6 | 52.3-59.5 47 .5-48.6 7-12 (54.7) (48.1) (9.5) P. spintpoller......... . ou 32 | 41.2 | 46.9-53.1 | 45.0-55.2 | 3-7 (49.0) (49.5) (4.9) Q 6 | 44.6 | 46.1-50.2 | 47.7-53.8 | 6-10 (48.8) (50.4) (7.6) P. schmidtorum schmidtorum...... rot 25 | 32.8 | 45.3-52.4 | 51.5-59.3 | 5-11 (48.1) (54.7) (6.2) 2 9 38.0 | 46.5-49.1 51.3-58.3 7-11 (47.7) (54.9) (8.7) P. schmidtorum chamulae......... of 40 | 30.5 | 46.0-51.9 48.2-65.6 | 4-6 (48.2) (54.9) (4.7) Q 4 | 31.8 | 48.1-52.4 | 51.4-61.7 | 4-9 (50.5) (55:7) (6.2) Pi 4QntC0lOr 0 os es. < of 13 | 30.5 | 45.9-52.2 | 37.1-47.1 | 3-9 (49.6) (43.2) (6.1) spinipollex (Fig. 1) and vary in number from 35 to 66 (average 47.4) on each thumb. In the other species of the Ptychohyla euthysanota group the spines are smaller and usually more numer- ous; the numbers of spines on each thumb (means in parentheses ) in members of this group are: P. euthysanota euthysanota, 44-143 (83.8); P. euthysanota macrotympanum, 40-110 (63.0); P. leonhard- MippLte AMERICAN TREE FROGS 305 schultzei, 24-80 (54.7). The species in the Ptychohyla schmidtorum group have no web between the first and second fingers and only a vestige of web between the other fingers. Furthermore, these species lack nuptial spines in breeding males. Like the usual horny excresences on the thumbs of many species of frogs, the horny spines on the thumbs of members of the Ptychohyla euthysanota group are seasonal in development. Fic. 1. Palmar views of right hands of (A) Ptychohyla spini- pollex (KU 58054) and (B) Ptychohyla a ana oe schmid- torum (KU 58048). x4 Many workers have used the presence of a bifid subarticular tubercle beneath the fourth finger as a diagnostic character of cer- tain species of hylids. Examination of the subarticular tubercles in Ptychohyla reveals considerable intraspecific variation. Bifid tubercles beneath the fourth finger are found in all species except P. ignicolor, which is known from only two specimens. In P. euthysanota euthysanota nearly 60 per cent and in P. schmidtorum schmidtorum about 90 per cent of the specimens have a bifid tubercle beneath the fourth finger on one or both hands. All specimens of P. leonhardschultzei have either a bifid or double tubercle beneath the fourth finger, and some have a bifid distal tubercle beneath the third finger. Fret.—Members of the Ptychohyla euthysanota group have a weak tarsal fold, whereas in the species comprising the Ptychohyla schmidtorum group the tarsal fold is absent. Webbing on the foot extends to the discs of the third and fifth toes and to the base of the penultimate phalanx of the fourth toe, except in P. ignicolor, which has less webbing. 306 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. VENTROLATERAL GLANDS.—Breeding males develop thickened, pigmented glandular areas on the sides of the body. In living specimens of P. schmidtorum and P. ignicolor the glands are not readily visible, but in preservative they show as distinctive orange- colored areas. These glands are most distinct in P. euthysanota; in many specimens of this species the glands are elevated above the surrounding skin. The extent of the glands is variable (Fig. 2); probably this variability is due to different degrees of develop- ment in individual frogs rather than to interspecific differences. All Ptychohyla ignicolor and some P. schmidtorum chamulae have a small, round glandular area on the chin; to my knowledge this does not occur in the other species. Superficial examination of micro- scopic preparations of the glands reveals no histological differences between species. The glands occupy most of the thickened area and have narrow ducts leading to the exterior. Detailed studies of the histology will be reported elsewhere. Since the glands are developed only in breeding males, it is surmised that the glands are associated with some phase of the breeding activity. aiaee YW vila Vian Fic. 2. Normal extent of ventrolateral glands in (A) Ptychohyla euthysanota euthysanota (KU 58008), (B) Ptychohyla schmidtorum schmidtorum (KU 58037), and (C) Ptychohyla ignicolor (UMMZ 119608). x 2h. MippLe AMERICAN TREE FROGS 307 ToncuE.—The shape of the tongue varies intraspecifically. Usually the tongue is ovoid; in some specimens it is barely notched posteriorly, whereas in others it is deeply notched, making the tongue cordiform. Deeply notched cordiform tongues are found in P. leonhardschultzei and P. schmidtorum; with the exception of these two species, some individuals of all species have emarginate tongues. Some individuals of all species have tongues that are shallowly notched posteriorly. Color and Pattern The dorsum in living frogs of the genus Ptychohyla is primarily yellowish or reddish brown, except in P. schmidtorum chamulae and P. ignicolor in which it is green. Usually there are some darker blotches or reticulations on the dorsum. The venter usually is white; in P. ignicolor it is yellow. The venter is spotted in all mem- bers of the Ptychohyla euthysanota group; the species, arranged from least to most spotting ventrally, are: P. euthysanota euthy- sanota, P. euthysanota macrotympanum, P. leonhardschultzei, and P. spinipollex. The last two species also have bo!d dark spots on the flanks. Ptychohyla schmidtorum lacks spots on the venter, whereas P. ignicolor has small dark flecks ventrally. Ptychohyla euthysanota and P. schmidtorum have white stripes on the upper lips and on the flanks. All species have some form of a pale stripe above the anus and usually rather distinct white or pale stripes along the ventrolateral edges of the tarsi and fore- arms. There are no bright or boldly marked flash-colors (colors that are revealed only when the hind limbs are extended), except in P. ignicolor, which has bright red flash-colors in the groin and on the thighs. In life the iris varies from several different shades of bronze color to deep red in P. schmidtorum schmidtorum. The degree of metachrosis is moderate. Usually any change of color in life consists only of change in the intensity of color. At times when the over-all coloration is darkened some markings are obscured. Osteology The following description of the skull, hyoid, sternum, and prepollex is based on a male specimen of P. spinipollex (KU 68632) that has been cleared and stained. The broad, flat skull (Fig. 3) has a large frontoparietal fontanelle. The ethmoid is large and has a flange laterally. The nasals are of moderate size and in broad contact with the ethmoid, but are separated from one another 308 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. medially. The anterior half of the maxillary bears a thin, high flange. The anterior process of the squamosal is short and widely separated from the maxillary. The quadratojugal is a small spine- shaped element projecting anteriorly from the ventral base of the quadrate; the quadratojugal does not articulate with the maxillary. Fic. 8. Dorsal aspect of skull of Ptychohyla spinipollex (KU 68632). Arrow indicates reduced quadratojugal. x 6. The posteromedian part of the hyoid plate is calcified; from this plate the long bony, posterior cornua (thyrohyales) extend pos- terolaterally. The omosternum is calcified, widest anteriorly, and has a convex anterior edge. The calcified xiphisternum is roughly bell-shaped having short lateral processes anteriorly and a deep notch pos- teriorly. The swollen thumb is supported by a dorsoventrally flattened spine that does not extrude through the skin. VARIATION.—In general, the skull varies little. Usually the quad- ratojugal is present only as a short element attached to the quadrate, but in one specimen of P. spinipollex the quadratojugal articulates with the maxillary and forms a complete quadratojugal-maxillary arch on each side of the skull. One specimen of P. leonhard- schultzei has a complete arch on one side and an incomplete arch on the other. Only P. spinipollex has lateral processes anteriorly on the xiphi- sternum; in the other species the xiphisterum is deeply bell-shaped. Ptychohyla schmidtorum and P. ignicolor have slightly longer MippLe AMERICAN TREE FROGS 809 premaxillaries than the other species. The longer premaxillary is reflected in the larger number of teeth on the bone—9 to 11 (aver- age 10) in four specimens of P. schmidtorum and 10 teeth in one P. ignicolor, as compared with 6 to 10 (average 7.9) in seven speci- mens of the other species. The number of maxillary teeth in the various species are: P. euthysanota euthysanota, 43; P. euthysanota macrotympanum, 38; P. leonhardschultzei, 38 and 40; P. spinipollex, 84 and 40; P. schmidtorum schmidtorum, 37 and 43; P. schmidtorum chamulae, 40 and 41; P. ignicolor, 43. The teeth on the premaxillary and anterior part of the maxillary are long, pointed, and terminally curved backwards. Posteriorly on the maxillary the teeth become progressively shorter and blunter. Variation in number of vomerine teeth is shown in Table 1. Fic. 4. Tadpoles of the Ptychohyla euthysanota group: (A) P. euthy- sanota euthysanota (KU 60042), (B) P. euthysanota macrotympanum (KU 60049), (C) P. leonhardschultzei (KU 68556), and (D) P. spini- pollex (KU 60053). 2%. 310 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Tadpoles Tadpoles of the genus Ptychohyla are adapted to live in moun- tain streams. The bodies are streamlined, and the tails are long and have low fins (Figs. 4 and 5). The mouths are large and directed ventrally. Tadpoles of the two groups of species have strikingly different mouthparts. Fic. 5. Tadpoles of (A) Ptychohyla schmidtorum schmidtorum (KU 60051), (B) P. schmidtorum chamulae (KU 58199), and (C) P. ignicolor (KU 71716). x 2h. Lips of tadpoles of the Ptychohyla euthysanota group (Fig. 6 A-D) are folded laterally; there are % or sometimes 4% tooth-rows. A lateral “wing” projects on either side of the upper beak. The beaks have blunt, peglike serrations. Lips of tadpoles of the Ptychohyla schmidtorum group (Fig. 6 E-G) are greatly expanded and form a funnel-shaped disc; there are *4 short tooth-rows. There is no lateral projection or “wing” on either side of the upper beak. The beaks have long, pointed serrations. Mippite AMERICAN TREE FROGS 311 ern NENA AA s\ A ©, * A oo . > KT > Gxeou Ue 2~? ey CWO Grain U yi fh y ? pe ¢ od od % ~ Bos - ¢ 2 Bee R ind : & = 3 = q coe an PN al A ® ay es tay PU UU Uw es Fic. 6. Mouthparts of tadpoles of Ptychohyla: (A) P. euthysanota euthy- sanota (KU 60042), (B) P. euthysanota macrotympanum (KU 60049), (C) P. leonhardschultzei (KU 68556), (D) P. spinipollex (KU 60053), (E) P. schmidtorum schmidtorum (KU 60051), (F) P. schmidtorum chamulae (KU 58199), and (G) P. ignicolor (KU 71716). x 10. Variation in certain structural details and in coloration is dis- cussed for each species and subspecies in the systematic accounts that follow. Sizes, proportions, and numbers of tooth-rows are tabulated in Table 2. 812 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. TABLE 2.—COMPARISON OF CERTAIN LARVAL CHARACTERS IN THE SPECIES OF PrycHoHyLa. (MEANs ARE IN PARENTHESES BELOW THE RANGES.) Number | Maximum! Head length Species of otal) | ee eothe specimens | length | Total length | TWS P. euthysanota euthysanota...... 23 40.8 30. 9-37 .3 4/6 (33.5) P. euthysanota macrotympanum. . 13 43.3 30. 6-33 .4 4/6 (32.7) P. leonhardschultzet............ 7 47.5 29 .2-32.7 4/6 (31.1) P. Spinipoller: sea waists te 30 32 45.0 30.2-35.9 4/7 (33.0) P. schmidtorum schmidtorum. .. . 14 42.5 28 .9-31.2 3/3 (29.9) P. schmidtorum chamulae....... 4 45.0 26 .9-29.3 3/3 (27.8) PPAgnacolones «scores icin Ne 2 39.6 29 .6-29.8 3/3 (29.7) Evidence on the pattern of development of tooth-rows indi- cates that the inner rows develop first. A small tadpole of P. euthysanota euthysanota has six lower rows and three fully de- veloped upper rows and only the beginning of the first (outer) upper row. A small tadpole of P. euthysanota macrotympanum has four upper rows and five lower rows. In a small tadpole of P. leonhardschultzei the three upper and four lower tooth-rows are well developed; the first upper and fifth lower rows are beginning to develop, and the sixth lower row is absent. In small tadpoles of P. spinipollex, the sixth lower row is poorly developed, and the seventh row is absent; large individuals normally have seven lower rows. A small tadpole of P. schmidtorum chamulae has % tooth- rows. Breeding Call Breeding calls of all species and subspecies of Ptychohyla were recorded in the field. Obtaining series of calls of Ptychohyla is difficult because these frogs call mostly from vegetation along roar- ing mountain streams and only by locating a calling frog some distance from the water or along a quiet stretch of the stream can good recordings be obtained. For example, four individuals of MrppLE AMERICAN TREE FROGS 818 P. spinipollex were recorded, but only one recording was sufficiently free of background noise to be analyzed. Analysis of breeding calls supports the division of the genus Ptychohyla into two groups of species. The call of each member of the Ptychohyla euthysanota group consists of a single long note, whereas the call of species in the Ptychohyla schmidtorum group consists of a series of short notes. Since no differences were found between the calls of subspecies of any given species, the following discussion of breeding calls pertains to species. These calls are de- scribed briefly below and at greater length in the systematic ac- counts farther on. Audiospectrographs of the breeding calls are shown in Plate 1, and comparisons of the characteristics of the calls are given in Table 3. TABLE 3.—COMPARISON OF THE BREEDING CALLS OF PTYCHOHYLA Notes Duration Pulses Frequency Dominant Species Number | per call of note per range frequency group (seconds) second (cps) (cps) Prt S PUTT POUCT oie 's) 2) eis /0.'s 1 1 .46 147 3000-5100 4300 P. euthysanota....... ai 1 .62 95.3 1800-4200 3070 (.60-.65) (91-102) (3000-3200) P. leonhardschultzei. . . 2 1 .79 77 1500-3500 2750 (.62-.95) (76-78) (2700-2800) P. schmidtorum....... 6 8.5* O84 110 1400-5800 3400 (8-9) (.054—.070) | (96-121) (3350-3450) PP SQnicolors).ciciee cls oe 2 12* .079 126 1000-5000 3150 (11-13) | (.078-.080) | (123-129) (3100-3200) *Only an analysis of the long series of calls is given here; see text for explanation. P. spinipollex (Pl. 1 A).—One long note is repeated at intervals of 45 seconds to four minutes and has an average dominant frequency of 4800 cycles per second. P. euthysanota (Pl. 1 B).—One long note is repeated six to nine times at intervals of 2.7 to 3.4 seconds and has an average dominant frequency of 3070 cycles per second. P. leonhardschultzei (Pl. 1 C).—One long note is repeated once after 10 to 13 seconds and has an average dominant frequency of 2750 cycles per second. P. schmidtorum (Pl. 1 D).—The complete call consists of one short series of notes alternating with two long series. Numbers of notes per series in one individual having a typical call were 5-8-8-3-9-9. The average dominant frequency of notes in the short and long series alike is 3400 cycles per second. P. ignicolor (Pl. 1 E).—The complete call consists of a short series of notes alternating with a long series. In one complete recording the numbers of notes in these series were 4-13-3-11. The notes in the short series have an average dominant frequency of 2100 cycles per second, whereas the notes in 314 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist. the long series have an average dominant frequency of 3150 cycles per second. The four series of notes were given in one minute and 15 seconds. SYSTEMATIC ACCOUNTS The museum catalogue numbers of the specimens examined, to- gether with the localities from which they came, are listed at the end of the account of each subspecies or monotypic species. The localities that are represented by symbols on the distribution map (Fig. 7) are in roman type; those that are not represented on the map, because overlapping of symbols would have occurred, are in italic type. Ptychohyla Taylor, 1944 Ptychohyla Taylor, Univ. Kansas Sci. Bull., 30:41, May 15, 1944. Type, Ptychohyla adipoventris Taylor, 1944 [= Hyla leonhardschultzei ( Ahl), 1934]. Diagnosis —Small hylids having stream-adapted tadpoles and differing from other hylid genera in having large ventrolateral glands in breeding males. Composition—Five species, two of which are made up of two subspecies, arranged in two groups of species on the basis of morphological characters of adults and tadpoles and on the basis of breeding calls. Distribution —NModerate elevations from southern Guerrero and northern Oaxaca, México, to northern El Salvador and central Honduras. Key To ADULTS 1. A weak tarsal fold; outer fingers one-third webbed; males having spiny nuptial tuberosities; color in life tan or brown with blotches or reticula- tions, never green; iris bronze color ......... P. euthysanota group—2 No tarsal fold; outer fingers having only vestige of web; males lacking nuptial tuberosities; color in life green or brown; iris red or golden COLOT pie ied ast hS0 A chee Pe Coe GENE cane P. schmidtorum group—5 2. Chest, throat, and flanks usually having brown or black spots; no dis- tinct white stripe on upper lip or on flanks; a faint white line usually present above anal opening; a rostral keel ...................+..---- 8 Chest, throat, and flanks usually unspotted; distinct white line on upper lip and on flank present or not; white line above anal opening faint or well. defined: noimostralkeel) 2st 15 o..,da0 Be neni oe ne 4 8. Interorbital region much wider than eyelid; spots on throat and chest black; spots only occasionally present on belly; flanks marbled with black and white; nuptial spines small, as many as 80 on one thumbs... ce Uh re Me ee AN) ee a P. leonhardschultzei Interorbital region about as wide as eyelid; spots on chest and throat brown or black; spots usually present on belly; flanks having round brown or black spots; nuptial spines moderate in size, conical, seldom more “than /60:on' one thumb.) 0.6 gets oe lah eee Pe P. spinipollex 4. A distinct, broad, white lateral stripe usually present; usually a dis- tinct white line above anal opening; a distinct white stripe on upper Hy 0 Ae, cudgel hess chan ee ML cate Aza P. euthysanota euthysanota No white lateral stripe; a faint white stripe above anal opening; no distinct white stripe on upper lip ........ P. euthysanota macrotympanum 5. A distinct, broad, lateral stripe; a white stripe on upper lip expanded to form a large spot below eye; hidden surfaces of thighs and webs of MippitE AMERICAN TREE FROGS 315 feet not red in life; internarial region slightly depressed; diameter of tympanum greater than one-half diameter of eye ................... 6 No lateral white stripe; no stripe on upper lip; in life dorsum green, hidden surfaces of thighs and webs of feet orange tan to bright red, and eye golden color; internarial region flat; diameter of tympanum less thanvoneshalf diameter iofveyes.....06 05208 eo: + eee ee ae eee P. ignicolor . Webs of feet and posterior surfaces of thighs pale cream color; dorsum in life reddish brown; iris bright red ........ P. schmidtorum schmidtorum Webs of feet and posterior surfaces of thighs pale brown; dorsum in life green; iris reddish bronze color ............... P. schmidtorum chamulae Kry To TADPOLES Lips greatly expanded forming a funnel-shaped mouth; tooth-rows %%4 ee it Se tess acer MO Ie P. schmidtorum group—2 Lips folded laterally, not forming a funnel-shaped mouth; tooth-rows WG IOLE MR EE OMe. ee he ek, Be EE ee P. euthysanota group—4 Belly and mouth mottled; tail cream color heavily blotched with Brown. Wetter See oe AL Nay Ge ee ee rs Se Belly dark gray; tail cream color with dense brown flecking, giving browneappearancey | hveke - 6 cde sce cree P. schmidtorum chamulae Belly cream color with brown mottling; no large tubercle at each end otsfirstilowerstooth-rows a). 22 nae dele soe P. schmidtorum schmidtorum Belly grayish green with brown mottling; a large tubercle at each end Ofthistwlowergtooth-row se! kc las a ee ae Bee P. ignicolor . Tooth-rows 4; cream-colored crescent-shaped mark on posterior part of body bordered posteriorly by large brown mark .................... Tooth-rows usually 44 (sometimes 46); cream-colored crescent-shaped mark on posterior part of body usually indistinct, not bordered pos- feriorly. byalarge brown Marks...) psec ee P. spinipollex Caudal musculature uniformly flecked with brown; lower tooth-rows 1-4 about equal in length to upper rows ...... P. euthysanota euthysanota Caudal musculature having brown square blotches dorsally on anterior one-half of tail; lower tooth-rows 1-4 usually slightly shorter than upper LOW SH ee Ce oR Teg Uy RS We Bir gelato ee eg Dorsal caudal blotches well defined and extending onto sides of tail; moderately large brown flecks on caudal fin; eye in life pale reddish PREG WING: eine 8 ee ek ne wk hk, nave Re eke P. leonhardschultzei Dorsal caudal blotches faint, not extending onto sides of tail; small brown flecks on caudal fin; eye in life silvery bronze................ P. euthysanota macrotympanum The Ptychohyla euthysanota Group Three species in group; adults having moderate amount of webbing be- tween fingers, and tarsal fold; breeding males having spinous, horny, nuptial tuberosities on pollex; mouths of tadpoles having lateral folds in lips and 4% or 44 tooth-rows; breeding call consisting of one long note. Ptychohyla euthysanota Diagnosis——Rostral kee] absent; nuptial spines in males small; interorbital region much wider than eyelid. Ptychohyla euthysanota euthysanota (Kellogg) Hyla euthysanota Kellogg, Proc. Biol. Soc. Washington, 41:123-124, June 29, 1928 [Holotype—USNM 73296 from Los Esemiles, Depto. Chala- tenango, El Salvador; Ruben A. Stirton collector]. Mertens, Sencken- 316 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist. bergiana, 33:169-171, June 15, 1952; Abhand. Senckenbergische Naturf. Gesell., 487:29, December 1, 1952. Stuart, Proc. Biol. Soc. Washington, 67:169, August 5, 1954. Hyla rozellae Taylor, Univ. Kansas Sci. Bull., 28:78-80, pl. 9, fig. 1, May 15, 1942 [Holotype—USNM_ 115089 from Salto de Agua, Chiapas, México; Hobart M. and Rozella Smith collectors]. Taylor and Smith, Proc. U. S. Natl. Mus., 95:587, June 30, 1945. Smith and Taylor, Bull. U.S. Natl. Mus., 194:86, June 17, 1948. Stuart, Proc. Biol. Soc. Wash- ington, 67:169, August 5, 1954. Ptychohyla bogerti Taylor, Amer. Mus. Novitates, 1437:13-16, fig. 5, De- cember 7, 1949 [Holotype—AMNH 51847 from Rio Grande, Oaxaca, México; Thomas MacDougall collector]. Stuart, Proc. Biol. Soc. Wash- ington, 67:169, August 5, 1954. Ptychohyla euthysanota, Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 13:351, April 27, 1961. Diagnosis.—Dorsum tan to reddish brown; venter white; rarely flecked with brown or black; a white stripe on upper lip, on flank, and usually above anus. Description.—The following description is based on KU 58008 from Finca La Paz, Depto. San Marcos, Guatemala (Pl. 2). Adult male having a snout- vent length of 35.0 mm.; tibia length, 16.5 mm.; tibia length/snout-vent length, 47.1 per cent; foot length, 14.2 mm.; head length, 11.0 mm.; head length/snout- vent length, 31.4 per cent; head width, 10.7 mm.; head with/snout-vent length, 80.6 per cent; diameter of eye, 3.3mm.; diameter of tympanum, 1.8 mm.; tympanum/eye, 54.5 per cent. Snout in lateral profile nearly square, slightly rounded above, and in dorsal profile bluntly rounded; canthus pronounced; loreal region moderately concave; lips thick, rounded, and slightly flaring; nostrils protuberant; internarial distance, 3.0 mm.; top of head flat; interorbital distance, 4.1 mm., and approximately a third broader than width of eyelid, 2.9mm. Moderately heavy dermal fold from posterior corner of eye above tympanum to point above insertion of forelimb, covering upper edge of tympa- num; tympanum round, its diameter slightly more than its distance from eye. Forearm moderately robust, having distinct dermal fold on wrist; dermal fold, but no row of tubercles along ventrolateral surface of forearm; pollex only slightly enlarged, bearing triangular shaped patch of small horn-covered spines (128 on right, 184 on left); second and fourth fingers equal in length; sub- articular tubercles round, distal one on fourth finger bifid; discs moderate in size, that of third finger equal to diameter of tympanum; no web between first and second fingers; other fingers one-third webbed. Heels broadly overlap when hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; low rounded tarsal fold; inner metatarsal tubercle large, elliptical, and flat; outer metatarsal tubercle small and round; low dermal fold from heel to disc of fifth toe; subarticular tubercles round; length of digits from shortest to long- est 1-2-5-3-4; third and fifth toes webbed to base of disc; fourth toe webbed to proximal end of penultimate phalanx; thin dermal fold from inner metatarsal tubercle to disc of first toe; disc smaller than on fingers. Anal opening at the level of the upper edge of thighs; anal flap short; anal opening bordered above by thin transverse dermal fold and laterally by heavy dermal fold. Skin of dorsum and ventral surfaces of forelimbs and shanks smooth; that of throat, belly, and ventral surfaces of thighs granular. Ventrolateral glands moderately developed, not reaching axilla or groin and broadly separated midventrally. Tongue ovoid, emarginate, and only slightly free posteriorly; vomerine teeth 2-2, situated on small triangular elevations between ovoid inner nares; openings MippLte AMERICAN TREE FROGS 317 to vocal sac large, one situated along inner posterior edge of each mandibular ramus. Dorsal ground-color of head, body, and limbs dull reddish brown with irregular dark brown reticulations on head and body and dark brown transverse bands on limbs; dorsal surfaces of first and second fingers and webbing on hand cream color; dorsal surfaces of third and fourth fingers dull brown; anterior surfaces of thighs dull creamy yellow; posterior surfaces of thighs dull brown; tarsi and toes tan with brown flecks; webbing of feet brown; faint creamy white stripe along lateral edges of tarsi and forearms; thin white line along edge of upper lip; distinct white stripe above and beside anal opening; axilla white; throat, chest, belly, and ventral surfaces of forelimbs creamy white; flanks white, separated from pale venter by a row of partly connected dark brown spots; ventral surfaces of thighs dull creamy yellow; feet grayish brown; ventrolateral glands pale grayish brown; small brown flecks on periphery of chin. In life the dorsal ground-color was pale reddish brown (Orange-Cin- namon); dorsal reticulations dark brown (Chocolate); dorsal surfaces of first and second fingers and webbing on hands creamy tan (Light Pinkish Cinna- mon); posterior surfaces of thighs reddish brown (Vinaceous-Tawny ); web- bing of feet gray (Deep Mouse Gray); throat and belly grayish white (Pale Gull Gray); ventral surfaces of hind limbs creamy white (Marguerite Yellow); spots on flanks dark brown (Warm Sepia); iris reddish bronze (Apricot Orange). Variation—No geographic variation in structural characters is discernible; variation in size and proportions is given in Table 1. Of 32 adults examined, seven have the tongue shallowly notched posteriorly; in the others the tongue is emarginate. Twenty specimens have a bifid subarticular tubercle beneath the fourth finger; in the others there are no bifid tubercles. The coloration described above is typical of the 16 specimens available from Finca La Paz. The living coloration at night, when the frogs were collected, was somewhat darker than the living colors described above, which were re- corded for the frogs the morning after collection, at which time one individual had a pale reddish brown dorsum (Orange-Cinnamon) with dull olive green (Deep Grape Green) reticulations on the back and transverse bands on the limbs; the dorsal surfaces of the first and second fingers and the discs on the third and fourth fingers were orange ( Mikado Orange). More than half of the specimens from Finca La Paz agree in all essential characters with the description given above. The distinctness of the white stripe on the upper lip is variable; in two individuals the stripe is barely dis- cernible. Likewise, in some individuals the white stripe on the flanks is not distinct, either because there are few or no brown spots separating the stripe from the pale venter, or because the ventrolateral gland has diffused the pale color on the flanks. There is some noticeable variation in dorsal coloration, either through a greater or lesser development of dark pigment. One speci- men (KU 58007) is grayish tan above with dark brown markings; the posterior surfaces of the thighs are dull grayish yellow; the first and second fingers and the webbing on the hands are pale yellowish gray; the belly and throat are dusty white; the flecks on the throat are gray; the ventral surfaces of the feet are grayish brown. Dark individuals, such as KU 58009 have a uniform dark brownish black dorsum; the belly is cream; the first and second 318 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. fingers and the webbing on the hands are dull creamy tan; the dorsal and ventral surfaces of the feet are dark brown. In KU 58013 there is a heavy suffusion of brown on the throat and flanks. Two specimens have scattered white flecks on the dorsum. The reddish brown dorsal ground-color with dark brown reticulations on the head and body and dark brown transverse bands on the limbs seems to be rather constant throughout the range of the subspecies. Likewise, the presence of the white stripe on the upper lip and the white stripe around the anal open- ing are present on most specimens. In breeding males having well-developed ventrolateral glands the lateral white stripe often is obliterated. Description of Tadpole-——The following description is based on KU 60042 from Finca La Paz, Depto. San Marcos, Guatemala (Figs. 4A and 6A). No limb buds; total length, 35.8 mm.; body length, 11.2mm.; body length/total length, 31.3 per cent. Body moderately depressed, slightly wider than deep, ovoid in dorsal profile; mouth directed ventrally; eyes small, directed dorso- laterally; nostrils slightly protuberant and directed anteriorly, closer to eye than snout; spiracle sinistral and posteroventrad to eye; anal tube dextral. Caudal fin low, rounded posteriorly; depth of caudal musculature about one-half greatest depth of caudal fin; musculature extends nearly to tip of tail. Mouth large; lips having deep lateral folds; two complete rows of papillae on lips; five to six rows of papillae laterally. Beaks moderately developed, bearing peglike serrations; slender lateral projections on upper beak; tooth- rows 4; upper rows subequal in length, second longest; fourth row interrupted medially; lower rows complete; lower rows 1-4 equal in length to upper rows; fifth lower row somewhat shorter; sixth lower row short. Body brown above; tip of snout cream color; grayish cream color below; caudal musculature creamy tan; caudal fin transparent; cream-colored crescent- shaped mark on posterior edge of body and anterior part of caudal musculature, bordered posteriorly by dark brown blotch; scattered brown flecks on caudal musculature and posterior part of caudal fin. Eye bronze color in life. Variation.—The variation in size and proportions is given in Table 2. In some specimens the first upper tooth-row is irregular, sometimes broken, and often shorter than other upper tooth-rows. Usually the fourth upper and first lower, and sometimes the sixth lower, tooth-rows are interrupted medially. One specimen has a short, irregular, seventh lower tooth-row; all others have six. The cream-colored crescent-shaped mark usually is distinct. The brown blotch posterior to this mark is variously shaped ranging from a narrow vertical bar to a triangular blotch. Brown flecks seldom are present on the anterior part of the ventral caudal fin. Comparisons.—Aside from the characters given in the diagnosis, P. euthy- sanota euthysanota can be distinguished from both P. spinipollex and P. leon- hardschultzei by the absence of bold black and white marbling on the flanks; furthermore, from the former it can be distinguished by having more and smaller horny nuptial spines and from the latter by having the snout, in lateral profile, rounded above and not acutely angulate. Ptychohyla euthy- sanota euthysanota differs from P. euthysanota macrotympanum by normally having a darker dorsal color, broader stripe on upper lip, and a distinct lateral stripe. Occurring sympatrically with Ptychohyla euthysanota euthysanota are several species of Plectrohyla, all of which differ in having a bony prepollex, rather MippLeE AMERICAN TREE FROGS 819 rugose skin on the dorsum, and more squat bodies. Other sympatric species are Ptychohyla schmidtorum schmidtorum, which lacks a tarsal fold and nuptial spines and has a red eye in life, Hyla salvadorensis, which has a green dorsum and lacks spinous nuptial tuberosities, and Hyla sumichrasti, a small yellow frog usually lacking vomerine teeth. Life History—This subspecies breeds in clear, swift mountain streams. Males call from stems and leaves of plants at the edge of, or overhanging, the streams. The breeding call consists of a soft “wraack,” repeated at intervals of three to four seconds. Each note has a duration of 0.60 to 0.65 seconds and has 91 to 102 pulses per second; the dominant frequency falls between 3000 and 3200 cycles per second. Tadpoles in various stages of development were found at Finca La Paz, Guatemala, in late July. This indicates that there is either extreme differential growth, or, more probably, an extended breed- ing season. A tadpole having a body length of 6.8 mm. and a total length of 19.1 mm. has a short median first upper tooth-row; lower tooth-rows 3-6 are only two-thirds as long as lower rows 1 and 2. Two recently metamorphosed young have snout-vent lengths of 14.2 and 14.8 mm.; they are colored like the adults. Remarks. — The type specimen of Hyla euthysanota Kellogg (1928:123) is a female; therefore, when Taylor (1944) proposed the name Ptychohyla for hylids having ventrolateral glands in breeding males, he was unaware that Hyla euthysanota was a member of this group. In his description of Hyla rozellae, Taylor (1942) did not compare his specimens with Hyla euthysanota, but instead placed H. rozellae with H. loquax and H. rickardsi. The type series of H. rozellae consists of one large adult female and several metamorphos- ing young. Taylor (1949:16) based the description of Ptychohyla bogerti on two males and compared these specimens with P. adipoventris Taylor [= P. leonhardschultzei (Ahl)]. Thus, in a period of 22 years the females of this species were given two names and the male another. Stuart (1954:169) suggested that Hyla euthysanota and Hyla rozellae were Ptychohyla. Now that sufficient specimens are available from throughout the range it is possible to determine that the various named populations are conspecific. Distribution.—This subspecies inhabits cloud forests at elevations of 660 to 2200 meters on the Pacific slopes of the Sierra Madre from extreme eastern Oaxaca and western Chiapas, México, through Guatemala to northern El Salvador; probably it occurs also in southwestern Honduras. Aside from the specimens listed below, three in the Frankfurt Museum from Depto. Santa Ana, El Salvador (44571, Hacienda San José; 48040, Hacienda Los Planes; 65119, Miramundo) are listed by Mertens (1952:29). 320 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Specimens examined.—MeExico: Chiapas: Cascarada, 30 km. W of Ciltapec, UMMZ 87851-2; Cerro Ovando, UMMZ 87853-4; Chicomuselo, UMMZ 94439- 40; Finca Juarez, 28 km. N of Escuintla, USNM 115052-5; Las Nubes, Cerro Ovando, USNM 115080-8; Salto de Agua, USNM 115089-51. Oaxaca: Cerro Pecho Blanco, UIMNH 40963; between La Gloria and Cerro Azul, UIMNH 40976-7; Rio Grande, AMNH 51847-8; Santo Tomd4s Tecpan, UIMNH 41071. GuaTEMALA: San Marcos: Finca La Paz, 2 km. W of La Reforma, KU 58001-14, 59937 (skeleton), 60042-3 (tadpoles), 60044 (4 young), MCZ 84997, UMMZ 107789, 123151-7 (tadpoles); Finca Pirineos, Rio Samala, CNHM 385066. Santa Rosa: Finca La Gloria, UMMZ 123148 (tadpoles), 123150 (tadpoles). Solold: Finca Santo Tom4s, UMMZ 123149 (tadpoles); Olas de Moca, near Moca, CNHM 20208. Ex Satvapor: Chalatenango: Los Esemiles, USNM 73296. Santa Ana: Miramundo, CNHM 65120. Ptychohyla euthysanota macrotympanum (Tanner) Hyla macrotympanum Tanner, Great Basin Nat., 17:52-53, July 31, 1957 [Holotype—AMNH 62141 (formerly BYU 13752) from 10 miles east of Chiapa de Corzo, Chiapas, México; Robert Bohlman collector]. Ptychohyla macrotympanum, Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 13:351, April 27, 1961. Diagnosis—Dorsum usually pale tan; venter white with scattered brown or black flecks; a thin white stripe on upper lip and another above anal opening; no distinct white stripe on flanks. Description—The following description is based on KU 58049 from Linda Vista, Chiapas, México (Pl. 3). Adult male having a snout-vent length of 38.0 mm.; tibia length, 19.5 mm.; tibia length/snout-vent length, 51.3 per cent; foot length, 15.7 mm.; head length, 11.8 mm.; head length/snout-vent length, 81.1 per cent; head width, 11.7 mm.; head width/snout-vent length, 30.8 per cent; diameter of eye, 3.8mm.; diameter of tympanum, 2.1mm.; tympa- num/eye, 55.2 per cent. Snout in lateral profile nearly square, slightly rounded above, and in dorsal profile bluntly rounded; canthus pronounced; loreal region concave; lips thick, rounded, and slightly flaring; nostrils pro- tuberant; internarial distance, 8.1 mm.; top of head flat; interorbital distance, 8.8 mm., and approximately a fourth broader than width of eyelid, 3.1 mm. A moderately heavy dermal fold from posterior corner of eye above tympanum to point above insertion of forelimb, covering upper edge of tympanum; tym- panum round, its diameter equal to its distance from eye. Forearm mod- erately robust, having a distinct dermal fold on wrist; dermal fold, but no row of tubercles along ventrolateral surface of forearm; pollex only slightly enlarged, bearing triangular patch of small horn covered spines (94 on right, 100 on left); fourth finger slightly longer than second; subarticular tubercles round, none bifid; discs moderate in size, that of third finger equal to diameter of tympanum; vestige of web between first and second fingers; other fingers one-third webbed. Heels broadly overlap when hind limbs adpressed; tibio- tarsal articulation reaches to middle of eye; weak tarsal fold on distal two- thirds of tarsus; inner metatarsal tubercle large, elliptical, and flat; outer metatarsal tubercle small and round; no dermal fold from heel to disc of fifth toe; subarticular tubercles round; length of digits from shortest to longest 1-2-5-3-4; third toe webbed to base of disc; fifth toe webbed to middle of penultimate phalanx; fourth toe webbed to proximal end of penultimate phalanx; no fold of skin from inner metatarsal tubercle to base of disc on first MipptE AMERICAN TREE FROGS 821 toe; discs much smaller than on fingers. Anal opening near upper edge of thighs; short anal flap bordered above by thin dermal fold; small tubercles and heavy dermal fold lateral to anal opening. Skin of dorsum and ventral surfaces of fore limbs and shanks smooth; that of throat, belly, and ventral surfaces of thighs granular. Ventrolateral glands weakly developed, not reaching axilla or groin and broadly separated midventrally. Tongue ovoid, shallowly notched posteriorly, and barely free behind; vomerine teeth 2-2, situated on small triangular elevations between ovoid inner nares; openings to vocal sac large, one situated along inner posterior edge of each mandibular ramus. Dorsal ground-color of head, body, and limbs pale pinkish tan with the greatest part of head and body covered by large gray interconnected blotches; black flecks over most of dorsum; grayish brown transverse bands on shanks; dorsal surfaces of first and second fingers pale grayish yellow; dorsal surfaces of third and fourth fingers and webbing on hand pale grayish tan; anterior surfaces of thighs pale flesh-color; posterior surfaces of thighs pale grayish yellow; dorsal surfaces of tarsi and toes pale grayish tan with black flecks; webbing of feet pale gray; faint creamy white stripes along ventrolateral edges of tarsi and forearms; a very thin white line along edge of upper lip; a narrow grayish white stripe above anal opening; axilla gray; throat, chest, belly, and ventral surfaces of forelimbs pale grayish white; ventral surfaces of hind limbs cream color; flanks gray flecked with brown; ventral surfaces of feet grayish tan; ventrolateral glands pinkish tan; anterior one-half of chin flecked with brown. In life the dorsum was pale tan (Pinkish Buff); the dark markings on dor- sum dull brown (Tawny-Olive); tarsi pale tan (Pale Pinkish Buff); flanks pinkish tan (Pale Cinnamon-Pink); iris coppery bronze (Capucine Orange). Variation—The few specimens from a limited geographic region preclude any analysis of geographic variation. All specimens, except the one described above, have the fifth toe webbed to the base of the disc. Many individuals have a bifid subarticular tubercle beneath the fourth finger. The shape of the posterior edge of the tongue varies from nearly straight and shallowly notched to bluntly rounded and emarginate. Two females (KU 58050-1) have more pointed snouts in dorsal profile than do males. Some specimens, such as KU 58048, are notably darker than the specimen described above; in dark specimens the dorsum is brown with dark brown markings; all fingers and the webbing on the hand are brown. The tarsi and feet are like those in the specimen described above, but the posterior surfaces of the thighs are yellowish tan heavily suffused with brown; the venter is cream color. In life KU 58048 had a brown (Verona Brown) dorsum with dark brown (Chocolate) markings. KU 58047 is slightly darker than KU 58048 and has scattered small white flecks on the dorsum. All specimens have the thin white line on the upper lip, but in some individuals it is in- distinct. The grayish white line above the anus is present in all specimens. Description of Tadpole—The following description is based on KU 60049 from Rio Hondo, 9.5 kilometers south of Pueblo Nuevo Solistahuacan, Chiapas, México (Figs. 4B and 6B). No limb buds; total length, 86.2mm.; body length, 11.1mm.; body length/total length, 80.6 per cent. Body moderately depressed, slightly wider than deep, ovoid in dorsal profile; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils slightly protuberant and 322 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. directed anteriorly, somewhat closer to eye than snout; spiracle sinistral and posteroventrad to eye; anal tube dextral. Caudal fin low, acutely rounded posteriorly; depth of caudal musculature slightly more than one-half greatest depth of caudal fin; caudal musculature extending nearly to tip of tail. Mouth large; lips having deep lateral folds; two complete rows of papillae on lips; five or six rows of papillae laterally. Beaks moderately developed, bearing small peglike serrations; moderately wide lateral projections on upper beak; tooth-rows 44; upper rows subequal in length; fourth row interrupted medially; length of lower rows 1-4 subequal to upper rows; fifth and sixth lower rows decreasing in length; first lower row interrupted medially. Body brown above; tip of snout cream color; venter creamy white; caudal musculature creamy tan; caudal fin transparent; cream-colored crescent-shaped mark on posterior edge of body and anterior part of caudal musculature, bordered posterodorsally by dark brown blotch; four dark brown blotches on dorsum of anterior part of caudal musculature; scattered brown flecks on caudal musculature and fin; eye silvery bronze in life. Variation —The variation in size and proportions is given in Table 2. All specimens have 4 tooth-rows; in some the first lower row is interrupted medially. Specimens from Jacaltenango and two kilometers west of San Pedro Necta, Depto. Huehuetenango, Guatemala, have weakly developed sixth lower tooth-rows. The cream-colored crescent-shaped mark is distinct in all specimens; the dorsal blotches on the anterior part of the caudal musculature are narrow and do not extend far onto the sides of the tail. The blotches are most distinct in small tadpoles and sometimes indistinct in large ones. Comparisons.—Ptychohyla euthysanota macrotympanum can be distinguished from both P. spinipollex and P. leonhardschultzei by the absence of bold black and white marbling on the flanks, as well as by the characters given in the diagnosis; furthermore, from the former it differs in having more and smaller horny nuptial tuberosities and from the latter by having the snout, in lateral profile, rounded above instead of angulate. Ptychohyla euthysanota macrotym- panum differs from P. e. euthysanota by normally having a paler dorsum, nar- rower stripe on upper lip, and no distinct lateral stripe. Ptychohyla euthysanota macrotympanum occurs sympatrically with Plectro- hyla guatemalensis and P. matudai matudai. Each of the last two has a bony prepollex, rather rugose skin on the dorsum, and more squat body. Other sympatric species are Hyla walkeri, which has a green dorsum with brown markings and a rather pointed snout, and Hyla sumichrasti, a small yellow frog usually lacking vomerine teeth. Life History—tThis species breeds in clear mountain streams in mixed pine and broad-leafed forest. Males call from trees and bushes along the streams. The breeding call consists of a soft “wraack,” repeated three to nine times with intervals of 2.7 to 3.4 seconds between notes. Each note has a duration of 0.60 to 0.65 second, and a rate of 92 to 100 pulses per second; the dominant frequency falls between 3000 and 3200 cycles per second (Pl. 1B). The call is indistinguishable from that of P. e. euthysanota. MippLeE AMERICAN TREE FROGS 323 Tadpoles in various stages of development were found in the Rio Hondo, Chiapas, on June 16, 1960. The smallest tadpole has a total length of 24.1 mm.; in this individual the sixth lower tooth row has barely started to develop. A metamorphosing frog taken at the same time has a snout-vent length of 19.8 mm., a short remnant of the tail, and the mouth and tongue developed, whereas another individual having a snout-vent length of 17.8 mm. and a tail 31.0 mm. in length still has larval teeth. Three completely metamorphosed juveniles collected by L. C. Stuart at Jacaltenango, Guatemala, on June 6 and 7, 1955, have snout-vent lengths of 16.0, 16.0, and 16.1 mm. Remarks.—Tanner (1957:52) based the description of Hyla macrotympanum on a single female, which, of course, lacked the characters diagnostic of Ptychohyla. On the basis of general ex- ternal characters Tanner suggested that Hyla macrotympanum was related to H. miotympanum, from which it differed in having a larger tympanum and a bifid subarticular tubercle beneath the fourth finger. The collection of additional females, together with males of the species, has shown that Hyla macrotympanum is a Ptychohyla. Intergradation between Ptychohyla euthysanota and P. macrotym- panum has not been demonstrated. Currently their ranges are separated by the dry valleys of the Rio Grijalva and Rio Cuilco. The similarity in structure of the adults and tadpoles and the in- distinguishable breeding calls are the basis for considering the two populations subspecies. Distribution——This species occurs in mixed pine and broad-leafed forests at elevations of 700 to 1700 meters on the southern slopes of the Chiapan High- lands and Sierra de Cuchumatanes, in Guatemala. These forests are on the south facing slopes north of the valleys of the Rio Grijalva and Rio Cuilco. Specimens examined.—MeExico: Chiapas: 6 km. NE of Chiapa de Corzo, TCWC 16188; 16 km. E of Chiapa de Corzo, AMNH 62141; Linda Vista, 2 km. NW of Pueblo Nuevo Solistahuacan, KU 58049-51, 59939 (skeleton); Rio Hondo, 9.5 km. S of Pueblo Nuevo Solistahuacédn, KU 58047-8, 60046-7, 60048-9 (tadpoles); San Fernando, MZTG 15, 17; Tonina (ruins), KU 41592. GuaTeMALA: Huehuetenango: Jacaltenango, UMMZ 123139 (tadpoles); 0.5 km. E of Jacaltenango, UMMZ 123142-3; 2 km. S of Jacaltenango, UMMZ 123141; 2 km. W of San Pedro Necta, UMMZ 123140 (tadpoles). Ptychohyla leonhardschultzei (Ahl) Hyla leonhard-schultzei Ah\, Zool. Anz., 106:185-186, fig. 1, April 15, 1934 [Holotype.—ZMB 34353 from Malinaltepec, Guerrero, México; Leonhard Schultze collector]. Smith and Taylor, Bull. U. S. Natl. Mus., 184:87, June 17, 1948. Hyla godmani, Ahl, Zool. Anz., 106:186, April 15, 1934. Hyla pinorum Taylor, Proc. Biol. Soc. Washington, 50:46-48, pl. 2, fig. 2, 824 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist. April 21, 1987 [Holotype—UIMNH 25049 from Agua del Obispo, Guer- rero, México; Edward H. Taylor collector]. Smith and Taylor, Bull. U. S. Natl. Mus., 194:87, June 17, 1948. Ptychohyla adipoventris Taylor, Univ. Kansas Sci. Bull., 30:41-45, May 15, 1944 [Holotype—UIMNH 25047 from Agua del Obispo, Guerrero, México; Edward T. Taylor collector]. Smith and Taylor, Bull. U.S. Natl. Mus., 194:91, June 17, 1948. Taylor, Amer. Mus. Novitates, 1437:16, December 7, 1949. Stuart, Proc. Biol. Soc. Washington, 67:169, August 5, 1954. Hyla milleri Shannon, Proc. U. S. Natl. Mus., 101:473-477, figs. 92b, 98a-c, May 17, 1951 [Holotype-—USNM 123700 from San Lucas Camotlan, Oaxaca, México; Walter S. Miller collector]. Ptychohyla leonhard-schultzei, Duellman, Herpetologica, 16:191-197, figs. 1-8, September 23, 1960; Univ. Kansas Publ. Mus. Nat. Hist., 13:351, April 27, 1961. Diagnosis.—Rostral kee] present; snout in lateral profile not rounded above; interorbital region much broader than eyelid; distal subarticular tubercle beneath fourth finger bifid or double; no white stripe on edge of upper lip; flanks white with black spots. Description——The following description is based on KU 64117 from Vista Hermosa, Oaxaca, México (Pl. 4). Adult male having a snout-vent length of 35.6 mm.; tibia length, 18.0 mm.; tibia length/snout-vent length, 50.5 per cent; foot length, 14.83mm.; head length, 10.7 mm.; head length/snout-vent length, 30.1 per cent; head width, 10.6 mm.; head width/snout-vent length, 29.8 per cent; diameter of eye, 3.6mm.; diameter of tympanum, 1.8 mm.; tympanum/eye, 50.0 per cent. Snout in lateral profile square, not rounded above, and in dorsal profile rounded with pointed tip resulting from vertical rostral keel; canthus pronounced; loreal region barely concave; lips thick, rounded, and barely flaring; nostrils protuberant; internarial distance, 3.2 mm.; top of head flat; interorbital distance, 3.8mm., and approximately a fifth broader than width of eyelid, 3.2mm. A moderately heavy dermal fold from posterior corner of eye above tympanum and curving ventrad to anterior edge of insertion of forelimb, covering upper edge of tympanum; tympanum round, its diameter equal to its distance from eye. Forearm moderately robust, hav- ing distinct dermal fold on wrist; row of small, low, rounded tubercles along ventrolateral surface of forearm; pollex only slightly enlarged, bearing triangular patch of small horn-covered spines (56 on right, 62 on left); second finger noticeably shorter than fourth; subarticular tubercles round, distal ones on third and fourth toes bifid; discs moderate in size, that of third finger slightly larger than diameter of tympanum; no web between first and second fingers; other fingers one-third webbed. Heels broadly overlap when hind limbs ad- pressed; tibiotarsal articulation reaches to middle of eye; a low rounded tarsal fold on distal half of tarsus; inner metatarsal tubercle elevated, flat, and ellipti- cal; outer metatarsal tubercle at base of fourth toe, round; row of low, some- times indistinct, tubercles from heel to base of fifth toe; subarticular tubercles round; length of digits from shortest to longest 1-2-3-5-4, third and fifth be- ing about equal in length; third and fifth toes webbed to base of disc; fourth toe webbed to base of penultimate phalanx; discs of toes much smaller than on fingers. Anal opening near dorsal surface of thighs; short anal flap; opening bordered laterally by heavy dermal fold and ventrolaterally by large tubercles. Skin of dorsum and ventral surfaces of forelimbs and shanks smooth; that of throat, belly, and ventral surfaces of thighs granular. Ventrolateral glands MippLE AMERICAN TREE FROGS 825 moderately developed, reaching axilla but not to groin and broadly separated midventrally. Tongue cordiform, shallowly notched behind and barely free posteriorly; vomerine teeth 4-3, situated on transverse elevations between ovoid inner nares; openings to vocal sac large, one situated along inner posterior edge of each mandibular ramus. Dorsal ground-color of head, body, and limbs pale tan with large inter- connected dark brown blotches on head and body and broad dark brown transverse bands on limbs; dorsal surfaces of first and second fingers and of webbing of hands pale brown; dorsal surfaces of third and fourth fingers dark brown; anterior surfaces of thighs flesh-color; posterior surfaces of thighs brown; dorsal surfaces of tarsi and feet dark brown; narrow white stripe along ventrolateral edges of forearms and tarsi; a faint creamy white stripe above anal opening; axilla white; flanks creamy white, boldly spotted with black; throat and chest white; ventral surfaces of tarsi and feet dark brown; other ventral surfaces dusty cream color; large brown spots on chin and anterior part of abdomen. In life the dorsum was reddish brown (Orange-Cinnamon) with dark brown (Chocolate) blotches; first and second fingers and webbing on hand pale reddish brown (Cinnamon); webbing on feet dark brown (Clove Brown); flanks pale creamy white (Pale Olive Buff) with dark brown (Bone Brown) spots; iris reddish bronze (Apricot Orange). Variation—No noticeable geographic variation is apparent in the few avail- able specimens; variations in proportions are given in Table 1. The distal subarticular tubercle of the fourth finger is either bifid or double in all speci- mens; that on the third finger usually is bifid, sometimes single. The vertical rostral keel is prominent in all freshly preserved specimens; in some older specimens it is indistinct. The tongue always is notched posteriorly; in some individuals the notch is shallow; in others it is deep. Some specimens are paler and less boldly marked than the specimen de- scribed above. All specimens from Agua del Obispo and some specimens from the northern slopes of the Sierra Madre Oriental in Oaxaca have a pale tan dorsum with brown markings. In most individuals the white color in the axilla extends onto the posterior edge of the upper arm. The creamy white color of the flanks is constant and usually extends slightly dorsad in the inguinal region. The white stripe above, and sometimes continuing down be- side, the anal opening varies from a thin indistinct line or row of flecks to a distinct continuous stripe. Two specimens have dark brown spots on the belly; in the others the spots are confined to the flanks and throat. Description of tadpole—tThe following description is based on KU 68556 from 7.5 kilometers south of Yetla, Oaxaca, México (Figs. 4C and 6C). No limb buds; total length, 37.3mm.; body length, 12.2mm.; body length/total length, 32.7 per cent; body slightly depressed, barely wider than deep, ovoid in dorsal profile; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils barely protuberant and directed anterolaterally, about midway between snout and eye; spiracle sinistral and posteroventrad to eye; anal tube dextral. Caudal fin low, bluntly rounded posteriorly; greatest depth of caudal muscula- ture about one-half depth of caudal fin; musculature extends nearly to tip of tail. Mouth large; lips having deep lateral folds; two complete rows of papillae on lips; five to seven rows of papillae laterally; beaks moderately developed, 326 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. bearing short peglike serrations; moderately wide lateral projections on upper- beak; tooth-rows 44; upper rows subequal in length; fourth row interrupted’ medially; length of lower rows 1-4 equal to upper rows; fifth and sixth lower rows shorter; first lower row interrupted medially. Body brown above; tip of snout brown; venter creamy gray; caudal muscula-. ture creamy tan; caudal fin transparent; cream-colored crescent-shaped mark on posterior edge of body; dark brown flecks on caudal musculature and all except anterior two-thirds of ventral caudal fin; large brown square blotches on dor-- sum of caudal fin; eye reddish brown in life. Variation.—The variation in size and proportions is given in Table 2. With. the exception of one specimen having a short, broken, seventh tooth-row, all specimens have 46 tooth-rows that are like those described above. In some: specimens the brown blotches on the dorsum of the caudal musculature are: fused and marked with cream-colored flecks. Comparisons.—Ptychohyla leonhardschultzei differs from all other members: of the Ptychohyla euthysanota group in having a square snout, and further: differs from P. spinipollex in more numerous and smaller nuptial spines and in transverse, instead of posteromedially slanting, vomerine processes between the inner nares. Ptychohyla leonhardschultzei differs from P. euthysanota im having a rostral keel and in having white flanks boldly spotted with black. All small hylids that are sympatric with Ptychohyla leonhardschultzei are- either yellow (Hyla dendroscarta and H. melanomma) or green (Hyla eryth- romma, which has a red eye, Hyla hazelae, which has a black line on the: canthus, and Ptychohyla ignicolor, which has red flash colors on the thighs). Life History—This frog has been found along streams in cloud’ forests and in pine-oak forest. Males call from vegetation along the stream or from rocks in and at the edge of the stream. The call is a single, long, soft “wraack,” repeated at intervals of anywhere: from several seconds to three or four minutes. Each note has a duration of 0.62 to 0.95 of a second and a rate of 76 to 78 pulses per second; the dominant frequency falls between 2700 and 2800: cycles per second (PI. 1C). Tadpoles were found in several streams in northern Oaxaca. A. tadpole having a total length of 21.1 mm. has three upper and four: lower tooth-rows well developed; the fourth upper and fifth lower- rows are weakly present, and the sixth lower row has not started to develop. Two metamorphosed young have snout-vent lengths of 15.2 and 15.5 mm. Remarks.—Four specific names have been applied to this species. Ahl (1934:185) based his description of Hyla leonhardschultzei on: a small, poorly preserved female. Taylor (1944:41) proposed the- generic name Ptychohyla for a species (named therein as P. adi-. poventris) of hylid having ventrolateral glands and horn-covered nuptial spines. Obviously, Taylor was unaware that Hyla leonhard- schultzei was the same species. Earlier Taylor (1937:46) described: MippLE AMERICAN TREE FROGS 327 Hyla pinorum. The types of all of these species came from the Pacific slopes of the Sierra del Sur in Guerrero. Examination of the types and other available specimens shows that they are repre- sentatives of a single species. The type of Hyla pinorum is an im- mature male having a snout-vent length of 26.7mm. All of these specimens have the square snout and black and white flanks char- acteristic of Ptychohyla leonhard-schultzei. Although Shannon (1951:473) based his description of Hyla milleri on a male having well-developed ventrolateral glands, he overlooked the presence of these glands in his description and discussion of relationships. The acquisition of more specimens from northern Oaxaca has shown that Hyla milleri is the same as Ptychohyla leonhardschulizei. Distribution —This species is known from pine-oak forest and cloud forest on the Pacific slopes of the Sierra Madre del Sur in Guerrero and Oaxaca and from the Atlantic slopes of the Sierra Madre Oriental in northern Oaxaca. Specimens have been collected at elevations between 700 and 1650 meters. Probably the species occurs in humid forests at similar elevations around the eastern end of the Mexican Highlands in Oaxaca. Specimens examined.—MEeExico: Guerrero: Agua del Obispo, CNHM 123489-90, 126651, 106300, MCZ 29639, UIMNH 25047, 25049, USNM 114551; Malinaltepec, ZMB 34351, 34353. Oaxaca: 2.5 km. N of La Soledad, KU 58061; San Lucas Camotlan, UIMNH 3201, USNM 128700-1; Vista Her- mosa, KU 64116-7, 64119, 68560 (tadpoles), 71344, 71717-8 (tadpoles), UMMZ 119604; 5 km. S of Yetla, KU 60045 (tadpoles); 7.5 km. S of Yetla, KU 64118, 68556-7 (tadpoles), 68559 (tadpoles), 68561 (2 young), 68630 (skeleton), UMMZ 115514-5, 118863 (tadpoles); 9 km. S of Yetla, KU 68558 (tadpoles ). Ptychohyla spinipollex (Schmidt) Hyla euthysanota, Dunn and Emlen, Proc. Acad. Nat. Sci. Philadelphia, 84:25, March 22, 1932. Hyla spinipollex Schmidt, Proc. Biol. Soc. Washington, 49:45-46, May 1, 1986 [Holotype—MCZ 21300 from the mountains behind Ceiba, Depto. Atlantidad, Honduras; Raymond E. Stadelman collector]. Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, 69:32-34, figs. 5-6, June 12, 1948; Contr. Lab. Vert. Biol. Univ. Michigan, 45:22, 52, 54, 57, May, 1950; Proc. Biol. Soc. Washington, 67:169, August 5, 1954. Ptychohyla spinipollex, Stuart, Contr. Lab. Vert. Biol. Univ. Michigan, 68: 48, November, 1954. Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 18:351, April 27, 1961. Diagnosis.—Rostral keel present; snout in lateral profile rounded above; eyelid nearly as wide as interorbital region; flanks white with brown spots; belly spotted; nuptial spines pointed and moderate in size. Description.—The following description is based on KU 58054 from Finca Los Alpes, Depto. Alta Verapaz, Guatemala (Pl. 5). Adult male having a snout-vent length of 37.7mm.; tibia length, 18.2mm.; tibia length/snout- vent length, 48.2 per cent; foot length, 15.8mm.; head length, 11.7 mm.: head length/snout-vent length, 31.0 per cent; head width, 11.7 mm.; head width/snout-vent length, 31.0 per cent; diameter of eye, 3.6mm.; diameter of tympanum, 1.9 mm.; tympanum/eye, 52.7 per cent. Snout in lateral profile 828 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist. nearly square, slightly rounded above, and in dorsal profile rounded with a pointed tip resulting from vertical rostral keel; canthus pronounced; loreal region barely concave; lips thick, rounded, and barely flaring; nostrils pro- tuberant; internarial distance, 3.0 mm.; top of head flat; interorbital distance, 3.7 mm., about equal to width of eyelid, 3.6mm. A heavy dermal fold from posterior corner of eye above tympanum to point above insertion of forearm, covering upper edge of tympanum; tympanum round, its diameter equal to its distance from eye. Forearm moderately robust, having faint dermal fold on wrist; row of low, rounded tubercles along ventrolateral edge of forearm; pollex only slightly enlarged, bearing triangular patch of moderate-sized, pointed, horn-covered spines (38 on right, 43 on left); second finger notice- ably shorter than fourth; subarticular tubercles round, distal one on fourth finger bifid; discs of fingers of moderate size, that of third finger slightly smaller than diameter of tympanum; vestigial web between first and second fingers; other fingers one-third webbed. Heels broadly overlapping when hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; dis- tinct, but low, tarsal fold extending length of tarsus; inner metatarsal tubercle elevated, flat, and elliptical; outer metatarsal tubercle small and round, near base of fourth toe; row of low indistinct tubercles from heel to base of fifth toe; subarticular tubercles round; length of toes from shortest to longest 1-2-3-5-4, the third and fifth being about equal in length; third and fifth toes webbed to base of disc; fourth toe webbed to base of penultimate phalanx; discs of toes slightly smaller than those on fingers. Anal opening near upper edge of thighs; opening bordered laterally by moderately heavy dermal folds and ventrolaterally by tubercles. Skin of dorsum and ventral surfaces of forelimbs and shanks smooth; that of throat, belly, and ventral surfaces of thighs granular. Ventrolateral glands barely evident. Tongue ovoid, barely notched behind and slightly free posteriorly; vomerine teeth 2-8, situated on \, /-shaped elevations between round inner nares; openings to vocal sac large, one situated along inner posterior edge of each mandibular ramus. Dorsal ground-color of head, body, and limbs grayish tan with dark brown reticulations on head and body and dark brown transverse bars or spots on limbs; first and second fingers cream color; third and fourth fingers and webbing on hands pale grayish brown; anterior surfaces of thighs reddish tan; posterior surfaces of thighs yellowish tan; tarsi and toes pale grayish tan with brown flecks; webbing on foot pale brown; faint white stripe along ventro- lateral edges of tarsi and forearms; narrow white line above and beside anal opening; no white stripe on edge of upper lip; axilla pale flesh-color; throat, chest, and ventral surfaces of limbs pale creamy gray; belly white with scattered brown flecks; flanks grayish white with dark brown flecks; ventral surfaces of tarsi dark brown; ventrolateral glands grayish tan. In life the dorsal ground-color of the head, body, fore limbs, and thighs was yellowish tan (Pinkish Buff); dorsal surfaces of shanks and tarsi pale yellowish tan (Pale Pinkish Buff); markings on head and back brown (Snuff Brown) to dark brown (Chocolate); dark bands on limbs brown (Tawny- Olive); first and second fingers creamy tan (Light Pinkish Cinnamon); posterior surfaces of thighs creamy tan (Light Pinkish Cinnamon); third and fourth fingers and webbing on hand grayish brown (Avellaneous); webbing on feet dark brown (Olive Brown); axilla pale pink (Hydrangea Pink); flanks buff PLATE 11 4 Mss se dalla COCO CC RAE RIAA PRT ata KIEOGY. CEES (PERS SECOND 0:2 0:4 TIME IN SECONDS Audiospectrographs of the breeding calls of the five species of Ptychohyla: (A) P. spinipollex (KU Tape No. 41), (B) P. euthysanota macrotympanum (KU Tape No. 48), (C) P. leonhardschultzei (UMMZ Tape No. 525), (D) P. schmidtorum chamulae (KU Tape No. 52), (E) P. ignicolor (UMMZ Tape No. 526). PEATE, 12 Ptychohyla euthysanota euthysanota (KU 58008 ). Sek, PICA EAS B34 & Ptychohyla euthysanota macrotympanum (KU 58049). > 2. PLATE 14 Ptychohyla leonhardschultzei (KU 64117). X 2. PLATE 15 Ptychohyla spinipollex (KU 58054). x 2. PLATE 16 a SS # ») Ptychohyla schmidtorum schmidtorum (KU 58043). x 2. PLAGE, 7, Ptychohyla schmidtorum chamulae (KU 58069). x 2. PLATE 18 Ptychohyla ignicolor (UMMZ 119603). Sez: MippLE AMERICAN TREE FROGS 829 (Cream-Buff) becoming yellow (Lemon Chrome) in groin; spots on flanks dark brown (Clove Brown); iris dull grayish bronze (Orange-Citrine). Variation —The distal subarticular tubercle beneath the fourth finger is bifid in about two-thirds of the specimens; in the rest it is round. The posterior edge of the tongue varies from being emarginate to shallowly notched. In most specimens the row of tubercles along the outer edge of the tarsus is made up of discrete tubercles, but in some individuals the tubercles form a nearly continuous dermal fold. Most specimens have the vomerine teeth situated on \_ /-shaped elevations, but in some individuals the elevations are more nearly transversely situated between the inner nares. All 42 specimens from Finca Los Alpes, Guatemala, have dark brown spots and flecks on the venter. Some individuals have only a few flecks on the throat and a few large spots on the flanks, as does KU 64125. Other specimens, such as KU 64132, have dense spotting over the entire venter. The color of the dorsum varies from pale tan to dark brown with darker brown markings; the white line above the anus is present in all specimens, but in some it is indistinct. KU 58058 and 64127 have a dark brown dorsum with large pale tan, square blotches; in life the blotches were pale tan (Pinkish Buff); the rest of the dorsum was dark brown (Sayal Brown). KU 58052 is dark brown with many small white flecks on the dorsum; in life the dorsum was deep olive brown (Dark Olive). Aside from the differences mentioned above, all specimens from Guatemala are similar in coloration. Three specimens from Honduras (MCZ 21300 and UMMZ 113102-3) have unspotted white venters. MCZ 21300, the holotype of P. spinipollex, lacks a white stripe above the anal opening, whereas the stripe is indistinct in UMMZ 113102-3. Description of tadpole—The following description is based on KU 60053 from Fina Los Alpes, Depto. Alta Verapaz, Guatemala (Figs. 4D and 6D). No limb buds; total length, 37.2 mm.; body length, 12.2 mm.; body length/total length, 32.8 per cent. Body rounded, not depressed, as wide as deep, ovoid in dorsal profile; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils barely protuberant and directed anterolaterally, somewhat closer to eye than snout; spiracle sinistral and posteroventrad to eye; anal tube dextral. Caudal fin low, bluntly rounded posteriorly; greatest depth of caudal muscula- ture about one-half depth of caudal fin; musculature extends nearly to tip of tail. Mouth large; lips having deep lateral folds; two complete rows of papillae on lips; six or seven rows of papillae laterally; beaks moderately developed, bearing fine blunt serrations; slender lateral projections on upper beak; tooth- rows 44; upper rows subequal in length; fourth row interrupted medially; lower rows 1-4 equal in length to upper rows; lower rows 5-7 decreasing in length; first lower row interrupted medially. Top of head and tip of snout brown; venter creamy gray; caudal muscula- ture tan; caudal fin transparent; faint cream-colored, narrow, crescent-shaped mark on posterior edge of body, not bordered posteriorly by dark brown mark; brown flecks scattered on caudal musculature and caudal fin; only a few flecks on anterior half of ventral caudal fin; eye bronze-color in life. Variation —The variation in size and proportions as compared with tadpoles of other species is given in Table 2. Of the 57 tadpoles of this species that I 2—6531 336 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. have examined, 21 have only six lower tooth-rows, although in some of these specimens a faint ridge for a seventh row is present. In those specimens hav- ing seven lower rows, the seventh often is broken. There is considerable variation in coloration. None has a distinct cream- colored, crescent-shaped mark bordered posteriorly by a dark brown bar or triangle, as in the other species in the Ptychohyla euthysanota group. Most specimens have a rather indistinct crescent; some have no crescent. In a few specimens there is a weakly outlined dark mark posterior to the crescent. Some specimens in a series of tadpoles from 32 kilometers north of Morazan, Baja Verapaz, Guatemala, have faint dorsal blotches on the dorsal musculature, much like those in tadpoles of Ptychohyla leonhardschultzei. Comparisons.—Ptychohyla spinipollex differs from all other species in the genus by having moderate-sized, instead of small, pointed nuptial spines; also it has fewer spines than the other species (see discussion of this character in Analysis of Data). The nearly equal interorbital breadth and width of the upper eyelid also is diagnostic of this species. Other hylids sympatric with Ptychohyla spinipollex include three species of Plectrohyla, each of which has a bony prepollex, heavy body, and rugose skin on the dorsum. The only other sympatric hylid that could be confused with Ptychohyla spinipollex is Hyla bromeliacea, which has a round snout and yellowish tan dorsum not marked with dark brown. Life History—At Finca Los Alpes, Guatemala, in July, 1960, and in August, 1961, calling males were found on bushes and trees along cascading mountain streams. The breeding call consists of a soft “wraack,” repeated at intervals of 45 seconds to four minutes. Each note has a duration of about .46 of a second and a rate of 147 pulses per second. The dominant frequency is 4300 cycles per second (RIVA), Tadpoles have been found in cascading mountain streams. Two metamorphosed young have snout-vent lengths of 15.0 and 15.5 mm. Remarks.—There is little doubt that all of the specimens herein referred to Ptychohyla spinipollex are conspecific. However, the three specimens from Honduras, including the type of Ptychohyla spinipollex, differ from Guatemalan specimens in lacking all dark spotting on the venter. Additional specimens from Honduras and eastern Guatemala may show that two subspecies are recognizable, in which case the nominal subspecies will be the population in Honduras. Distribution—tThis species lives in cloud forests at elevations of 800 to 1700 meters on the Atlantic side of the Guatemalan Highlands from the Sierra de Cuchumatanes in western Guatemala eastward to central Honduras. Specimens examined.—GuaTEMALA: Alta Verapaz: Finca Chichén, UMMZ 90876 (tadpoles); Finca Los Alpes, KU 58052-60, 59939 (skeleton), 60053 (tadpoles), 64122-41, 68562, 68563 (tadpoles), 68631-2 (skeletons), MCZ 35000-1, UMMZ 90873, 90874 (3), 90875 (tadpoles); La Primavera, UMMZ 90877 (tadpoles); Panzamali, UMMZ 90878 (tadpoles). Baja Verapaz: 82 MippLE AMERICAN TREE FROGS 331 km. N of Morazdn, KU 68564 (tadpoles); Santa Elena, UMMZ 98119, 98120 (2). Huehuetenango: 1 km. E of Barillas, UMMZ 123136-7 (tadpoles ). Progreso: Finca Bucaral, UMMZ 106783 (3), 123138 (tadpoles), S-1292 (skelton). Honpuras: Atlantidad: Mountains behind Ceiba, MCZ 21300. Morazan: Cerro Uyuca, UMMZ 123102-3. The Ptychohyla schmidtorum Group Two species in group; adults having only vestige of web between fingers and lacking tarsal fold; pollex of breeding males lacking spinous, horny, nuptial tuberosities; mouth of tadpole greatly expanded, funnel-shaped, lacking lateral folds, and having 34 tooth-rows; breeding call consisting of series of short notes. Ptychohyla schmidtorum Diagnosis—Diameter of tympanum more than half of diameter of eye; internarial region depressed; toes three-fourths webbed; no red flash-colors on thighs. Ptychohyla schmidtorum schmidtorum Stuart Ptychohyla schmidtorum Stuart, Proc. Biol. Soc. Washington, 67:169-172, August 5, 1954 [Holotype—CNHM 27055 from EI] Porvenir (17 kilo- meters air-line west of San Marcos), Depto. San Marcos, Guatemala; Karl P. Schmidt collector]. Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 13:351, 355, April 27, 1961. Diagnosis—Vomerine teeth 5-11; dorsum dark brown; white spot below eye; eye red in life. Description—The following description is based on KU 58043 from Finca La Paz, Depto. San Marcos, Guatemala (Pl. 6). Adult male having snout- vent length of 31.6 mm.; tibia length, 15.0 mm.; tibia length/snout-vent length, 47.5 per cent; foot length, 12.5 mm.; head length, 10.2 mm.; head length/snout- vent length, 32.3 per cent; head width, 9.9mm.; head width/snout-vent length, 81.3 per cent; diameter of eye, 3.4mm.; diameter of tympanum, 1.8 mm.; tympanum/eye, 52.9 per cent. Snout in lateral profile nearly square, slightly rounded above and below, and in dorsal profile bluntly squared; canthus pro- nounced; loreal region concave; lips thick, rounded, and flaring; nostrils pro- tuberant; internarial distance, 2.2mm.; internarial region depressed; top of head flat; interorbital distance, 3.4mm., much greater than width of eyelid, 2.5mm. Thin dermal fold from posterior corner of eye above tympanum to insertion of forelimb, covering upper edge of tympanum; tympanum round, its diameter equal to its distance from eye. Forearm slender, lacking distinct dermal fold on wrist; row of low rounded tubercles along ventrolateral edge of forearm; pollex slightly enlarged; no nuptial spines; second and fourth fingers about equal in length; subarticular tubercles small and round, distal one beneath fourth finger bifid; discs small, that of third finger noticeably smaller than tympanum; no web between first and second fingers; vestige of web between other fingers. Heels overlap when hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; no tarsal fold; inner metatarsal tubercle large, flat, and elliptical; outer metatarsal tubercle small, ovoid, slightly more distal than inner; subarticular tubercles round; length of digits from shortest to longest 1-2-5-3-4; third and fifth toes webbed to base of discs; fourth toe webbed to base of penultimate phalanx; discs of toes smaller 302 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. than on fingers. Anal opening directed posteriorly at upper edge of thighs; no anal flap; pair of large tubercles below anal opening and smaller tubercles farther below. Skin of dorsum and ventral surfaces of forelimbs and shanks smooth; that of belly and ventral surfaces of thighs granular. Ventrolateral glands well developed, reaching axilla and groin and narrowly separated on chest. Tongue ovoid, emarginate posteriorly, and only slightly free behind; vomerine teeth 3-3, situated on small triangular elevations between ovid inner nares; openings to vocal sac large, one situated along inner posterior edge of each mandibular ramus. Dorsum of head, body, and limbs reddish brown with indistinct, irregular darker brown markings on body and dark brown transverse bands or spots on limbs; first and second fingers creamy white; third and fourth fingers brown; dorsal surfaces of tarsi and third, fourth, and fifth toes tan with brown spots; first and second toes and webbing on feet creamy tan; enamel-white stripe along edge of upper lip continuing over, and on posterior edge of, forearm to groin, expanded to form spot below eye; belly white, unspotted; ventrolateral glands pale brown; ventral surfaces of hind limbs and anterior and posterior surfaces of thighs cream color; enamel-white stripe on heel; creamy white stripe along ventrolateral edges of tarsi and forearms. In life dorsum reddish brown (Terra Cotta) with dark brown (Burnt Umber) markings; first and second fingers and first and second toes orange- yellow (Light Orange-Yellow); posterior surfaces of thighs pale reddish tan (Ochraceous-Salmon); webbing on feet yellowish tan (Deep Colonial Buff); belly white; iris red (Nopal Red). Variation.—Little variation in structural characters was observed. All but five specimens have bifid subarticular tubercles beneath the fourth finger. Three specimens have cordiform tongues, and in four others the tongue is ovoid and shallowly notched behind; all other specimens have an emarginate ovoid tongue. Some individuals when active at night had a pale brown (Ochraceous- Tawny) dorsum with dull olive green (Dark Olive Buff) markings. Other- wise there was no noticeable variation in color. Description of tadpole—rThe following description is based on KU 60051 from Finca La Paz, Depto. San Marcos, Guatemala (Figs. 5A and 6E). Small hind limbs; total length, 37.9mm.; body length, 11.6 mm.; body length/total length, 30.6 per cent. Body only slightly depressed, nearly as deep as wide, in dorsal profile ovoid, widest just posterior to eyes; in lateral profile snout rounded; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils barely protuberant, directed anteriorly, somewhat closer to eye than snout; spiracle sinistral and posteroventrad to eye; anal tube dextral. Tail long and slender; caudal fin low and rounded posteriorly; depth of caudal musculature one-half greatest depth of caudal fin; musculature not extending to tip of tail. Mouth large; thin fleshy lips greatly expanded and forming large funnel- shaped disc; width of mouth two-thirds greatest width of body; outer edge of lips having one row of small papillae; inner surfaces of mouth smooth; scattered large papillae forming one nearly complete row around teeth; other large papillae laterally; beaks moderately developed, bearing long, pointed denticulations; no lateral projections on upper beak; tooth-rows %, all short; second and third upper rows subequal in length; first upper row shorter; first MippLe AMERICAN TREE FROGS 333 and third upper rows interrupted medially; first lower row interrupted medially, equal in length to second and third upper rows; second lower row slightly shorter; third lower row shortest. Body mottled brown and creamy gray above and below; mouth colored like body; caudal musculature creamy tan; caudal fin transparent; dark brown streak mid-laterally on anterior third of caudal musculature; rest of tail and all of caudal fin heavily flecked with brown; eye red in life. Variation.—The third upper tooth-row is interrupted in all specimens; in some individuals the first upper and first lower rows are complete. The varia- tion in size and proportions is given in Table 2. The dark brown lateral streak on the anterior part of the caudal musculature is distinct on most speci- mens; the only other variation in coloration is in the amount of brown fleck- ing on the caudal musculature and fin. Comparisons.—Ptychohyla schmidtorum schmidtorum differs from P. schmid- torum chamulae as stated in the diagnosis and in having pale creamy tan, as opposed to dark brown, webbing on the feet; and from P. ignicolor in having a depressed, as opposed to a flat, internarial region. Tadpoles of P. s. schmid- torum have a mottled appearance, as opposed to the more uniform brown color of P. s. chamulae. Ptychohyla schmidtorum schmidtorum and several species of Plectrohyla are sympatric. All species of the latter genus have a bony prepollex, rugose skin on the dorsum, and heavy body; also sympatric is Ptychohyla e. euthy- sanota, which has a tarsal fold and in breeding males spinous nuptial tuberosi- ties. Life History—This species breeds in clear mountain streams where males call from vegetation along the stream. The call con- sists of series of short notes, three to nine notes per series, sounding like “raa-raa-raa.” The duration of each note is approximately .065 of a second, and has a rate of 96 to 119 pulses per second; the dominant frequency is about 3400 cycles per second. The call is almost indistinguishable from that of Ptychohyla schmidtorum chamulae. Tadpoles and metamorphosing young were found at Finca La Paz, Guatemala, in late July, 1960. Two young lacking tails but not having completely developed mouths have snout-vent lengths of 14.2 and 146mm. L. C. Stuart collected four metamorphosing young at Finca La Paz on May 6, 1949. By May 10 the frogs were completely metamorphosed, at which time they had snout-vent lengths of 15.5 to 17.0 (average 16.1) mm. Remarks.—There is no doubt that this frog is most closely re- lated to Ptychohyla schmidtorum chamulae, even though the ranges of the two subspecies are separated by the interior depression of Chiapas. Since at least at Finca La Paz, Guatemala, P. s. schmid- torum occurs with P. e. euthysanota, it is surprising that the former species has not been found at more localities along the Pacific slopes 334 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. on northern Central America. At Finca La Paz in July, 1960, P. s. schmidtorum was more abundant than P. e. euthysanota. Distribution —This species is known only from a limited area at eleva- tions between 1300 and 2200 meters on the Pacific slopes of the Sierra Madre in extreme eastern Chiapas and western Guatemala. Specimens examined.—MeExico: Chiapas: Finca Irlandia, UMMZ 105429-30. GuaTEMALA: San Marcos: El Porvenir, CNHM 20755, 20761, 69904, UMMZ 80918; Finca La Paz, 2 km. W of La Reforma, KU 58016-44, 59940-2 (skeletons), 60050 (3 young), 60051 (tadpoles), 60052 (4 young), MCZ 84998-9, UMMZ 123144-7 (tadpoles). Ptychohyla schmidtorum chamulae Duellman Ptychohyla chamulae Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 13: 354-357, pl. 25, fig. 2, April 27, 1961 [Holotype—KU 58063 from 6.2 kilometers south of Rayén Mescalapa, Chiapas, México; William E. Duell- man collector]. Diagnosis—Vomerine teeth 4-6; dorsum bright green; white lateral stripe; eye reddish bronze in life. Description —The following description is based on KU 58069 from 6.2 kilometers south of Rayén Mescalapa, Chiapas, México (Pl. 7). Adult male having a snout-vent length of 27.6mm.; tibia length, 13.0 mm.;_ tibia length/snout-vent length, 47.1 per cent; foot length, 10.8 mm.; head length, 9.6mm.; head length/snout-vent length, 34.7 per cent; head width, 9.2 mm.; head width/snout-vent length, 83.1 per cent; diameter of eye, 3.0mm.; diameter of tympanum, 1.6mm.; tympanum/eye, 53.3 per cent. Snout in lateral profile nearly square, slightly rounded above and below, and in dorsal profile blunt, almost square; canthus pronounced; loreal region concave; lips thick, rounded and flaring; nostrils protuberant; internarial distance, 2.5 mm.; in- ternarial region slightly depressed; top of head flat; interorbital distance, 3.3 mm., much greater than width of eyelid, 2.5mm. Thin dermal fold, from posterior corner of eye above tympanum to insertion of fore limb, covering upper edge of tympanum; tympanum nearly round, its diameter equal to its distance from eye. Forearm slender, lacking distinct fold on wrist; row of low, rounded tubercles on ventrolateral surface of forearm; pollex slightly en- larged, without nuptial spines; second and fourth fingers equal in length; sub- articular tubercles round, that under fourth finger bifid; discs small, that of third finger noticeably smaller than tympanum; no web between first and second fingers; vestige of web between other fingers. Heels overlapping when hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; no tarsal fold; inner metatarsal tubercle large, flat, and elliptical; outer metatarsal tubercle small, elliptical, slightly more distal than inner; subarticular tubercles round; length of digits from shortest to longest 1-2-5-3-4; third and fifth toes webbed to base of disc; fourth toe webbed to base of penultimate phalanx; discs smaller on toes than on fingers. Anal opening directed posteriorly at upper edge of thighs; no anal flap; pair of large tubercles below anal opening. Skin of dorsum and of ventral surfaces of forelimbs and shanks smooth; that of throat, belly, and ventral surfaces of thighs granular. Ventrolateral glands well developed, not reaching axilla or groin and broadly separated mid- ventrally. Tongue cordiform, shallowly notched behind and only slightly free posteriorly; vomerine teeth 2-3, situated on small triangular elevations be- MipptE AMERICAN TREE FROGS 335 tween ovoid inner nares; openings to vocal sac large, one situated along inner posterior edge of each mandibular ramus. Dorsum of head, body and limbs reddish brown with dark purplish brown markings on back and shanks; first finger creamy tan; other fingers pale brown; dorsal surfaces of tarsi, third, fourth, and fifth toes dull tan with brown spots; first and second toes creamy tan; webbing on feet brown; anterior and posterior surfaces of thighs tan; faint creamy white stripe along ventrolateral edges of tarsi and forearms; enamel-white stripe on heel; axilla and groin gray; enamel-white stripe on edge of upper lip, continuing onto proximal upper surfaces of forelimb and on flanks to groin, widened under eye to form large spot, and bordered below on flanks by dark brown stripe; white stripe above and white spots below anal opening; throat and chest white; belly and ventral surfaces of limbs cream color; brown dash on either side of chin and brown spot on throat near angle of jaws; few brown flecks on belly; ventrolateral glands orange-tan; ventral surfaces of tarsi and feet brown. In life, dorsal surfaces of head, body, and limbs bright green (Shamrock Green); first and second fingers pale orange (Apricot Yellow); stripe on upper lip and spot below eye enamel-white; stripe on flanks silvery white, bordered below by brown (Saccardo’s Umber) brown; anterior and posterior surfaces of thighs yellowish brown (Old Gold); webbing of feet dull brown (Brownish Olive); belly deep yellow (Amber Yellow); iris reddish bronze (English Red). Variation —Tubercles beneath the fourth fingers are bifid in 20 specimens and rounded in all others. The tongue is emarginate in 12 specimens and cordiform in all others. In most specimens the white stripe on the upper lip continues onto the flanks and to the groin; in five specimens the stripe termi- nates above the forearm, and in three it terminates at mid-flank. The lateral stripe is absent in two specimens. All specimens were uniform green above when found at night; later some changed to pale green (Light Oriental Green) on the dorsum with irregular yellowish tan (Naples Yellow) blotches. Most males have brown flecks on the throat and ventrolateral gland, but some specimens are immaculate below, and one has dark brown mottling on the throat. Several males have a round, orange-tan glandular area on the chin, as does P. ignicolor. Description of tadpole—The following description is based on KU 58199 from 6.2 kilometers south of Rayén Mescalapa, Chiapas, México (Figs. 5B and 6F). Hind limbs small; total length, 39.0mm.; body length, 10.5 inm.; body length/total length, 26.9 per cent. Body barely depressed, only slightly wider than deep, widest just posterior to eyes; in dorsal profile ovoid; mouth directed ventrally; eyes small, directed dorsolaterally; nostrils barely protu- berant, directed anterodorsally, slightly closer to eye than snout; spiracle sinistral and posteroventrad to eye; anal tube dextral. Tail long and slender; caudal fin low, rounded posteriorly; depth of caudal musculature one-half greatest depth of caudal fin; musculature not extending to tip of tail. Mouth large; thin fleshy lips greatly expanded and forming funnel-shaped disc; outer edge of lips having one row of small papillae; inner surfaces of mouth smooth; scattered large papillae forming nearly one complete row around teeth; other papillae laterally; beaks moderately developed, bearing long, pointed denticulations; no lateral projections on upper beak; tooth-rows 34, all short; second and third upper rows subequal in length; first upper row shorter; first and third upper rows interrupted medially; first lower row inter- 336 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. rupted medially, equal in length to second and third upper rows; second lower row slightly shorter; third lower row shortest. Body dark brown above and dark gray below; fleshy part of mouth creamy gray mottled with dark brown; caudal musculature pale tan with heavy suf- fusion of brown flecks; caudal fin transparent with brown spots; dark brown streak mid-laterally on anterior one-fifth of caudal musculature, bordered be- low by cream-colored spot; eye brown in life. Variation.—The third upper tooth-row is interrupted in all specimens, but in some individuals the first upper row and first lower row are complete. The only noted variation in color is the intensity of brown pigmentation on the caudal musculature, which in most specimens is sufficiently dense to make the tail look brown. In some specimens the mid-lateral streak is indistinct, and the pale spot below the streak is absent. Comparisons.—Aside from the characters listed in the diagnosis, Ptychohyla schmidtorum chamulae differs from P. schmidtorum schmidtorum by having dark brown webbing on the feet, instead of pale creamy tan webbing, and in having in life a yellow venter, instead of a white venter. Ptychohyla ignicolor also is green in life, but has red flash-colors on the thighs, red webbing on the feet, and lacks the white lateral stripe diagnostic of P. schmidtorum chamulae. Plectrohyla matudai matudai and P. guatemalensis are sympatric with Ptychohyla schmidtorum chamulae. Each of the first two has a bony pre- pollex, rugose skin on the dorsum, and heavy body. Also living with Ptychohyla chamulae are Hyla chaneque, a large species having a tuberculate dorsum and webbed fingers, and Hyla bivocata, a small yellow species having a broad, flat head, small indistinct tympanum, and an axillary membrane. Life History—Calling males were found on leaves of herbs and bushes by cascading streams in cloud forest. The call consists of series of short notes, three to nine notes per series, sounding like “raa-raa-raa. The duration of each note is .054 to .070 of a second, and has a rate of 96 to 110 pulses per second. The dominant frequency falls between 3350 and 3450 cycles per second (Pl. 1D). The call is almost indistinguishable from that of Ptychohyla schmid- torum schmidtorum. Tadpoles were found in the cascading streams; the smallest tad- pole has a total length of 17.2 mm. and has only % tooth-rows. At a stream 6.2 kilometers south of Rayén Mescalapa, Chiapas, meta- morphosing young were found on June 16 and August 5, 1960. Each of two completely metamorphosed young has a snout-vent length of 15.7mm. Another having a snout-vent length of 16.2 mm. has a tail stub 2mm. long and a completely metamorphosed mouth. Two others have snout-vent lengths of 13.6 and 14.1 mm. and tail lengths of 11.5 and 8.1 mm. respectively; in these the mouth parts are incompletely metamorphosed. Remarks.—Duellman (1961:354) described Ptychohyla chamulae MippLeE AMERICAN TREE FROGS 337 and stated that it probably was most closely related to P. schmid- torum, Further study has revealed additional resemblance in mor- phological and behavioral details. It is concluded that the two populations are more realistically treated as subspecies than as species. The geographic ranges, as now known, are disjunct. Ptychohyla schmidtorum chamulae inhabits cloud forest on the Atlantic slopes of the Chiapan Highlands, whereas P. s. schmid- torum lives in cloud forest on the Pacific slopes of the Sierra Madre in Chiapas and Guatemala. Between their known geographic ranges are the pine clad Sierra Madre and Chiapan Highlands, and intervening sub-humid Grijalva Valley. Distribution.—This species is known only from elevations between 1500 and 1700 meters on the Atlantic slopes of the Chiapan Highlands; it is to be ex- pected in cloud forests on the northern slopes of the Sierra de Cuchumatanes in Guatemala. Specimens examined—Mexico: Chiapas: 15 km. N_ of Pueblo Nuevo Solistahuacén, UMMZ 123825 (4); 16.5 km. N of Pueblo Nuevo Solistahuacdn, UMMZ 123322 (10); 18 km. N of Pueblo Nuevo Solistahuacan, UMMZ 121395-9, 123324 (8), 123326 (5); 18.6 km. N of Pueblo Nuevo Solistahuacdn, UMMZ 123328 (4); 5.6km. S of Rayén Mescalapa KU 58062, 58200 (tadpoles); 6.2 km. S of Rayén Mescalapa, KU 58063-74, 58199 (tad- pole), 58234-8, 59936 (skeleton). Ptychohyla ignicolor Duellman Ptychohyla ignicolor Duellman, Uni. Kansas Publ. Mus. Nat. Hist., 13:352- 853, pl. 95, fig. 1, April 27, 1961 [Holotype—UMMZ 119603 from 6 kilometers south of Vista Hermosa, Oaxaca, México; Thomas E. Moore collector]. Diagnosis —Diameter of tympanum less than one-half diameter of eye; internarial region flat; 3-7 vomerine teeth; toes one-half webbed; no white spot below eye; no lateral white stripe; in life dorsum green; groin and thighs having bright red flash-colors. Description.—The following description is based on UMMZ 119603 from 6 kilometers south of Vista Hermosa, Oaxaca, México (Pl. 8). Adult male having a snout-vent length of 30.0 mm.; tibia length, 14.6 mm.; tibia length/ snout-vent length, 48.7 per cent; foot length, 12.3 mm.; head length, 9.2 mm.; head length/snout-vent length, 30.7 per cent; head width, 9.3mm.; head width/snout-vent length, 31.0 per cent; diameter of eye, 3.2 mm.; diameter of tympanum, 1.8 mm.; tympanum/eye, 40.6 per cent. Snout in lateral pro- file square, and in dorsal profile rounded; canthus pronounced; loreal region slightly concave; lips moderately flaring; top of head flat; nostrils protuberant; internarial distance, 2.8mm.; internarial region flat; interorbital distance, 8.38 mm., much broader than width of eyelid, 2.8mm. A heavy dermal fold from posterior corner of eye above tympanum to insertion of forelimb, cover- ing upper edge of tympanum; tympanum elliptical, its greatest diameter equal to its distance from eye. Forearm moderately robust having distinct dermal fold on wrist; pollex moderately enlarged without nuptial spines; second and fourth fingers equal in length; subarticular tubercles round, none is bifid; 338 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. discs on fingers moderate in size, that on third finger slightly larger than tympanum; no web between first and second fingers; vestige of web between other fingers. Heels overlap when hind limbs adpressed; tibiotarsal articula- tion extends to anterior corner of eye; no tarsal fold; inner metatarsal tubercle large, flat, and elliptical; outer metatarsal tubercle near inner one and tri- angular in shape; subarticular tubercles round; length of digits from shortest to longest 1-2-5-3-4; third toe webbed to proximal end of penultimate phalanx; fourth toe webbed to distal part of antepenultimate phalanx; fifth toe webbed to middle of penultimate phalanx; discs on toes smaller than on fingers. Anal opening directed posteriorly at upper edge of thighs; no anal flap; pair of large tubercles below anal opening; small tubercles ventral and lateral to these. Skin of dorsum and ventral surfaces of limbs smooth; that of throat and belly granular. Ventrolateral glands noticeably thickened, extending from axilla nearly to groin and only narrowly separated midventrally on chest; skin of anterior part of chin thickened and glandular. Tongue cordiform, shallowly notched behind and only slightly free posteriorly; vomerine teeth 0-3, situated on rounded elevations between somewhat larger, round inner nares; openings to vocal sac large, one situated along posterior margin of each mandibular ramus. Dorsal ground-color of head, body, and limbs dull brown with dark brown reticulations on head and body and dark brown transverse bands or spots on limbs; first and second fingers cream color; third and fourth fingers dull tan; anterior surfaces of thighs pale brown; posterior surfaces of thighs cream color with heavy suffusion of brown; dorsal surfaces of tarsi and third, fourth, and fifth toes dull brown with dark brown spots; first and second toes creamy white; webbing on foot brown; axilla and groin cream color; flanks brown; no white stripes on edge of upper lip or on flank; faint, barely discernible tan streak above anal opening; faint creamy tan line on ventrolateral edges of tarsi; throat, belly, ventral surfaces of limbs, inner edges of tarsi, and first toes cream color; outer ventral surfaces of tarsi and other toes brown; chest and throat spotted with brown; ventrolateral and chin glands orange-brown. In life the dorsum was uniform green (Cosse Green) becoming paler green (Bright Green-Yellow) on flanks, later changing to paler green (Javel Green) on dorsum with irregular darker green (Lettuce Green) markings and greenish yellow (Green-Yellow) on flanks; anterior and posterior surfaces of thighs, ventral surfaces of shanks, anterior surfaces of tarsi, and upper proximal sur- faces of first, second, and third toes red (Coral Red); venter pale creamy yellow (Sulfur Yellow); iris pale golden color (Aniline Yellow). Variation.—Of 13 specimens, six have a cordiform tongue; the others have an emarginate tongue. Five specimens have round subarticular tubercles beneath the fourth fingers; six specimens have a bifid tubercle on one hand, and two specimens have bifid tubercles on both hands. 2 jy P90. eer ae I a ee er A OP a a 456 Introduction Throughout much of the United States the racer is abundant and is one of the snakes best known to man. Its active diurnal habits and its preference for a habitat in meadows, pastures, and hayfields rather than in remote wilderness areas, result in frequent encoun- ters with humans. The racer is a predator on many kinds of small animals, both vertebrates and invertebrates; it takes as food chiefly animals that are agricultural pests but also destroys some beneficial kinds. Yet, in general, the attitudes of rural people toward the racer are little influenced by these economic and ecologic considerations, but rather, are dominated by an unreasonable fear, despite the racer’s inoffensive disposition, and inability to inflict any harm on humans. Although an extensive literature exists regarding the racer, no thorough study of the species’ natural history has been made here- tofore. Obviously such study is needed. Few species of vertebrates having comparable economic bearing have been similarly neglected. In 1948, undertaking a program of ecological research on the recently created University of Kansas Natural History Reservation, I included the racer among the many common species studied to gain insight into the functioning of the local ecosystem. Live- trapping of snakes on the area was begun in 1949, and these opera- tions were greatly intensified in the years 1957 through 1962, with efforts concentrated on the study of the racer in the 1960, 1961 and 1962 seasons. Thus my study is based upon 14 consecutive years’ records on the Reservation, the northeasternmost section in Douglas County, Kansas, six and one-half miles north northeast of the Uni- versity of Kansas campus at Lawrence. After the acquisition in 1956 of the 160-acre Rockefeller Tract adjacent to the Reservation on the north in Jefferson County, field work was extended to this new area, which, because it was superior habitat, in the final years of the study produced more records than the Reservation. An important but relatively minor segment of the data originated from Harvey County Park, 13 miles west of Newton, Kansas, where lines of live-traps were maintained in 1959, 1960, 1961, 1962. Smaller collections of data were obtained from the Lalouette Ranch in the Flint Hills, three miles northeast of Florence, Marion County, Kansas, and from Cedar Bluff Reservoir, 23 miles west and seven miles south of Hays in Trego County, western Kansas, where live- trapping was carried on in 1959 and 1960. Additional data were (355) 356 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. obtained on numerous field trips to various collecting localities in northeastern Kansas. My first-hand knowledge of the species is also based, in part, on many years of field experience with the far western subspecies C. c. mormon in western Oregon and California, and on similar experience in 1947 and 1948 with the southern sub- species, C. c. anthicus in central Louisiana. This varied field experience with the species at localities well scattered throughout its geographic range has added perspective to the study even though most of the records were collected within a radius of three quarters of a mile. No one locality can be re- garded as entirely typical of a species’ habitat over its range as a whole. According to my philosophy, the ecological niche of a species is subject to geographical variation analogous to the varia- tion to be seen in the morphological characters of the animal itself. Different community associates, including different competitors, prey, and predators, and different physical factors enforce a some- what different way of life on a species in geographically remote parts of its range. When analyzed these differences often turn out to have a genetic basis. Thus, limits of tolerance to heat, cold, and drought often vary geographically, and the population density, reproductive potential, seasonal cycle, and other properties of populations may be altered either by the direct effect of the en- vironment, or through its effect on the genetic constitution, pro- duced by natural selection. The local population of racers studied was near the center of the species’ geographic range, and is to some extent representative of the species as a whole, though differing in its ecology from other populations in proportion to their remoteness and the distinctness of their habitats. It has not been demonstrated that ecological traits of populations change in a discontinuous manner or corre- spond in their limits with those of named subspecies. More likely geographical variation is continuous and parallels morphological variation only in a general way. Certainly the boundaries of sub- species’ ranges should not be accorded undue emphasis in an ecological study. My investigation of the blue racer under natural conditions, com- bined with a compilation and analysis of published literature, has re- sulted in a fairly satisfactory understanding of some phases of the species’ ecology and natural history, such as the food habits, the growth rate, the extent of home range and of seasonal movements. However, relatively little was learned concerning some phases of NATURAL Hisrory OF THE RACER 357 the life history. Unfortunately, the traps used did not catch young of the smaller sizes. Facts concerning egg-laying, incubation, and hatching therefore are known chiefly from snakes kept in confine- ment. Although first-year young were captured by hand from time to time they were obtained in relatively small numbers, and little was learned regarding their population density, movements, or mortality factors. Of course, such hiatuses are to be expected; even in man such enigmas as the disparate sex ratio still challenge the investigator. For the subspecies of racer involved in my field study the widely used vernacular “blue racer” has been adopted in this report. In general I advocate conformity with the vernacular names pub- lished by the Committee on Herpetological Common Names (1956). However, in this list, the name blue racer was assigned to Coluber constrictor foxi, an invalid subspecies of the Prairie Peninsula that has been relegated (Auffenberg, 1955:92; Smith, 1961:196) to the synonymy of C. c. flaviventris. It therefore seems appropriate that the book name “yellow-bellied racer” applied to flaviventris by the Committee should be abandoned for this subspecies, and that the name blue racer be applied officially, as it is in actual practice by both laymen and herpetologists, to all populations of this subspecies. Acknowledgments Financial assistance from the National Science Foundation in 1957 through 1962 is acknowledged. Although none of the three separate grants involved was made specifically for the autecologi- cal study of the racer, all three contributed to the support of the extensive program of live-trapping for snakes, which yielded most of the records upon which this report is based. Student assistants who were employed on these projects include James W. Bee, William N. Berg, Donna M. Hardy, Robert M. Hedrick, Dale Hoyt, Robert M. Packard, Robert G. Webb, and Wayne Wiens, at the Reservation; Roy Henry, Dale Horst, Dwight R. Platt, and Howard L. Schrag at Harvey County Park, and Gilbert L. Adrian at Cedar Bluff Reservoir. Dr. Edwin P. Martin, formerly of Fort Hays, Kan- sas State College, was helpful in planning and carrying out the field work at Cedar Bluff Reservoir. Eric Shulenberger assisted with field work and processing of data in 1962 under the National Science Foundation program for Undergraduate Research Participa- tion. Robert Miner assisted with the examination of specimens in 1960. Mr. August Lalouette of Florence, Kansas, permitted field 358 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. work on his ranch and contributed information and materials to expedite this work. Mr. and Mrs. Harold Brune of Route 3, Lawrence, Kansas, kindly contributed several clutches of racer eggs found on their farm in Jefferson County, and also made available significant information accompanying them. Dr. William H. Stickel kindly made available at my request records of predation on racers from the food habits files of the U.S. Fish and Wildlife Service. Dr. William E. Duellman of the University of Kansas Museum of Natural History and Dr. Robert C. Stebbins of the University of California Museum of Vertebrate Zoology kindly permitted examina- tion and dissection of specimens in the collections under their care. Dr. George W. Byers of the University of Kansas Department of Entomology identified numerous insects eaten by racers. My daughter, Alice V. Fitch, often assisted me with the field work and the processing of data. My wife, Virginia R. Fitch, read the manu- script critically, assisted me with the examination of museum speci- mens, and with typing, and helped in various other ways. Methods and Materials This investigation was based primarily on the capture in live- traps, marking, release, and recapture of blue racers in their natural habitat. On the combined area of the Reservation and the Rocke- TABLE 1. NUMBERS AND DISTRIBUTION OF CAPTURES AND RECAPTURES OF BLUE RACERS ON THE RESERVATION AND ROCKEFELLER TRACT Span of years within Number of separate Number of times which each individual | years within which each each individual was captured individual was captured was captured Years | Individuals | Years | Individuals | ,oguesq | Individuals 1 749 1 749 1 679 2 137 2 197 2 181 3 56 3 51 3 93 4 32 4 15 4 31 5 19 5 6 5 a 6 13 6 1 6 14 a 7 a 0 7 8 8 2 8 1 8 4 9 3 9 1 10 0 10 0 11 1 11 0 12 1 12 1 13 0 14 0 15 0 16 1 NATURAL HIsToRY OF THE RACER 359 feller Experimental Tract, 1020 blue racers were recorded a total of 1688 times from August 30, 1948, to October 27, 1962. At Harvey County Park 361 blue racers were marked, and were captured a total of 467 times from May 6, 1959, to September 14, 1962, and at Cedar Bluff Reservoir 42 were captured from May 11, 1959, to June 30, 1960. The traps used were cylinders of galvanized wire, “hardware cloth” (Fitch, 1951:77; 1960:77), having funnels opening into each end, or having a funnel at one end and a plug at the other. The traps, open at both ends, were used along hilltop rock ledges where an exposed vertical rock face provided a barrier along which a snake might travel and where it could be easily intercepted by the trap without any accessory equipment (see Pl. 21, Fig. 1). Where such natural barriers were lacking, as in level fields, barriers con- sisting of boards, screens or sheet metal were installed to guide the racer toward the trap and into a funnel entrance. Two such barriers at each end of a trap forming a V to guide the snake into the funnel were used in 1956 and 1957, when trapping at places away from the hibernation ledges was undertaken. Later it was found simpler and more effective to use a single barrier with a trap at each end. The barrier extended up into the funnel entrance, and usually the racer, following along the barrier on either side, would pass into the funnel and through its apex. However, it was possible for a racer to travel around the end of the barrier without entering the trap, and perhaps some did so. The need for making the barrier and trap a tight unit impassable to the snake, was somewhat counterbalanced by the need for hav- ing the whole installation loosely constructed so that it could be easily altered, opened, adjusted, and cleaned. Since the traps were kept set in large numbers, and the task of checking them was time- consuming, speed of operation was more important than the perfect functioning of any one trap. Approximately 200 traps were kept set when operations were at their maximum. No record was kept of the number of “trap days” involved in the study, but the total was well over 100,000 for the Reservation and the Rockefeller Tract. When traps were set at both ends of a barrier, the outer end of each trap was closed with a plug. No bait was used in the traps. Occasionally small vertebrates and insects of kinds used as food by the racers may have gotten caught first and served to attract the snakes. Best catches of racers were made in the breeding season, since males were attracted by females already in the traps, and 360 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. several males might be captured simultaneously with one female. Occasionally as a person approached or handled a trap, a racer darted out of it, displaying a perception, acuity of vision, and skill in avoiding the inward projecting wire prongs encircling the small funnel opening, that were exceptional among the several species of snakes trapped. Doubtless many other racers that were caught in traps escaped before they were discovered. If the funnel entrance of the trap was of the same diameter as the snake itself, or only a little larger, there was little likelihood of the trapped racer escaping. However, funnel openings were usually adjusted at a diameter of approximately 1% inches, allowing an ample margin for even the largest racers, though inadequately small to permit ingress of a few of the largest black rat snakes, bull snakes and timber rattlers oc- curring locally. Ordinarily the snakes trapped were processed in the field and re- leased immediately. The method of marking was essentially that of Blanchard and Finster (1933:334). Two subcaudals, one on the right side and one on the left, were clipped on each snake, and when these marks healed they left permanent scars. In the racer, as in most other colubrines, the subcaudals are divided into a double series, one on the left and one on the right. Scales of the left and right sides are placed alternately. At the base of the tail one or more undersized scales usually are present on each side, and there might be some question as to precisely where the count should be- gin. The rule followed was to exclude from the count any small basal scales on either side that did not extend medially to contact at least one scale of the opposite side. The scale designated as “one left” (or “one right”) was the first to contact one of the opposite series, regardless of whether the former was of normal size or (as was usually the case) smaller and narrower than those following it. In marking, this “number one” scale was never clipped but was left as a point of reference since a base mark was needed from which to begin the count. The marks were read from left to right, for example U 51 2r, the “U” referring to the subcaudals or “urosteges,” the “SI” indicating the fifth on the left side, and “2r” indicating the second on the right. The subcaudals clipped were the first 19 following the basal scale. When the 361 possible combina- tions all had been used, ending with U 201 20r, a new series was begun duplicating the first except that on each snake the first ventral (or “gastrostege”) anterior to the anal plate was clipped on the left side (GIL) to distinguish these snakes from the series NATURAL History OF THE RACER 361 previously marked. Later, a third series, “G2R” was marked, and eventually a fourth series, “G3L” was started. There were many borderline instances in which the basal scale barely contacted one of the opposite side. In such instances the formula was written U 51,,, 2r, the subscript ISB signifying “in- cluding small basal.” In other instances a basal subcaudal barely failed to contact a scale of the opposite side and this condition was indicated by the subscript NSB—“not including small basal.” The condition might be so nearly equivocal that on successive occa- sions the same formula might be read U 51,,, 2r and U Al opt Occasional misidentifications of individuals that resulted from such discrepancies were in most instances readily detected when the field records were transferred to individual file cards where the sex, size, and location of the snake at its previous captures were shown. In some instances racers recaptured after periods of years re- tained conspicuous scars where scales had been clipped, but in other instances the marks had become obscure, and in fact the only trace of a mark might be a slight narrowing or notching of part of the scale originally clipped. Snakes caught and marked early in life probably retained more power of regeneration than those clipped after the attainment of maturity, but otherwise the basis for dif- ference in extent of regeneration was not evident. In the same snake, three scales, all clipped on the same day, might show much different degrees of regeneration after the lapse of a year or more. In general, obscuring of marks by regeneration was a source of in- convenience rather than of error; only a negligible percentage of the recaptured racers had marks so obscure that their identities might have been seriously questioned, and it is doubtful that any marks were lost completely by regeneration. Racers found in traps were removed, measured (snout-to-vent length, tail length), weighed in a cloth bag suspended from spring scales, and marked. The mouth was forced open and the snake was examined for flukes. Enamel paint of a bright color, red, green, yellow, blue or orange was smeared on the snake to gain informa- tion regarding the time of molt. The stomach was palpated for recently ingested food items, and any detected were forced up into the gullet to be identified, then were squeezed back into the stomach. The rear part of the body was palpated to detect un- digested material in the gut, and if any was present, an attempt was made to squeeze out the fecal material, using only light pres- 362 UNIVERSITY OF KANnsAS Pusts., Mus. Nat. Hist. sure, with care not to injure the racer. The inside of the trap and the ground beneath it were inspected for fecal material that might have been voided while the snake was confined. Any scatological material obtained was wrapped in a paper towel, labelled and brought back to the laboratory where it was stored. Eventually each scat was soaked for a day or more in a detergent solution, rinsed in running water in a fine gauze bag, dried, and placed with its label in a cellophane envelope for subsequent microscopical study. Various items concerning reproductive condition were also routinely recorded. In females the ventral surface was palpated at the rear end of the body to detect the genital bursa or vagina, which in sexually mature individuals has a much thickened wall, and can be felt as a distinct lump. Males were likewise tested for sexual maturity by pipetting a small amount of fluid from the cloaca into a vial and returning it to the laboratory where it was examined microscopically for motile sperm. Also, sperm samples were often taken from males at different times throughout the season of activity, and cloacal samples from females occasionally were checked for sperm as evidence of recent copulation. In the summer of 1962 an outdoor enclosure of 100-foot cir- cumference was constructed, of galvanized sheet iron, with wall three feet high, set on a concrete base extending to a depth of two feet. A two-foot-deep concrete basin inside the enclosure served as a water container. The enclosure was partly shaded by a large walnut tree and the area enclosed had lush vegetation, including brome grass, various shrubs, and young trees up to 15 feet high, thus including most features of the racers’ habitat, and it was situated in an area frequented by the snakes. Throughout the summer several racers were kept in the enclosure, and frequent observations on them yielded much information concerning time of activity, temperature preferences, and social and sexual behavior that could not have been obtained readily either from racers con- fined in small cages or from those free under natural conditions. Description Color Pattern Hatchling racers differ much in appearance from adults; whereas the latter are of dull uniform coloration dorsally, the hatchlings have a checkered pattern of alternating blotches in several rows, in- cluding a middorsal row, with blotches much larger than those of NATURAL History OF THE RACER 363 the other rows. This basic pattern is perhaps the most common one in all snakes, and is found in the young of various other genera (notably Elaphe) which lose or alter their markings during develop- ment. In these genera and in the racer, the juvenal checkered pattern may represent recapitulation of an ancestral condition. The adaptive significance of having a blotched, checkered pattern in the young, and uniform coloration in the adult is not evident. I have rarely seen the hatchlings under natural conditions except by finding them hiding beneath flat rocks. Their concealing pat- tern must be fully as effective as that of adults and the young them- selves are more secretive than the adults. A hatchling of 240 millimeters snout-vent length was described as follows on September 22, 1962: Ground color pale olive anteriorly, gradually darkening posteriorly, marked with chocolate blotches; middorsal blotches largest averaging about five scales long and seven or eight scales wide anteriorly; posteriorly, blotches become less distinct (tending to blend into progressively darkening ground color), and width-to-length ratio increases; gradual ob- scuring of blotches proceeds posteriorly, until on tail they can no longer be distinguished, and color is uniformly olive; on each side, row of lateral blotches alternates with that of middorsal blotches; lateral blotches average somewhat less than two scales long, and ap- proximately two scales wide and are of irregular shape, sometimes subdivided; farther down on sides, lower row of lateral blotches alternates with upper lateral row; this lower lateral row, approxi- mately same size as upper lateral row, and situated at level of first scale row, overlapping onto ventrals, where it pales to reddish brown; almost every ventral scute has one pair or more of dark reddish brown spots, tending toward semicircular shape, the arc of each projecting forward, but posteriorly on body these spots be- come increasingly obscure, and are indiscernible on posterior end of body and on tail; ventral surface white on chin and throat, gradually assuming suffusion of pale greenish gray posteriorly; eye dark with narrow yellowish margin around pupil; top of head gray- ish olive, mottled with faint and irregular dark markings; supra- labials whitish, with chocolate markings, mostly wedge-shaped, in their posterior parts; rostral and internasals edged with dark posteriorly; posterior upper corner of loreal and adjacent corner of prefrontal dark; temporals having dark markings; chin almost immaculate, but with narrow black posterior edges on some of the infralabials (see Plate 19, Figs. 1 and 2, and Plate 20, Fig. 2). 364 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. The checkered pattern of the juvenile fades gradually as develop- ment proceeds. Persistence of the juvenal markings varies greatly in individuals and probably is subject to geographic variation also. By the time sexual maturity is attained the dorsal pattern often is indiscernible or represented only by faint traces. The ventral speckling is more persistent. A female of 602 millimeters snout-vent length in mid-July 1962, at a probable age of 11 months, was described as follows: Over- all dorsal color olive gray, but with remnants of juvenal pattern dis- cernible; dark dorsal blotches have almost faded, but their edges, about one scale wide, are still distinct; low on sides, color fades to pale bluish gray, and to pale greenish blue on edges of ventrals; on anterior one-third of body midventral surface is lemon yellow; farther posteriorly it fades to ivory, almost white on tail; chin white, except for reddish brown posterior edges of last infralabials, and streak of same color on each antepenultimate infralabial; top of head olive gray with irregular scattered dark marks; preoculars pale centrally with olive brown edges; supralabials white with reddish brown triangular marks; postnasals white anteriorly, gray pos- teriorly; uppermost postocular brownish orange, paler centrally; two rows of semicircular spots on belly, distinct anteriorly, but fading posteriorly until indiscernible on posterior part of belly; numerous small black spots scattered irregularly over dorsal and lateral surfaces. A male racer retained more of the juvenal pattern at an approxi- mate age of 15 months, when he was described on November 30, 1961: Dorsal surface dark grayish brown, the large juvenal dorsal blotches (each about eight scales wide) easily discernible, but faint; top of head dark olive brown, mottled with black, paling in rostral region; supralabials white on their anterior and lower portions, marked with brown and blue-gray on their upper and posterior portions; chin white, but with rusty markings on last four infralabials; ventrals ivory-yellow with rusty spots of which the largest are approximately three-fourths of the ventrals’ breadth; first row of scales bluish gray, or with greenish suffusion (in neck), most of scales having indistinct dark spots; second row of scales similarly colored but more suffused with dusky pigment, blending into the darker duller color of the dorsal surface. A female of 720 millimeters snout-vent length, presumably about 20 months old, but not gravid, on June 18, 1962, was described as follows: Juvenal pattern no longer clearly discernible but scattered NATURAL History OF THE RACER 365 traces of it remain; dorsal color predominantly grayish olive, with occasional small black spots in streaks scattered irregularly over dorsal and lateral surfaces; at anterior end of body lateral scales have bluish green edges but this shading becomes less noticeable posteriorly; head mostly olive dorsally with marking obscure; parietals have a large faint blotch; supralabials retain faint dark, brownish markings on their upper parts; dark spot on median edge of each prefrontal; supraoculars slightly darker than adjacent scutes; chin mostly white, with yellowish suffusion at edges of scales; ventral surface predominantly yellow, but fading to grayish white posteriorly; remnants of juvenal ventral spots faintly discernible as tan or whitish areas on yellow ventrals. Munro (1950b:124) mentioned a blue racer of 749 millimeters total length (hence probably having a snout-vent length of 600 millimeters, or a little less), which retained faint juvenal markings when caught on June 23, 1948, even though it was sexually mature, since it laid eggs on the night of July 4, 1948. During several weeks of captivity this snake’s markings faded perceptibly. In fully adult blue racers, those more than three years old, the juvenal markings have become completely obliterated. In those from the area of my study, the dorsal coloration is subject to much individual variation, typically olive brown, but ranging from pale bluish gray to dark brown, dark blue, or slaty. In most, the dorsal color is uniform, but in some there are streaks and isolated scales of black. The dark dorsal color extends down the sides onto the lateral corners of the ventrals and the subcaudals. The chin is white but the remainder of the ventral surface is ivory colored. Bodily Proportions The slender and streamlined bodily proportions of the racer are subject to change through allometric growth. The head, and es- pecially the eyes, are relatively large in the hatchling, and become relatively smaller as growth proceeds. The relative tail length seemingly increases in the growing young and then decreases slightly in adults. Snout-vent length and tail length were recorded in almost all the racers examined, but other measurements were recorded in relatively few. In many racers, especially the larger adults, tails had been damaged and lacked their terminal parts. Often only the tip was missing, but, of course, such individuals were not usable in the study of proportions of the tail. Table 2 summarizes information concerning relative tail length in 935 racers of both sexes and 366 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. various sizes, from northeastern Kansas. Nearly all measurements are from the live snakes; a few are from recently killed individuals. In hatchlings, the proportions of the tail are not noticeably dif- ferent in males and females, but data indicate that the snakes’ tails are approximately seven per cent longer in males than in females; as growth proceeds, the tails become relatively longer in proportion to the body, in both sexes. The ratio reaches its maximum in young adults, having increased from approximately 28 per cent of the snout-vent length in males and 26 per cent in females, to 31 per cent in males and a little more than 28 per cent in females. In the largest racers, of both sexes, these percentages are slightly reduced. Tail-length is subject to a fairly wide range of individual variation, which tends to obscure the trends determined by sex and size. TABLE 2. RELATIVE TAm LENGTH IN MALE AND FEMALE RACERS OF DIFFERENT SIZE Groups Males Females Snout-VENT LENGTH IN Number Mean ratio of Number Mean ratio of MILLIMETERS of tail-length to of tail-length to racers snout-vent length racers snout-vent length 150-200 .... ie 27.3 .833 2 26.3 201-250 .... 24 27.8+ .490 23 25.9% .479 251-300 .... 9 28.8 + .634 4 26.6 301-350 .... 20 29.8 + .246 11 25.0+1.060 351-400 .... 8 28.8 + .530 12 27.5+ .404 401-450 .... 6 27.8 .775 5 26.3= .357 451-500 .... 12 29.7 += .434 6 26.8+ .858 501-550 .... 45 30.5 .283 15 27.6 .284 551-600 .... 76 31.0+.218 35 27.1 .254 601-650 .... 45 29.6 .313 64 27.8 .163 651-700 .... 50 30.5 .241 36 27:1) 2350 701-750 .... 72 30.5 .177 45 27.6 .373 751-800 .... 45 30.3 .373 38 27.8 .325 801-850 .... 48 29.7 .274 50 27.5 .205 851-900 .... 18 29.1+.519 35 28.6 .422 901-950 .... 5 29.8 + .672 31 26.7 .252 951-1000.... 1 29.5 19 26.9+ .390 TOOTH1O5002 Mote rca ha ck a,.: eee 15 25.9 .413 NO SIA=TMOO «oe tuts cere ecehel iiss sce Dehecvevsae ca neater 6 26.4 .725 TOUS TT SO Sail. et eehicistolc, 5 nebo seen eee 1 28.5 PU 5 I=—T ZOO Bae EE Se | te. Pe eee 1 25.0 In 88 racers caught in the summer of 1962 the following measure- ments were recorded: Head length, from tip of snout to angle of jaw; maximum head width; greatest diameter of eye; circumference of neck; circumference at mid-body; circumference at posterior end NATURAL History OF THE RACER 367 TABLE 3. BopiLy PROPORTIONS (EXPRESSED AS RATIO OF SNOUT-VENT LENGTH) IN RACERS OF DIFFERENT SIZES Large Medium Small S1zE Group (more than 800] (500 to 800 (less than 500 millimeters) millimeters) millimeters) Length, of heads 5 csr: mcrae: os 3.61 + .036 3.82 .025 5.39 Width of head 2442... 1.93 = .049 2.02 + .023 2.53 Diameter of eye............. .56 = .008 .63 = .009 1.00 Circumference of neck....... 4.71+ .082 5.05 .052 6.64 Circumference at mid-body...| 7.11+.238 7.66 += .082 8.58 Circumference at posterior endiof bodynis iG. 2230: 5.06+ .113 5.03 + .061 5.90 Circumference of tail at base..| 4.23+.1131 | 4.22+.075} 4.66! 4.47+.171? | 4.66+.043? 1. Females. 2. Males. of body; and circumference of tail-base. Because the measurements were small, and were made in the field on active, struggling snakes, a high degree of precision could not be attained, and the range of error was several per cent, with occasional relatively large errors. Nevertheless, ontogenetic trends are clearly indicated. Most of the racers measured were adults of small to medium size—in the range 500 to 799 millimeters, snout-vent length. Twelve females and seven males ranged from 800 to 1035 millimeters, and seven young (all females) were less than 500 millimeters. In measure- ments other than circumference of tail-base, significant differences could not be found between males and females of the same size group; therefore the sexes were combined to obtain larger series. — Table 3 shows that as compared with adults, the small young racers have stouter, stubbier bodies, relatively large heads, and, especially, large eyes. Allometric growth seems to continue through- out life and the changed proportions of the adults are accentuated in the largest and oldest individuals. Lepidosis Scalation that of typical colubrid (see Pl. 19); rostral large, ex- tending back onto dorsal surface of snout, bluntly pointed behind; paired internasals considerably wider than long, convex anteriorly, almost straight-edged posteriorly, each extends laterally to naris; 2—7864 368 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. paired prefontals approximately twice size of internasals, and wider than long, extending laterally on each side to level of nostril; frontal convex anteriorly, concave on each side, bluntly pointed behind, nearly twice as wide anteriorly as posteriorly; parietals large; angle formed between them by frontal slightly more than 90 degrees; nostril large, situated between almost equal sized anterior nasal and posterior nasal plates; loreal slightly smaller than nasals, its anterior edge inclined forward superiorly; two rows of temporals on each side; in upper row, first one narrow and elongate, second much shortened, third intermediate in shape; in lower row all three ap- proximately alike in size and shape; two postoculars, the lower larger; seven supralabials, first small and low, longer along upper edge than along lower, second slightly longer than high, third higher than long, contacting eye; fourth largest, contacting posterior part of eye, and lower postocular; fifth nearly as large, pointed above; sixth also large, pentagonal; seventh low and rectangular; on chin first pair of infralabials separate mental from anterior genials; second infralabial minute; third approximately twice its size; fourth much smaller, rhomboidal, fifth also large, pentagonal; sixth smaller, rhomboidal, bluntly pointed behind; seventh smaller, narrow behind; eighth small and elongate; second pair of genials longer and narrower than those of first pairs, separated from each other by smaller scales; genials in approximately five rows, but somewhat irregular in arrangement, mostly smaller and narrower than body scales; latter all smooth, arranged in 17 rows for about two-thirds of body length, then, by loss of third row on each side, reduced to 15; scales of neck region rounded and relatively small, one-third to one-fourth size of larger body scales; lowest scale row on each side largest with its scales much wider and less symmetrical than others; most of body scales of approximately hexagonal shape; on forebody they average approximately twice as long as wide, but farther posteriorly on body, width-length ratio gradually increases and some of scales, notably those of lowest row, approximately as wide as long; regularity of scale rows broken on sides just above vent by presence of many small additional scales; on tail scale rows drop out posteriorly in rapid succession, until on posterior third only four are present; ventrals strongly convex posteriorly, with free posterior edges, nearly half length of scales; anal plate divided, with diagonal suture; subcaudals in double series, those of right and left sides alternating; several minute subcaudal-like scales on each side of vent. NATURAL History OF THE RACER 369 Dentition In the racer the maxillary, palatine, pterygoid, and dentary bones bear teeth (Fig. 1). The teeth are all much alike in size and shape, small, sharp, and recurved, typically at an angle of approximately 50 degrees. The number of teeth present is variable. Because the teeth are small and loosely attached to the jaw bones, and often are broken off in the capture and ingestion of prey, each bone usually lacks part of its complement of teeth. Even the sockets vary some- what in number between individuals, and between the left and right sides in some snakes. Most of the skulls that I examined were not thoroughly cleaned, and the adherent dried tissues made it dif- ficult to obtain accurate counts of the sockets. In ten skulls from Kansas and Nebraska, most frequently occurring numbers of sockets for each of the dentigerous bones were: maxillary, 15; palatine, 11; pterygoid, 18; dentary, 18. Fic. 1. Lateral view of right side of skull of adult female blue racer, x 4. University of Kansas Museum of Natural History no. 18305, from Greenwood County, Kansas, Hemipenis Penial characters have proven to be useful in the classification of snakes, providing bases for separating subfamilies, genera, and species. In the racer even the subspecies have trenchant penial characters by which they may be spearated in some instances. The hemipenis is roughly cylindrical, but widest at the base (Fig. 2). The sulcus spermaticus is unbranched. Approximately the basal one-third of the hemipenis has a smooth surface, broken only by the sulcus spermaticus and by three greatly enlarged spines, which form 370 UNIVERSITY OF Kansas PusBts., Mus. Nat. Hist. hooks—one anterior, one posterior, and one dorsal. The dorsal hook is the largest of the three. Distal to the smooth part is a zone of small spines, each recurved and mounted on a fleshy tubercle. The bee eee oe Sot to SED ase as 2 oy ee De oe ia Fic. 2. Lateral view of injected and everted left hemipenis (slightly en- larged) of a blue racer from the Rocke- feller Tract, Jefferson County, Kansas, showing heavy spines at base of organ, small spines of central zone and la- mellae of terminal part. This hemi- zone of spines is poorly de- veloped on the anterior side and is interrupted on the posterior side in the vicinity of the sulcus spermaticus but is best developed on the pos- terior side a short distance above and below the sulcus spermaticus. The spines are arranged in several oblique rows. Those of the proximal row are best developed, and there is rapid diminution in the size of those situated farther distally. | Approxi- mately the distal two fifths of the hemipenis forms a third zone, lacking distinct spines, but having numerous deep longitudinal grooves, alternating with lamel- Jae which have fimbriated edges, and which fuse with each other and divide to form a reticulated pattern. penis is not fully engorged. Relationships The large genus Coluber is much in need of revision. Its many species, perhaps more than a score in all, occur in North America from southern Canada south to Guatemala, in eastern and south- western Asia, in southern Europe, and in North Africa. All are active, slender-bodied snakes having smooth scales in few rows, and having large eyes with well developed vision. The North American species fall into two natural groups, the typical racers, and the whip snakes, often assigned to a separate genus, Masticophis (Ortenburger, 1928). The whip snakes are more specialized than the typical racers in having the eyes more enlarged, and the body form more slender and attenuate, with number of scale rows more reduced. The racers of the Old World are more diverse. Inger and Clark (1943) suggested a partitioning of the genus Coluber on the basis of the pattern by which scale rows are reduced, from the maximum number on the forebody to the minimum number at the posterior end of the body, supplemented by certain characters of the hemipenis and of the cephalic scutellation. Besides Coluber NatTurAL History OF THE RACER 371 and Masticophis these authors recognized within the group the genus Platyceps with several species in southern Europe and south- western Asia; Zamenis with several species in the same region and in North Africa, and Haemorrhois, a monotypic genus of Spain, North Africa and several Mediterranean islands. Although apparently valid in principle, this arrangement has not been generally followed. Exclusive of those species groups whose assignment to the genus Coluber are somewhat questionable, the remaining species in the genus are: C. constrictor occurring throughout most of the United States and south along a narrow Atlantic coastal strip of Mexico to Guatemala; C. oaxacae of southern Mexico; and C. spinalis of north- ern China. C. oaxacae is poorly known as only a few specimens have been collected, but seemingly it is a near relative and deriva- tive of C. constrictor, especially of that species’ southernmost popu- lation. C. spinalis is much more distinct, as might be expected from its geographical remoteness. It is a slender, active snake, of olive coloration dorsally with 17 scale rows and a bright yellow, black- edged dorsal stripe and yellow ventral surface. It is relatively small (up to 755 millimeters snout-vent length) and is partial to riparian habitats but is also found in forests and in dry and barren regions (Pope, 1935:224-226). It is known to feed upon lizards. Range The common racer has been recorded in each of the 48 states of the mainland of the United States, also in New Brunswick, Nova Scotia, southern British Columbia, and southward through Mexico where it is limited to a narrow strip of east coast lowlands but ex- tends as far as Guatemala. C. c. constrictor occupies the northeast- ern states and extends south into the Appalachian and Piedmont. C. c. priapus with its associated races paludicola, helvigularis, and anthicus has an Austroriparian distribution, occupying the Atlantic Coastal plain and the Gulf Region, and extending north in the Mis- sissippi Valley to southern Illinois and Indiana. C. c. paludicola is localized with two disjunct populations—in the Everglades and on Cape Canaveral, Florida. C. c. helvigularis is even more restricted in range and is known only from the Appalachicola region of the Florida Panhandle and the adjacent corners of Alabama and Georgia. C. c. anthicus occupies much of central and western Louisiana and adjacent Texas. C. c. flaviventris occurs throughout the Great Plains, east in the “Prairie Peninsula” through Michigan and northern Ohio and west to the Rocky Mountains. C. c. 372 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. stejnegerianus occurs from Matagorda Bay in Texas southward through eastern Mexico, with a seemingly isolated population in the Sierra del Carmen region of northern Coahuila. C. c. mormon occurs in the Pacific Coast states and Great Basin. Actually, the range limits and the continuity of distribution within the area outlined are still poorly known. The species has not been recorded from the northern parts of Maine, Vermont, New Hamp- shire, Michigan, Wisconsin, or Minnesota, nor from northeastern New York. It is generally absent from southwestern desert areas. Records are particularly scarce and scattered in the Rocky Mountain states, suggesting that the distribution in this area may be discon- tinuous. In a large area comprising all of New Mexico and Arizona, the western half of Colorado, and the southern halves of Utah and Nevada, records are so scarce as to indicate that the species is there represented by only a few well isolated relict colonies. The type locality of mormon is “Valley of the Great Salt Lake,” and there are numerous records from the northern part of Utah east of Great Salt Lake (Woodbury, 1931:75), but a record from Moab is the only one known to me from the southern half of the state. The only records from western Colorado are from three miles east of Fruita and two miles west of Grand Junction, Mesa County (Maslin, 1959:56). Apparently the only valid record from Arizona is that of Shannon (1950:59) from Eagar, Apache County, in the east- central part. Shannon also recorded the racer from Boulder Dam in extreme southern Nevada. Brattstrom (1955:152) has recorded the species from the lower Pleistocene of southeastern Arizona (Curtis Ranch), bearing out the idea that the racer has partly withdrawn from a range formerly occupied in the Southwest at a time when cooler and moister climate prevailed. Other fossil oc- currences are of late Pleistocene age—Vero Beach and Seminole, Florida (Brattstrom, 1953a:245) and, doubtfully, Rancho LaBrea, California (Brattstrom, 1953b:376). The range of mormon has been mapped (Wright and Wright, 1949:134) as extending east to south-central Montana on the basis of one specimen allocated on the basis of two characters. Otherwise the range of mormon seems to be entirely west of the Continental Divide, well separated from that of flaviventris by desert and mountain barriers. The con- specificity of mormon with the other subspecies needs to be more thoroughly investigated, and geographic variation within mormon also merits study. NATURAL History OF THE RACER 373 Geographic Variation The common racer and the several species of whip snakes (Masticophis) were revised by Ortenburger (1928). More recently with much larger series of specimens, Auffenberg (1955) again re- vised the classification of C. constrictor, but his study was concen- trated in Florida and neighboring southeastern states with relatively little attention devoted to populations of the western and central United States. As the species occurs throughout most of the United States and south through the coastal lowlands of eastern Mexico to Guatemala, it is found over a wide range of environmental condi- tions. Various characters are subject to geographic variation, and some of them follow clines that are maintained over extensive areas. Such characters as the number of hemipenial spines, and the enlargement of one or more basal spines into hooks, the shape of the premaxillary bone, the number of maxillary teeth, the num- bers of ventrals and caudals, color of eye, number of dorsal saddle- marks and of ventral spots in juveniles, and ratios of body propor- tions including tail length to total length have been used to characterize the subspecies. Also important is the shade of coloration of adults. The sub- species constrictor, priapus and helvigularis that are characteristic of forested habitats in the eastern United States are black dorsally and have their ventral surfaces suffused to a large extent with dark or dusky coloring. Farther westward the ground color becomes progressively paler, greenish, grayish or light brown, and the ventral surface is yellow (white on the throat and neck). The same tendency appears in C. c. paludicola of the Everglades. The popu- lations of arid climates in southern Texas and in the far western states are relatively pale colored. The species thus conforms to Gloger’s Rule in changing from a pallid coloration in arid climates to a dark pattern with eumelanins predominating in a humid climate. Perhaps the most important character that is subject to geo- graphic variation in the racer, and certainly one of the most neg- lected, is body size. With information now available it is not pos- sible to compare the sizes of different populations except in a general way. The best sources of information concerning size in several subspecies other than flaviventris, are the publications of Auffenberg (1949 and 1955). Comparison of these data with my own is not entirely satisfactory because Auffenberg did not indicate clearly differences in size between the sexes, nor indicate the 874 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. boundary line between young and adults. Also, his measurements are of overall length. For the purpose of comparison I have sub- tracted 22.5 per cent, an approximation of ratio of average tail length, from Auffenberg’s figures. He stated (1955:98) that the series of specimens on which measurements were based were those “with a uniform coloration,” that is to say they had lost the juvenal pattern and were probably sexually mature. Whether the same statement applied to the large series of stejnegerianus in the same author’s earlier paper (1949:55) is doubtful. C. c. constrictor: 84 New York specimens averaged 806 millimeters ( Auffen- berg, 1955:96). C. c. flaviventris: 100 Kansas specimens averaged 791 millimeters (males, 746; females, 836). . c. priapus: 171 Florida specimens averaged 713 millimeters ( Auffenberg, G 1955:96). C. c. stejnegerianus: 291 Texas specimens averaged 664 millimeters (Auffen- C C berg, 1949:54). . c. mormon: 94 West Coast specimens (Museum of Vertebrate Zoology ) averaged 563 millimeters (males 515, females 600). . c. anthicus: 35 northern Louisiana specimens estimated to average 582 millimeters (Clark, 1949:249—the author did not present individual measurements, but indicated the numbers in several size groups in his sample). The 100 flaviventris in the above list were recorded in June, July and August, a season when the young of the preceding year are still small, and these young were not included. In a fall sample 63 males averaged 729 millimeters and 65 females averaged 886 milli- meters, but with the nearly grown young (44 males and 40 females) included, the averages were changed to 651 and 768 respectively. Maximum length perhaps express differences between the subspecies as well as, or even better than, the averages listed above. The fol- lowing figures indicate some of the maximum overall length meas- urements in inches that have been published by various authors. These measurements pertain to females unless otherwise indicated. C. c. constrictor: 74%, 74%, 73, 65% (Auffenberg, 1955:96). . c. flaviventris: 71 (Pope, 1944:172), 72 (Conant, 1958:147). .c. priapus: 52% ( 6) (Auffenberg, 1955:98). . c. stejnegerianus: 87 (Auffenberg, 1949:54), 40 (Conant, 1958:148). .c. mormon: 51% (Museum of Vertebrate Zoology ). .c. anthicus: 70 (Conant, 1958:149). In my own study, the largest racers recorded from the Reservation and Rockefeller Tract had the following overall lengths: 59, 57%, 55%, 58 ¢ (projected from stub tail). QQLALAKA NATURAL History OF THE RACER 375 These sets of figures show that compared with the blue racer in Kansas, with which my own study was concerned, the black racer of the northeastern states reaches a larger size, while the racers of the Southeast and especially those of the far West and of southern Texas, are dwarfed. These size differences are almost certainly correlated with behavioral and ecological differences among the snakes involved. Adaptations to exploit certain types of prey and to utilize most efficiently certain habitats and types of cover, have led to divergent trends in different parts of the range. Habitat The racer dwells primarily in open situations, but as might be expected from its extensive geographic range, bringing it under the influence of diverse climates and habitats, its populations have diverged somewhat in adaptation to different environmental con- ditions. The eastern blacksnakes (subspecies constrictor and priapus) seem to prefer woodland and forest edge. In central Louisiana, anthicus occurs chiefly in an open type of woodland. The subspecies stejnegerianus is found chiefly in brushland and thorn forest. The western mormon is found in varied habitats, in- cluding moist streamside meadows, and chaparral. Published state- ments of herpetologists, based upon studies in limited areas, are briefly quoted below to show the trend of geographic change. C. c. constrictor: This snake “occurs chiefly in fields” (Atkinson, 1901:148; Pennsylvania); “in more or less wooded regions and along hillsides and among bushes” (Surface, 1906:167, in Pennsylvania); “abundant, especially in wooded regions” (Hibbard, 1936:28, in Kentucky); “dry and more or less open places” (Conant, 1938:52, in Ohio); “old fields and areas about buildings” (King, 1939:572, in Tennessee). C.c. priapus: In “drier parts of the [Okefinokee] swamp . . . seems to prefer blueberries and saw palmettos” (Wright and Bishop, 1915:159 in Georgia); “common in grassy fields and in upland woods” (Allen, 1932:18, in Mississippi); “abundant along fence rows . . . in dry pine-oak forest and in bottomland forest” (Trowbridge, 1937:296, in Oklahoma); “probably most abundant in open upland hammock or in old fields; limestone flatwoods” (Carr, 1950:80, in Florida); Oak and oak-hickory forest and small hill prairies in southern Illinois (Rossman, 1960:219). C. c. paludicola: In “all pet of the freshwater Everglades, in cabbage palm hammocks, in salt marshes, and in mangrove swamps. On Key Largo - . . in mesophytic hammock” (Duellman and Schwartz, 1958:296, in southern Florida). C. c. anthicus: In “wooded areas in the vicinity of briar patches or other brushy undergrowth” (Clark, 1949:249, in northern Louisiana); “especially grassy uplands” (Fitch, 1949:88, in central Louisiana). C. c. stejnegerianus: Of 291, 94 were in scattered brush, 92 in sparse brush, 41 in lightly wooded areas, 26 in grassy areas, 24 in heavy brush, eight in semi- arid places and six in heavily wooded situations (Auffenberg, 1949:55, in southern Texas). 376 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. C. c. mormon: In “thin brush skirting open prairie land” (Lord, 1866: 304, British Columbia); cottonwood-willow and water margin habitats in prairie (Dice, 1916:310-312, in eastern Washington); “grass; amid water cress; bank of small ditch near meadow; barley field; sandy ground covered with rocks and driftwood; among sedges; in sagebrush; swimming in irrigation ditch” (Grinnell, Dixon and Linsdale, 1930:149, in northeastern California); low foothills, around the fields, and in the timber and brush along the canyon bottoms (Woodbury, 1931:75, in Utah); “open woods of Garry oak and poison oak, on grassy slopes, in chaparral, and in grain or hay fields” (Fitch, 1936:644, in southwestern Oregon); “low hot canyons where it was found to occupy areas having rather dry, rocky hills” (Ferguson, 1952:68, in northeastern Oregon). C. c. flaviventris: Pastures, meadows, and fields (Hurter, 1911:170, in Missouri); “usually frequents dry open fields, although it is often found in bushes or cut-over land which has grown up into thickets” (Ortenburger, 1928:181); “pasture lands and on hill sides” (Peters, 1942:183, in Illinois); “along the levees in the salt marshes” (Liner, 1954:82, in southern Louisiana); “common in both prairie and woodland habitat” (Smith, 1947:34, in east- central Illinois); Flood plain, sand around sage-sumac brush, rocky slopes (Fouquette and Lindsay, 1955:411, in northwestern Texas). Several observers have described the habitat in Kansas as follows: “grassy valleys and thinly wooded hillsides” (Burt, 1927:5); “moist and dry habitats, in wooded areas, and in prairies” (Smith, 1956:237); Oak-walnut hillside forest, cultivated field, buckbrush-sumac, prairie (Clarke, 1958:22). Every part of the 590-acre Reservation is used to some extent by blue racers living on this area. Home ranges of most individuals are so large as to include a variety of habitats, both woodland and grassland. The habitat preferences vary according to season. In autumn some racers migrating to rock ledges to hibernate are found in mesic forest, but at that time of year leaves have begun to fall and the forest floor is less dark and humid than in summer. In spring also racers not yet back on their summer ranges are often seen either along the hilltop ledges, or moving downhill through woods toward bottomland meadows. However, in summer, the finding of a racer in mesic woodland is a rare event. Occasionally the snakes make trips across such blocks of woodland hundreds of feet wide, but they do not linger in the woodland. In decreasing order of preference the habitats used by racers on my study area may be classified as follows: 1. Tall-grass prairie, (Plate 22) either original or regenerated, dominated by native perennial grasses in stands three feet to seven feet high, including big blue-stem (Andropogon gerardi), little blue-stem (A. scoparius) Indian grass (Sorghastrum nutans), and switchgrass (Panicum virgatum). 2. Pastureland, with introduced brome grass (Bromus inermis) and as- sociated weedy vegetation. 8. Brush, in ravines, at woodland edge, and in riparian thickets. 4. Weedy fields, dominated by such pioneer plants as ragweeds, sunflowers, horseweed, milkweed, wild lettuce, aster and goldenrod. 5. Open type of woodland dominated by such trees as honey locust and osage orange. NaATuRAL History OF THE RACER Sm These habitat types are interspersed on the study area, and each racer has a wide choice of habitats without travelling beyond the limits of its own chosen area. Grassland that has been closely grazed, mowed or burned does not provide entirely adequate food or shelter, and under such con- ditions clumps of brush or other dense vegetation may be of critical importance. Throughout the racer’s extensive range, fields of grain and hay at times provide suitable habitat, and may support large populations, but in spring, before the young cultivated plants have made much growth, or later in the season, after they have been cut, the racer may need to depend on adjacent areas of pasture, thicket, or woodland edge and the availability of such refugia to a large ex- tent determines the numbers of racers on cultivated areas. Temperature Relationships In the locality of my study racers spend approximately half the year in winter dormancy. Earliest spring records and latest fall records for 13 years are shown in Table 4. The spring records in nearly all instances pertain to snakes found in the open or beneath flat rocks warmed by sunshine, usually at or near the rock ledges where hibernation occurs. Juveniles are especially well represented in these earliest spring records, and it seems that they tend to emerge a little earlier, on the average, than the adults, either because they have hibernated in more superficial and less well insulated situations or because their lesser body mass permits more rapid warming to activity than can occur in the adults. The latest fall records all pertain to racers trapped along the rock outcrops, and none was a young of the year. TaBLE 4. EARLIEST AND LATEST RECORDED ANNUAL Dates WHEN BLUE Racers WERE ACTIVE ON THE RESERVATION OR ROCKEFELLER TRACT Earliest Latest Earliest Latest YEAR spring fall YEAR spring fall record record record record 1950. April 16 | November 4 BY (aie cide April 30 | November 2 1ObIR ae April 19 | November 13 || 1958..... April 25 | November 20 1952. April 23 | November 12 || 1959..... April 6 November 10 1953 .cmat April 8 November 12 || 1960..... April 22. | November 6 WQ54:. cies April 20 | November 12 || 1961..... April 19 | November 1 O55 cae April 15 | November 2 196223200 April 23 | October 27 1956. May 11 | November 14 378 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. Most of the population undoubtedly emerged somewhat later than the average date of April 16 indicated by the records in Table 4, and retired somewhat earlier than the average date of November 8. However, a small percentage of the population probably emerged even earlier each year than my records indicate, and re- tired into hibernation later than my records indicate. In a typical year, temperatures in April and early May are only occasionally above the level at which racers are able to become active, but are below this threshold most of the time. The same statement ap- plies to an autumn period of late October and November. Most racers are dormant in their hibernacula during these transitional periods of spring and autumn, but some—those that have emerged early in spring, or those that have not yet retired (in fall)—retreat to temporary shelters and revert to a semi-torpid state when tem- peratures fall below the critical level. Fig. 3 shows the relative extent of acnvity along the hilltop out- crops, as reflected by numbers of racers caught at different times during the autumn. Data from 14 years are combined, and the large composite sample indicates that in an average year there is relatively little activity along the hilltop outcrops in early Septem- ber, but that activity rapidly increases to a peak in mid-October and then tapers off rapidly, usually ending in mid-November, but occasionally ending as early as late October or as late as late No- vember. The racers recorded in traps had, in many instances, been con- fined in them for from one to three days before they were found. For any one year records are not sufficiently numerous to show the trend as well as Fig. 3, but Fig. 4 shows year-to-year differences; 1958 was a fairly typical year, and also was the year in which the largest sample was obtained; in 1949 the largest catches were made earlier than usual, and the racers retired early into hibernation; in 1954 warm weather persisted until unusually late in autumn, and racers remained active beyond the time when they ordinarily would have been hibernating; in 1955 and 1961 the most concentrated activity along the outcrops, as reflected by day to day catches, came later than usual, but unseasonably cold weather ended all activity abruptly, earlier than usual. The racer’s annual cycle of activity is, of course, controlled pri- marily by the weather, and is much delayed or accelerated in some years. But certain stabilizing factors cause the racer’s annual cycle to be less variable than that of the weather. For example, in spring NATuRAL History OF THE RACER FEMALES RECORDS NUMBER OF DEGREES C SEPT. OCT. NOV. Fic. 8. Records of blue racers trapped along hilltop limestone outcrops in autumn, a composite sample of 14 years (1949 through 1962) from the Reservation and Rockefeller Tract, showing the catch grouped in ten-day intervals, beginning with September 1 to 10 and ending with November 20 to 29. Averages of the maxima, means, and minima of daily temperatures for each period are shown. 379 380 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. when persistently cool weather delays emergence from hibernation beyond the normal time, the increasing azimuth of the sun, and more intense sunshine cause the soil to warm, despite low air tem- perature, until emergence is finally triggered. Having once emerged, the racer is able to control its bodily temperature to a large extent by basking in sunshine to gain warmth, or by seeking shade or underground shelter to escape overheating. By such behavioral thermoregulation extremes of weather are neutralized, or at least buffered to some degree. In the course of my study no racers were found in their actual hibernacula. Insofar as known, these were always in deep crevices in strata of limestone near hilltops, and were inaccessible except by removal of the solid rock. The crevices where racers hibernate are known to be several feet deep in some instances, extending well below the frost line. Periodic temperature readings in two such crevices at depths of 12 inches and 30 inches, taken in the winter of 1954 (Fitch, 1956:471) showed that temperatures during dor- mancy are usually well within the range 0° C to 10°. Whether the oN asa cad - eee? —o" << NUMBER OF RECORDS Fic. 4. Yearly variation in catch of blue racers along hilltop outcrop in autumn on the Reservation and Rockefeller Tract, grouped in same ten-day intervals indicated for Fig. 8. NATURAL HisTrory OF THE RACER 381 racers congregate in hibernating masses in regular “dens” on the Reservation has not been definitely determined, but if so, most of the hibernating groups must be small, because those trapped along the ledges are well scattered, and, in fact, give the impression of being rather uniformly distributed along them. However, ledges of northward exposure are not used as hibernation sites, so far as known, and those of full southward exposure are perhaps pre- ferred, especially where the hilltop has a southward projecting spur, and the exposed rock face is extensive, with many cracks and fissures. I have been unable to detect differences in types of hibernation sites preferred between the racer and the copperhead, which is somewhat more numerous on the same area. Several authors have contributed to knowledge of hibernation in the racer. Boyer and Heinze in Missouri (1934:195) noted that blue racers often were associated with copperheads in the vicinity of places chosen for hibernation. Burt (1935:329) in Kansas found blue racers emerging from dens among rocks on a prairie hillside, associating with other snakes, Diadophis punctatus, Elaphe guttata, and Pituophis melanoleucus. In the Chicago region, Pope (1944: 173) reported scores of blue racers aggregating in October on and around a sand dune with oak woodland. In Ohio, Conant (1938: 55) found three blue racers hibernating together about three feet underground in a small hole. One found at another locality had apparently hibernated in company with a massasauga (Sistrurus catenatus). In Maryland Cohen (1939:187) saw racers (C. c. constrictor) in the act of emerging from an old vole burrow that was a communal hibernaculum, on April 6, 8, and 10. Air tem- peratures at the time of emergence were 12.5° and 18.5° Centigrade. In Illinois, Schroder (1950:1-2) found seven blue racers hibernating in masses, intertwined with each other and with bull snakes at depths of 36 inches and 42 inches in an abandoned mammal burrow in a sand dune area in early February. In the course of routine field work I often carried a Schultheis quick-reading thermometer, and from time to time I had opportuni- ties to take the body temperatures of blue racers newly captured by hand. The trend of these records indicated the temperature range within which the snakes normally limit their activity, and the preferred temperature. In an earlier publication (Fitch, 1956: 459-460) based on a few more than half the number of records of temperature now available, I discussed responses of the blue racer to temperature. The newer data bear out the trends previously 382 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. revealed; of 60 records, 39 are within the six-degree range from 29° to 35°, and records are most concentrated in the one-degree range, 34° to 35°. Racers were found active at air temperatures between 15.5° and 32.4°, with the greatest concentration between 26° and 27°. Compared with most other kinds of North American snakes, the racer is remarkably tolerant of heat, and often is several degrees warmer than the level that those of other genera will normally tolerate. Racers have been seen crawling in the sun- shine, or basking on days that were uncomfortably hot for humans. For example, on August 28, a large female racer released from a trap was followed and her behavior observed; after crawling some 50 feet through the grass she climbed from a ditch bank onto sun- flower stalks and elm saplings, and came to rest among the stalks, NUMBER OF RECORDS 25 30 35 40 DEGREES CENTIGRADE Fic. 5. Bodily temperatures of blue racers kept in a large outdoor enclosure and checked from time to time when they were active and the sun was shining, in June and July, 1962. NaTuRAL History OF THE RACER 383 NUMBER OF RECORDS 20 25 30 35 DEGREES CENTIGRADE Fic. 6. Bodily temperatures of blue racers captured by hand in their natural surroundings. The preferred level is approximately the same as indicated by Fig. 5 (between 29° and 86° Centigrade), but some of the racers caught were not fully active and had lower temperatures. Some bias results from the fact that those having the lowest temperatures were the least active and hence were most easily caught. NUMBER OF RECORDS 15 20 25 30 DEGREES CENTIGRADE Fic. 7. Air temperatures recorded at captures of the racers whose records were used in Fig. 6. An active racer typically maintains, by basking, a bodily tem- perature several degrees warmer than the air. 38—7864 384 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hisr. NUMBER OF RECORDS 30 DEGREES CENTIGRADE Fic. 8. Bodily temperatures of blue racers found in live-traps at Harvey County Park. Opportunity to regulate temperature by behavior was limited in these snakes in traps, which tended to match ambient temperatures. in a spot strategically situated for catching grasshoppers. More than half her body was exposed to sunshine and air temperature was slightly above 34° Centigrade, yet the snake showed no sign of dis- comfort, and for the several minutes that she was kept under ob- servation, did not attempt to withdraw into the shade. At the Harvey County study area, bodily temperatures were re- corded in many of the racers that were caught in traps. These records are much less significant than the records obtained from racers caught by hand and promptly checked for bodily tempera- tures. The temperatures of the trapped snakes may, to a large extent, reflect the temperatures of air and soil at the time. How- ever, despite their confinement, the trapped racers probably were able to exercise some control over their temperatures by shifting from shade to sunshine, or from the top of the trap to its bottom, where they would be in contact with the substrate. In most of the racers removed from traps, as in those caught by hand, bodily temperatures were somewhat above air temperatures, but the dif- ference was less in the former group of snakes. Figure 8 shows the bodily temperatures of these snakes removed from traps. Oc- casionally racers died in the traps from overheating. On July 2, 1960, each of two racers in traps had temperatures of 39.4°. One of these was especially vicious and frantic in its attempts to escape, but otherwise seemed unharmed. The second racer was dead, seemingly having succumbed just before it was found. Probably NATURAL History OF THE RACER 385 prolonged exposure to temperature in excess of 39° would always result in death of the snake. Racers and other snakes that had become overheated in the traps and were nearing exhaustion had a characteristic limp feel when they were handled. In June, 1960, heat tolerance of a halfgrown racer was compared with that of several other snakes including a copperhead, garter snakes, and ringneck snakes. Each snake in turn was enclosed in a plastic tube plugged with cotton at one end, the snake having a quick-reading thermometer taped in place for a rectal reading. The tube was then placed in sunshine. Over periods of minutes the enclosed snake passed through a characteristic cycle. Soon it would begin to register discomfort as its temperature rose rapidly. Its struggles would become increasingly violent, then would cease abruptly. The snake would suddenly collapse, its body mostly limp, but knotted in slow contortions, its mouth gaping widely. Within a few seconds all movements would cease, but in each instance the seemingly dead snake was soon revived by holding it in cold run- ning water. The copperhead, garter snakes, and ring-necked snakes all collapsed at temperatures near 41°. At this same temperature the racer showed signs of acute discomfort, but did not collapse even after many minutes of exposure. Probably more protracted exposure at this level would have been fatal to the racer as well as to the other kinds tested. Home Range and Movements Blue racers that were recorded on more than one occasion were rarely caught again at the original location. For different indi- viduals, distances between capture points ranged from zero up to a little more than three-fourths of a mile. The area of concentrated study was a mile and a half in greatest diameter; there was scant opportunity for capturing racers that moved greater distances. Even those that moved as far as a mile would have passed beyond the boundaries of the study area in most instances. Many of the marked racers that disappeared from my records probably moved beyond the limits of the study area. Nevertheless, in the great majority of instances, the distances between successive capture points for the same individual were relatively short, indicating that each racer tends to remain permanently in a restricted area. Most captures were made in the type of grassland or brush that provides favorable habitat for the racer during the season af activity, but many other captures were made in woodland along the rock 386 University OF Kansas Pusts., Mus. Nar. Hist. ledges where the snakes come to hibernate. Four different types of movement may be recognized: 1) those in the rock ledge area where hibernation occurs; 2) those between the area where the summer is spent and the hibernation ledge—an actual small scale seasonal migration which takes place in spring and autumn—3) those within a home range, which are part of the day-to-day activi- ties of the racer, and, 4) wandering movements by which the racer shifts its activities, perhaps permanently, from one area to another. In the records of any one snake these different types of movements cannot always be sorted with certainty. Each type will be discussed separately. Relatively few movements along the ledges were recorded. It seems that having migrated to a ledge, the racer promptly finds its hibernaculum and retires for the winter. In spring there is equally prompt scattering of the emerging racers, which no longer find the ledge attractive. Most recorded movements along the ledges were short. Of 76 movements, nine exceeded 1000 feet, and only four others exceeded 500 feet. Most of the shorter movements were re- corded within an autumn season, but several were recorded after the lapse of one or more seasons of activity. The longer move- ments were as follows: 1250 feet after 8 seasons (male); 1300 feet after three seasons (female); 1600 feet after one season (female); 2000 feet after one season (female); 2280 feet after seven seasons (male); 2200 feet in same season (female); 2410, 2600, and 3200 feet, each after one season (all males). The trend of these records suggests that the tendency to return year after year to the same hibernaculum is not strong; after using one for a period of years, the racer may abandon the stretch of ledge and, starting out in the opposite direction from its summer range, find a new hibernaculum as much as half a mile from the old one. Records of distances be- tween capture points on the ledges for individual racers are shown in Fig. 12. A total of 124 movements between summer ranges and ledges were recorded, and the distances averaged 1309 feet—approxi- mately a quarter mile. Some racers living in hilltop fields may have had home ranges that included rock ledges, or at least were adjacent to them. In such instances no seasonal migrations would have been necessary to reach hibernacula in the autumn and sum- mer ranges in the spring. Several short movements—100 feet, 150 feet, and 200 feet—can be explained on the basis that home ranges and hibernation ledges overlapped or were near at hand, but most NATURAL History OF THE RACER 387 of the movements were longer. The longest movement was 4020) feet, after a lapse of four seasons. Twenty-four movements exceed- ing 2000 feet were recorded. For these the intervals between captures averaged more than double the time for the remaining movements, indicating that the longtime permanent shifts were in- volved in many instances. For all the racers living in bottomland, ranges were separated from ledges by areas of wooded hillsides averaging approximately 700 feet across. These relatively unfavorable areas had to be traversed in the course of the semi-annual migrations. Even some 0 1000 2000 DISTANCES OF MOVEMENTS: FEET Fic. 9. Histogram of movements of blue racers between hilltop rock outcrops used for hibernation, and summer habitat on the Reservation and Rockefeller Tract. Movements of females tend to be somewhat shorter than those of males, of the racers that lived in hilltop fields apparently crossed wooded slopes in order to reach distant hibernation ledges, or else each reached the ledge by a roundabout route although it could have found a ledge much nearer its summer range. For the 124 ledge- to-field and field-to-ledge movements, the median distance was 1030 feet. The sexes were almost equally represented in this sample but the average distance for the 55 males—1425 feet—notably ex- ceeded that for the 69 females—1220 feet. These movements are shown in Fig. 9. McCauley (1945:76) in Maryland described what seemed to be incipient territoriality in a large male racer that remained several hours in a small area, crawling about conspicuously with head 388 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. raised, seemingly on patrol. When an even larger male racer in- truded, the first one aggressively drove him away, but neither paid any attention to a king snake that was also on the area. Other authors have noted the attachment of a racer to a small familiar area. Conant (1938:53) wrote that many of the racers he saw sought shelter in definite retreats. One of these racers was seen resting on top of a brush pile four times in a single afternoon, and each time it followed the same route to the same inaccessible spot beneath the brush. My own observations do not bear out the idea that racers main- tain regular territories, since several males may be present within a small area, even in the breeding season. Hostile behavior be- tween males has not been observed by me under natural conditions, and in confinement has been seen only in instances of self defense. Like the racer Conant observed on a brush pile, individuals may linger in the vicinity of a favored shelter or foraging area for periods of hours, but such associations are ephemeral, and soon the snake moves on. In a uniformly favorable habitat a racer may cruise about freely in tall grass or brush. Individuals that I have at- tempted to follow, after flushing them or releasing them from traps, often covered distances of 100 to 300 feet within periods of a few minutes before I lost them. In such instances I maintained suf- ficient distance between myself and the snake so that the latter was not actively escaping. Probably the snake was not aware of pur- suit in most instances, although I was able to glimpse it through the stems of grass, weeds, or shrubs, or was informed of its course by the swaying tops of grass and other vegetation. For many of the racers captured over periods of years it was possible to plot “minimum home ranges” in the areas that they oc- cupied. One caught 12 times in five consecutive years will serve as a typical example. There were seven locations involved; three captures were made at one point and two captures at each of two others; the other five locations were each represented by a single capture. One of the seven locations was for a capture made at a rock ledge in October, and hence can be eliminated from considera- tions of home range. The other six locations are based upon captures made from late May to early August, and they form a rhomboid pattern, with three locations in alignment on one side and two others inside the quadrangular figure formed by the five outlying points. Obviously such a group of records gives some idea of the location and extent of the snake’s activities but the in- NATURAL Hisrory OF THE RACER 389 formation is far from complete. As shown by Odum and Kuenzler (1955), a much larger series of records, usually several dozen, with eight or more marginal locations, is necessary to illustrate even an approximation of the actual home range. Under the conditions of my study such a series of records was unattainable. Few if any of the racers recaptured had more complete series of records than the one mentioned above. For 20 racers the records were sufficiently numerous and well distributed to permit plotting of minimum home ranges. One of these ranges was hexagonal, nine were pentagonal, eight were rhomboidal and three were triangles. In four instances the area encompassed was broken by woodland, indicating that the home range comprised two or three disjunct segments. In all instances the smaller segments were triangular. The 20 minimum home ranges averaged 6.6 acres (3.2 to 12.8). The 15 ranges of males averaged 7.3 acres, whereas the five ranges of females averaged only 4.5 acres, but the sample is too small to be relied upon for differences in the sexes. In an earlier publication (Fitch, 1958:73) I discussed an alterna- tive method for determining size of home range in animals that NUMBER OF RECORDS ld ai Ais nance, 0 | 500 1000 1500 2000 2500 3000 DISTANCES OF MOVEMENTS: FEET Fic. 10. Movements of blue racers within or between areas of summer habitat on the Reservation and Rockefeller Tract. The trends are much alike for males and females. 390 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. move about freely within a chosen area, not having their move- ments restricted by attachment to a specific home base. Ordinarily any two records of the animal within its home range will be separated on the average by a distance equal to half the diameter of the area. Assuming that home ranges in general tend to have a circular shape, except as restricted by limiting environmental fac- tors, the area can be easily computed from the average recorded movement—the home range radius. It is necessary, of course, to have a sufficiently large number of records of movements to ob- tain an average that is statistically reliable. A major problem is that of recognizing movements that involve an extension of the original range or a shift away from it to a new area. A few exceptionally long movements were recorded. If these are included in the computations of home range, they greatly increase the average distance, probably introducing error. Also, the number of exceptionally short movements was greater than might have been expected if all locations of capture are at random to each other. In some instances a racer newly released may have blundered into the same trap again, or into the trap at the opposite end of its drift fence. In other instances traps may have been so strategically situated with respect to preferred travel routes that they caught the same snakes repeatedly. In still other instances, the range of an individual might have been mostly outside the study area, with only one end or corner overlapping the trap sites. A total of 471 records for consecutive captures in field areas is available, 305 for males and 166 for females. In 20 instances suc- cessive sites of capture were the same and movement was recorded as zero. Of the 471 records, 207 involved a relatively long time span, including at least one hibernation period; the remaining 264 were based upon successive records within the same season of activity. The trends were much the same in the records involving a longer time span (up to four years) as in those records involving captures made in a single season, but for the longer periods there were some exceptionally long movements, and relatively few short movements of less than 100 feet. Records of male racers and those of females were used for separate computations. For each series, the ten per cent of move- ments that were longest and the ten per cent that were shortest were eliminated from consideration in calculation of the average distance between points of capture. For the remaining 244 records of males an average movement of 595 feet was calculated, and for NATURAL HisToRY OF THE RACER 391 132 records of females an average movement of 574 feet. These distances, if accepted as typical home range radii, would represent home ranges of 26.3 acres for males and 23.8 for females. In an earlier discussion of spatial relationships in the racer (Fitch, 1958: 119), based upon relatively scanty data, I estimated the home range to be approximately 23 acres in males. But with only nine records for female racers I calculated the home range to be 9.7 acres. The disparate figures obtained from plotting minimum home range and from calculating average home range radius are not ir- reconcilable, since a minimum home range based on only four or five points would ordinarily include only a fraction of the actual range. Distances up to 1500 feet are included in the calculation of home range. It seems that home ranges often have a diameter of this magnitude or a little larger, although the estimated average diameter is 1140 feet. Home ranges probably most often deviate from circular shape to form an ellipse, with one diameter markedly exceeding the other. Woodland, water, roads, buildings, or cul- tivated fields, or other areas that are unfavorable or uninhabitable often form the boundary of a home range and influence its shape. Many of the longer movements constituted clear-cut shifts in range. In one exceptional instance a large adult female captured in the northeastern part of the Reservation on June 22, 1950, was released 21 days later at a point 3900 feet southwest of the place of capture. On May 27, 1960 she was caught within 600 feet of the original location, seemingly having made a homing movement. Among the nine racers recorded to have made longest movements (exclusive of those movements made to or from hibernacula) four were recorded also to have made later long movements in the re- verse direction, probably returning, each to its original home range, although in every instance the return movement was somewhat less than the original. A female of two-year-old size when first cap- tured on September 2, 1957, was recaptured 3100 feet southeast on May 10, 1958. On August 7, 1959, she was recaptured again 2400 feet from the second location in the direction of the original capture. Similarly, in a three-year-old female a shift of 2730 feet was re- corded at the second capture after 21 months, and at the third capture 14 months after the second, a return trip of 2360 feet had been made. A second-year female made a trip of 2640 feet between May 17 and October 1, 1960; by May 1961 she had returned 2000 feet to the vicinity of her original capture. From one year to the next an adult male shifted 2450 feet; after another year he had 392 UNIVERSITY OF KANSAS PuBLis., Mus. Nat. Hist. moved back 1550 feet. Most of the longer movements recorded were those between home ranges in fields and hibernacula along ledges, but in this class of movements, distance was somewhat proportional to elapsed time. For 59 such movements exceeding 2000 feet the average was 3.1 years, whereas for 114 field-to-ledge movements of less than 2000 feet, average elapsed time was 1.6 years. This trend suggests that over periods of years a racer is likely to shift its range or its hibernaculum or both. s 3 TS tT 24754 ¥ eaten Ree 3 Bo treet ‘ WSO SSI ZF OrCe ss e* &.) SXeHILLTOP ROCK ¢ INTERMITTENT 2).0UT CROP _—7 STREAM ; Fic. 11. Map showing home ranges of five blue racers, as indicated by numer- ous captures in successive summers, in small valley where Reservation head- quarters are located, and spatial relations of their hibernacula, as represented by points of capture along hilltop limestone outcrops. In spring and autumn, traveling to and from hibernacula, the snakes migrate across wooded slopes. Each “minimum home range” is enclosed in a dotted line, and a distinctive symbol is used to show successive points of capture for each snake. NATURAL History OF THE RACER 393 Average elapsed time between captures was 7.7 months. In the 471 field-to-field movements recorded, 53—slightly more than eleven per cent—exceeded 1500 feet and can reasonably be considered shifts of home range. The average elapsed time between captures for this group of snakes was 9.5 months. The evidence suggests Fic. 12. Map of 600-acre area of Reservation and Rockefeller Tract, including pas where field study was most concentrated, showing movements of the 20 lue racers recorded to have shifted over the longest distances. The figure following the sex sign of each individual indicates number of months elapsed between the captures at the localities represented by the dots. 894 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. that, even in an area of favorable habitat, somewhat more than ten per cent of the racers in a population annually shift their home ranges somewhat, but that many stay in the same home range for periods of years or perhaps throughout life. Shifts in range were especially noticeable where availability of suitable habitat underwent seasonal change. Along the north edge of the Reservation, prairie adjoined cultivated fields where grain or hay was grown. Until late May, the cultivated crops made little growth and the fields were almost bare. They provided insufficient shelter for the racers, which tended to keep to the prairie, where old grass of the previous year’s growth furnished them with ample cover. Later in the season, crops of oats, wheat, and alfalfa con- stituted suitable cover for the racers, and many of them shifted their ranges to the cultivated fields, but corn and milo crops were much less adequate for their needs. After harvesting of crops, cover in the fields was again inadequate for the racers’ needs, and they tended to retreat to edge situations, or to adjacent prairie. Food Habits Methods of Obtaining Prey The racer hunts by stealth, but actively, obtaining its prey by keen eyesight and swift movements. Wright and Bishop (1915: 160) wrote that because of its great speed it can catch anything that moves on the ground. As a racer moves stealthily through dense vegetation, its dull, uniform dorsal color blends well with the sur- face litter of dead plant material. In prowling, the snake glides along rapidly and alertly, in a jerky fashion, with frequent momen- tary pauses and changes of direction. Because of its inconspicuous- ness, it is not likely to be detected by the prey until it is close at hand. The snake is ready to dash in pursuit of any small animal that flies, jumps or runs to escape. On August 27, 1955, my daughter observed a large racer hunting among tall weeds at the edge of the pond on the Reservation. Several times in the course of its movements, it flushed small frogs (Rana pipiens) and each time the snake darted in unsuccessful pursuit of the rapidly hopping frog. On several occasions I have been led to a blue racer by the distressed croaking of a frog that the snake had captured. In each instance, despite my cautious ap- proach, the racer saw me before I detected it, and then darted away, abandoning its prey. On one occasion, while I was still a few yards from the racer, and before the latter had detected me, the NATURAL History OF THE RACER 895 frog broke free and hopped away rapidly through tall grass and weeds, and after several leaps, hid, concealed by dense screening vegetation. The racer darted in pursuit but could not find the frog. For several minutes the snake persisted in an active search; with forebody elevated and head held high, it would turn first in one direction and then in another, with nervous, jerky movements, obviously keyed up to a high pitch of excitement. Then it became aware of my presence, lowered its head, and glided away rapidly, abandoning the search. Although the racer depends to a large extent on sight to find its prey, scent may play some part also, as indicated by the presence in the food of young mammals taken from nests, some probably found underground. Near Garnett, Kansas, on May 4, 1952, Richard B. Loomis found a racer attacking a collared lizard (Crotaphytus collaris) beneath a large flat rock. The lizard was retaliating by biting the snake’s neck. The posterior part of the snake protruded into the open, and its thrashing had directed the attention of the observer to it. Whether the racer first found the lizard under the rock, or followed it there after flushing it in the open is unknown. An encounter between a large blue racer and an adult Great Plains skink (Eumeces obsoletus) on August 30, 1948, was de- scribed as follows: “The skink, grasped by one flank, had twisted back and seized the skin of the snake’s neck in a bulldog grip, and they lay interlocked, motionless except for their rapid panting, and occasional straining of the skink to bite harder or of the snake to shift its grip and work its jaws toward the skink’s head. The racer broke the skink’s grip, and began to swallow it head first. When only the hind legs and tail of the skink still protruded from the racers mouth, I lunged forward in an attempt to catch both reptiles. With a sudden movement the snake disgorged the skink, which darted away into the grass and escaped” (Fitch, 1955:78). Composition of Food Many authors have contributed to knowledge of the racer’s food habits. In most instances the records have been few or casual, but several intensive studies have been made, notably by Surface (1906) in Pennsylvania, Ortenburger (1928) for the species as a whole, Uhler, Cottam and Clark (1939) in Virginia, Clark (1949) in Louisiana, Auffenberg (1949) in southem Texas, Hamilton and Pollack (1956) in Georgia, and Klimstra (1959) in southern 396 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. Illinois. However, the findings of different authors are not strictly comparable; some have made general statements concerning the food habits but have mentioned specific items only when these were considered unusual. Certain authors have listed individual prey animals eaten; others have indicated the percentages (in bulk or in frequency) that the different kinds of prey comprised. Some writers have identified food animals only in broad categories such as “insect,” “beetle” or “snake” while others have undertaken specific determinations for all the prey or for certain taxonomic groupings that were subjects of special interest. For the eastern black racer (C. c. constrictor) the following food items have been recorded: 1 robin (Turdus migratorius, Storer, 1839:226); 1 copper- head (Agkistrodon contortrix, Verrill, 1869:158); 1 weasel (Mustela sp.— presumably the diminutive M. rixosa—Atkinson, 1901:148); 3 undetermined mammals, 1 rabbit, 1 undetermined mouse, 7 voles (2 Microtus sp., 4 M. pennsylvanicus, 1 Clethrionomys gapperi), 1 undetermined bird, 2 robin eggs, 2 garter snakes (Thamnophis sirtalis), 1 water snake (Natrix sipedon), 1 grass snake (Opheodrys vernalis), 1 green frog (Rana clamitans), 1 wood frog (R. sylvatica), 1 grasshopper (Melanoplus femur-rubrum) 2 camel crickets (Ceuthophilus sp.), 5 moths (cecropia, regal, imperial), 4 beetles, 1 currant worm, 1 ichneumonid wasp (Nematus ribesii), 1 currant worm (Surface, 1906: 170); 1 ribbon snake (Thamnophis sauritus, Ditmars, 1907:282); 8 snakes (1 Liopeltis vernalis, 1 Storeria occipitomaculata, 1 undetermined), 6 white- footed mice (1 Peromyscus leucopus, 5 P. nuttalli), 1 vole (Microtus penn- sylvanicus), 16 crickets (9 Gryllus pennsylvanicus, 4 G. assimilis, 2 Miogryllus verticalis, 1 Nemobius fasciatus), 2 grasshoppers ( Dissosteira sp.), 1 lepidop- teran, 3 elaterid beetles (Ortenburger, 1928:200). Richmond and Goin (1938:310) recorded finding the stomach of a black racer crammed with June beetles (Phyllophaga). Conant (1938:58) recorded a black racer from Ohio that had a smaller individual of its own species in its stomach. The smaller snake contained a caterpillar. Uhler, Cottam and Clark (1939:34) found food in 16 of 34 black racers from Virginia. Mammals, including a shrew (Blarina brevicauda), a mole, a flying squirrel (Glaucomys volans), a microtine, and a mouse (Peromyscus sp.) made up 26 per cent, 2 worm snakes (Carphophis amoenus), 2 ring-necked snakes (Diadophis punctatus), and 1 water snake (Natrix sipedon) made up 25.6 per cent, 5 birds including a warbler and a sparrow, made up 17.75 per cent; 2 frogs (Rana sp.) made up 9.38 per cent, 1 fence lizard (Sceloporus undulatus) made up 6.25 per cent, and insects, including cicadas (Tibicen sp.) and larval lepidopterans, made up 15.09 per cent. In Indiana, Minton (1944:457) examined 11 food-con- taining stomachs; there were rodents in six, snakes in five, a tree frog in one, and insects (cicadas, large grasshoppers) in four, and another black racer was found swallowing a small box turtle (Terrapene carolina). In Maryland, McCauley (1945:75) examined eight digestive tracts and recorded a shrew (Blarina brevicauda) in one, an unidentified mammal in one, 2 small cicadas in one, 2 small chickens in one, a fence lizard (Sceloporus undulatus) in one, and frogs and toads (including Hyla crucifer) in one; a ninth snake had eaten a half grown rat. In Connecticut, Finneran (1948:124) observed a large black racer eating a 21-inch garter snake (Thamnophis sirtalis). Duell- man (1951:338) recorded a black racer in Greene County, Ohio, swallow- ing a large garter snake (Thamnophis sirtalis). In Kentucky, Barbour (1950: 104) recorded remains of an unidentified snake in one stomach. Many authors likewise have recorded food of the southern black racer (C. c. priapus). In Georgia, Wright and Bishop (1915:160) recorded finding 2 racerunners (Cnemidophorus sexlineatus), a skink (Lygosoma laterale), 4 green tree frogs (Hyla cinerea) and 1 pine woods tree frog (H. femoralis) in NATURAL History OF THE RACER 397 stomachs. They also stated that the toad (Bufo lentiginosus [= terrestris] ) was the most important article of food. Burt and Hoyle (1934:205) wrote that a racer from Rogers County, Oklahoma, had eaten an adult male collared lizard (Crotaphytus collaris). In Florida, Carr (1950:80) found one of these black racers eating a leopard frog (Rana pipiens). Hamilton and Pollack (1956:523) examined digestive tracts of 62 and found food in 57, comprising the following percentages by volume: Lygosoma laterale, 34.2; Eumeces fasciatus and E. egregius, 11.8; Cnemidophorus sexlineatus, 8.8; Sceloporus undulatus, 3.5; undetermined lizard, 3.5; Opheodrys aestivus, 6.6; Diadophis punctatus, 8.1; Storeria dekayi, 1.6; Coluber constrictor, 1.8; Heterodon platyrhinos, 1.8; Masticophis flagellum, 1.8; Rana sp., 5.3; Hyla cinerea, 1.8; nue versicolor, 1.8; Peromyscus, 1.8; undetermined rodent, 1.8; lepidopterous Vaile In southern Illinois in an intergrading population of racers intermediate between C. c. priapus and C. c. flaviventris, Cagle (1942:188) examined several stomachs and found 1 chipmunk (Tamias striatus), 2 voles (Microtus sp.), 2 mice (Peromyscus sp.), 2 green snakes (Opheodrys sp.), 1 water snake (Natrix sipedon) and grasshoppers. From this same population Klimstra (1959:212) examined 137 digestive tracts of which 115 contained food as follows: 194 locustids, 118 gryllids, 17 undetermined beetles, 13 carabids, 6 scarabaeids, 10 pepidontetans, 9 hemipterans, 1 hymenopteran, 2 homopter- ans, 1 dipteran, 17 undetermined insects, 73 Peromyscus sp., 19 Microtus ochrogaster, 9 M. pinetorum, 12 Sylvilagus floridanus, 8 Scalopus aquaticus, 3 Rattus norvegicus, 4 Mus musculus, 2 Tamias striatus, 2 Synaptomys cooperi, 16 Rana pipiens, 8 Acris crepitans, 2 Rana clamitans, 2 R. palustris, 1 R. catesbeiana, 4 Hyla crucifer, 8 Pseudacris nigrita, 4 Lampropeltis calligaster, 4 Sceloporus undulatus, 4 Chrysemys picta, 1 Heterodon platyrhinos, 1 un- identified reptile, 4 Sturnella magna, 1 Otocoris alpestris, 4 unidentified birds. Percentages by volume of the various categories in this sample were: insects, 89.1; mammals, 32.9; amphibians, 10.8; reptiles, 8.8; birds, 6.3; miscellaneous, 6: Food of the “buttermilk snake” (C. c. anthicus) is known only through Clark’s study (1949:249). In an unstated number of examinations he found “mice” in 25, “rats” in five, lizards (Sceloporus undulatus and perhaps others) in eight, frogs (Rana pipiens) in seven, and birds in three. The food of C. c. stejnegerianus is known only from the work of Auffenberg (1949) but his sample was based on 206 racers that had food, among the total of 291 recorded. Unfortunately, he did not present actual numbers of the various prey animals, but divided the food into seven categories and listed these as percentages. He did not indicate whether the percentages represented volumes or numbers of individual occurrences, and evidently there was some error in computation since his combined percentages totalled 111. The categories and their percentages were as follows: grasshoppers, 42.5; crickets, 18.5; miscellaneous insects, .6; earless lizards (Holbrookia sp.), 40.1; scaly lizards (Sceloporus sp.) 2.1; frogs (Rana sp.) 10.0; rodents, 2.2. Auffenberg divided his sample of racers into five size classes, and showed that the smaller snakes fed chiefly on insects whereas vertebrates were increasingly prominent in the food of the larger snakes. The food of C. c. mormon is known chiefly through the work of Ortenburger (1928:228) who cited instances of a skink (Eumeces skiltonianus) and a young garter snake (Thamnophis sirtalis) being eaten, and listed the following items from 24 stomachs that he examined: 7 decticids, 8 acridids, 5 oedipines, 1 tryxaline, 6 Melanoplus sp., 8 M. mexicanus, 2 M. devastator, 1 M. bivittatus, 2 Dissosteira carolina, 1 Chortophaga viridis, 3 Neduba carinata, 8 Trimero- tropus sp., 7 Hippiscus sp., 2 Steiroxys sp., 3 Canoula pellucida, 2 Stenopnel- matus fuscus, 2 S. pictus, 4 Gryllus assimilis, 4 Ceuthophilus sp., 1 Pristoceutho- philus pacificus, 6 Gammarotettix bilobatus and 2 cicada nymphs. Grinnell, Dixon and Linsdale (1930:149) found that one of these racers had eaten a cricket. Fitch (1936:644) found another in the act of swallowing an adult vole (Microtus californicus), and recorded (1935:18) that two alligator lizards (Gerrhonotus multicarinatus) were found in the stomach of still another. 398 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. Woodbury (1931:75) recorded that a racer from Utah had a sagebrush scaly lizard (Sceloporus graciosus) in its stomach. Of the specimens examined in the University of California Museum of Vertebrate Zoology, no. 17256 from the Mad River, Trinity County, California, had eaten an alligator lizard (Gerrhonotus coeruleus), and no. 10120 trom Yolla Bolly Mountain in the same county had eaten a bird (unidentified) and a Jerusalem cricket (Steno- pelmatus sp.). Several authors have published specific information regarding the food of C. c. flaviventris. Hurter (1911:171) caught a blue racer in the act of swallow- ing a copperhead (Agkistrodon contortrix). Taylor (1892:331) recorded find- ing garter snakes in several large racers. Pope and Dickinson (1928:53) re- corded instances of blue racers feeding on racerunners (Cnemidophorus sex- lineatus). Ortenburger (1928:181) examined 22 stomachs and recorded: 1 large garter snake (Thamnophis sirtalis), 1 vole (Microtus pennsylvanicus), 1 frog (Rana sp.) 31 crickets (Gryllus assimilis), 4 decticines, 2 acridids, grass- hoppers (1 Hippiscus, 2 Melanoplus sp., 1 M. confusus, 1 M. differentialis, 1 Dissosteira carolina, 1 Sphargemon collare, 1 Trimerotropus sp., 1 Orphulella sp., 1 Chloealtis conspersa, 1 Chortophaga viridifasciata, 1 Omaseus sp., 1 Pedocetes sp.), and 2 caterpillars (1 noctuid, 1 sphingid). Gloyd (1928:123) recorded a hatchling glass lizard (Ophisaurus attenuatus) in the stomach of a juvenal racer. Force (1930:31) found a racer eating eggs from the nest of a cardinal (Richmondena cardinalis) and another racer eating eggs of a red- wing (Agelaius phoeniceus). Gloyd (1932:408) recorded an abies of a racer overpowering and swallowing a copperhead. Anderson (1942:210) recorded remains of crickets and grasshoppers in feces. Hudson (1942:55) recorded a racerunner (Cnemidophorus sexlineatus) in the stomach of a juvenile and recorded an earless lizard (Holbrookia maculata) 3 lizard eggs, and 14 grasshoppers (Melanoplus differentialis and others) in the stomach of another. Marr (1944:484) found a harvest mouse (Reithrodontomys monta- nus) in one. Breckenridge (1944:118) recorded stomach contents including a garter snake (Thamnophis sirtalis), a frog (Rana pipiens), 8 crickets and 2 moths. Mossimann and Rabb (1952:27) recorded that a racer disgorged several grasshoppers. Fouquette and Lindsay (1955:411) recorded that a blue racer had eaten a harvest mouse (Reithrodontomys sp.). Carpenter (1958:114) recorded that one blue racer had eaten a green snake (Opheodrys aestivus) and another had eaten a grasshopper and a camel cricket. Even though the sets of data cited above are not entirely com- parable, certain trends are evident. The black racers of the eastern states (especially C. c. constrictor of more northern regions) take a high proportion of vertebrates in their prey. Among these verte- brates snakes especially are well represented and the black snake would seem to be of some importance as an ophiphagous predator. The birds and mammals taken include some that are bulky (robin, cottontail, and even a weasel—the most formidable prey eaten). Presumably the rabbits that were eaten were young. In samples from the eastern United States insects made up small to insignificant parts of the food; they were lacking entirely or at least were not mentioned in the samples examined by McCauley and Wright and Bishop. In the blue racer of the central states, insects (mostly grasshoppers and crickets) are much more prominent in the food and vertebrates correspondingly less prominent. The vertebrates eaten are largely lizards, small snakes and mice. C. c. stejnegerianus is much like flaviventris in the trend of its feeding. C.c. mormon NATURAL HisTORY OF THE RACER 399 is less known than these subspecies in its feeding, but indications are that it takes a higher proportion of orthopteran insects and smaller proportions of mammals and snakes than do any of the other subspecies. In my own field study a total of 1357 food records were ac- cumulated, one of the largest samples known for any kind of snake. Most of these records were from the small area where my popula- tion study was carried on, and studies of other kinds of animals, in- cluding those that were the racer’s prey, were simultaneously in progress. Because large collections of reference materials were available, it was possible to identify to species many of the prey items found, even though they were incomplete and highly frag- mented because most of them were recovered from fecal material. The prey is, of course, swallowed entire, and the recently swallowed items squeezed from the stomachs provide the best ma- terial for the study of food habits. However, relatively few racers had detectable food items in their stomachs; digestion is rapid and often the snake was in a trap for a day or more before it was found. Therefore the greater number of records were obtained from scats. The residue in scats consisted entirely of hard and indigestible parts such as the chitin of insects’ exoskeletons and the hair, feath- ers, scales, teeth and occasional bone fragments of the vertebrate prey. The insects eaten could usually be counted individually by sorting parts, such as heads or hind legs. With mammals, birds and reptiles the hair, feathers, or scales did not permit counting of individuals—each occurrence was assumed to represent one in- dividual but in some instances two or more may have been present. Amphibians, lacking indigestible dermal structures were in most instances not represented at all in the scats, since their tissues were more or less completely dissolved by the digestion of the snakes. Soft-bodied larvae of insects and other invertebrates conceivably could be likewise completely digested, but such occurrences must be rare, as most of the invertebrates known to be eaten have the mouth parts, at least, heavily chitinized. Admittedly the factors discussed above would cause some bias in the percentage composition of the food determined from scats, but I believe that the amount of error introduced was slight, be- cause, judging from the records of items from stomachs, amphibians are not eaten frequently, and even mammals are not eaten fre- quently enough so that there is much chance of a snake taking two or more individuals at the same meal, unless it is robbing a nest containing a litter of young. 4—7864 400 UNIVERSITY OF KANSAS PuBsis., Mus. Nat. Hist. The largest sample, based on 1008 food items, was obtained from 479 scats collected from the Reservation and Rockefeller Tract over the period 1949 through 1961. Items recorded were: 183 gryllid crickets (144 Gryllus assimilis, 86 Gryllus sp., 3 unspecified); 353 locustid grasshoppers (41 unspecified, 73 Arphia simplex, 67 Mela- noplus femur-rubrum, 66 M. bivittatus, 39 M. differentialis, 17 Melanoplus sp., 15 Dissosteira carolina, 8 Chortophaga viridifasciata, 6 Syrbula admirabilis, 6 Sphargemon equale, 2 Melanoplus scud- deri, 2 Schistocerca obscura, 1 S. americana; 94 camel crickets (Ceuthophilus sp.), 93 katydids (36 Neoconocephalus robustus, 15 Orchelimum vulgare, 15 O. nigripes, 6 Conocephalus sp., 4 Orcheli- mum sp., 2 Amblycorypha inasteca, 1 Neoconocephalus sp., 1 Daihinia brevipes); 7 cicadas (5 Tibicen sp., 1 T. pruinosa, 1 T. CEUTHOPHILID AND TETTIGONIID 177% GRYLLID 178% LocUuSsTID 34.9% Fic. 18. Diagram showing percentage frequency of occurrence of various categories of prey in a sample of 1008 food items identified from scats and stomachs of blue racers from the Reservation and Rockefeller Tract. Insects, especially, orthopterans, made up the great majority of prey items taken. NaTuRAL History OF THE RACER 401 lyrica); 45 unidentified insects; 17 beetles (including 1 Phyllophaga, 1 Calosoma scrutator, and 2 other carabids); 2 noctuid moths (Mocis latipes) and 1 caterpillar; 2 homopterans, 1 bee, 1 ant, 1 spider; 69 voles (59 Microtus ochrogaster, 9 Microtus sp., 1 M. pinetorum); 31 white-footed mice (15 Peromyscus leucopus, 14 Peromyscus sp., 1 P. maniculatus); 36 miscellaneous small mammals (6 Cryptotis parva, 4 Sigmodon hispidus, 4 Reithrodontomys mega- lotis, 3 Blarina brevicauda, 2 each of Scalopus aquaticus, Sylvilagus floridanus, and 1 unspecified shrew); 50 snakes (16 Coluber con- strictor, 15 Diadophis punctatus, 14 Thamnophis sirtalis, 4 Elaphe obsoleta, 1 Natrix sipedon); 7 lizards (5 Eumeces fasciatus, 1 E. MAMMAL ORTHOPTERAN 14.3% MISCELLANEOUS INSECT. 1.9% Fic. 14. Diagram showing estimated percentage by weight of various cate- gories of prey in a sample of 1351 items, including all those represented in Fig. 18 and others from various parts of Kansas. Since the vertebrate items are on the average much bulkier than the insects eaten, vertebrates comprise most of the food, even though insects are eaten in much larger numbers. 402 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. obsoletus, 1 Cnemidophorus sexlineatus); 8 unspecified “reptiles”; 5 birds (none identified to genus); 3 bird eggs, 1 narrow-mouthed toad (Gastrophryne olivacea). Over the same period that the sample of scats was collected, a much smaller food sample of 73 prey items was collected by squeez- ing recently eaten food from the racers’ stomachs, or by finding the snakes actually swallowing their prey. These items from stomachs are listed separately because they include relatively more verte- brates than do the items from scats. A grasshopper or cricket eaten by a large racer might have passed undetected, while a relatively LIZARD 61.5% Fic. 15. Diagram showing estimated percentages by weight of various cate- gories of prey in a sample of 69 food items squeezed out of stomachs of the blue racers captured at Harvey County Park. Most of the items were verte- brates, and lizards (Cnemidophorus) were especially prominent in the food at this locality. Samples of prey from scats (included in Figs. 18 and 14) and from stomachs show somewhat different trends, and neither is entirely repre- sentative of the actual feeding. Also, local differences in food sources are important. NaturRAL History OF THE RACER 403 large item such as a vole or lizard would have produced a con- spicuous bulge in the snake that ate it, and would have excited the curiosity of the investigator. A second difference is that the items from stomachs included several frogs, whereas amphibians were absent from the much larger sample from scats. A third difference is that the many insects found in stomachs were all orthopterans with the exceptions of three noctuid moths and the larva of a moth. Miscellaneous insects, such as beetles, bees and ants recorded from scats were not found in stomachs. Amphibians eaten are digested so completely that no recognizable parts of them are to be found in scats, but remains of the insects previously eaten by amphibians are to be seen in racers’ scats. If not recognized as secondary items, such remains might lead to erroneous conclusions regarding the racer’s food. The items from stomachs were as follows: 21 grasshoppers (5 oedipines, 4 tryxalines, 5 Melanoplus bivittatus, 8 M. differentialis, 1 M. femur-rubrum and one each of Chortophaga viridifasciata, Dissosteira carolina, and Sphargemon equale); 8 crickets (Gryllus sp.), 8 atydigs: 8 camel crickets (Ceuthophilus sp.), 3 noctuid moths, 1 larva of a moth; 10 voles (Microtus ochrogaster), 6 white-footed mice (5 Peromyscus leucopus and 1 P. maniculatus), 4 harvest mice (Reithrodontomys megalotis); 1 shrew (Cryptotis parva); 4 snakes (3 Thamnophis sirtalis, 1 Storeria dekayi); 4 lizards (2 Eumeces obsoletus, 1 Cnemidophorus sexlineatus, 1 Ophisaurus attenuatus); 4 frogs (Rana pipiens), 1 tree-frog (Hyla versicolor). Records from the Harvey County, Kansas study area include a series of 69 food items from 55 stomachs (of living snakes) and 210 food items from 118 scats. There is a relatively high proportion of vertebrates, including some frogs, in the stomachs, and with no frogs but more miscellaneous insect material in the scats. But, for the sake of brevity, the two categories of items are combined in the following list: 55 grasshoppers (12 unspecified, 1 “locustid,” 81 “oedipines,” 7 “tryxalines,” 5 “locustines,” 2 Melanoplus bivittatus and one each of M. femur-rubrum, M. scudderi, M. differentialis, and Arphia simplex); 48 crickets (31 Gryllus assimilis, 17 unspecified); 14 katydids (11 Daihinia brevipes, one each of rhadiphorine, conocephaline and Neoconocephalus sp.); 9 noctuid moths and 1 moth larva; 26 miscellaneous insects (including 13. “beetles,” 1 elaterid, 1 curculionid, 1 lygaeid bug, 1 ant, 1 wasp); 1 spider, 7 mice (5 Peromyscus maniculatus, 2 unspecified), 4 unidentified mammals, 1 vole (Microtus ochrogaster), 1 shrew (Cryptotis parva), 84 lizards (77 Cnemidophorus sexlineatus, 6 Sceloporus undulatus, 1 unspecified), 6 snakes (4 “natricines,” 1 Thamnophis sp., 1 Pituophis melanoleucus), 1 “reptile,” 1 “bird,” 9 frogs (4 unspecified, 1 Rana catesbeiana, 4 Rana pipiens, 1 Rana sp., 1 Pseudacris triseriata). Kinds of Prey Throughout the range of the racer small mammals make up an important portion of the food, and the bulk of those eaten are voles (Microtus sp.) and white-footed mice (Peromyscus sp.). The voles being diurnal, and having habitat preferences similar to those of the racer, are especially subject to attack, but only large adult racers are capable of swallowing a full grown vole. Probably most of the voles eaten are immature. Of the white-footed mice, P. 404 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. maniculatus especially prefers a grassland habitat, and is usually found in situations frequented by the racer. Being mainly nocturnal and crepuscular, it is usually in hiding at times when the racer is prowling, but may be flushed from its nest in a shallow burrow or beneath a sheltering object, and overtaken by the snake. Other mammals that are important in the food are harvest mice and other mice, shrews, and young cottontails. The latter are small enough to be eaten by racers only in the early stages of their life in the nest before weaning. Rats (Rattus, Sigmodon), moles, sciurids, and weasels are less frequent prey, ordinarily too large to be eaten by racers and taken chiefly as defenseless juveniles. Predation on birds is relatively uncommon, and in most instances it involves the eggs or nestlings, or fledglings still slow and clumsy and incapable of sustained flight, or, occasionally, injured adults. Nests that are vulnerable are chiefly those of ground nesting species, or of kinds that nest near the ground in grass or thickets. Many of the birds recorded have not been identified to species, but those identified have included a variety of small passerines and also domestic chicks. Lizards figure prominently in most of the food samples, but only a few species, those that live on or near the ground in grassy places, have been recorded. Most of the records pertain to scaly lizards (Sceloporus undulatus and S. graciosus), earless lizards (mainly or entirely Holbrookia maculata), racerunners (Cnemidophorus sex- lineatus) and skinks (Lygosoma laterale and Eumeces sp.). Snakes are important in the racer’s food in most parts of the range, but the large racers of the Northeast are those most inclined to ophiphagous habits. The common garter snake (Thamnophis sirtalis) is the species most commonly eaten. Probably this is a matter of availability rather than preference, since the garter snake is one of the commonest and most widely distributed of North American snakes, occurring throughout most of the racer’s range. The green snakes (Opheodrys) also are represented frequently. The other snakes eaten are mostly medium-sized to small colubrids, of a variety of kinds. However, there are three records (from Connecticut, Missouri and Kansas) of the racer preying on the venomous copperhead. There are many records of the racer prey- ing on smaller individuals of its own species. In my own records racer remains appeared 18 times, equalling in frequency those of the common garter snake and exceeding all other kinds. In four of these instances the scale remains were relatively few and the NATURAL History OF THE RACER 405 scales were relatively large, suggesting as an alternative to actual predation that a racer may have eaten part of its own sloughed skin, or that patches of shed skin may have adhered to the scat after its deposition in the trap. However, in the remaining 14 instances the remains of racer found in scats clearly indicated cannibalism, since the scales found were small and numerous and often were associated with bone. Cannibalism seems to occur frequently enough to be a significant factor in the reduction of the first year young. Liner (1949:230) described two instances of cannibalism in a litter of blue racers hatched in captivity. In one instance two young had seized the same lizard, and one having swallowed the lizard, con- tinued to engulf the other snake, although it was of a size approxi- mately equal to that of the first snake. Nevertheless, swallowing was completed, with the snake eaten pressed in a series of curves. A second instance of cannibalism occurred when one young racer attempting to catch a lizard struck another racer by mistake, then retained its hold and commenced swallowing. A similar instance was observed in a brood that I kept in 1962 after hatching had oc- curred in the laboratory. Hatchling turtles of two kinds (Chrysemys picta, Terrapene carolina) have been reported in the racer’s food. Probably other kinds are eaten also. However, the awkward shape and almost in- flexible shell of the prey on the one hand, and the slender form of the racer, with limited distensibility of the gullet on the other, would limit this type of predation to occasional instances involving an unusually large racer and a small turtle. There seem to be no records of the racer preying on salamanders. Many kinds of frogs are eaten, chiefly ranids and hylids, and the leopard frog (Rana pipiens) is the most frequent victim. Wright and Bishop (1915:160) stated that the toad (Bufo terrestris) oc- cupied first place in the racer’s food in the region of Okefinokee Swamp, Georgia, but they mentioned no specific instances of this species being eaten. Klimstra (loc. cit.) found only four toads in his large sample of digestive tracts from Illinois. Because of their virulent dermal secretions, bufonid toads are avoided by many kinds of snakes and predation on them by the racer probably is unusual. Most authors who have written concerning the food habits of the racer have mentioned insects as part of the diet. Statements in the literature have often seemed to imply that the racer feeds on in- sects in general, according to their availability. However, the large number of records now available demonstrate that the racer 406 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. is highly selective in choice of its insect food, that soft-bodied orthopterans, chiefly crickets, grasshoppers and katydids, are the usual insect prey, with occasional predation on moths and their larvae. Eating of other insects such as cicadas and June beetles, is a rarity, but on occasion a racer may be tempted to sample such prey when it finds the newly emerged imago before its exoskeleton has hardened. I am convinced that such rarely occurring items as carabid beetles, hemipterans, homopterans, diplopods and spiders are secondary prey items, eaten by frogs that later were eaten by the snakes, in most instances if not in all. It is noteworthy that several of the same genera of grasshoppers and crickets are promi- nent in the food samples collected in widely separated parts of the racer’s range. As might have been anticipated, different species of prey were not utilized by the racers to the same extent throughout the snakes’ season of activity. Grasshoppers, for instance, fluctuated from a low of 25. 3 per cent (frequency) in the May sample to a high of 41.4 per cent in the September sample. Availability of prey, rather than any change of preference on the part of the racer, explains this trend. Thus, the locust, Arphia simplex, which, unlike most local grasshoppers, overwinters in the adult stage, is most prominent in the food in May, represented by 15.7 per cent, but it decreases progressively to a low of 1.8 per cent in September. The common grasshoppers of the genus Melanoplus show just the opposite trend, increasing during the summer, from a low of 2.62 per cent in May (when all are nymphs and most are too small to constitute a meal worthy of a racer’s attention) to a high of 31.5 per cent in September. Mammals are best represented in the food in May, when they collectively comprise nearly 80 per cent of the items taken, and they are progressively less well represented as the summer advances. Both Microtus and Peromyscus conform to this trend, but the relative numbers of Peromyscus rise again abruptly in October. The general trend may be explained by the fact that in May most small mammal populations have a high proportion of young of the year, and these young are especially vulnerable to predation by the snakes. Also, insects in general are less available in spring, and this may force the racers to utilize vertebrates to a greater extent than at other seasons. Actually, the seasonal changes in food sources are not especially striking, and it seems that each important prey species is utilized more or less throughout the sea- son of the racer’s activity. NATURAL HisToRY OF THE RACER 407 TABLE 5. DisTRIBUTION BY MONTHS OF VARIOUS CATEGORIES OF PREY ITEMS RECORDED FROM BLUE RACERS FROM KANsAs, CHIEFLY FROM THE RESERVA- TION AND ROCKEFELLER TRACT @nicketa(233) Reese Ceuthophilus (94)...... Katy did’ (96))sa-. 6 a6 -c Melanoplus (188)...... All grasshopper (388)... WERDER) Beano deb ee Peromyscus (39) eesciree < All mammal (162)..... TaizardediO) eee ccc Snake: (Gil) si44s ee ee Api (2) acres e ee ise: 17 37 .28 .O1 05 12 or 01 19 .30 14 13 23 Bae 03 06 .08 .25 15 43 wieate 03 12 27 15 19 24 rats 15 17 21 16 12 19 Sees 45 32 13 .02 07 01 ater 41 18 08 .08 05 20 sueusrs 41 25 14 .03 05 12 tehene 07 46 .29 15 01 02 yc Yake 37 18 .08 13 09 15 Bae 50 25 12 04 03 06 TABLE 6, DisTRIBUTION OF VARIOUS COMMON PREY ANIMALS IN A SAMPLE OF 625 AMONG RACERS OF DIFFERENT SIZE GROUPS Snout-VENT LENGTH (MM.) OF RACERS IN SAMPLE gryllid cricket...... WM CLONGDUUS Soci. oles Ceuthophilus....... Orchelimum........ tettigonideewsscs. “other grasshopper” and miscellaneous orthopteran........ Arphia simpler..... miscellaneous insect PEROMYSCUS... 10k “other mammal’’.. . Kinds of prey and their percentage frequencies in samples Total prey items for size group 200 | 300 | 400 | 500 | 600 | 700 | 800 | 900 | 1000 to to to to to to to to to 299 | 399 | 499 | 599 | 699 | 799 | 899 | 999 | 1099 ———$—<— _|J |__| | SEE | ES -03+)-.02-1- 063031" 30271205 .03 |} .02 | .05| .08 | .08 | .04 | .09 .04 | .04} .09 | .07 | .04 |] .08 | .05 | .04 .20 07 | 06% 205.) 04. (2050708) | 12 04° |,:.02.0 08). 213, 09" a2) jho2o 09 02 | .05 | .06) .03 | .05 | .04 408 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. The wide disparity in size between young and adult racers also results in utilization of different food sources to some extent. In some kinds of snakes adults and young draw their food from en- tirely different sources, but in the racer there is broad overlap, as shown in Table 6. The samples from the largest and smallest size groups of racers are relatively small. Two important kinds of prey —voles and grasshoppers of the genus Melanoplus—were not found at all in the smallest size groups of snakes and comprised increasing percentages in the food of the larger size groups. A large adult vole is too large to be swallowed except by an unusually large racer, and a young vole old enough to leave its nest is far too large for a hatchling racer. Grasshoppers of the genus Melanoplus are rela- tively large and heavily armored, and so are relatively immune to attacks from the smaller snakes. Small soft-bodied orthopterans in- cluding Gryllus, Ceuthophilus and Orchelimum, and also lizards and snakes, are best represented in the food of the smaller racers. Other types of prey showed no definite correlation with size of the racer taking them. Reproduction Sexual Behavior Many observers have published accounts of the courtship and/or mating of the racer, but all of these are, to some degree, incomplete. Because of the widely different circumstances, and the different viewpoints of the observers involved, the several accounts give much different impressions of sexual behavior in this species. Either singly or combined, the published accounts do not provide an adequate description of the process. My own observations, made both under natural conditions and in large outdoor enclosures, are likewise somewhat incomplete, but indicate that the whole sequence of courtship and mating is divisible into the following well-defined stages: 1) the finding of a recep- tive female by the male; 2) the persistent following of the female by the male, who courts her by lying extended along her body and performing writhing movements, with periodic interruptions during which he momentarily leaves the female and courses rapidly through the grass around her; 3) the acceptance of the male by the female, signalled by the raising of her tail and the almost in- stantaneous intromission; 4) the dragging of the passive male by the female while he is firmly attached to her during the period of coitus; 5) separation of the pair and involution of the male’s hemipenis. NATURAL History OF THE RACER 409 Even in the breeding season, racers that were confined in en- closures usually were either indifferent to each other or responded with reactions of fear or hostility. In moving they tended to fol- low the edges, and often two moving in opposite directions would approach each other; when this occurred, one snake might strike at the other with a short jab that seemed to be mostly bluff, and then would dart away. The males, being smaller, were usually the more wary. Sexual behavior was noticed on only a few occasions. Several large adult males were less wary than others and usually manifested curiosity or interest toward other racers. My most complete ob- servations of sexual behavior were made on May 18, 1962, when a newly caught adult male was added to an enclosure of 100-foot circumference already containing several racers, two of which were large adult females. Within half an hour the male was found court- ing one of the females. She was lying in a loose coil, with the male extended along her. At my approach the female darted way in alarm for approximately three feet, and the male moved with her, so swiftly and adroitly that he maintained contact and was in ap- proximately his original position with respect to the female when she stopped. Spasmodic rippling movements passed down the body of the male as he lay in contact with the female. These movements lasted several seconds, increasing in intensity, alternating with longer periods of little or no movement. As each period of vigorous writhing reached its climax, the male’s head jerked forward and backward several times in seeming excitement. The female’s be- havior was mostly passive. She seemed to be receptive, but from time to time, without any noticeable warning, she darted away for several feet as she had when the pair was first discovered. Each time the male darted forward with her, maintaining contact while she moved. These swift movements of the female seemed to be spontaneous, at least in most instances there was no evident cause for alarm. The female’s movements seemed to stimulate the male’s interest rather than to discourage him. In most instances the female moved only four to five feet, then stopped abruptly or turned back. She would stop in a loose resting coil, in thick grass, with the male lying over her. Often she coiled in such a way that the posterior end of her body was beneath her forebody, but this did not seem to deter the male from moving the posterior end of his body into position beside hers. After a sudden change in the 410 UNIverSITY OF Kansas Pusts., Mus. Nat. Hist. female’s position, the rear of the male’s body would perform groping movements along that of the female until his cloacal region was approximately opposite hers. The male sometimes had his chin pressed against the female’s back, especially when he was moving forward along her, but more often his head was raised, and freqently was as much as 18 inches from the female's head. At intervals averaging approximately ten minutes, during a little more than an hour of observation, the male would suddenly dart away from the female, and with unusually rapid and animated movements, he would move around her in an irregular and devious course, sometimes as far as five feet away, but usually within 18 inches. Usually on each such expedition several or many circuits were made; then the male would return to the female and would glide rapidly along her until he attained the mating position. A period of especially vigorous courting movements would follow. At 12:55 p.m. it was necessary for me to discontinue observa- tions, and I left the female confined in a cloth bag. Returning at 1:20 p.m. I found that the male was not displaying interest in the female confined in the bag, nor in the other female loose in the en- closure. The first female was released from the bag, and was out of sight for approximately four minutes. When relocated she was again attended by the male, who was carrying on courtship even more vigorously than he had before. At 1:35 p.m. the male achieved intromission. Although the pair was under observation at the time intromission occurred, the actual eversion of the hemi- penis was not seen because the snakes were partly concealed by dense vegetation. There was a sudden flurry of movement, the male’s head waving and his body thrashing. In an instant these violent movements subsided, and after a few seconds the female be- gan to crawl forward slowly. The male had relaxed, and re- linquished his contact with the female anteriorly. As she moved away he was dragged after her tail-first. He made slight back- ward wriggling movements that perhaps aided in maintaining sexual contact. The female’s restlessness increased, and in eight minutes she dragged the male in a circuitous course a distance estimated to be between 20 and 30 feet. At 1:40 p.m. the pair was ten feet from the point where copulation had begun. The female showed increasing inclination to climb, raising her head and forebody against the trunks of saplings, and finally reaching up one to a branch 20 inches above the ground, and climbing first along the branch and then farther up the main trunk. As she progressed the NATURAL HisToRY OF THE RACER 411 male was lifted from the ground, dangling limply suspended by his hemipenis and its base had become exposed. At 1:43 p.m. separa- tion occurred and the male dropped into the grass. Semen dripped from the cloacae of both snakes. That from the female was tinged with blood. The individuals involved in this observation were kept in the enclosure subsequently but no further sexual behavior was noted. Contrary to the popular belief that these racers have permanent mates, all available evidence indicates that they are promiscuous, and two or more males may simultaneously court the same female in the brief spring breeding season. On May 24, 1960, while I was walking in a hilltop field of brome grass, a sudden movement attracted my attention to three racers lying alongside each other. Only the posterior parts of their bodies and their tails were visible. Two were males and were performing the characteristic slow writh- ing movements against the body of the female from either side. Although the heads were not in view, the snakes may have been able to see me through the screening vegetation; after I had watched for approximately 20 seconds, all three suddenly took alarm, for no apparent cause, and scattered. Further evidence of promiscuity is provided by the account of Ellicott (1880:207) who wrote regarding the eastern subspecies: “I noticed a ball of black snakes (Bascanion constrictor L) rolling slowly down a steep and stony hillside . . . about two miles above Union Factory, Baltimore County, Md. . . . kept to- gether by procreative impulses.” It was stated that this observa- tion was made in early spring. “Snake balls” have often been ob- served, and described in the literature; usually the snakes involved were garter snakes (Thamnophis) or water snakes (Natrix). Seem- ingly, typical aggregations consist of a single adult female and several or many males attempting to mate with her. There is a distinct possibility that the snakes involved in Ellicott’s observations were misidentified. Sexual behavior of the racer is in most respects remarkably similar to that of the common garter snake, Thamnophis sirtalis, well known through the work of Blanchard and Blanchard (1942). In studying sexual behavior of racers, several observers have failed to differentiate between the different stages of the mating process, and have assumed that copulation was occurring when actually only the precopulatory behavior was observed. In an early descrip- tion of courtship in this racer in Kansas, Brons (1882:365) stated 412 UNIVERSITY OF KANSAS PuBts., Mus. Nat. Hist. that the female “at times, seems to toy with the male, indisposed to yield to his importunities, though pressed with ardor. To avoid his suit, at times, she will dart through grass, among stones, or enter a crevice. Should he be able to reach his mate while within a hole, he is not slow in bringing her to the surface, again to be repulsed. Upon an unbroken ground the sexual union is less pro- longed. Here she is unable to free herself from his quick and effectively directed moves. In case she attempts to quit him, a coil is thrown about her body, and his head laid flat upon her neck, and replaced as promptly as dislodged, evidently in the endeavor to propitiate her.” Another account probably based on courtship rather than copula- tion is that of Wright and Wright (1957:135), who described the behavior of a pair of C. c. priapus on Billy Island, Okefinokee Swamp in southern Georgia, on May 8, 1921, as follows: “They were stretched out, more or less coiled . . . the rear parts of the bodies from the vent were entwined. The female, or smaller one seemed to have its tail around that of the male. There were contortions or quiverings from time to time. . . . May 8, 1921: Jackson Lee saw black snakes entwined, the male seizing the female by the top of the neck.” Blanchard and Blanchard (op. cit.) have described the dragging of the male by the female during coitus in the garter snake, and the temporarily inseparable bond formed between members of a pair by the recurved spines of the engorged hemipenis, but it has not been generally recognized that the process is much the same in other colubrines. Cottam (1937:229) described and photographed mat- ing in a pair of C. c. mormon in Utah. The copulating racers were shown in a loose coil lying alongside each other with tails inter- twined. However, when disturbed by the observers, these racers made frantic efforts to escape, crawling in a spiral course, while remaining attached and intertwined, “with no evident attempt to separate” during approximately a quarter hour of observation. The racer is notorious for its aggressive behavior and occasional alleged attacks on humans in the breeding season. The tendency has doubtless been much exaggerated, especially in the verbal second- or third-hand accounts based on the alleged observations of eye-witnesses. Nevertheless, the supposition that large adults will sometimes pursue or attack humans when disturbed is well substantiated. In most of the instances known to me, it is the large eastern subspecies, C. c. constrictor, involved in these incidents, and NATURAL History OF THE RACER 413 seemingly the smaller racers of the Middle West, far West and South are less inclined to behave aggressively. In May 1958 two pairs of large racers were confined in a semicircular wire enclosure thirty feet across and open on top, and with natural vegetation, at the Reservation headquarters. Often in approaching the cage I saw two or more racers in close association, but because of shelter- ing vegetation, and the snakes’ timidity observation was difficult. On May 19 a pair were lying partly extended in loose coils, but immediately the female took alarm and darted away, breaking loose from the male; his hemipenis was exposed, and underwent involu- tion and retraction in approximately 30 seconds. Unlike the female, the male on this occasion did not attempt to escape, but turned to face me with a show of aggressiveness. Probably copulation was in its final stages when the disturbance occurred. Circling of the female racer by the male from time to time in the course of courtship has not been recognized by previous observers as a part of the mating pattern, but Pope (1944:171) described somewhat analogous behavior, probably modified by unnatural con- ditions of captivity and the crowding of many racers in one cage. Pope, citing earlier observations by Noble, wrote: “When sexually excited, the male blacksnakes dash wildly about before paying court to individual females. In captivity these dashes excite all specimens confined together. A male, after picking out a mate, moves his chin lightly along her back, while undulations run forward along his sides and he extends his tongue now and then. Later he throws the part of his body near his vent over the corresponding part of the female, the two tails sometimes becoming loosely intertwined.” Recorded dates of mating for the species are all in spring, but indicate a span of many weeks for the breeding season, and this spread results in part from geographical differences. Published records are as follows: Subspecies constrictor May 12, 1930, in Ohio (Conant, 1938:55) Subspecies priapus May 8, 1921, in Georgia (Wright and Wright, 1957:135) May 9, 1921, in Georgia (Wright and Wright, 1957:135) Subspecies flaviventris May 8, 1931 (two pairs) in Missouri (Boyer and Heinze, 1934:195) April 18, 1986, in Missouri (Anderson, 1942:210) May 12, 1928, in Kansas (Gloyd, 1928:128) Subspecies mormon June 10, 1927, in Utah (Cottam, 1937:229) July 7, 1938, in California (Cunningham, 1959:17) 414 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. In the course of my live-trapping, I occasionally found more than one racer in a trap. As might be expected from the low yield per trap, such double or multiple captures were relatively rare. Chance, and unusually strategic placement of certain traps were doubtless contributing factors. May and October, being the most productive months for trapping, yielded a high proportion of these combined captures. Some involved an adult and an immature snake, or two adults of the same sex. Eliminating all these, there remain 44 heterosexual captures of adults. These latter captures are signifi- cantly concentrated in their seasonal distribution and indicate a spring breeding season; 34 were in May, six were in June and four were in October. Eight of the May records and one June record each involved a trio of snakes—two males and a female in every instance. Distribution of the spring records, grouped in five-day intervals, was as follows: May 11 to May 15: 14 June 5 to June 9: 0 May 16 to May 20: 5 June 10 to June 14: 2 May 21 to May 25: 8 June 15 to June 19: 2 May 26 to May 80: 13 June 20 to June 24: 0 May 31 to June 4: 1 June 25 to June 30: 1 Approximately 87 per cent of the records fell in the twenty-day interval, May 11 to 30, which is regarded as the main breeding season. Presumably males continue to be at the peak of breeding condition and continue to search for females after the latter have become unreceptive, partly explaining the scattering of records through most of June. Several of the females found in traps with males in May had abundant active sperm in their cloacae and oviducts and probably had been inseminated within a few hours of the time they were checked. Others lacked sperm, but the cramped quarters inside the traps may have effectively prevented the consummation of court- ship, especially when two males were confined with the same fe- male. None of the females trapped with males in October was found to be inseminated, and it seems doubtful whether copulation ever occurs at that time of year, although males have motile sperm and seem to be in breeding condition then. Cycle of the Male Cloacal smears indicate that males mature sexually and first pro- duce sperm in August and September when they are a little more than a year old. Insofar as could be determined, there was no sexual activity at this time of year, and actual breeding of the NATURAL History OF THE RACER 415 adolescent racers was postponed until the following May. By this time at an average age of 20 months, the snakes had made further growth. Mr. Dwight R. Platt studied the changes in the male reproductive organs during the annual cycle at Harvey County. In racers recently emerged from hibernation he found the seminiferous tubules filled with Sertoli syncytium, but containing few germ cells. Spermatogonia proliferate in May and June. During the first half of July primary spermatocytes are the dominant cells in the semi- niferous tubules. By early August spermatids are dominant and the first free spermatozoa are present. In late October spermio- genesis is essentially complete and the tubules are relatively empty before the snakes hibernate. During the season of activity the seminiferous tubules increase to approximately double their mini- mum diameter, reaching the maximum in August. Cyclic changes NUMBER OF RECORDS APR. MAY JUN. JUL. AUG. SEP OCT. NOV. Fic. 16. Catches, in semi-monthly periods, of racers in their summer habitats, at Harvey County Park (upper) and at Reservation and Rockefeller Tract (lower). Intensive activity in spring (the breed- ing season), tapering off rapidly as the season advances, is well shown by the larger sample, but in the years of trapping at Harvey County Park operations usually were not fully underway until the latter part of May. 5—7864 416 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. in size and secretory activity of the ductus deferens, ductus epididymis, and sexual segment of the renal tubules occur, with maximum size and secretory activity coinciding with the time of movement and storage of the spermatozoa. The latter are stored in both ductus deferens and ductus epididymis. Despite the short breeding season, a male racer has active sperm at all seasons. Eggs In accounts of the racer in the humid southeastern United States, Brimley (1903:261), Wright and Bishop (1915:160) and Tinkle (1959:195) mentioned the ease with which the eggs might be found and the superficial situations in which they were sometimes deposited. Both Wright and Bishop, and Tinkle made field studies in swamps, where presumably the subsoil was saturated with moisture and too wet for the eggs. Tinkle mentioned finding one clutch beneath a discarded newspaper and another beneath a small, thin board. Surface (1906:167) stated that in Pennsylvania the eggs were to be found in loose soil, in sawdust piles, or in de- caying wood of hollow logs or trees. Clark (1949:249) stated that in northern Louisiana the eggs are laid in soft, moist soil such as may be found beside decaying logs. Minton (1944:457) found two clutches under flat stones on hillsides in Indiana. In the more arid climates of the far western states the species’ habits are much dif- ferent in this regard. Through many years of familiarity with C. c. mormon, I have never seen its eggs. Presumably nests in this part of the range are deep underground, most often in old burrows of the pocket gopher (Thomomys), which are so abundant that in many areas the soil is riddled with them. Burrows of the ground squirrels (Spermophilus sp.) and other small digging mammals also provide potential insulated nest sites with the favorably moderate tempera- tures and high humidities that the eggs of snakes require. On the morning of July 10, 1962, I was directed to the sites of two clutches recently plowed up, 1% miles north of the Reservation (Pl. 21, Fig. 2). The eggs were in a fallow field having a stand of sunflowers three to five feet high. The plow blades turned the soil at a depth of approximately seven inches. In each instance only a few eggs were visible. They were well scattered in the loose soil turned up by the plow; 21 were found in one clutch and 10 in the other. All the eggs were intact except two that had minute punc- tures from which liquid oozed. Seemingly the eggs in situ had been well above the level of the blade—at depths of four to five NATURAL Hisrory OF THE RACER 417 inches. No nest cavities were discernible where the eggs were found, but elsewhere in the field tunnels of moles (Scalopus aquaticus) and prairie voles (Microtus ochrogaster) were exposed by the plow. Presumably the eggs had been in such tunnels, which had disappeared as the loose soil crumbled. Another clutch was discovered in an adjoining field on July 16. The nine eggs were at depths ranging from 6% to nine inches, and only the two top- most eggs had been turned up by the plow. All three clutches were within a few feet of the edges of the fields. On the Reservation and nearby areas I have seen remains of an estimated 20 clutches that have been destroyed by predators. The remains in every instance consisted of an excavation, and the strewn torn and empty eggshells. Nests were at depths of four to eight inches in old tunnels, which most often seemed to be those of moles but also included some of the prairie vole, and perhaps some of the pine vole (Microtus pinetorum). All these nests were in open sunny places in prairie or pasture habitat. Many observers have described the eggs of the racer, which are white, elliptical, somewhat elongate, with tough, leathery, some- what flexible, shells, and a granular surface. Like other snake eggs, those of the racer gradually absorb moisture during incubation. They become more turgid and increase in weight and dimensions, especially in breadth, and by the time of hatching are nearly twice their size at laying. Between different clutches and even within the same clutch there is notable variation in the size of the newly laid eggs. Munro (1948:199) noted that in a small adult racer TABLE 7. NUMBERS AND SIZES OF Eccs IN CLUTCHES OF THE BLUE RACER FROM EASTERN KANSAS NUMBER) — Lengths Widths Weights | Snout-vent OF ( length of Ecas mm.) (mm.) (grams) female (mm.) ile 26.5 (29-24) 17.0(19.0-16.5) 5.5(6.0-4. 4) 892 8 33.3(39-31) 16.3(17.5-14.0) | 6.0(6.7-5.6) 899 12 29.1(32-25.5) 17.1(18.5-16.0) 4.9(5.2-4.4) US) 14 26.9(30-24) 19. 2(20-18) 5.2(6.2-4.4) 772 10 31.7(33-29) 16.5(18-15) 6.0(6.5-5.6) 807 ul 29 .7(33-27) 16.7(18-15) 5.4(5.8-5.0) 858 PA 28 .9(32-27) 18.4(19.5-18) 5.9(6.3-5.6) 1038 13 30.7 (34-28) 19.3(20-18) 6.8(7.5-6.1) 907 18 29 .0(30-26) 17.9(19-16) 4.9(5.5-4.3 911 12 30.3(34-28) 17.8(19-15) 5.2(6.1-3.8) 843 14 30. 9(36-29) 19.4(21-18) 6.8(7.6-6.2) 846 418 UNIVERSITY OF KANSAS PuBts., Mus. Nar. Hist. kept by him, the eggs laid were larger but less numerous than those produced by a large adult. Munro noted also that shape of eggs in the two clutches differed; the smaller snake produced more elongate eggs of smaller diameter. The idea that eggs laid by the smaller females are more slender and elongate is not supported by my own data. For 11 clutches of eggs examined soon after laying, dimen- sions, weights, and the lengths of the females are shown in Table 7. In a clutch of eggs beginning to hatch on September 3, 1958, dimensions and weights were as follows: length 31.8 (36-30), diameter 22.0 (24-21), weight 9.7 (10.3-9.3). Gravid females that were kept in captivity in anticipation of their laying usually produced their clutches within a few days. The lay- ing dates of such individuals are shown in the following list. Those with asterisks were from the Harvey County study area, others were from the Reservation and Rockefeller Tract. June 19, 1961 July 1, 1961 July 12, 1961 June 21, 1959 July 4-5, 1962* July 15, 1961* June 23, 1961* July 6, 1955 July 18, 1961* June 26, 1959 July 7, 1959* July 20, 1961 June 29-30, 1962° July 7, 1959* August 8, 1960* A further indication of the period when laying occurs was pro- vided by the appearance of females gravid and progressively more swollen with eggs, then their abrupt disappearance and replace- ment by thin and wrinkled individuals that obviously were recently parturient. The following records show the course of these events on the Reservation in the years when summer trapping was done with sufficient consistency. These dates provide a rough approxi- mation of the time when laying occurs locally. They indicate a laying season concentrated in a period of approximately three weeks in this locality. Records from published literature also indicate that laying occurs in late June and early July at the latitude of Kansas, but somewhat earlier in the southern United States. ‘TABLE 8. DATES WHEN PARTURIENT AND GRAVID RACERS WERE CAPTURED ON RESERVATION AND ROCKEFELLER TRACT IN SEVERAL YEARS, INDICATING TIME OF OVIPOSITION First parturient female........ June 18 | June 17 | June 22 | June 20 | June 13 Last gravid female........... July 8 | July 11] July 5 | July 12 | July 16 NATURAL History OF THE RACER 419 Many authors have made statements regarding the size of the clutch in the racer, on the basis of those found in the field, those laid after capture, or those dissected from gravid females. Some of the statements were based upon small but unspecified samples, and are far from the mark. From records accumulated in the course of my own field work, and a summarization of those in published accounts a substantial sample is available showing the usual size of clutch in the area of my study, and the trends of geographic variation in some parts of the range. C. constrictor (all combined ) 151 clutches averaged 10.61 (2 to 81) eggs. C. c. constrictor ....... 14 clutches averaged 16.80 (7 to 31) eggs. CANCE PTADUR oo Sek 11 clutches averaged 12.60 (7 to 21) eggs. CC MOTMon) nig oe 32 43 clutches averaged 5.79 (2 to 18) eggs. C. c. flaviventris ....... 82 clutches averaged 11.78 (5 to 26) eggs. C. c. stejnegerianus, one clutch contained 10 eggs. In the foregoing list the sample of C. c. flaviventris may be divided as follows: Reservation and vicinity: 86 clutches averaged 11.65 (6 to 21) eggs. Harvey County study area: 21 clutches averaged 12.0 (5 to 18) eggs. Museum specimens from Kansas: five clutches averaged 9.2 (6 to 14) eggs. Published records (Kansas, Indiana, Iowa, Louisiana, Missouri, Oklahoma, Texas): 20 clutches averaged 12.5 (5 to 22) eggs. TABLE 9. PUBLISHED RECORDS INDICATING DATES OF LAYING IN DIFFERENT POPULATIONS OF COLUBER CONSTRICTOR ne ahs fb Subspecies Area Authority June 9, 1952 | flaviventris SE Texas Guidry (1953:50) July 1-2, 1920 | flaviventris Texas Ortenburger (1928:183) June 1, 1920 | flaviventris Texas Ortenburger (1928:183) June 9, 1926 | flaviventris Oklahoma Force (1930:31) June 24, 1926 | flaviventris Oklahoma | Force (1930:31) July 19, 1928 | flaviventris Oklahoma Force (1930:31) July 16 flaviventris Oklahoma Carpenter (1958:114) July 4-5, 1948 | flaviventris Kansas Munro (1948:199) July 4, 1948 flaviventris Kansas Munro (1948:199) June 26, 1930 | flaviventris Ohio Conant (1938:55) July 3, 1961 mormon NW Oregon | Van de Velde, Martan and Risley (1962:212) June 7, 1957 priapus S Illinois Rossman (1960:219) June 19, 1912 | priapus S Georgia Wright and Bishop (1915:160) July 10, 1940 | priapus S Illinois Cagle (1942:187) July 6 constrictor New York | Ditmars (1907:284) June 27- July 5, 1951 constrictor Virginia Werler and McCallion (1951:251) June 5, 1947 stejnegerianus | S Texas Auffenberg (1949:54) 420 University oF Kansas Pusts., Mus. Nat. Hist. Published records of clutches laid by racers, from which figures used in the foregoing account were obtained, include the following: C. c. constrictor: 22 (Barbour, 1950:104); 21, 18 (Brimley, 1903:261); 25 (Conant, 1938:55); 8 (Ditmars, 1907:284); 7, 12, 14, 16, 19, 20, 31 (McCauley, 1945:76); 14 (Wright and Wright, 1957:136). C. c. flaviventris: 7 (Anderson, 1942:210); 22 (Brumwell, 1951:205); 11 (Carpenter, 1958:114); 8, 9, 9 (Force, 1930:31); 10 (Guidry, 19538: 50); 18 (Liner, 1949:280); 5, 8, 17 (Marr, 1944:484); 5, 14 (Munro, 1948:199); 18, 19 (Ortenburger, 1928:188); 6, 15 (Tinkle, 1959:195); 15, 19 (Wright and Wright, 1957:141). C. c. priapus: 20, 21 (Cagle, 1942:187); 7 (Conant, 1938:55); 19 (Ross- man, 1960:219); 5, 9, 11, 14 (Wright and Bishop, 1915:160); 16 (Wright and Wright, 1957:147). C.c. mormon: 18, 9, 8, 5, 5,5, 4, 4 (Cunningham, 1959:17); 5, 6 (Stebbins, 1954:374); 6 (Van de Velde, Martan and Risley, 1962:212); 3, 6 (Wright and Wright, 1957:144). C. c. stejnegerianus: 10 ( Auffenberg, 1949:54). In general, the number of eggs in the clutch is proportiona! to the size of the female producing them. The larger and bulkier females produce more eggs. Geographic trends in number of eggs produced are perhaps controlled by differences in size between different popu- lations; thus, the large eastern constrictor produces nearly three times as many eggs per clutch as does the small western mormon, whereas the centrally located flaviventris is somewhat intermediate in size and in numbers of eggs produced. In most reptiles growth in length and bulk continues after attain- ment of sexual maturity. For many kinds including Eumeces fas- ciatus, Crotaphytus collaris, Cnemidophorus sexlineatus, Agkistro- don contortrix (Fitch, 1954:60; 1956:236; 1958:36; 1960:174), and Sceloporus olivaceus (Blair, 1960:94), it has been shown that the larger and older females in a population produce more offspring than do the smaller and younger individuals. This situation ap- plies in the racer, as shown by the clutches of 52 females cor- related with their sizes and presumed ages (Fig. 17, Table 10). The two-year-olds contribute a relatively small quota to the annual brood, partly because their clutches are small, but more especially because many of them fail to attain sexual maturity in time to breed. Many of the female racers that are more than two years old also fail to produce an annual clutch of eggs. The 24-day period May 28 to June 20 inclusive is judged to comprise the period when eggs have generally enlarged sufficiently to be detected in gravid females, but still have not been laid in most instances. In this period, in 1960, 1961 and 1962, ratios of gravid females to those NATURAL HisTORY OF THE RACER 421 not detectably gravid in several supposed age groups arbitrarily established on the basis of size, were as follows: Two-year-olds .......... 2 gravid, 13 apparently not gravid Three-year-olds ........ 5 gravid, 4 apparently not gravid Four-year-olds ......... 8 gravid, 2 apparently not gravid Five-year-olds .......... 4 gravid, 3 apparently not gravid Six-year-olds (or older) .. 8 gravid, 2 apparently not gravid From the appearance of these snakes it is reasonably certain that none had already laid eggs when it was recorded, but there is some NUMBERS OF EGGS eT Te aetna ey AGE IN YEARS Fic. 17. Graph showing number of eggs per clutch, and correlation with supposed age (as deduced from length of body) in female blue racers from the Reservation, Reckefelr Tract, and Harvey County Park, 422, UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. possibility that a few individuals not noticeably gravid at the times they were examined, produced eggs subsequently. However, these meager data do seem to indicate that most of the two-year-old fe- males and a minority of older individuals fail to produce clutches in the annual breeding season. Tasie 10. FECUNDITY OF FEMALE RACERS IN VARIOUS AGE-SIZE CLASSES, ALL FROM THE RESERVATION AND ROCKEFELLER TRACT Most PROBABLE AGE Number Snout-vent length Number IN YEARS OF FEMALES of females in | of females; average of eggs in (AS INDICATED BY SIZE) sample and extremes clutch Atos etnae 10 688 (589-748) 9.2 (6-12) Dihiwak ater es 19 789 (756-840) 9.9 (5-14) A cert ects 7 856 (850-861) 10.8 (8-12) i ee ee 6 907 (892-933) 13.0 (8-17) 6 or more.... 10 1005 (955-1088) 15.7 (11-19) Under unfavorable conditions eggs can be resorbed, but prob- ably this can occur only if initiated before ovulation. A racer in which six small eggs were palped on June 28, 1960, was kept until July 23 but did not oviposit. It no longer appeared gravid and the ova could not be detected by palpation. Another female had 13 eggs on June 21, 1960, but by July 23 when the snake was re- leased the eggs had not been laid and no longer could be detected. Both snakes refused to feed throughout their confinement. Like other reptilian eggs, those of the racer are dependent upon the warmth of their surroundings for incubation. They are toler- ant of a wide range of environmental temperatures, but the higher the temperature the more rapidly incubation proceeds. Under natural conditions there may be much difference in hatching time in two clutches laid at the same time and in the same locality. Site of the nest—deep and well insulated, or shallow; in a well shaded situa- tion or one exposed to maximum amounts of sunshine—would largely control rates of development. Clark (1949:249) writing of the subspecies anthicus in north-central Louisiana, stated: “eggs are laid about the first of June. . . . young begin to make their appearance at about . . . July 1.” Even for the southern states these dates of laying and hatching seem somewhat too early to reconcile with the records published by other observers, and are in need of verification, especially since they seem to be based upon vaguely remembered observations rather than upon written records. At the other extreme Surface (1906:167) wrote of constrictor in NATURAL History OF THE RACER 423 Pennsylvania that hatching may occur as late as October, and that there is evidence some young may even remain in the egg over winter before hatching occurs. Several incubation periods are on record for clutches laid and hatched in captivity, as follows: C. c. mormon, Oregon, 47 and 51 days (laid July 3, 1961, hatched August 19 and 28; Van de Velde, Martan and Risley, 1962:212). C. c. stejnegerianus, Texas, 78 days (laid June 5, 1947, hatched August 17, Auffenberg, 1949:54). C. c. priapus, S. Illinois, 58 and 59 days (laid July 10, 1940, hatched Sep- tember 6 and 7, Cagle, 1942:187). C. c. flaviventris, Kansas, 50 days (laid July 4 and 5, hatched August 28 and 24; Munro, 1950:124). G: a rene te Texas, 48 days (laid June 9, 1952, hatched July 22; Guidry, No incubation periods for eggs in natural nests have been recorded. In the course of my study, eggs obtained from 12 captive females were hatched in confinement, with an average incubation period of 51 days (43 to 68) as follows: Laid July 6, 1955, hatched August 20. Laid July 8, 1958, hatched September 3 and 4. Laid June 21, 1959, hatched August 17, Laid June 26, 1959, hatched August 17. Laid July 7, 1959, hatched August 28 to 25.* Laid June 28, 1960, hatched August 20.* Laid June 80, 1961, hatched August 30. Laid July 15, 1961, hatched September 1 and 2.* Laid July 18, 1961, hatched September 2 and 8.* Laid July 4 and 5, 1962, hatched August 15 and 16.® Laid June 29 and 80, 1962, hatched August 14 and 15.* Laid July 6, 1962, hatched August 16 to 20.* In the foregoing list those entries marked with asterisks were ob- tained from the Harvey County study area; all others were from the Reservation and Rockefeller Tract. Hatching Detailed observations on hatching were made on a clutch of eggs laid on June 29 and 30, 1962, by a female caught in Harvey County. The first egg in the clutch had already been laid in the trap when the female was found at 11:30 a. m., June 29. Two of the eggs were abnormal, with thin transparent shells, and were found to lack embryos when they were opened on July 7. Later, two other eggs were attacked by mold and the embryos died early in development. The clutch was kept in a can of slightly damp soil. At 2:30 p.m. on August 13, when the clutch was examined, egg no. 6 was found to have hatched. The young snake had made a 21-millimeter slit in the shell. At 12:50 a.m. on August 14, it was discovered that eggs 1, 4, 5 and 7 each had been slit. No. 4 had two parallel slits separated by a two-millimeter strip of shell, and the young racer 424 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. could be seen inside. At 1:05 a.m. this young snake had changed position and was lying upside down in the egg, his snout protruding slightly through one of the slits. At 1:45 a.m. he was again right side up, still in the shell. At this time each of the slit eggs showed the protruding snout of a young snake. Occasionally the viscous liquid egg white would be blown into a large bubble on the surface of the shell as the young snake exhaled. A third slit, parallel to the others, had appeared in egg no. 4. A hatchling emerged from egg no. 1 between 2:20 and 3:20 a.m., and another from no. 7 between 7:00 and 9:00 a.m. The hatchling struck vigorously many times, and vibrated his tail when he was disturbed. Egg no. 3 was first slit between 7:00 and 9:00 a.m., and three more slits appeared in it between 9:15 and 10:15 a.m. At 12:45 p.m. a hatchling was found in the act of emerging from egg no. 4, and approximately the anterior one-fourth of its body protruded. Disturbed by the move- ments of the observer, the little snake drew back into its shell. This hatchling began to emerge again at 12:50 and his hatching was com- pleted at 1:00 p.m. Between 5:40 and 6:20 p.m. a hatchling emerged from egg no. 3 (for several hours this hatchling had been lying on its back inside the egg, with only its snout protruding); two slits appeared in egg no. 2 and three slits appeared in egg no. 8. At 6:50 p.m. the hatchling in the latter thrust his snout through the slit in this eggshell. This hatchling was lying on its back at first but by 10:50 it had shifted to a normal position. It emerged from the shell between 2:35 and 2:50 a.m. Egg no. 2 was the last to hatch. At 7:05 p.m. the hatchling inside made two additional small slits in the shell, and at 7:30 p. m. thrust its snout through one of them, while lying on its back. At 1:45 a.m. it was right side up, but at 3:00 a. m. had reverted to its previous position. At 4:40 a.m. it was again right side up, and it emerged from the shell at 5:55 p.m. On August 17, at 11:00 a. m., hatchlings no. 5 and no. 6 had lost their egg teeth. All others still had their egg teeth then, but by 10:00 p.m. that of no. 8 was missing, and that of no. 4 was loose and dropped out while the snake was being handled. On August 20 at 9:00 a. m., hatchling no. 7 had lost its egg tooth; nos. 2 and 3 retained theirs only in part, and no. 1 had its egg tooth intact. By noon on August 22 no trace of an egg tooth remained on any of the hatchlings. In the same group of hatchlings sign of impending molt was first noticed on the morning of August 17, when no. 6 was noted to have NATURAL History OF THE RACER 495 its eyes clouded and milky in appearance. By evening no. 1 had attained the same stage and no. 7 was beginning to show it. On the morning of August 20, shedding had begun in no. 6, while no. 2 and no. 8 had milky eyes. The eyes had cleared in no. 1 and no. 7, and were still clear in the remaining hatchlings. On August 22 shedding had been completed by no. 1 and no. 8, and all others were in the process of shedding. Another clutch of 14 eggs from a recently captured female was found freshly laid in a cage on July 6, 1962. Hatching of 13 oc- curred August 16 to 20, as shown in Table 11. Growth Hatching usually occurs in late August or early September, and the disparity in size between hatchlings and adults is greater than in some other kinds of snakes. In 76 young from ten clutches of eggs incubated in the laboratory, averages and extremes for meas- urements and weights were as follows: snout-vent length, 214.5 (186 to 244) millimeters; tail, 59.3 (44 to 73) millimeters; weight, 4.16 (2.4 to 5.8) grams. In each brood the size tended to be fairly uniform, except that there were usually one or more stunted individuals markedly smaller than the others. However, there were striking differences in size between the young of different broods. None of the young captured was as small as the average hatchling from the clutches incubated in captivity, but in the 14 years of my study only four young were captured in August. The hatchlings TABLE 1]. Times oF HATCHING IN A CLUTCH OF RACER Eccs FRoM Harvey County Park Time at which shell Time that hatchling NUMBER was slit by hatchling emerged OF Eac Day Hour Day Hour 1 Aug. 17 10:50 a.m. Aug. 18 3:00 to 6:00 p.m. 2 Aug. 17 12:15 to 12:50 p.m.| Aug. 18 3:25 to 4:25 p.m. 3 Aug. 17 1:40 p.m. Aug. 17 10:45 p.m. 4 Aug. 18 3:00 to 6:00 p.m. Aug. 19 before 7:00 a.m. 5 Aug. 16 10:45 a.m. Aug. 17 | 6:30 to 7:55 a.m. 6 Aug. 17 | 9:00 a.m. Aug. 18 | 6:30 to 7:30 a.m. a Aug: V7, \\ 4:15 p.m. Aug. 20 1:00 to 7:00 a.m. 8 Aug. 17 | 3:40 p.m. Aug. 18 | 5:00 to 6:00 p.m. 9 Aug. 17 | 7:30 p.m. Aug. 18 | 6:30 to 7:30 a.m. 10 Aug. 16 10:48 a.m. Aug. 17 11:05 a.m. 2 ie no record Aug. 18 | 6:00 to 6:30 a.m. 13 Aug. 17 8:00 to 8:20 a.m. Aug. 17 5:30 to 5:50 p.m. 426 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. are relatively secretive and elusive, and the lush vegetation of late summer provides them with abundant hiding places. Neverthe- less it is remarkable that the hatchlings are so seldom seem, when their probable abundance is taken into account. Probably all of those captured had already made some growth after hatching. By early November or the last week of October, racers have almost or quite completed their season of activity, and are at the hilltop ledges, preparing to hibernate, if they have not already retired into dormancy. For 25 young of the year captured in this period at the end of the growing season, measurements and weights were as follows: snout-to-vent, 827 (273 to 418) milli- meters; tail 93 (72 to 114) millimeters; weight, 12.3 (7 to 19) grams. In the ten-week period between hatching and hibernation these young had already passed through their period of most rapid growth, having added, on the average, more than 50 per cent to their original lengths, and almost tripled in weight. In these young about to enter their first hibernation, variation in size and weight is much greater than in the hatchlings; some have fared much better than others, and there are significant age differences. Within any one year the time of hatching is spread over several weeks because of differences in the time of laying, and differences in nest sites, with variation in heat received, which promotes or delays the rate of incubation. Year to year differences in the trend of weather in- crease the dispersion as the incubation time is shortened in hot, dry summers and lengthened in those that are relatively cool and moist. TABLE 12. GrowTu OF FimsT-YEAR RACERS Original record Record(s) of recapture ; Snout- Snout- EX vent . vent : Date length, Weight, Date length, Weight, mili- | Ses milli- | 87@™s meters meters fof Aug. 17, 1959 223 5.3 Sept. 12, 1960 503 42 ce) Oct. 26, 1959 278 10 Oct. 26, 1959 566 55 e) Nov. 2, 1959 340 18 May 11, 1961 582 65 ie) Nov. 10, 1959 348 13.7 June 20, 1961 733 127 ie) Oct. 19, 1952 361 15.3 Oct. 9, 1953 620 68 fof May 9, 1953 368 17 May 13, 1954 609 84 9 April 16, 1950 328 142 Oct. 10, 1950 603 78.2 °) May 17, 1956 358 15.2 Oct. 6, 1956 575 52 fof Nov. 1, 1953 330 11 May 21, 1955 674 92 NATURAL History OF THE RACER 427 Though covering a wide size-range, the young of the year enter- ing hibernation are still a distinct size group, not yet overlapping that of the next older group of young. Growth during the first year of life is best shown by the individuals in Table 12, all of which were marked either before their first hibernation or soon after emerging from it, and were recaptured either the following autumn, or in spring soon after emerging from a second hibernation. These records indicate that the young racers at the time of their second hibernation have grown to a snout-vent length of well over 500 millimeters, but less than 700 millimeters, and a weight of more than 40 grams but less than 100 grams. Other racers marked in the first few weeks of life were recaptured after two or more seasons of growth, and indicate the sizes that may be expected in young adults from two to five years old, as shown in Table 183. TasLe 18. GrRowTH OF RACERS MARKED AT AN AGE OF LESS THAN ONE YEAR AND RECAPTURED IN THEIR SECOND, THIRD OR FouRTH YEARS Original record Record(s) of recapture . n « m E/E Z| 2 LD» a fD) 2s| 2! 23 eq) “| 23 Sex Date Gl oul) cae leg. Date Dict au monel Ponoile oue lic o a) a0) one $80/ Gia é $6) 2 I'38 Sa] | 2 28 0 | 2 Oo} o |} o.4 SEE eG) ||) a |Eela ae lems co | Nov. 12, 1952 342 |....| 2144 | May 20, 1955 702 | 110 | 33 o' | Nov. 1, 1953 SoU n |) Lhe Aug. 16, 1956 780 | 162 | 354% & | Nov. 12, 1953 293 |....| 214 | May 14, 1956 634 | 96 | 3214 Oct. 18, 1956 689 | 110 | 37% o | April 15, 1955 320 | 13 | 944 | June 8, 1958 740 | 118 | 45 & | Oct. 2, 1955 348 | 18 | 1 | May 22, 1958 728 | 120 | 33 May 21, 1960 795 | 130 | 57 o | June 25, 1959 378 | 10 | 10 June 30, 1960 590 60 | 22 June 7, 1961 4O5S: |) 1238133 &# | Nov. 1, 1953 330 | 11 | 2 | Aug. 25, 1957 805 | 183 | 48 Unlike young of the year, racers in their second autumn were trapped in large numbers. By this time all were large enough to be caught in the traps of quarter-inch wire mesh, and they were the most abundant size group. Many that were marked at this stage were recaptured after intervals of months or years, showing the trend of growth. Some of these snakes in their second autumn already had overtaken the more retarded third-year individuals. 428 University OF Kansas Pusts., Mus. Nar. Hist. The two age classes cannot be separated with certainty. Selected records of individuals that were almost certainly second-year young at the time they were marked are presented in Table 14. TABLE 14. GrowrH oF YOUNG RAcERS THAT WERE MARKED NEAR THE TIME OF THEIR SECOND HIBERNATION Original record Record(s) of recapture E| 3 Al & u o o Ya oo wa »¢ bb Efe Sex Date Die |ees od Date eee rs oa a) 2 faye) PS 2 Zone oo] a 3S oO} a-| 3c oo o\ 3°98 oO 26) SS ne °.5 ox ‘S 0.8 Nn = A, Nn S AY o | Oct. 6, 1960 595 : July 23, 1961 o' | Oct. 13, 1950 | 525 ‘ Oct. 2, 1951 o' | Nov. 2, 1950 | 545 L Oct. 24, 1951 o | Sept..2,. 1957 |522 7 Oct. 14, 1958 o' | Oct. 17, 1953 | 558 May 10, 1955 o' | Nov. 14, 1956 | 587 4 | May 20, 1958 June 17, 1959 o | Sept. 7, 1959 | 533 June 6, 1961 o | Oct. 17, 1959 | 558 July 21, 1960 June 7, 1961 o | Oct. 16, 1952 | 577 E May 31, 1955 May 11, 1956 o' } Oct: 11, 1950: | 570 ; July 11, 1956 o' | Oct. 14, 1953 | 560 | 5: June 29, 1958 oa | Oct. 6, 1950 563 : Sept. 28, 1958 o' | Oct. 14, 1953 | 523 May 19, 1959 May 17, 1960 o | Oct. 13, 1953 | 521 : May 13, 1958 June 3, 1959 o' | Nov. 5, 1953 | 512 May 22, 1958 9 | Aug. 11, 1953 | 534 : Sept. 18, 1954 9 | Oct. 14, 1949 | 588 : : Oct. 11, 1950 9 | Oct. 6, 1950 570 ; Oct. 11, 1951 9° | ;Oct.31, 58953 | 582 Oct. 5, 1954 9 | Oct. 21, 1959 | 588 4 | May 7, 1961 9 | May 14, 1960 | 506 Oct. 26, 1960 | 690 o' | Oct. 22, 1960 | 527 Oct. 7, 1961 620 o | May 3, 1960 | 530 May 16, 1961 | 700 o | May 27, 1961 | 535 Z Oct. 25, 1961 | 672 From the records in Table 14 and many more like them, average and extreme sizes for progressively older age groups were esti- mated. Even racers that were already of adult size when they were marked were tentatively identified with one or another age group, and their records of subsequent growth were used. Most of the NATURAL History OF THE RACER 429 records show that the females grow more rapidly than the males, and are, on the average, larger at any given age. Relatively few individual racers were recaptured after periods of several years. Each of the eight listed in Table 15 is among those that were captured in four or more different years, and their records are significant in revealing the trend of growth after sexual maturity has been attained. These snakes, one to three years old at the time they were marked, show well the persistent but decreasing growth, and the fluctuating weight that is characteristic of this and other species. Some of the largest racers recorded had already reached un- usually large size when they were first captured, so there was no opportunity to determine their ages. Several others, originally captured as small- or medium-sized adults, subsequently grew to a size approaching the maximum, and thus provided a basis for IN MILLIMETERS SNOUT-VENT LENGTH AGE IN YEARS Fic. 18. Graph from records of blue racers marked early in life and recaptured, showing average and extreme snout-vent lengths for males and females of various ages. Growth is especially rapid in the first year, but continues, with gradual slowing, throughout life. In hatchlings, the sexes are of approximately equal size, but females grow to be markedly larger than males. 430 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. TABLE 15. CHANGES IN LENGTHS AND WEIGHTS IN E1icuHT Racers RECAPTURED REPEATEDLY OVER PERIODS OF YEARS Snout-vent | Tail length Weight Probable DaTE oF CAPTURE length in in in age in millimeters | millimeters grams months No. 1, male October 21, 1953... 676 216 85 25% October 12, 1954... 733 238 128 37% July 31, 1955. ....%: 755 243 143 46 May 20, 1956...... 785 250 153 55% July 05; VOT 36 oo 788 252 162 694% June 30, 1958...... 820 264 200 81 May 15, 1959...... 828 263 195 91144 May 14, 1960...... 840 265 195 103% No. 2, male May 9, 1955...... 678 212 105 32 May 26, 1958...... 827 248 195 69 May 3, 1959. . 2... 865 262 218 80 May 14, 1960...... 877 264 218 9214 July 16, 1960...... 890 257 190 9414 No. 8, male May 26, 1955... .... 565 171 53 21 August 28, 1957.... 798 245 155 48 May 13, 1958...... 814 249 137 5614 May 4, 1959...... 848 252 180 68 June 12, 1961...... 896 262 198 934% No. 4, male May 19, 1956...... 632 208 78 20% August 2, 1957..... 740 238 123 35 May 20, 1959...... 796 246 132 56% May 23, 1960...... 813 280 140 6814 No. 5, male June 30, 1956...... 594 188 73 22 October 12, 1956... 658 200 80 27% July 14, 1957...... 690 220 102 34% June 23, 1958...... 749 226 136 46 July 24, 1959...... 751 239 133 59 May 27, 1960...... 765 235 130 69 No. 6, male October 19, 1956... 518 167 43 13% July 19, 1957...... 673 218 96 221% May 5, 1960....... 780 250 175 56 May 9, 1961....... 840 272 272 68 No. 7, female May 12, 1957...... 790 238 175 32144 September 3, 1957.. 810 243 208 36 October 1, 1958.... 883 259 206 49 July 11, 1959...... 915 247* 190 5814 September 12, 1960 930 249* 238 72% September 14, 1961 1012 268* 332 8414 No. 8, female June 23, 1955...... 601 175 67 22 August 3, 1958.... 920 255 300** 60 July 23, 1959...2.4.. 955 281 211 Cal May 18, 1960...... 962 262 260 81 * Incomplete. ©* With food. NATURAL HisToRY OF THE RACER 431 FEMALES IN GRAMS {MALES WEIGHT [Ole we igputl gi wig, Of EHO 7) ge eg) MO AGE IN YEARS Fic. 19. Graph from records of blue racers marked early in life and recaptured, showing average and extreme weights in samples of different ages. Early in life the females outstrip males in growth and the size differential increases throughout life. estimating the ages of large individuals. The records of nine such racers are listed in Table 16. A few adult racers recaptured, including different individuals of small, medium, and large size, failed to make any measurable gain in length over periods of months, or even over several years. Most often the large individuals were those that failed to grow or made only slight gains. Some of the snakes that failed to increase in length suffered dramatic weight losses, perhaps as a result of in- jury, disease, or parasitism. However, other individuals that failed to gain appreciably in length did gain in weight. Doubtless both 6—7864 432 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. TasLE 16. Growru IN SEVERAL RAcERS THAT WERE ALREADY ADULT WHEN ORIGINALLY CAPTURED AND THAT SUBSEQUENTLY ATTAINED UNUSUALLY LARGE SIZE Original record Record(s) of recapture . @m . nm a| 2 A] 8 ler Pe © es| >| #3 2a)| oes Sex Date Degel) te © a Date Se a= oe PS a ae rs ~~ ron 4 | 38 eo | 38 38 op 2 34 0) ro Cf! & 0.8 ox oD °.8 a eS | a wa a 3 | May 14,1956 | 815 | 191 | 5614 | May 15,1959 | 905 | 243 | 9214 May 16, 1961 | 945 | 287 | 10414 May 22, 1958 | 825 | 251 | 68!4 | June 25, 1959 840 | 205 | 82 May 27, 1960 880 | 210 | 93 Oct. 14, 1961 900 | 223 | 10914 o' | May 19, 1959 705 | 105 | 321% | May 28, 1960 S28 45 July 3, 1961 900 | 265 58% 9 | Nov. 6, 1954 823 | 190 | 38 June 30, 1958 1030 | 350 | 82 9 | June 6, 1955 810 | 190 | 33 May 21, 1959 1087 | 345 80% 9 | Aug. 29, 1957 885 | 265 | 48 Aug. 6, 1960 1041 | 235 83 Q | May 1, 1958 | 633] 84|20 | May 30,1961 | 1088 | 320] 57 9 | July 20, 1958 862 | 203 | 461% | June 30, 1960 1020 | 238 70 g | Oct. 18, 1956 845 | 185 | 3714 | May 20, 1958 905 | 240 56% Oct. 22, 1959 1085 | 375 74 genetic and environmental factors were involved. A few racers gave the impression of being stunted by adversity. All records ob- tained of growth (or failure to grow), throughout the period of my study were used in compiling Table 17. Mortality Factors and Adaptations for Survival Defense, and Escape The behavior patterns that are associated with defense and escape in snakes are widespread. The common racer is typical of the more generalized snakes. Almost all elements of the racer’s behavior are found in snakes of other genera and perhaps, of other families. The racer differs from these only in minor details of its behavior, in the circumstances under which various behavioral traits are elicited, and their relative importance. The common racer is one of the swiftest of North American snakes, and it usually depends upon speed to avoid enemies. Racers crossing roads may detect approaching automobiles at a distance, and thereby may be stimulated to accelerate their movements and so escape onto the NATURAL HisTORY OF THE RACER 433 shoulders rather than becoming traffic casualties as many indi- viduals of most other kinds of snakes do under the same circum- stances. The chances of such escape are of course much better on dirt roads that are relatively narrow and have a rough surface per- mitting efficient traction, than on wide smooth, paved highways. The snakes are somewhat reluctant to venture onto open expanses of pavement. A racer suddenly startled by close approach of a human usually flees at high speed. The first part of its course, up to 20 feet or more, is marked by a violent lateral thrashing of the body and tail, which may help the snake to gain speed, but which seem chiefly TABLE 17. Srzes OF RACERS OF DIFFERENT AGE GROUPS IN MAY AND OCTOBER Males AGE Snout-vent length in mm. Weight in grams IN eaaee Ae ] Ob d Ob ypica serve served Average range range Average range 1+ 539 432-609 2— 615 500-676 560-674 68.2 51-92 2+ 668 610-700 620-710 107.4 63-134 3— 706 677-730 648-755 102.1 65-129 3+ 740 701-754 667-780 147.0 93-216 4— 757 731-772 725-809 139 95-251 4+ 785 755-793 720-850 167.4 128-225 5— 805.5 773-810 743-855 152.4 110-198 5+ 810 801-831 773-858 163.9 89-211 6— 827 811-831 765-883 175.9 130-230 7— 845 832-850 788-900 181.2 125-210 g— 868 851-873 740-890 217.5 194-225 8+ 870 861-884 Females i 581 415-658 2— 644 520-739 580-738 83.5 52-127 2+ 743 685-804 670-826 135.2 73-200 3— 810 740-829 730-880 149.4 98-219 3+ 836 804-869 736-915 181.2 120-268 4— 866 830-880 791-920 212.3 175-243 4+ 883 870-914 810-952 191.2 143-300 5— 914 880-929 833-1088 209.6 136-275 5+ 932 915-954 883-990 250.4 195-336 6— 965 930-961 892-1020 245.9 218-283 6+ 970 955-980 885-1003 271 243-336 i— 974 962-990 919-1050 251.3 150-330 far 1000 980-1015 930-1085 295.6 235-375 434 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. designed to draw attention of a potential attacker to the spot that the snake is leaving. In any event, the mode of progression abruptly changes to a swifter and much smoother travel, and the snake may seem to vanish completely or as it glides rapidly away, its course may be marked by the slowly waving tops of tall grass. The trend of travel away from the site of disturbance is in an irregularly zig-zag course. The method of crawling, with lateral undulations of the body, supported against scattered pivot-points along the way, tends to prevent the snake from moving more than a short distance in a straight line. In checking traps along hilltop rock ledges in October, I have often flushed racers that had come to the ledges searching for hibernation dens. In such an instance the startled racer would usually dart away down the steep slope at unusually high speed, and within a few seconds might have covered 100 feet or more, progressing with a minimum of lateral undulations, and seeming to flow or coast over the ground surface with scarcely any effort. Although coursing downhill in this manner provided an effective method of escape, racers startled at the ledges or on other sloping ground sometimes followed a course parallel to the ledge, or even turned uphill, with disastrous slowing. Often I have accomplished capture by hand, by the strategy of running downhill to get below the escaping snake, causing it to turn back up the slope, there to be slowed to the extent that it could be overtaken. A racer that has been flushed, and has disappeared after creat- ing a commotion by its violent thrashing, may, within a period of minutes, return surreptitiously to the vicinity, gliding back slowly and silently from the direction opposite that in which it departed. It may climb into a tree or bush where, lying stretched on outer twigs and concealed in dense foliage, it will sometimes permit close ap- proach, rather than reveal its presence by moving. If too much dis- turbed however, such a snake will move away, higher, or to the opposite side of the clump, with a slow and stealthy gliding motion which is likely to escape detection. The sudden stops and abrupt changes in direction make it difficult to follow the course of a climb- ing racer, even though it is moving slowly. The tendency to climb to escape danger seems to be especially strong in the western sub- species, C. c. mormon. In my years of experience with the racers in the chaparral belt of western Oregon and California, I found that one of the most characteristic responses to danger was to climb into a bush or low tree and seek concealment in the thick foliage. NATURAL History OF THE RACER 435. In most instances the snake was from one to eight feet above the ground; usually it did not attempt to climb high in a tree even when there was opportunity to do so. The racer’s prowess as a climber is limited. In thick brush where its weight is supported by numerous stiff twigs simultaneously, it is at home and can travel rapidly, but it cannot cling effectively to rough bark of a vertical tree trunk as can the rat snakes (Elaphe sp.) and others that are more specialized in their arboreal adaptations. Chaparral-type shrubs such as Ceanothus cuneatus and Arctostaphylos viscida were those in which western yellow-bellied racers were seen climbing most frequently. Crab-apple (Pyrus ioensis) and haw (Crataegus sp.) were the shrubs most frequently utilized by blue racers on the Reservation. Carr (1950:80) noted that Florida racers were much less given to tree-climbing than were those of more northern regions. C. c. stejnegerianus of southern Texas is reputed to have arboreal tendencies highly developed (Mulaik and Mulaik, 1942:14). The racers kept in an outdoor enclosure in the summer of 1962 spent much of their time above the ground, climbing in several walnut saplings. Usually in the warmer part of the day three or four of the five racers in the pen were climbing. In the trees a racer usually lay extended or in a loose coil among foliage in the outer twigs, at a height of five to 12 feet. Such a snake might rest for hours in almost the same position. Racers do not ordinarily enter the water voluntarily, but they are swift and efficient swim- mers when the need arises. On one occasion a large female, seen near the edge of the pond on the Reservation, and approached from the landward side, took to the water without hesitation and swam strongly to the opposite shore more than 100 feet away. Other observers have recorded similar incidents. A racer that is suddenly startled at close range may make no attempt to escape, but instead may coil and perform slow writhing movements, with the head pressed to the ground and concealed beneath part of the body. Musk, exuding from the anus is spread over the body surface as the movements continue. Only a small percentage of the racers found free in their natural habitats reacted in the manner described. In each such instance, the circumstances were such that the racer was prevented from making its usual rapid getaway, either because temperature was unusually low for activity, or because the snake was away from suitable cover. After removal from a live-trap, with handling, and especially clipping of 436 UNIVERSITY OF Kansas PuBts., Mus. Nat. Hist. scales, racers were much more likely to behave in this passively defensive manner, which seemingly constitutes a second line of defense in snakes which have been prevented from escaping— either cornered, captured, or injured by a predator. The musk is creamlike in color and consistency, and is secreted from sausage- shaped glands in the base of the tail. Its odor is rather disagree- able, but less so than that of musks of various other genera. Also, it is less penetrating and lasting than many other musks. In han- dling the racers removed from traps I attempted to avoid being smeared with the musk by grasping the snake by the tail and neck and keeping its body stretched out. While the snake was being examined, and measured a droplet of musk would form at the anal orifice and would begin to flow down the snake’s body. To avoid contact with the musk I would wipe away the droplet with a paper towel or leaf, but the musk droplet might be replaced several times within the few minutes that the snake was restrained. A racer that is confronted with an object arousing its suspicion or indicating possible danger, may move away slowly with a char- acteristic “threat display.”. The forebody and neck are held rigid, well off the ground, and slightly arched, with the neck flattened in a vertical plane—causing this part of the snake to appear from lateral view larger than it actually is. The tongue is protruded frequently and waved slowly. A racer that has been disturbed and is trying to gain shelter in a direction that brings it closer to the danger usually will adopt this tense attitude, but a single threaten- ing movement will cause it to abandon its circumspect pose, and panic in an attempt to reach the nearest shelter in the shortest pos- sible time. A common response to an alarm is to vibrate the tail. The tail being long and slender is vibrated much less rapidly than the short, muscular organ of a rattlesnake, or even the medium-short tail of a bull snake. The rapid twitching produces a characteristic sound in dry vegetation. A racer that vibrates its tail is fully active and aroused; the sound is heard as the snake pauses before it makes a dash for shelter. At times the sound probably functions as a decoy to distract the attention of potential predators, affording the racer an opportunity to escape. A racer that is cornered or captured usually puts up a spirited struggle, striking vigorously and repeatedly at its tormentor. If only cornered, it will make slashing strokes at the enemy, jerking back to a coil from each stroke in a manner that causes the teeth NATURAL History OF THE RACER 437 to lacerate the enemy’s skin in long scratches if the stroke finds its target. The rapid recoil often causes teeth to be jerked from their sockets and left embedded in the wound inflicted by the bite. On many occasions in sustaining a bite from a racer I have received such teeth which have remained undetected for a day or more until soreness and festering led to their discovery and removal. A racer that is grasped may deliver several bites within a few seconds, chewing vigorously to imbed its teeth to the maximum at each bite. If grasped by the tail and held clear of the ground, the racer would swing its body with a rapid whirling and twisting motion, which in a few seconds would twist off the end of the tail, unless countermeasures were promptly taken. Many racers, especially the larger and older ones, have parts of the tail missing, as a result of such escapes, and perhaps also from freezing in hibernation or from certain injuries and infections. If grasped by the body, the racer struggles with a violent lateral thrashing to break the grip of its captor, at the same time striking to bite and discharging musk, urinary wastes and feces. Many of the racers examined bore scars from wounds that were probably inflicted by predators. A few had survived severe skeletal injuries, involving deformation of the spinal column or extensive tearing of muscle and connective tissue layers of the body wall, altering the normal body shape. A more frequent type of injury involved chiefly the integument, which had been ripped open by the teeth, claws, or bills of adversaries, despite the fact that the skin is remarkably tough and leathery. Patches of scar tissue with scales in irregular sizes, shapes and patterns characterized such in- juries. The most frequently observed type of injury involved loss of part of the tail. Usually only a small terminal part was missing, but occasional stub-tailed individuals had lost as much as three- fourths of the tail. No consistent trend of difference between the sexes in incidence of injuries to the tail was noticeable, but there was definite correla- tion with age. In the entire sample from the Reservation and Rockefeller Tract the percentages in each supposed age group (actually size group) lacking part of the tail were as follows: hatchlings, 2.9 per cent; one-year-olds, 9.5 per cent; two-year-olds, 15.8 per cent; three-year-olds, 14.3 per cent; four-year-olds, 15.7 per cent; five-year-olds, 23.2 per cent; six-year-olds, 30.9 per cent; seven- year-olds, 28.9 per cent; those eight years old or older, 21.1 per cent. The seeming reversal of trend in the older racers is difficult 438 UNIvERSITY OF KANSAS Pusts., Mus. Nat. Hist. to explain, but probably results from inadequate numbers in this part of the sample. Although the racer’s most characteristic response to any disturb- ance is to flee at high speed, certain individuals are inclined to be- have aggressively under exceptional circumstances. Records of such aggressive behavior nearly all pertain to large individuals of the northeastern black racer (C. c. constrictor) in the breeding season. Cope (1900:794) wrote: “it is courageous and will sometimes at- tack, moving forward with the head raised from 1 to 2 feet above the ground.” Ditmars (1944:13) wrote: “Occasional specimens in the breeding season . . . willactually attack . . . glide to- ward an intruder . . ._ striking madly at one’s feet or legs.” Woods (1944:257) quoted a 18-year-old amateur herpetologist, Leon Gonthier, regarding the latter’s encounter with an aggressive black racer on May 7, 1944, as follows: “When I ran toward the snake about 20 feet away, it turned and came for me. As I bent over to catch it, the snake grabbed me by the shirt and hung on held it off with a stick. It jumped twice more at me and came clear off the ground.” Finneran (1948:124) describing the species’ habits in Connecticut, stated: “A farmer . . . con- stantly warned me away from Coon Ledge during the spring, say- ing the snakes were breeding and would ‘chase you.’ This very thing happened. In 1943 a blacksnake followed me for approxi- mately ten feet, and, in 1946, a male aggressively attacked me for a period of three minutes. There was ample opportunity for escape.” Natural Enemies Little has been published concerning the natural enemies of the common racer. The king snake (Lampropeltis getulus) is notorious for ophiphagy, and doubtless preys upon the racer at times. Wright and Bishop (1915:169) wrote of the king snake in Okefinokee Swamp, in Georgia, that “all the smaller snakes suffer, and of the larger species, the blacksnake [racer] and spreading adder are the commonest prey.” However, Clark (1949:252) examined 301 stomachs of king snakes of this species in Louisiana and found no racers, although many other kinds of snakes were represented, and, collectively, made up the greater part of the food. A rat snake (Elaphe obsoleta) found in Leon County, Florida, on June 10, 1924, had a racer in its stomach, according to the Patuxent food habits file of the U. S. Fish and Wildlife Service. A California garter snake (Thamnophis elegans terrestris) was found NATURAL HIsTORY OF THE RACER 439 to have a juvenal racer in its stomach (Fitch, 1940:96). A large alligator lizard (Gerrhonotus multicarinatus) was found swallow- ing a small racer that had been confined with it in a bag (Fitch, 1935:12). A few records of predation on blue racers by other reptiles on the Reservation and Rockefeller Tract were obtained. In 25 scats of the prairie king snake (Lampropeltis calligaster), there were 29 vertebrate prey items of which one was a blue racer. On September 15, 1962, an adult male prairie king snake 49% inches in length was found to have a 27-inch yearling racer in its stomach. In 254 scats of the slender glass lizard (Ophisaurus attenuatus) vertebrate re- mains were rare, but there were scales and bones of one hatchling blue racer. Among 21 vertebrate prey items in 14 scats of the timber rattlesnake (Crotalus horridus) there were remains of one blue racer. Among 589 prey items of copperheads there were two juvenal blue racers (Fitch, 1960:200). In contrast to these scarce records of racers in the prey of other snakes, there were more frequent records of the blue racer preying on its own young. The 1008 food items from 479 racer scats contained remains of 16 racers, small young in most instances. Raptorial birds are known to be important predators on snakes. Breckenridge (1944:118) reported finding remains of a blue racer in the pellet of a marsh hawk (Circus cyaneus). Many pellets of the marsh hawk collected on the Reservation were all found during the colder half of the year, and they contained no remains of reptiles. The same seasonal restrictions applied to the many pellets of four species of owls that were collected, and these also lacked remains of reptiles. One of the predators whose food habits have been most thoroughly investigated on the area is the red-tailed hawk (Buteo jamaicensis). Over the period 1955 through 1962, 1131 pellets of these hawks were collected, many from the Reserva- tion, but more from localities scattered throughout the eastern one- fourth of Kansas. The pellets were those of nestlings and fledglings, nearly all collected from beneath the nests, in late May, June, or early July. Some 49 different nests were represented, and remains of 43 blue racers were found. In one nest which yielded a total of 191 food items, racer remains occurred 13 times, but no other nest yielded records of more than three racers, and some other species of reptiles, notably the black rat snake (Elaphe obsoleta) comprised much more important components of the food. Since only one meal in 26 contained remains of racers, it seems that a red- 440 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. tailed hawk would destroy only a few racers in the course of a year on its territory of perhaps half a square mile, and that its preda- tion would not be a major factor in the racer’s ecology. That the red-tailed hawk is a natural enemy to be reckoned with throughout the racer’s extensive range was demonstrated by the analysis of contents of 116 stomachs from localities well scattered over the United States and Canada. Among the 152 vertebrate prey items represented there were three racers: a juvenile C. c. constrictor from Stag Lake, New Jersey, October 16, 1927; a C. c. mormon from Weiser, Idaho, April 11, 1930, and a C. c. stejne- gerianus from San Roman, Cameron County, Texas, in the spring of 1938. Because of the widely scattered geographical origin of these stomachs, a remarkably large number of species of prey were represented, and no one species predominated. Small mammals of the genera Sylvilagus, Spermophilus, and Thomomys comprised the most important component of the food; the only other reptile so well represented as the racer was the common garter snake (Thamnophis sirtalis) which likewise had three occurrences. In June and July, 1954, a nest of broad-winged hawks (Buteo platypterus) on the Reservation yielded 71 prey items of which seven were blue racers. At least 19 species of prey were repre- sented, although specific determinations were not possible in some instances. The prairie vole with eight records was the most frequent prey, and the racer and cardinal (nestlings) each had seven, while all other species were represented by fewer occurrences. Six of the racers were first-year young but the remaining one was thought to be an undersized second-year individual. Although an adult racer, especially a large one, would probably be an adversary too powerful to be killed and eaten by a broad-winged hawk, this hawk is perhaps one of the more important natural enemies of the first- year young in the eastern United States. Further records of predation on racers by raptors were obtained from the U. S. Fish and Wildlife Service files through the kind- ness of Dr. Wm. H. Stickel. There were three records of such predation by red-shouldered hawks (Buteo lineatus) from Lunen- berg, Massachusetts, May 1, 1896; Portland, Maine, March 16, 1906; and 65 miles northeast of Sarasota, Florida, June 10, 1918. There were two records of predation by broad-winged hawks from Catlettsburg, Kentucky, July 26, 1910, and Portland, Con- necticut, May 18, 1912. There were four records of predation by marsh hawks on racers—Peck’s Island, Maine, September 18, 1908; NATURAL HIsToRY OF THE RACER 441 Edgartown, Massachusetts, April 12, 1912; West [Tisbury?], Massa- chusetts, July 31, 1912; and Okanagan Landing, British Columbia, June 5, 1918. A sparrow hawk (Falco sparverius) from De Ranch, Wyoming, May 27, 1910, had eaten a racer, as had a barn owl (Tyto alba), from Franklin County, Kansas, November 16, 1922, and a crow (Corvus brachyrhynchos, nestling ) from Onaga, Kansas, May 16, 1914. Perhaps certain mammalian predators are even more important natural enemies than are raptorial birds, but records of predation on racers by mammals are few. Substantial samples of scats of opossums and coyotes from the Reservation have contained no re- mains of racers. The Fish and Wildlife Service files include a record of an opossum from Adrian, Michigan, on April 26, 1934, that had racer remains in its stomach. No racers were specifically recorded from a collection of 820 scats and 22 stomachs of raccoons (Procyon lotor) from Douglas County, Kansas, reported upon by Stains (1956:43), but occurrences of unidentified snake may have per- tained in part to the racer. Skunks are probably more important natural enemies, but food habits data from the Reservation are lacking for the two kinds of skunks occurring there. Crabb (1941: 356) in a study of the food habits of the spotted skunk (Spilogale putorius) in southeastern Iowa, did not report any reptiles among the many kinds of prey found in scats. Several times in many years of residence near Medford, Jackson County, Oregon, I saw remains of racers which appeared to be victims of striped skunks (Mephitis mephitis). Tracks and other sign of the skunks were often noticed along a little-used road on a hillside, passing between a pasture and a hay field, with an oak grove, high weeds, and brush, bordering the road. The racers sometimes found as victims along this road were small- or medium-sized individuals. In every instance the predator had begun eating on the tail end of the snake, and later had abandoned the remains leaving the head and part of the fore- body still intact. Predation had occurred at night. Skunks foraging mostly in twilight or darkness, probably find racers inactive beneath flat rocks or in shallow burrows. The skunks would be too slow and clumsy to catch the snakes in the open when they were fully active. The funnel traps used for catching the racers also caught many other kinds of animals. Often a racer and another kind of snake were caught together, but, ordinarily, in these instances no damage to either resulted, although racers, copperheads, king snakes, and garter snakes are all known to eat each other’s young. At times, 442 UNIVERSITY OF KANSAS PuBts., Mus. Nat. Hist. large predatory ground beetles (Calosoma scrutator and Pasimachus sp.) were so abundant that a dozen or more were caught together in a trap, and several times such groups attacked and killed and partly ate young racers caught in the same traps. White-footed mice (Peromyscus leucopus), deer mice (P. maniculatus), harvest mice (Reithrodontomys megalotis), short-tailed shrews (Blarina brevicauda), and least shrews (Cryptotis parva) all were caught rather frequently in the traps, and each, on one or more occasions, gnawed and killed or severely injured a racer trapped with it. The attacks probably were motivated by hunger in all instances, the snake in some instances failing to defend itself or escape because of low temperature. Occasionally such encounters might occur even under natural conditions, the mouse or shrew finding and attack- ing the snake while the latter was torpid and helpless in its hiber- naculum or in a more superficial temporary shelter. Disease Relatively few of the racers examined showed signs of disease. The most common type of affliction was an infection of the skin causing cankerlike sores, chiefly on the ventral plates, sometimes on the head or tail. These were seen most often in snakes recently emerged from hibernation, but persisted later in the season in years of exceptionally wet weather. Some of the racers most severely afflicted appeared to be in debilitated condition. The causal organism was not determined, nor was it definitely determined whether this type of disease causes mortality. Parasites The ectoparasites of the racer are chiefly chiggers, the parasitic larvae of mites. Loomis (1956) in his study of the chigger mites of Kansas examined many of the racers captured in the early years of my field work. He checked a total of 130 racers, mostly from the Reservation, and found four different species of chiggers, all of the genus Trombicula: T. alfreddugési, T. lipovskyana, T. kansensis and T. sylvilagi. The common pest chigger, T. alfreddugési, para- sitizes most of the species of reptiles, birds, and mammals occurring in Kansas, and it was by far the most common kind on the racers. The numbers per racer in different months were as follows: June, 81; July, 285; August, 482; September, 123; October, 15. Many of the racers were collected in the relatively cool and moist summers of 1950 and 1951. In a year of typical weather, heaviest infesta- NATURAL History OF THE RACER 443 tions occur in early summer, June or the first half of July, and the numbers taper off rapidly in the hot but often dry weather of late July, August, September, and October. In hot, humid weather of early summer a racer may have several hundred chiggers attached to it, filling most of the areas of exposed skin between the scales. The chiggers are conspicuous because of their bright orange color. Like T. alfreddugési, T. lipovskyana was also found in large num- bers on racers and is found on many kinds of hosts. Loomis (op. cit.:1281) recorded it from one kind of frog, one kind of toad, one kind of turtle, two of lizards, six of snakes, 19 of birds, and nine of mammals from eastern Kansas. Five larvae of T. sylvilagi were recorded from a racer captured in October. That kind of chigger is primarily a parasite of small mammals, and perhaps cannot develop successfully when it attaches to a snake. Unlike most other kinds of chiggers, this species is most in evidence in autumn and winter. A single larva of T. kansensis was found on a racer in October. This relatively rare kind of chigger has been found on several kinds of snakes and small mammals (including pocket gophers) and is known from hot and dry rocky places. Even the racers that were heavily loaded with chiggers showed no obvious ill effects, but the chiggers are potentially vectors of various diseases. Of the many endoparasites found in racers, the lung fluke, Neorenifer lateriporus was the only one identified and frequently observed in my study (Stewart, 1959). This is a digenetic trema- tode of the subfamily Reniferinae. The racer is its specific host. The life history is still unknown, but in other members of the sub- family, all of which parasitize snakes, an aquatic snail and a frog are required as hosts at different stages of the life cycle. Presum- ably N. lateriporus has similar requirements. The two common local water snails, Heliosoma trivolvis and Physa anatina, are both potential hosts. By far the most probable frog host is the leopard frog. In wet weather of July the recently metamorphosed leopard frogs leave the water and disperse to all habitats, probably carry- ing with them the parasites acquired in the tadpole stage. The racers in turn probably acquire their flukes by eating the young frogs in summer. In any case, the adult racers are nearly all parasitized, but the flukes have not been found in those racers that were less than one year old. During their first few months, the racers are too small to swallow leopard frogs, even the young. The flukes have been seen in the live racers mostly in May, when most adults are infested with the flukes. Seemingly at this season the 444 UNIVERSITY OF KANSAS PuBLis., Mus. Nat. Hist. flukes migrate forward into the mouth of the host. Probably this is the time when the flukes breed and lay eggs; if so, the eggs would pass through the digestive tract of the snake and escape with its feces. The latter are usually left in terrestrial situations unfavor- able for the development of an aquatic stage, but perhaps some of the eggs are washed into ponds by heavy summer rains. In late summer and fall the flukes are not to be found in the mouths of the live snakes. Most complete records of the flukes present in racers were kept in 1959. The following table shows the numbers of racers examined and the percentage having flukes in that year. Ortenburger (1928:182) recorded lung flukes ( Renifer ellipticus) from blue racers. In Maryland, McCauley (1945:76) also recorded numerous lung flukes (Pseudorenifer sp.) in an immature racer 490 millimeters in total length. Parker (1941:34) recorded Neore- nifer septicus from racers collected at Reelfoot Lake, Tennessee, Greensboro, Georgia, and Kissimmee, Florida; also he recorded N. georgianum from racers collected at Reelfoot Lake. N. septicus was recorded by the same author from the water moccasin (Agkis- trodon piscivorus) and N. georgianum was also recorded from the king snake (Lampropeltis getulus). Cloacal smears from the racers examined usually showed an abundance of ciliate protozoans, either parasites or commensals, and occasionally nematode worms. Harwood (1933:66) examined two racers from the vicinity of Houston, Texas, and found four kinds of helminths: Kalicephalus agkistrodontis, K. rectiphilus, Ophidascaris sp., and Polydelphis sp. Each parasite was found in only one of the two snakes. McCauley (loc. cit.) recorded nema- todes (Physaloptera obtusissima) from black racers in Maryland, and Ortenburger (loc. cit.) recorded Physaloptera sp. from the blue racer. TABLE 18. SEASONAL INCIDENCE OF THE FLUKE NEORENIFER LATERIPORUS, IN MourTHS OF BLUE RACERS ON THE RESERVATION AND ROCKEFELLER TRACT Apr. | May | May | June | June 15-30) 1- 15 15-81] 1-15 |15-30 July | Aug. | Sept.| Oct. Number of racers checked). << caus > 1 28 25 24 17 30 5 9 2 Percentage of semple having flukes. . ol OO 79 72 50 18 0 0 0 0 NATURAL History OF THE RACER 445 Populations Composition Since there is a brief annual breeding season, any local popula- tion of racers consists of a series of discrete annual age groups. The population reaches its annual maximum in early September, after undergoing sudden increase by the addition of the annual crop of hatchlings. Throughout the remainder of the year numbers of racers undergo gradual reduction as a result of the many combined mortality factors that affect them. This mortality is distributed among all the age classes, but the heaviest losses, both percentage- wise and in actual numbers, are sustained by the first-year young. Being by far the most numerous group, these young suffer more mortality than all the other age classes combined. Presumably much of this mortality is concentrated in the early weeks of life, while the young are still near their minimum size; the rate of loss is gradually reduced as larger size is attained and some of the early hazards are outgrown. In the adult age classes also, the larger and older snakes live in greater security, and the rate of mortality is higher in the smaller and younger snakes. Even before hatching, the eggs are subject to heavy losses from predators, and probably from drying, flooding, and other unfavorable climatic factors. Un- fortunately it was not possible to obtain definite figures on any of these losses since the eggs were never found under natural condi- tions and the hatchlings were seen only in relatively small numbers. The records obtained from trapping racers in late spring and summer in fields provided a somewhat different picture of the population from the sample obtained along the ledges in autumn. In the former sample there were 400 males to 257 females, but in the latter sample there were 355 males to 379 females. I regard the summer sex ratio as a distorted one, brought about by the greater activity of the males in the breeding season. Racers are caught most easily in May, and the fact that two or more males often were trapped with the same female, while the reciprocal combinations did not occur, demonstrates the increased activity of the males in their search for mates at this season. In autumn there is no sexual activity; both sexes probably are equally active in seek- ing places to hibernate when they are trapped along the hilltop outcrops. The ratio of 51.6 per cent females in my sample of 734 may indicate that in the males greater activity at other seasons re- sults in a somewhat higher mortality. This idea is borne out by 446 UNIvERSITY OF Kansas Pusts., Mus. Nat. Hist. FLAVIVENTRIS 3Il MALES MORMON 57 MALES uJ a | a = <— w ie (eo) uJ oO = FLAVIVENTRIS ea 211 FEMALES O ~ [oa ld a MORMON 68 FEMALES 1000 SNOUT-VENT LENGTH IN MILLIMETERS Fic. 20. Histograms comparing snout-vent length in random sam- ples of Coluber c. flaviventris (those captured on the Reservation and Rockefeller Tract in 1960 and 1961) and C. c. mormon (speci- mens in the Museum of Vertebrate Zoology, University of Cali- fornia). In the museum sample, collected by conventional methods, the first-year young constitute a prominent and fairly distinct size group, whereas in the sample of racers from the Reservation and Rockefeller Tract, mostly caught in wire traps of quarter-inch mesh, relatively few young were secured. It is demonstrated that hatch- lings are approximately the same length in both populations, but flaviventris grows much longer, and that the differences in length between the sexes is approximately the same in each population. NATURAL HIsToRY OF THE RACER 447 the fact that for the supposed two-, three- and four-year-olds com- bined, females comprise 51.2 per cent, but they comprise 55.6 per cent of those more than four years old and 61.3 per cent of those more than five years old. By assigning to each racer caught an arbitrary age, on the basis of size according to Table 17, I calculated the population (exclusive of those snakes in their first year of life) to have the composition shown in Table 19. TaBLE 19. PERCENTAGES OF ADULT POPULATION OF BLUE RACERS COMPRISED BY Each ANNUAL AGE GROUP YEARS OF AGE | 2 3 4 5 6 a 8 9 10 11 or Percentage of population...) 41.5/17.8/12.6] 9.5 | 6.1 | 4.3 | 2.7 | 2.4/1.2] 1.9 Figures are completely lacking to show the relative numbers of juveniles, until, already approaching adult size, the young are about to enter their second hibernation. My combined fall samples in- clude 303 of these latter young, as against 142 racers about to enter their third hibernation. Thus, after having nearly attained adult size, these adolescent snakes sustained a loss of 53 per cent in a year. Losses must occur at an even more rapid rate in the younger and smaller snakes. It may be speculated that of the approximately 300 eggs produced by a population of 100 adult racers, 150 are lost before or during the period of incubation which lasts nearly two months. Of the 150 hatchlings emerging in early September, at least one-third probably are eliminated by the following breeding season in late May, leaving 100. The 100 survivors at this stage are still small juveniles, but by autumn they have attained adoles- cent size. By this time, if they had undergone a further reduction by 53 per cent, only 47 would remain—approximately the number to be expected if the population were stable from year to year. A notable difference between the fall sample and the spring sample that I obtained was the higher proportion of large and old racers in the former sample. This difference can be attributed to the year-to-year changes in the population during the 14 consecu- tive years spanned by my field work. The fall sample of 734 racers represented the combined catch of the years 1949 through 1962, rather evenly distributed, but the spring sample included few snakes from the years 1949 through 1957; most were from the years 1958 7—7864 448 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. through 1961. In 1949 when the study was begun, the Reserva- tion was being protected for the first time, and formerly overgrazed pastures or cultivated fields were acquiring a rank growth of grass and weedy vegetation, and thus becoming favorable habitat. The abundant new habitat promoted rapid increase in the population of racers until the newly available areas were filled to their “carrying capacity.” Table 21 shows the changing trends of the different age groups. Although the separate annual samples are perhaps too small to show the composition of the population accurately, it is significant that in the fall of 1949 an unusually high proportion of the racers caught were one-year-olds, hatched in September, 1948. Table 20 shows that in a typical group of 100 subadult and adult racers (second year and older) only a little more than one-fourth are productive females. The largest females, six years old and older, making up less than ten per cent of the adult population, contribute nearly half the total complement of eggs. The calculated number of eggs pertains to a stage before ovi- position, and subsequent losses through resorption under unfavor- able conditions, through inviability of embryos and through deaths of some of the gravid females, are to be expected. In the weeks of incubation further losses are sustained. Although these losses cannot be measured, they must be severe as on numerous occasions scattered and torn eggshells representing entire clutches dug out and destroyed by predators, have been found. Probably other clutches are destroyed underground by such predators as moles and egg-eating snakes, and still others by insects. Tinkle (1959:195) wrote that in a clutch of 15 eggs found under a board, four were parasitized and had small perforations. Molds destroy a high per- centage of all reptilian eggs that are incubated artificially and doubt- less destroy many under natural conditions also. Excessive heat or moisture, or desiccation, resulting either from climatic extremes or from poor choice of a nest site by the female, would cause further loss. In four different years, Blair (1960:108) found that losses of eggs between laying and hatching in the Texas spiny lizard (Sceloporus olivaceus) ranged from 69 per cent to 86 per cent; no other comparable study of the extent of egg losses in a species of reptile is known to me. The racer is somewhat less prolific than the spiny lizard, and potentially longer lived; the racers eggs are larger and thicker-shelled, and they are deposited in deeper burrows. It might be expected that losses during incuba- tion would be somewhat less in the racer than in the spiny lizard. NaATuRAL History OF THE RACER 449 Although figures for the youngest age groups—one-year-olds and hatchlings—are missing, approximations of them may be fur- nished by extrapolation, from the information available regarding the productivity of the population. Some factors involved in pro- ductivity are that the sex ratio deviates from parity, slightly in favor of the females in the adolescents but more markedly in favor of the females among the older age groups; that some adult females apparently fail to produce eggs in the breeding season, but the percentage decreases in the older snakes; and that number of eggs TABLE 20. CALCULATED PRopuctTivity IN A HypotuHetTicaL Group or 100 SUBADULT AND ADULT BLUE RACERS OOS |r|. .aT.. . TTOTOTOOTOOO TT oOVOO—eneree————— Percentage | Percentage | Percentage Eggs Ps of — of fecund | productive per_ umber ( ) population | females in females in | productive e ane d EATS in age-group | age-group age-group female produce 2 41.5 51.2 13 9.2 26 3 17.8 51.2 56 9.9 51 4 12.6 51.2 60 10.8 42 5 9.5 55.0 57 13.0 39 6 and over 18.6 61.3 80 15.7 143 TABLE 2]. PERCENTAGES OF RACERS IN EACH ANNUAL AGE Group (EXCLUSIVE oF HaTCHLINGS) IN AUTUMNAL SAMPLES AT DIFFERENT STAGES OF THE FIELD Stupy, SHOWING SHIFT TOWARD OLDER AGE-GROUPS IN THE LATER YEARS —S—SSSSSS=S=S=S=S=SS9DnBnanmna==S=SSS=[=|[=[=[=[=—__—_—_—_—_— Year or combination of years represented by each sample YEARS OF AGE 1949 1950 1953 1956 1959 1951 1954 1957 1960 1952 1955 1958 1961 1962 1 54 44 39 39 41 2 14 24 21 19 19 3 18 iL 23 16 13 4 4 10 10 11 9 5 6 6 3} 8 8 6 2 1 3 1 3 7 2 4 4 2 3 Shek oe SSS. ces cll othe. eecdinds heen 82 5 2 Qe en” tclacic Sok 1b A Als) Z, 1 Oe 1D A Laciak | Pearse gl aehven| | ah eta ale ake 4 2 ] Olderithan 10" We. f=s 5a). ose leeesorets: Seales wget. Number in sample 49 126 174 194 242 450 UNIVERSITY OF KANSAS PusLs., Mus. Nat. Hist. per clutch increases in proportion to the size and age of the female producing them. Too few figures are available concerning most of these factors to indicate more than the trends; nevertheless the available figures have been used in Table 20 in an attempt to esti- mate the productivity of a hypothetical population. Numbers Conant (1938:178, Pl. 7) published a photograph of 106 blue racers killed in February, 1932, by farmers near Bellville, Ohio, and Pope (1944:173) mentioned that scores of blue racers ag- gregated in October around an oak-covered dune near Chicago. In both these instances large hibernating aggregations were in- volved, and the areas represented by them are unknown; nothing has been recorded regarding population densities. The records obtained through my fall trapping, along hilltop rock outcrops, yielded no information concerning population densities, but those obtained in fields in summer did provide significant in- formation in this regard. Even after years of trapping on the same area, the catch still consisted largely of new individuals; the method was not sufficiently effective to catch all racers present at any one time, and the total catch for a season therefore provided only a crude index of the minimum number present. The summer trapping was carried on in three separate areas. One of these was the area of bottomland pastures and formerly cultivated fields where the Reservation headquarters are located, a block of 39 acres bounded on three sides by woodland, and on the fourth by cultivated farm land. Effective trapping in this area was carried on through the years 1955 to 1961 inclusive. A second area, of 48 acres, was one of upland fields, mostly covered with re- established prairie grasses, in the northeastern part of the Reserva- tion. A third area, of 137 acres, also upland, was that of the Rocke- feller Tract, cultivated through 1956 and sown to prairie grasses the following year, and the adjacent northwestern hilltop portion of the Reservation. Effective trapping on these two latter areas was carried on in 1958 through 1962. For the seven years of trapping in the House Field area, the catch was as follows: 30, 33, 38, 38, 34, 24, and 20. In four years of trapping, the northeast field area yielded 42, 28, 37, 59, and 19 blue racers, and the Rockefeller Tract yielded 52, 67, 67, 126, and 106. The actual catch was hence less than one per acre in nearly all instances, but the year-to-year differences in catch are believed NATURAL History OF THE RACER 451 to be caused chiefly by differences in numbers of traps used and in trapping effort, rather than by changes in the numbers of racers present. Best index to the number of racers actually present is provided by the number of recaptures, and their ratio to first captures. The population of course, undergoes alteration from year to year, with many racers eliminated and replaced by others. In 1955, 26 racers were caught in the May-June-July period, in the headquarters field area. In the August-September-October period of the same year five racers were caught of which only one was a member of the original 26. The five-to-one ratio indicates that the original 26 may have represented an actual population of 130, but of course the single recapture is much too small a sample to provide a reliable ratio. Some of the racers caught in May were recaptured in June, others in July, and still others not until late sum- mer or early autumn. Somewhat different estimates can be ob- tained for the population depending on how the season’s records are divided. For instance, in the headquarters field area in May 1955, eleven racers were caught; in the remainder of the season 20 were caught, of which two were members of the original group of eleven. The 20 to 2 ratio indicates that the 11 caught in May represented an actual population of 110. In the period May-June, 18 racers were caught, and in July-August-September-October, 15 were caught, including four of the original 18, hence indicating a popula- tion of 67. Although obviously the population underwent some change during the course of the season, the three sets of census figures apply essentially to the same population, and the divergence in them illustrates the wide range of error arising from insufficiently small samples. Common sources of error in the censusing of natural populations of animals by the capture-recapture method (“Lincoln Index” or “Petersen Index”) arise from the fact that the composition of a local population often changes between two sampling periods, or even within them. Some of the animals marked may move elsewhere, to be replaced by unmarked immigrants, or they may die and be replaced by unmarked maturing young. First-year racers that could pass through the quarter-inch mesh of the traps in spring and early summer became too large to escape in this way in the latter half of the summer, but these young were excluded from the census com- putations. There was doubtless some shifting of marked individuals away from the study areas and shifting of new individuals onto 452 UNIvERSITY OF Kansas Pusts., Mus. Nat. Hist. these areas in the periods of weeks between successive samplings. Jackson (1939) has explained a method of correcting census com- putations based on capture-recapture ratios when there is a con- sistent trend of diminishing recaptures with increase in elapsed time. However, in my records no such trend is discernible; furthermore it has been demonstrated that individual racers tend to stay within the same home range area throughout most of their season of activity. Therefore, I conclude that shifts of individuals away from the study areas or into them, in the intervals between samplings constitute only a minor source of error. A combination of the figures from the three samples listed above provides an intermediate “smoothed” figure that can be accepted with somewhat more confidence than any one of the separate censuses because it is based on more records. The combined ratios indicate a total of 105 racers in the headquarters field area. The figures obtained in the different sampling periods, and the census figures derived from their ratios are shown in tables 22 and 23. Differences from year-to-year in the census figures for any one area show no consistent trends and their variation is similar to that shown in different samples for the same season. Probably populations were fairly stable throughout the periods involved. If such stability is assumed, the samples from different years may be combined, and the composite figures derived from them may be accepted with more confidence. For the headquarters field area, for instance, 419 records of racers were gathered in all the pre- liminary sampling periods of the seven seasons involved; the rec- ords totalled 263 for all the secondary sampling periods, with 70 recaptures in secondary periods of the snakes recorded in the cor- responding preliminary periods. A population of 75 racers is indi- cated—1.9 per acre. Corresponding figures for the northeast field area are: preliminary samples 453, secondary samples 163, re- captures 39, calculated number 135 (2.82 per acre). For the Rockefeller Tract the figures are as follows: Preliminary samples 807, secondary samples 476, recaptures 126, the ratio indicating a population of 158, or 1.11 per acre. These figures represent the number of adults present in early summer when the population is near its annual low point. The first-year young, excluded from this census because they cannot be caught in representative numbers, perhaps approximate the num- ber of adults, in May, so the figures obtained would need to be ap- proximately doubled to be representative of the entire population. NATuRAL History OF THE RACER 453 By late summer the adults, and especially the yearlings, have under- gone substantial reduction in numbers, but in late August and early September the hatching of a new crop of young increases the popu- lation to its annual maximum. The maximum numbers probably are about three times those obtained by censusing adults in early summer. The peak population of late summer or early autumn is estimated to consist of hatchlings, comprising somewhere near 50 per cent; adults, comprising a little more than 25 per cent; and yearlings comprising a little less than 25 per cent. Densities in early summer of one to three adult blue racers per TABLE 22. CApTuRES RECORDED AND POPULATION CALCULATED FROM THEM on Hitttor GRASSLAND AREAS OF ROCKEFELLER EXPERIMENTAL TRACT AND ADJACENT RESERVATION IN Four DIFFERENT YEARS Four- 1959 1960 1961 1962 year average First Census: arly MERA yeh nakane es choy ahs 0 8 21 32 61 Mate MMA WE che slow malt ease 3 13 33 35 84 IRecaptures scsi tre ct ciclo sie 0 1 2 7 10 Estimated population......]........ 104 346 160 128 Second Census: BY ee eee tom eon 3 25 54 59 141 DUNG ete Neiais si states ek 2 9 6 25 40 80 Recapturestsrwaeic sci - 16 «6 - 0 3 6 24 33 Estimated population......]........ 50 165 99 85 Third Census: May-June-July........... 25 34 94 100 253 Sept-Oct... 6. atoe oer 8 13 20 13 54 Recaptures® 4. scccm. tee 1 3 5 4 13 Estimated population...... 200 182 376 324 263 Fourth Census: NayAJUNE inmates eros 31 30 1 76 210 July-Aug.-Sept.-Oct........ 24 15 34 35 108 Recaptunestts. sere ent 3 2 14 12 31 Estimated population...... 104 225 177 222 183 Fifth Census: May pri racer ne sc mcrae 3 25 54 60 142 June to October........... 23 20 67 40 150 Recapturesneayabecoecee 0 5 14 20 39 Estimated population......]........ 100 258 120 162 Five Sets Combined: Combined first samples... .. 62 122 296 327 807 Combined second samples. . 67 67 179 163 476 Recaptures; poco see e : 4 14 41 67 126 Estimated population...... 208 117 258 159 153 454 UNIVERSITY OF KANSAS PuB3s., Mus. Nat. Hist. acre probably are typical of the better types of habitat in the region of my study. The upland field area estimated to have 2.82 racers per acre was better habitat than the other two study areas. Prior to 1948 it had been cultivated and severely eroded. In 1949 most of it was sown to seeds of prairie grasses, and by 1958 dif- ferent parts of it were dominated by different species of native perennial tall grasses interspersed with areas that supported a weedy type of vegetation, and other areas that supported dense thickets of sumac, dogwood, elm saplings, or other woody plants. The abundance and diversity of dense cover and of small animals made this area especially favorable habitat for the racer. Some local areas probably support higher populations of racers TABLE 23. CAPTURES RECORDED AND POPULATIONS ESTIMATED FROM THEM IN HEADQUARTERS FIELD AREA OF RESERVATION Seven- 1955.| 1956 | 1957 | 1958 | 1959 | 1960 | 1961 year average First Census: Maya. Sette 11 21 7 24 17 13 6 99 June-July-Aug.| 20 11 Do 20 22 12 14 124 Recaptures.... 2 1 4 5 5 2 0 19 Estimated population..| 110 | 231 44 96 75 US ¥||Segentors 92 Second Census: May-June..... 18 25 17 28 27 15 13 143 July-Aug.- Sept.-Oct...} 15 8 17 15 15 11 8 89 Recaptures.... t 1 7 5 9 3 1 30 Estimated population..| 68 | 200 41 84 45 55 | 104 61 Third Census: May-June-July | 26 28 22 31 31 20 19 177 Aug.-Sept.- Ochs. artes 5 7 13 2, 6 5 2 50 Recaptures.... 1 2 f 5 3 1 1 20 Estimated population. .} 130 128 47 87 63 75 148 63 Three Sets Combined: Combined first samples....| 55 74 46 83 75 48 38 419 Combined sec- ond samples} 40 26 55 A7 43 28 24 263 Combined recaptures. . 7 4 18 15 17 6 3 70 Estimated population. .| 105 160 47 87 63 75 101 75 BEALE AS Fic. Fic. Fic. Fic. — to (os) 4, mately Head of hatchling blue racer, dorsal view, September 1962, approxi- mately x 3. Head of hatchling blue racer, lateral view, September 1962, approxi- mately x 3. Head of yearling male blue racer, lateral view, August 1, 1961, a little less than twice natural size. Head of adult male blue racer, lateral view, July 16, 1961, approxi- er All three snakes from Rockefeller Experimental Tract, Jefferson County, Kansas. PLATE 20 1. Abandoned limestone quarry on a hilltop of southward exposure on University of Kansas Natural History Reservation, in late autumn of 1951. crevices along the base of the ledge provided favorite hibernating sites for blue racers. 2. Hatchling blue racer and eggshell from which it had recently emerged, in early September, 1962; * approximately 175. PLATE 21 > pe - ‘ FP aX «! hk oan Fic. 1. Wire funnel trap set at base of hilltop limestone outcrop in a spot strategically located for interception of blue racers searching for deep crevices in which to hibernate, October 15, 1949. Fic. 2. Large clutch of 21 blue racer eggs, recently plowed out, at Harold Brune farm, Jefferson County, Kansas, July 10, 1962. PLATE 22 Fic. 1. Habitat of blue racer, bluestem prairie on Botany Bluff at northwest corner of the University of Kansas Natural History Reservation, looking south. Trees and brush in background are along limestone outcrop at top of slope. Mowed area in foreground is southwest corner of Rockefeller Experimental Tract, a privately owned farm at the time this photograph was taken in the summer of 1951. Fic. 2. Habitat of blue racer, bluestem prairie on south slope of Botany Bluff, looking north along west edge of the Reservation, summer of 1951. By 1962, with exclusion of fire, and protection from mowing, prairie vegetation had largely disappeared from this slope, and had been replaced by trees and brush. As a result of these successional changes racers no longer found this slope a suitable habitat in summer, but they continued to resort to the hilltop rock outcrop to hibernate in autumn. NATURAL History OF THE RACER 455 than do areas where censuses were made, but under modern con- ditions, situations that offer near optimum habitat are not likely to be extensive or to persist long. On land that is capable of produc- ing a good crop of vegetation, the crop is usually harvested either by grazing of livestock or by using the land for cultivation, with the result that the racers are, at least in some seasons, forced into marginal situations. More than 50 years ago in Missouri, Hurter (1911:170) wrote that the racer “was quite common 20 years ago in pastures, meadows and fields but as cultivation has advanced it is becoming quite rare.” In 1962 the widespread and adaptable blue racer is still common in many parts of its range, including Mis- souri, but in most places its population densities probably are lower than formerly. Reduction since 1911 has probably been far more drastic than the reduction that had occurred up to that time. Schmidt and Necker (1935:69), writing of the racer in the Chicago region, noted “the snakes which raise their heads and face mowing machinery tend to be exterminated in agricultural areas.” They stated that in the Chicago region the racer had been exterminated by the advance of agriculture except in two extensive sand dune areas. In July 1962, Mr. V. B. Howell, a progressive farmer of the Great Bend area in central Kansas, told me that the kinds of snakes inhabiting cultivated land—blue racer, bull snakes, prairie king snakes, hog- nosed snakes, and others—had undergone great reduction in num- bers during the period of his farming. He estimated that in a forty- year period the numbers had declined to perhaps five per cent of their level in the area most familiar to him, centering at his farm 11 miles northwest of Great Bend, Barton County. In accounting for this change in population density Howell pointed out the relative destructiveness to small animals of modern farm machinery as con- trasted with horse-drawn equipment or that used with tractors of earlier models. Modern tractors move forward so rapidly that there is little opportunity for snakes or other small animals to avoid them, and the plows and disks cut wide swaths penetrating more deeply into the soil than did older types. On July 10, 1962, in searching the furrows of a freshly plowed small field on the Harold Brune farm, for turned-up nests of the snakes, I found two adult blue racers that had been struck and killed by the plow, possibly while they were underground. In fields that are plowed or cultivated between the times of egg-laying and hatching, the eggs are de- stroyed. Because of its rapid movements and alertness, the racer 456 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. is more likely to escape farm machines than are most other kinds of snakes. Nevertheless, it is vulnerable and survives in cultivated areas only when they are interspersed with pastures, woodlots, or streamside thickets where at least part of the population may find refuge. Summary Field study of the blue racer was carried on in several localities in Kansas, but chiefly at the University of Kansas Natural History Reservation (the northeasternmost section of land in Douglas County), and the adjacent 160-acre Rockefeller Experimental Tract in Jefferson County. By October 26, 1962, after 14 years of field work, a total of 1423 racers had been captured some 2197 times. The locale of the present study was near the geographic center of the blue racer’s range. The range, chiefly in the Mississippi Valley and Great Plains region, is centrally situated with respect to the other seven subspecies. An extensive but scattered literature concerning the ecology of the species as a whole, and its several geographic races, has been reviewed and utilized for comparison with my own field data. Blue racers were caught in wire funnel traps set in prairie and pastureland habitat in summer, and along hilltop limestone out- crops in woodland in autumn. The autumn trapping along rock outcrops was carried on each year from 1949 to 1962, but effective summer trapping was carried on only in the last six years of the study. Each racer caught was individually and permanently marked by scale clipping. More than half were caught only once, but many were recorded repeatedly, with a maximum of 16 captures. The racer occurs throughout most of the United States, and its populations are subject to much geographic variation. The snakes are largest in the northeastern part of the range, with clines of de- creasing size toward the southeastern, southern and western parts of the United States. There are somewhat parallel trends in colora- tion; the black racer of the northeastern states grades into paler, gray or light brown subspecies in southern Florida, Texas, and the far western states. Accompanying these changes in color and size are minor morphological changes and major ecological changes. The black racers of the eastern states often inhabit forest or forest- edge habitats while the paler and smaller snakes of more southern or western areas typically inhabit scrub, chaparral, or prairie. The large, dark-colored racers of the eastern and northeastern states are especially inclined to attack larger prey including small verte- NATURAL History OF THE RACER 457 brates, even weasels, rabbits, and chipmunks, whereas the smaller and paler racers of more southern and more western areas take a higher proportion of insects and rarely attack vertebrates other than small reptiles. On the area where field work was carried on in northeastern Kan- sas, tall-grass prairie habitat is preferred, but fields of grain or alfalfa, grazed pasture, brush, woodland edge, groves or open wood- land, and weedy fields are all utilized to some extent. The racer is strictly diurnal and largely terrestrial but it may climb through bushes or small trees in foraging or escaping. The blue racer is a typical colubrine snake of slender build, with large eyes, and vision plays an important role in finding prey and detecting enemies. In the adult blue racer the dorsal color is vari- able, pale brown or gray, bluish, greenish or slaty. In the hatchling, however, there is a distinct pattern of a type widespread among colubrines and also among snakes of other groups—a series of middorsal blotches on an olive ground color, with alternating rows of smaller spots on each side. The ventral surface is pale, with dark speckling. The pattern is sharply defined on the anterior part of the body, but markings become progressively more obscure pos- teriorly and are scarcely discernible on the tail. The juvenal pattern fades gradually as growth proceeds, and there is much individual variation in the rate of its loss. Some racers still retain the juvenal pattern faintly discernible after attain- ment of sexual maturity. There are also striking ontogenetic changes in the proportions of the head, body and tail. The diameter of the eye is approximately one per cent of the snout-vent length in hatchlings, but is only a little more than half that relative size in the largest adults. In the course of allometric growth other parts of the head also enlarge less rapidly than the body, but more rapidly than the eye. In hatchlings there is a slight average dif- ference between the sexes in relative tail length, with males’ tails the longer. Relative tail length increases slightly in both sexes up to the time of sexual maturity, and then decreases slightly with ad- vancing age. Racers in northeastern Kansas spend nearly half the year in hiber- nation, with average recorded emergence date April 16, and average date of retirement into hibernation November 8. Hibernacula are usually in crevices in hilltop limestone outcrops with south ex- posures. Winter temperatures within the hibernacula are usually well within the range 0 degrees to 10 degrees Centigrade. Spring 458 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. emergence has been recorded at an air temperature of only 12.5 degrees Centigrade. Racers bask in sunshine frequently even in warm weather, and the temperature preferendum is several degrees higher than in most other kinds of snakes. Bodily temperatures ob- tained from blue racers that were fully active, either under natural conditions or in a large outdoor enclosure, were concentrated in the neighborhood of 34 and 35 degrees Centigrade. For short periods racers can survive temperatures up to 45 degrees without damage, but more prolonged exposure to temperatures of slightly less than 40 degrees can be fatal. In hibernation, racers can withstand temperatures slightly below freezing, but they cannot survive being frozen solid. Blue racers tend to limit their activities to familiar areas or home ranges; some individuals may live out their entire lives within the same home range, but others shift from time to time. Average home ranges of approximately 26 acres for males and 24 acres for females were calculated. The racers’ preference for hibernacula in a habitat different from that to which summer activities are con- fined necessitates spring and fall migrations between the limestone outcrops where hibernation occurs and the grasslands where the snakes stay in summer. The average spring or fall migration is approximately a quarter of a mile, but an individual racer does not consistently return to the same hibernaculum. Many racers were recorded to have made movements of 2000 to nearly 4000 feet, in- volving shifts in home range, but some later shifted back to their original areas. Some may have made even longer shifts but their movements would not have been recorded since they would have gotten beyond the limits of the study area. Blue racers hunt by various methods, often by coursmg through dense vegetation in active search in which vision is of primary im- portance in locating the prey. Almost any small animal that moves nearby may be overtaken and caught with a sudden dash. From analysis of scats and prey from stomachs, a total of 1357 food items of more than 50 species was compiled. Favorite prey species were the cricket (Gryllus assimilis), grasshoppers (Arphia simplex, Mela- noplus femur-rubrum, M. bivittatus, M. differentialis), camel crickets (Ceuthophilus sp.), katydid (Neoconocephalus robustus), vole (Microtus ochrogaster), white-footed mouse (Peromyscus sp.), racerunner, (Cnemidophorus sexlineatus), and leopard frog (Rana pipiens). The insects taken greatly outnumbered the vertebrates, but the vertebrates made up most of the actual bulk of prey eaten. NATURAL History OF THE RACER 459 Crickets, grasshoppers, and katydids comprised most of the insect prey. Beetles, moths and cicadas were rarely taken. Vertebrate prey included miscellaneous small snakes (some of them juvenal racers), mammals, and birds. Seasonal change in the composition of the food is slight, but vertebrates figure more prominently in early summer, and insects comprise increasing percentages later in the season. Composition of the food differs according to size of the snake; gryllid and ceuthophilid crickets are best represented in the food of juveniles whereas small mammals, and grasshoppers of the genus Melanoplus are best represented in the food of large adults. The breeding season is mainly in May. Mating is promiscuous and two or more males may court the same female simultaneously. A courting male lies on or alongside a receptive female, with spasmodic rippling abdominal movements, and with his vent ad- pressed to hers. At intervals in the courtship period the female moves swiftly for a few feet or a few yards shifting to a new spot, and during her activity the male strives to maintain contact with her. From time to time the male leaves the female briefly and courses rapidly around her in a devious route. Courtship is con- summated when the female raises her tail in acceptance of the male and intromission is effected. During coitus, which lasts for periods of minutes, the female moves forward slowly, dragging the passive male, tail-first behind her. Ovulation normally is in late May. Eggs from 29 to 39 milli- meters in length and 14 to 21 millimeters in breadth are laid, from mid-June to early August, usually in tunnels of fossorial mammals such as voles or moles, at depths of five to 12 inches. Clutches of the blue racer average 11.8 eggs but the number is correlated with age and size of females; two-year-olds average only 9.2 eggs, whereas those females that are six years old or more average 15.7 eggs. Also, there is geographic variation in size of clutch, from only 5.8 eggs in C. c. mormon of the West Coast to 16.8 eggs in C. c. constrictor of the northeastern states. In each breeding season some females of adult size do not produce clutches. Only about 13 per cent of the two-year-olds in a small sample were fecund, but the ratio increased to 80 per cent in old adults. Incubation aver- ages 51 (43 to 63) days. Hatchlings usually make several longitudinal slits in the egg- shell with the egg tooth before emerging, and often require a day or more to emerge after the first slit is made. Hatchlings average 460 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. 10% inches and 4.16 grams. By late October when these young are ready to enter hibernation, they have grown to 16% inches and 12.3 grams. Typical October lengths (overall) in inches for males and females, respectively, after successive seasons of growth, are as follows: yearlings, 27% and 29; two-year-olds, 34% and 37%; three- year-olds, 37% and 41%; four-year-olds, 40 and 44%; five-year-olds, 41% and 46é; six-year-olds, 42% and 48M; seven-year-olds, 43% and 50; eight-year-olds, 44 and 51%. Judging from trends in the small samples available, sex ratio in hatchlings is approximately 1 : 1. In the summer trapping of blue racers, males are caught in larger numbers than females, but seemingly this is because of their greater activity. In fall along the limestone outcrops where the racers hibernate females were caught in slightly greater numbers than males among the young adults, and made up a little more than 60 per cent of the old adults. Presumably the males are eliminated more rapidly, because of their greater activity, their smaller size, or a combination of both factors. The composition by age groups of the adult racers captured was as follows: two-year-olds, 41.5 per cent; three-year-olds, 17.8 per cent; four-year-olds, 12.6 per cent; five-year-olds, 9.5 per cent; six- years-olds, 6.1 per cent; seven-year-olds, 4.3 per cent; eight-year- olds, 2.7 per cent; nine-year-olds, 2.4 per cent; ten-year-olds, 1.2 per cent; more than ten years old, 1.9 per cent. Attempts to census blue racers on the study plots, from the capture-recapture ratios, yielded highly variable figures for dif- ferent areas and even for the same area at different times. Vari- ability is believed to result mostly from the small sizes of the samples, none of which is large enough to yield a census figure that is statistically reliable. Combined samples yielded figures indicat- ing populations of one to three adults per acre in early summer in areas of favorable habitat. To represent the entire population in late summer—the time of its annual maximum—these figures would need to be approximately tripled. The blue racer depends mostly on speed to escape its enemies. In escaping it often takes advantage of a downhill slope to increase its speed. A racer startled at close range often sets out with a violent thrashing that attracts attention to a given spot, then glides away so silently and rapidly that its course is not detected. The snake may circle back and approach the scene of disturbance slowly and stealthily, from the direction opposite to that taken in depart- ing. Often a racer seeks concealment by climbing into a bush or NATURAL HistoRY OF THE RACER 461 tree, and occasionally escape is effected by swimming. A racer that is caught or cornered usually makes a spirited defense by striking and biting. An alternative reaction, seen most typically when the snake is handicapped by injury or by low temperature, is to coil with the head concealed, and, with writhing movements, to smear the surface of the body with musk discharged from glands in the base of the tail. Many natural enemies prey upon the racer, but in the present study the majority of records pertained to the red-tailed hawk. The broad-winged hawk, marsh hawk, red-shouldered hawk, sparrow hawk, and barn owl also are among the raptors that feed on this kind of snake. Among mammals the striped skunk is the only species definitely recorded to prey on the racer, feeding on the eggs as well as upon the young and adults. The common king snake, prairie king snake, timber rattlesnake, copperhead, and even the slender glass lizard have been recorded to prey on the blue racer, but probably all are of minor importance as natural enemies. Shrews (Blarina, Cryptotis) and mice (Peromyscus, Reithrodon- tomys) have sometimes killed and eaten racers confined in traps, and under natural conditions they possibly prey upon snakes that are immobilized when torpid from cold, in their hibernacula and temporary shelters. Chiggers are the most common ectoparasites of the racers. Four species of the chigger genus Trombicula have been recorded on racers from the Reservation. The racer is an important host of the common pest chigger, Trombicula alfred- dugési, which often attacks humans. 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Kansas Acad. Sci., 39:277-281. Hupson, G. E. 1942. The amphibians and reptiles of Nebraska. Nebraska Conservation Bull. no. 24, 146 pp. Hupson, R. G. 1949. A record length milk snake. Herpetologica, 5:47. Hurter, J. H. 1911. Herpetology of Missouri. Trans. Acad. Sci. St. Louis, 20:59-274. INGER, F. and Cuark, P. J. 1948. Partition of the genus Coluber. Copeia, 1943(3):141-145. Jackson, C. H. N. 1939. The analysis of an animal population. Jour. An. Ecology, 8:238-246. Kinc, W. 1939. A survey of the herpetology of Great Smoky Mountain National Park. Amer. Midl. Nat., 21(3):531-582. Kurmstra, W. D. 1959. Food of the racer, Coluber constrictor, in southern Illinois. Copeia, 1959(3):210-214. LANGBARTEL, D. A. 1947. Snakes collected at Camp Eastman, Hancock County, Illinois. Herpetologica, 4:27-28. Liner, E. A. 1949. Notes on the young of the blue racer, Coluber constrictor flaviven- tris. Copeia, 1949(3):230. 1954. 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Notes on amphibians and reptiles from the central United States. Amer. Midl. Nat., 32:478-490. MartTIN, E. P. 1956. A population study of the prairie vole (Microtus ochrogaster) in northeastern Kansas. Univ. Kansas Publ. Mus. Nat. Hist., 8(6): 361-416. MAsLIN, T. P. 1959. An annotated check list of the amphibians and reptiles of Colorado. Univ. Colorado Studies, Biol. Series, No. 6, 98 pp. McCau.ey, R. H. 1945. The reptiles of Maryland and the District of Columbia. Published by the author, Hagerstown, Maryland, 194 pp. Minton, S., JR. 1944. Introduction to the study of reptiles in Indiana. Amer. Midl. Nat., 32:488-477. 1949. An ectopic egg in Coluber. Herpetologica, 5:96. MossiMan, J. E. and Rass, G. B. 1952. The herpetology of the Tiber River area, Montana. Copeia, 1952(1):23-27. Mu ark, S. and Mutaix, D. 1942. A neglected species of Coluber. Copeia, 1942:13-15. Munro, D. F. 1948. Comparative sizes of two female blue racers and their eggs. Her- petologica, 4:199-200. 1949. Eating habits of young Coluber constrictor flaviventris. Herpe- tologica, 5:72-78. 1950a. Incubation and hatching of eggs of Coluber c. flaviventris. Herpe- tologica, 6:122-124. 1950b. Juvenile pattern in a mature Coluber c. flaviventris. Herpetologica, 6:124. NEILL, W. T. 1948. Hibernation of amphibians and reptiles in Richmond County, Georgia. Herpetologica, 4:107-114. Ovum, E. P. and KuENz_er, E. J. 1955. Measurement of territory and home range size in birds. Auk, 72:128-137. ORTENBUBGER, A. I. 1928. The whip snakes and racers, genera Masticophis and Coluber. Mem. Univ. Michigan Mus., vol. 1, pp. xviii + 247. OwsEns, V. 1949. An overwintering colony of Coluber c. constrictor (Say) and Elaphe o. obsoleta ( sayy Herpetologica, 5:90. PARKER, M. V. 1941. The trematode parasites from a collection of amphibians and rep- tiles. Jour. Tennessee Acad. Sci., 16(1):27-45. Peters, J. A. 1942. Reptiles and amphibians of Cumberland County, Illinois. Copeia, 1942(3):182-183. Popr, C. H. 1935. The reptiles of China. Amer. Mus. Nat. Hist., Nat. Hist. Cent. Asia, v. 10, lii +- 604 pp. 1944. Amphibians and reptiles of the Chicago area. Chicago Nat. Hist. Mus. Press, 275 pp., 12 pls. Pore, T. E. B. and Dickinson, W. E. 1928. The amphibians and reptiles of Wisconsin. Bull. Pub. Mus. City of Milwaukee, 8(1):1-138. Natura History OF THE RACER 467 Ricumonp, N. D. and Gor, C. J. 1938. Notes on a collection of amphibians and reptiles from New Kent County, Virginia. Ann. Carnegie Mus., 27:301-310. RossMan, D. A. 1960. Herpetological survey of the pine hills area of southern Illinois. Quart. Jour. Florida cae Sci. (1959), 22(4):207-225. Scumipt, K. P. and NEcKErR, W. 1935. Amphibians and veriles of the Chicago region. Bull. Chicago Acad. Sci., 5(4):57-77. SCHRODER, R. C. 1950. Hibernation of blue racers and bull snakes in western Illinois. Nat. Hist. Misc., 75:1-2. SHANNON, F. A. 1950. Coluber constrictor mormon—a state record for Arizona. Herpe- tologica, 6:59. . M. 1956. Handbook of amphibians and reptiles of Kansas. Univ. Kansas Mus. Nat. Hist. Misc. Publ., 9:357 pp. Smitu, P. W. 1947. The reptiles and amphibians of east central Illinois. Bull. Chicago Acad. Sci., 8(2):21-40. 1961. The amphibians and reptiles of Illinois. Bull. Illinois Nat. Hist. Surv., 28(1):1-298. Strains, H. J. 1956. The raccoon in Kansas. Univ. Kansas Mus. Nat. Hist. Misc. Publ., No. 10, pp. 1-76, pls. 1-4. STEWART) 2. EE: 1959. Lung flukes of Thamnophis and Coluber in Kansas. Unpublished thesis University of Kansas Library, pp. 1-62. StIcKEL, W. H. and Cops, J. B. 1947. The home ranges and wandering of snakes. Copeia, 1947:127-136. Storer, D. H. 1839. Reports on the ichthyology and herpetology of Massachusetts. Comm. Zool. and Bot. Surv. Mass., Boston, 201-258. SuRFACE, H. A. 1906. The serpents of Pennsylvania. Pennsylvania State Dept. Agr. Monthly Bull., Div. Zool., 4 (4 and 5):118-208. TINKLE, D. W. 1959. Observations of reptiles and amphibians in a Louisiana swamp. Amer, Midl. Nat., 62(11):189-205. Trowsriwnce, A. H. 1937. Ecological observations on amphibians and reptiles collected in southeastern Oklahoma during the summer of 1934. Amer. Midl. Nat. 18(2):285-808. Unter, F. M., Corram, C. and Ciarxe, T. E. 1939. Food of snakes of the George Washington National Forest, Virginia. Trans. 4th Amer. Wildlife Conf., Amer. Wildlife Inst., Washington, D. C., 1939:605-632. Van DENBURGH, J. 1923. The reptiles of western North America. Occas. Papers California Acad. Sci., 10:1-1028. VAN DE VELDE, R. L., MARTAN, J. and RIsLey, P. L. 1962. Eggs and hatchlings of the snake Coluber constrictor mormon from Oregon. Copeia (1):212-218. VERRILL, A. E. 1869. The biter bitten. Amer. Nat., 3:158-159. 468 UnrversiTy OF Kansas Pusts., Mus. Nat. Hist. WELTER, W. A. and Carr, K. 1939. Amphibians and reptiles of northeastern Kentucky. Copeia (3): 128-130. WERLER, J. E. and McCattion, J. 1951. Notes on a collection of reptiles and amphibians from Princess Anne County, Virginia. Amer. Midl. Nat., 45(1):245-252. Woopsury, A. M. 1931. A descriptive catalog of the reptiles of Utah. Bull. Univ. Utah, 21 (no. 5): x + 129 pp. Woops, G. T. 1944. Black snake attacks a boy. Copeia, 1944(4):257. Wricut, A. H. and Bisuop, S. C. 1915. A piles reconnaissance of the Okefinokee Swamp in Georgia. II. Sn Proc. nae Nat. Sci. Philadelphia, 67:189-192. Wricut, A. H. and Wricut, A. 1957. Handbook of snakes fe the United States and Canada. Comstock Publ. Associates, Cornell Univ. Press, vol. I, xviii + 564 pp. Transmitted June 3, 1963. 18. 19. 20. 21. 22. 23. (Continued from inside of front cover) Conspecificity of two pocket mice, Perognathus goldmani and P. artus. By E. Raymond Hall and Marilyn Bailey Ogilvie. Pp. 513-518, 1 map. Janu- ary 14, 1960. Records of harvest mice, Reithrodontomys, from Central America, with de- scription of a mew subspecies from Nicaragua. By ‘Sydney Anderson and J. Knox Jones, Jr. Pp. 519-529. January 14, 1960. Small carnivores from San Josecito Cave (Pleistocene), Nuevo Leén, México. By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, 1960, Pleistocene. pocket gophers from San Josecito Cave, Nuevo Leén, México. By Robert J. Russell, Pp. 539-548, 1 figure in text. January 14,1960. Review of the insectivores of Korea. By J. Knox Jones, Jr., and David H. Johnson. Pp, 549-578. February 23, 1960. Speciation and evolution of the pygmy mice, genus Baisnoys. By Robert L, Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960. Index. ; Pp. 671-690 Vol. 10. 1. 2. 8. *6. Studies of birds killed in nocturnal migration. By Harrison B. Tordoff and Robert M. Mengel. Pp. 1-44, 6 figures in text, 2 tables: September 12, 1956. Comparative breeding behavior of Ammospiza caudacuta and A. maritima. By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956. The forest habitat of the University of Kansas Natural History Reservation. By Henry S, Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures in text, 4 tables. December 31, 1956. Aspects of reproduction and development in the prairie vole (Microtus ochro- gaster)., By Henry S. Fitch.. Pp. 129-161, 8 figures in text, 4 tables. Decem- ber 19, 1957. Birds found on the Arctic slope of northern Alaska. By James’ W. Bee. Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958. The wood rats of Colorado: \distribution and ecology. By Robert B. Finley, Jr. Pp. 213-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958. » Home_ranges and movements of the eastern cottontail in Kansas. By Donald W. Janes. Pp. 553-572, 4 plates, 3 figures in text, May 4, 1959. Natural history of the salamander, Aneides hardyi. By Richard F. Johnston and Gerhard A. Schad. Pp. 573-585. October 8, 1959: A new subspecies of lizard, Cnemidophorus sacki, from Michoacan, México. f By William E, Duellman. Pp. 587-598, 2 figures im text. May 2, 1960. 10. A taxonomic study of the~middle American snake, Pituophis deppei. By William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960. Index. Pp. 611-626. Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960. Vol. 12. 1. m2. Functional morphology of three bats: Sumops, Myotis, Macrotus. By Terry A. Vaughan. Pp. 1-153, 4 plates, 24) figures in text. July 8, 1959. The ancestry of modern Amphibia: a review of the evidence. By Theodore H. Eaton, Jr.” Pp. 155-180, 10 figures in text. July 10, 1959. The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49 figures in text. February 19, 1960. A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in text. May\2, 1960. j Natural history of the bell vireo: By Jon C, Barlow. Pp. 241-296, 6 figures in text. March 7, 1962. Two new, pelycosaurs from the lower Permian of Oklahoma. By Richard C. Fox. Pp. 297-307, 6 figures in text. May 21, 1962. 7. Vertebrates from. the barrier island of Tamaulipas, México. By Robert K. Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 809- 345, pls..5-8. June 18, 1962. Teeth of Edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 fig- * ures in text. October 1, 1962. 9. Variation in the muscles and nerves of the leg in two genera of grouse Tympanuchus and Pedioecetes). By E. Bruce Holmes. Pp. 863-474, 20 gs. October 25, 1963. A new genus of Pennsylvanian Fish (Crossopterygii, Coelacanthiformes) from Kansas. By Joan Echols. Pp. 475-501, 7 figures. October 25, 1963. Observations on the Mississippi Kite in southwestern Kansas. By Henry S. Fitch. Pp. 508-519. October 25, 1963. Jaw musculature of the Mourning and White-winged doves. By Robert L. Merz. Pp. 521-551, 22 figures. October 25, 1963. More numbers will appear in volume 12. Five hatural hybrid combinations in minnows (Cyprinidae). By Frank B. * Cross and W. L. Minckley. Pp. 1-18. June 1, 1960. A distributional study of the amphibians of the Isthmus of Tehuantepec, México. By William’ E. Duellman. | Pp. 19-72, pls. 1-8, 3 figures in text. August 16, 1960. i A new subspecies of the slider turtle (Pseudemys scripta) from Coahulia, anes By John M. Legler. Pp. 73-84, pls. 9-12, 8 figures in text. August Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, pls. 18-20, 26 figures in text. November 30, 1960. Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and Rocky Mountains. By Henry S. Fitch and T. Paul Maslin. Pp. 289-808, 4 figures in text. February 10, 1961. (Continued on outside of back cover) 10. (Continued from inside of back cover) Fishes of the Wakarusa river in Kansas. By James E. Deacon and Artie L. ~ Metcalf. Pp. 809-322, 1 figure in text. February 10, 1961. Geographic variation in the North American cyprinid fish, Hybopsis gracilis. By Leonard J. Olund and Frank B. Cross. Pp, 323-348, pls. 31.54, 3 asaree in text. February 10, 1961. Decriptions of two species of frogs, genus Ptychohyla; studies of Ameri- can hylid frogs, V. By William E. Duellman. Pp. 349-357, pl. 25, 2 figures ~ in text. April 27, 1961. Fish populations, following a_drought, in the Neosho and Marais des Cygnes rivers of Kansas. By James Everett Deacon. Pp. 859-427, pls. 26-30, $ figs. August 11, 1961. Recent ‘soft-shelled turtles of North America (family Trionychidae). By gee Webb. Pp. 429-611, pls. 31-54, 24 figures in text. February Index. Pp. 613-624. Vol. 14. I. 2. 3. Neotropical bats from western México. By: Sydney Anderson. Pp. 1-8. October 24, 1960. Geographic variation in the harvest mouse. Reithrodontomys megalotis, on the central Great Plains and in adjacent regions. By J. Knox Jones, Jr., and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961. Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson. Pp. 29-67, pls..1 and 2, 8 figures in text. July 24, 1961. A ‘new subspecies of the black myotis (bat) from eastern Mexico. By E. berg Hall anad Ticul Alvarez. Pp. 69-72, 1 figure in text. December North American yellow bats, “Dasypterus,’’ and a list of the named kinds of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr. Pp. 73-98, 4 figures in text. December 29, 1961. , Natural history of the brush mouse (Peromyscus boylii) in Kansas with description of a new subspecies. By Charles'A. Long. Pp. 99-111; 1 figure in text.. December 29, 1961. Taxonomic status of some mice of the Peromyscus boylii group in eastern Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 118- 120, 1 figure in text. December 29, 1961. A new subspecies of ground squirrel (Spermophilus spilosoma) from Ta- maulipas, Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962. Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J. ras Jones, Jr., and Ticul Alvarez. Pp. 125-183, 1 figure in text. March 7, Anew doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene, of Texas. By E. Raymond Halland Walter W. Dalquest. Pp. 185-138, 2 figures in text. April 30, 1962. : A new subspecies of wood rat (Neotoma) from northeastern Mexico. By Ticul Alvarez. Pp, 189-148. April 30, 1962. Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul a are and M. Raymond Lee. Pp. 145-159, 1 figure in text. -May 18, A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, 1 figure in text.. May 21, 1962. The mammals of Veracruz. By E. Raymond Hall anad Walter W. Dalquest. Pp. 165-362, 2 figures. May 20, 1963. The recent mammals of Tamaulipas, México. By Ticul: Alvarez. Pp. 3638- 473, 5 figures in text. May 20, 1963. ; More numbers will appear in volume 14, The amphibians and reptiles of Michoacan, México. By William E. Duell- * man. Pp. 1-148, pls. 1-6, /11 figures in text. December 20, 1961. Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox Jones, Jr., and. George W. Byers. Pp. 149-178. January 81, 1962. A new species of frog (Genus Tomodactylus) from western México. By Robert G. Webb. Pp. 175-181, 1 figure in text. “March 7, 1962. Type specimens of amphibians and reptiles in the Museum of Natural His- tory, the University of Kansas. By William E. Duellman and Barbara Berg. Pp. 183-204. October 26, 1962. Amphibians and Reptiles of the Rainforests of Southern El Petén, Guatemala. By William E. Duellman. Pp. 205-249, pls. 7-10, 6 figures in text. October 4, 1968. A revision of snakes of the genus Conophis (Family Colubridae, from Middle were) By John Wellman. Pp. 251-295, 9 figures in text. October 4, 68. ‘ A review of the Middle American tree frogs of the genus Ptychohyla. By Wea Duellman. Pp. 297-349, pls. 11-18, 7 figures in text. October 18, 1963. Natural history of the racer, Coluber constrictor. By Henry S. Fitch. Pp. 351- 468, pls. 19-22, 20 figures. December 30, 1963. More numbers will appear in volume 15. ey wi eat sentiewss tis ig a a <4 es ee ae ar em ct =} LS mea ae ee ee r ® en eet apertg Moeg Re 5k RES A + = ai ee ee a Te ee ee om Oe ae H A i V A I x D UNIVERSITY OF KANSAS PUBLICATIONS MusEuUM OF NATURAL HIstTory Volume 15, No. 9, pp. 469-491, 4 figs. March 2, 1964 A Review of the Frogs Of the Hyla bistincta Group BY WILLIAM E. DUELLMAN UNIVERSITY OF KANSAS LAWRENCE ~ 1964 Unrversiry oF KANsAS PUBLICATIONS, MusEUM OF NATURAL HiIsTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 15, No. 9, 469-491, 4 figs. Published March 2, 1964 UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY THE STATE PRINTER TOPEKA, KANSAS 1964 MUS. COMP. zoo) LIBRARY JUL 21 1964 A Review of the Frogs ae Of the Hyla bistincta Crop eaeD VERSITY, BY WILLIAM E, DUELLMAN CONTENTS PAGE NEN ERODUCTION ict fh tide coetbylett oitdha to's es Seca esis ONO OBI eres ae 471 INC mow ecient ss went cs deh ec ats d a wscanepeeel einai ret a ere 471 REESE YEA DISTINGTAT GROUP Ws © cylin cy aie Giemsa sei ac rt icwene wera 472 Amalysissote Gharacters scenic cits ir ee eee aoe 473 Key to the Species of the Hyla Bistincta Group ............ 474 INCCOUNTSIOF) THE SPECIES 13/557. 2 ies divas eal cede eM Ie 475 ig as biStinet a, Copan weno radey sale Shey aad. Ae ee reo 475 Hula charadricola mew SPeCiIes, «. 64. eee ee ae 478 LAU LG TODETESOMUM MAYO ons lates AeA yar pe nee 481 ALG NAChYAeTINGMaylOn ak acy 6 ace eet 485 PAU IG Crassa (BYOCCHINE och oie calla sie sete eas ae ete Tee 486 FUIEATION SEEPS icc ep tn Me alee a ih ny aye sgn sacl Ry ST Ree RO 489 HETEERAT URE 6 CURED 0. ci G.ch eee s 2 nv mute sua eRe eae ogee: 491 INTRODUCTION In the mountainous regions of Middle America there are several groups of hylid frogs that inhabit mountain streams. Some of these groups, such as Plectrohyla and Ptychohyla, have been elevated to generic rank, whereas others are retained in the large and complex genus Hyla. In the mountains of México five species of hylids that seem to compose a phyletic unit are herein referred to as the Hyla bistincta group. Since 1955 I have been accumulating specimens of, and data on, this group with the result that all specimens known to me, including the types of all named taxa, have been studied. Detailed observations have been made on the ecology and life his- tories of three of the species; the other two species are known to me only from preserved specimens. Acknowledgments For permission to examine specimens in their care I am indebted to Charles M. Bogert, American Museum of Natural History (AMNH); Doris M. Coch- ran, United States National Museum (USNM); Jean Guibé, Museum National d'Histoire Naturelle, Paris (MNHN); Robert F. Inger, Chicago Natural His- tory Museum (CNHM); Hobart M. Smith, University of Illinois Museum of Natural History (UIMNH); Charles F. Walker, University of Michigan Museum of Zoology (UMMZ). (Abbreviations of institutions given above in parentheses are used throughout; the Museum of Natural History, University of Kansas is abbreviated KU.) (471) 472, UNIVERSITY OF KANSAS PuBts., Mus. Nat. Hist. For their willing assistance in the field I am grateful to Ann S. Duellman, Dale L. Hoyt, and John Wellman. Permits for collecting in México were generously issued by the late Ing. Luis Macias Arellano, Departamento de la Fauna Silvestre, Direccién General de Caza. The drawings in figures 1 and 8 were executed by Gail Selfridge. This research has been supported by the National Science Foundation (NSF G-9827). THE HYLA BISTINCTA GROUP The five species comprising the Hyla bistincta group are mod- erate-sized hylids having rather blunt heads and robust bodies. The fingers are long and have little webbing (Fig. 1). The skin of the dorsum is thick and glandular, but not tuberculate. An anal sheath is present. The skull is rather broad, flat, and solidly roofed. The ethmoid is broad, curved downward laterally, and solidly sutured to the frontoparietal. The nasals are broad, sutured for their entire width with the ethmoid, and broadly in contact medially. The premaxillaries are in contact medially; each has a long, flat nasal process. The quadratojugal is absent, and the maxillary tapers to a point posteriorly. There is no squamosal-maxillary con- nection. The maxillary and premaxillary teeth are rather long, bifid, and moderately spatulate. Some teeth on the premaxillary and anterior part of the maxillary are hooked. The vomerine teeth are spatulate and bifid. A broad, flat, ossified prepollex is present but does not project as a spine. The known tadpoles have ventral mouths, 2/3 tooth-rows, two or more rows of labial papillae, and long tails with low fins. a Fic. 1. Palmar view of right hand of: a.—Hyla robertsorum (KU 57661), b.—Hyla charadricola (KU 58414). x 3. FRrocs OF THE HyLA BISTINCTA GROUP 473 As thus defined the Hyla bistincta group can be distinguished from all other groups of Middle American frogs by the combination of absence of the quadratojugal, non-projecting prepollex, long fingers with little webbing, and stream-inhabiting tadpoles having 2/3 tooth rows and two or more rows of labial papillae. Possibly Hyla arborescandens and Hyla hazelae belong in this group. Because these species are somewhat different from the in- cluded species and because their tadpoles are as yet unknown, I have refrained from including these two species in the Hyla bistincta group. Taylor (1948:261) assigned Hyla proboscidea (= H. dal- questi) and (1949:272) Hyla cyclomaculata to this group, but be- cause these two species have a quadratojugal and notably different tadpoles, they are excluded from the group. Frogs of the genus Plectrohyla closely resemble species in the Hyla bistincta group but differ principally in having a projecting prepollex. In the highlands of Costa Rica a group of species, of which Hyla moesta is best known, resembles species in the Hyla bistincta group. At present insufficient information is available on the Costa Rican species to determine their affinities. Analysis of Characters The characters used in the systematic study of the frogs in this group are those usually employed in anuran systematics. Of the various measurements and proportions, the snout-vent length and the relative size of the tympanum to the eye apparently are of more taxonomic importance than the others (Table 1). In all of the species the tympanum is at least partially covered by a heavy, dermal supratympanic fold, and in some specimens of H. pachy- derma the tympanum is completely obscured. In two species (H. bistincta and H. charadricola) the snout is square, whereas in the other species it is round. The fingers are long and slender in H. crassa, pachyderma, and robertsorum and somewhat shorter with more webbing in H. bistincta and charadricola. Breeding males of Hyla pachyderma have moderately large nuptial spines; the other species have small spines, except H. charadricola in which spines apparently are absent. A well-defined thoracic fold is present in H. pachyderma, and a weak fold is present in H. robertsorum; the other species lack folds. In all species there is an anal sheath; this sheath is longest in H. bistincta, in which the anal opening is directed ven- trally at the level of the lower edge of the thighs. AT4 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. TABLE 1.—CoMPARISON OF CERTAIN MEASUREMENTS AND PROPORTIONS IN THE SPECIES OF THE HyYLA BISTINCTA Group. (Data FrRoM ADULT MALES; Means Are GIVEN IN PARENTHESES BELOW THE RANGES. ) < s eae =a =| gE Sec oe eel vahe oe ~ » o SPECIES N oo = 5 ie 5 ef a) AS =| a) fl S 1 = AH] 6 ° BE 2) R RH TE ADIStiNClas © sie ces a 38 | 43.0-53.8 | 0.47-0.52 | 0.32-0.37 | 0.35-0.48 (46.3) (0.49) (0.34) (0.42) H. charadricola....... 10 | 35.3-44.4 | 0.50-0.54 | 0.31-0.33 | 0.30-0.37 (40.4) (0.52) (0.32) (0.34) H. robertsorum....... 26 | 39.9-47.9 | 0.48-0.51 | 0.30-0.36 | 0.36-0.47 (43.1) (0.49) (0.32) (0.41) H. pachyderma....... 1 39.9 0.53 0.32 ICTASSO Se Se eee 1 Dana 0.50 0.33 0.28 Frogs in this group are rather drab in appearance. The dorsal color varies from dull green to various shades of brown. The most distinct aspect of the coloration is the different color patterns on the flanks and posterior surfaces of the thighs. The flanks in all species are marked with spots or reticulations. Hyla bistincta differs from other members of the group in having vocal slits and a distensible vocal sac. Only this species has been reported to call (Shannon, 1951:473). Insofar as is known, the other species are mute. Examination of skeletal preparations of H, bistincta, charadricola, and robertsorum and X-rays of the other species shows no notable specific differences in the osteology. Since the tadpoles of only H. bistincta (Duellman, 1961:47) and H. robert- sorum (Rabb and Mosimann, 1955) are known, larval characters are of limited use in intra-group systematics. Key to the Species of the Hyla bistincta Group 1. Snout, in dorsal profile, short and bluntly rounded; canthus rounded, sometimes indistinct, vocal slits "absent?") {yo nes se oe ee ee Snout, in dorsal profile, longer, squared; canthus distinct, vocal slits present or absent 2. Feet webbed to base of discs; dorsum dull olive green; flanks having cream-colored:spots.2 4.8. em St sie acre eae eae Gee H. crassa, p. 486 Feet not webbed to base of discs; dorsum brown ...............- 8 8. Tarsal fold strong; thoracic fold heavy; webbing on feet extending to middle of penultimate phalanx of fourth toe; distinct white stripe above anus; cluster of large spines on thumb in breeding males, H. pachyderma, p. 485 © Yel jeyie) 8) 0] ela: erie joo sale /eviel ie ollie elie 'eheie ue jes) (es \e\e ies e) sile).e) ¢,e) 0/\e\e-[e FROGS OF THE HyLA BISTINCTA GROUP 475 Tarsal fold weak and short; thoracic fold absent or barely indicated; webbing of feet extending to base of penultimate phalanx of fourth toe; no distinct white stripe above anus; nuptial tuberosities in breeding males consisting of small spines ................ H. robertsorum, p. 481 4, Tarsal fold strong; anal flap elongate; anus opening at lower edge of femur; dorsum tan or brown; flanks mottled with cream and brown; venter immaculate creamy white; no anal stripe; vocal slits present, H. bistincta, p. 475 Tarsal fold faint and short; anal flap not elongate; anus opening at middle of femur; dorsum olive-green with black reticulations; flanks greenish gray with brown or black spots; a row of white flecks above and below anus, vocal slits absent .............. H. charadricola, p. 478 ACCOUNTS OF THE SPECIES In the following accounts complete synonymies are given for each species. In each account one specimen is described in detail; after this description the variation is discussed. In the list of specimens examined, localities are arranged alphabetically within the states, which also are given in alphabetical order. Localities given in italics are not shown on the accompanying maps (Figs. 2 and 4) due to crowding of symbols. Hyla bistincta Cope Hyla bistincta Cope, Proc. Amer. Philos. Soc., 17:87, August, 1877 [Holo- type.—USNM 32261 from “most probably Veracruz,” México; Francis Sumichrast collector]. Brocchi, Etude des batraciens de lAmérique Centrale, p. 43, 1881. Boulenger, Catalogue Batrachia Salientia, 2nd Ed., p. 401, February 1, 1882. Sumichrast, La Naturaleza, 6:81, 1882. Cope, Bull. U. S. Natl. Mus., 82:14, 1887; Bull. U. S. Natl. Mus., 34:85], 1889. Giinther, Biologia Centrali-Americana, Reptilia and Batrachia, pp. 265-6, June, 1901. Diaz de Ledén, Indice de los batracios que se en- cuentran en la Republica Mexicana, p. 17, June, 1904. Nieden, Das Tierreich, Amphibia, Anura I, p. 247, June, 1923. Kellogg, Bull. U. S. Natl. Mus., 160:163-164, March 31, 1932. Taylor, Proc. Biol. Soc. Wash- ington, 50:50-58, April 21, 1937; Univ. Kansas Sci. Bull., 26:889, No- vember 27, 1940. Taylor and Smith, Proc. U. S. Natl. Mus., 194:87, June 7, 1948. Taylor, Univ. Kansas Publ. Mus, Nat. Hist., 1:261, August 16, 1948; Copeia, no. 4:272-278, December 15, 1949. Smith and Taylor, Univ. Kansas Sci. Bull., 33:346, March 20, 1950 [Type locality restricted to Acultzingo, Veracruz, México]. Rabb and Mosimann, Occas. Papers Mus. Zool. Univ. Michigan, 563:6-9, March 29, 1955. Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 15:47-49, December 20, 1961. Hyla bistincta labeculata Shannon, Proc. U. S. Natl. Mus., 101:470-473, figs. 92a, 93d, May 17, 1951 [Holotype—USNM 123689 from San Lucas Camotlan, Oaxaca, México; Walter S. Miller collector]. Smith and Wil- liams, Herpetologica, 19:28, April 11, 1963. Hyla bistincta bistincta, Shannon, Proc. U. S. Natl. Mus., 101:472, May 17, 1951. Shannon and Werler, Herpetologica, 11:85, July 15, 1955. Smith and Williams, Herpetologica, 19:28, April 11, 1963. Diagnosis—Maximum snout-vent length in males 54mm.; snout in dorsal profile truncate; tarsal fold strong; inner metatarsal tubercle large, high, and elongate; outer metatarsal tubercle absent; webbing on foot extending to mid- dle of antepenultimate phalanx of fourth toe; nuptial spines on thumb small; thoracic fold absent; anal opening at level of lower edge of femur; dorsum 476 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. brown or tan; belly cream-color; flanks creamy yellow with brown reticulations or spots; anal stripe absent; vocal slits present. Description—The following description is based on KU 68078 from Uruapan, Michoacan, México. Adult male having a snout-vent length of 43.2 mm.; tibia length, 22.1mm., 51.1 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe), 20.8mm.; greatest width of head, 15.4mm., 35.6 per cent of snout-vent length; head length, 13.9 mm., 32.1 per cent of snout-vent length; diameter of eye, 5.1mm.; diameter of tympanum, 2.4mm., 47.1 per cent of diameter of eye. Snout in lateral profile bluntly rounded, in dorsal profile truncate; canthus pronounced, rounded, not angular; loreal region slightly con- cave; lips thick, round, not flaring; nostrils slightly protuberant; internarial distance, 3.4 mm.; interorbital distance, 4.5 mm., somewhat broader than width of eyelid, 3.5mm. A heavy dermal fold from posterior corner of eye above tympanum and curved downward to insertion of forearm; tympanum round, its diameter slightly more than its distance from eye. Forearm not robust; row of small pustules on ventral surface of forearm; fold on wrist; prepollex moderately enlarged, covered with small, horny, nuptial spines continuous on edge of digit; row of spines on inner surface of second finger; subarticular tubercles large, round, none bifid; supernumerary tubercles rather large, round; palmar tubercles rather small, flat, elliptical; fingers long, moderately slender; length of fingers from shortest to longest, 1-2-4-3; discs moderately large, that on third finger slightly larger than tympanum; rudimentary web between first and second and between second and third fingers; web between third and fourth fingers extending about one-fourth length of fourth finger. Heels over- lap by about one-third length of shank when hind limbs adpressed; tibiotarsal articulation extends to anterior edge of eye; tarsal fold strong, extending to heel; inner metatarsal tubercle large, high, and elongate; outer metatarsal tubercle absent; subarticular tubercles moderately large, round; supernumerary tubercles small, in single rows on proximal segments of digits; toes moderately short; length of toes from shortest to longest, 1-2-3-5-4, third and fifth toes about equal in length; toes about two-thirds webbed; web extending to middle of antepenultimate phalanx of fourth toe, to discs of first, second, and fifth toes, and to base of penultimate phalanx of third toe; discs rather small, about two-thirds size of those on fingers. Anal opening at level of lower edge of thighs; anal sheath elongate, deeply creased medially; transverse dermal fold above anus. Skin of dorsal surfaces of head, body, and limbs faintly areolate; skin of chin, belly, and ventral surfaces of thighs granular, that of ventral sur- faces of limbs, except thighs, areolate; thoracic fold absent. Tongue nearly round, slightly longer than wide, shallowly notched behind and barely free posteriorly. Vomerine teeth 4-4, situated on rounded vomerine ridges between rather small ovoid inner nares; vocal slits present, situated along posterior edge of each ramus. Color (in alcohol) pale brown on dorsal surfaces of head, body, and limbs; flanks and anterior surfaces of thighs creamy white with dark brown reticula- tions; posterior surfaces of thighs tan with creamy white spots; belly cream- color; anal stripe absent. Color (in life) pale tan on dorsal surfaces; flanks and anterior surfaces of thighs pale creamy yellow with purplish brown reticulations; posterior surfaces of thighs tan with yellow spots; ventral surfaces yellow; iris pale copper-color. FROGS OF THE HYLA BISTINCTA GROUP 477 Variation.—There is little variation in structure. The total number of vomerine teeth varies from 6 to 14. In some individuals the supratympanic fold covers the upper part of the tympanum, but at least the lower part of the tympanum is always visible. The extent of the webbing between the toes varies from three-fourths to two-thirds complete. Usually the web extends to some point on the antepenultimate phalanx of the fourth toe, but in some specimens the web extends to the base of the penultimate phalanx. In the large series of specimens from Uruapan, Michoacan, the coloration of the flanks and anterior surfaces of the thighs varies from nearly uniform creamy yellow with only fine dark reticulations to bold reticulations enclosing yellow spots. Some specimens from Oaxaca and Veracruz have slightly different markings on the flanks; in these the dark pigment is in the form of irregular spots or dashes, instead of reticulations. There is considerable variation in color in the living frogs. The dorsum varies from greenish tan and pale yellowish tan to reddish brown, and some individuals are dark chocolate brown. Remarks.—Shannon (1951:470) named Hyla bistincta labeculata on the basis of a single male from San Lucas Camotlan, Oaxaca; he diagnosed the subspecies as differing from Hyla bistincta bistincta by having “the gray reticulation of the sides entirely broken up into elongate biack blotches; tarsal fold moderately elevated.” The condition of the tarsal fold is characteristic of the species. The dis- persion of dark pigment on the flanks is variable. The type of Hyla bistincta labeculata (USNM 123689) is extreme in the de- velopment of dark dashes on the flanks, but this condition is ap- proached in several specimes from Oaxaca and Veracruz. For example, in some specimens from Cumbres de Acultzingo, Veracruz, the mottling on the flanks is bold; in others the flanks are reticulated. The specimen from San Vicente, Oaxaca, has black dashes on the flanks (Smith and Williams, 1963:23), whereas a specimen from Cerro San Felipe, Oaxaca, has no pattern on the flanks. In general, specimens from western México have reticulate mottling on the flanks as compared with the marbling on the flanks in specimens from eastern México. On the basis of available data, the recognition of subspecies in Hyla bistincta is unwarranted. The tadpoles of this species described by Duellman (1961:47) are like those of Hyla robertsorum in having 2/3 tooth-rows, peglike serrations on the beaks, and long, rounded tails. At Uruapan tad- poles were found in a rocky stream on April 24, 1956, and metamor- phosing young were found there on August 2, 1956. A completely metamorphosed juvenile has a snout-vent length of 24.8 mm. Hyla bistincta is found only along streams, where individuals can be seen clinging to vines and other vegetation closely over-hanging fast-moving parts of the stream. 478 UNIVERSITY OF Kansas Pusis., Mus. Nat. Hist. Distribution —Hyla bistincta occurs at elevations from 1400 to 2600 meters in the mountains of the Sierra Madre Occidental in western Jalisco southward through the Cordillera Volcanica in Michoacan, México, and Morelos, the Sierra de Coalcomaén in Michoacan, and the Sierra Madre del Sur in Guerrero and Oaxaca, and thence northward in the Sierra Madre Oriental to central Veracruz (Fig. 2). Specimens examined.—GuErRRERO: Omiltemi, UIMNH 38023-5, Jaxisco: 25 km. SE Autlan, UMMZ 102076. Mexico: 19 km. W Villa Victoria, UIMNH 28162, USNM 114513. Moretos: Cuernavaca, USNM 121523; 3 km. N Cuernavaca, UIMNH 28168-70. MicHoacan: Cerro San Andrés, UMMZ 102075; Dos Aguas, UMMZ 119193; 12.5 km, ENE Dos Aguas, UMMZ 119194; Los Conejos, UMMZ 94238-40; Uruapan, KU 68077-8, 69093 (skele- ton), UIMNH 20457, 28167, UMMZ 85452-3, 112838, 112839 (16), 115231 (tadpoles), 115232, 115233 (12), 121515, S-1699 (skeleton), S-1826 (skele- ton), USNM 114514-5, 114517-24. Oaxaca: Cerro San Felipe, UIMNH 28163; Pluma Hidalgo, AMNH 13447; San Lucas Camotlan, USNM 123689; San Vincente, UIMNH 51346 (Smith and Williams, 1963:23). VERACRUZ: no specific locality, USNM 32261; Cumbres de Acultzingo, CNHM 105482-3, UIMNH 28164-6, 49133-4, USNM 114525. Kilometers ® H. bistincta O #. charadricola 104° 100° $6° Fic. 2. Map showing locality records for Hyla bistincta and Hyla charadricola. Hyla charadricola new species Holotype.—University of Kansas Museum of Natural History No. 58414 from the Rio Totolapa, 14.4 kilometers by road west of Huachinango, Puebla, México, 2280 meters; obtained by John Wellman on June 8, 1960. Paratypes.—KU 58415-58423, same data as holotype, and UIMNH 50966, obtained at the type locality by William E. Duellman on February 11, 1961. Diagnosis—Maximum snout-vent length in males, 45 mm.; snout in dorsal profile truncate; tarsal fold short and weak; inner metatarsal tubercle moder- ately large, elliptical, and flat; outer metatarsal tubercle minute, round, and indistinct; webbing extending to base of terminal phalanx of fourth toe; nuptial FROGS OF THE HYLA BISTINCTA GROUP 479 spines on thumb absent; thoracic fold absent; anal opening at level of middle of thigh; dorsum olive-green with black reticulations; venter cream color with brown flecks on chin; flanks pale grayish green with brown spots; anal stripe absent; row of white flecks above and a row below anus; vocal slits absent. Description of Holotype.—Adult male having a snout-vent lenth of 44.4 mm.; tibia length, 22.2 mm., 50 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe), 20.4 mm.; greatest width of head, 14.0mm., 31.5 per cent of snout-vent length; head length, 13.6 mm., 30.6 per cent of snout-vent length; diameter of eye, 5.0 mm.; diameter of tympanum, 1.6 mm., 32.0 per cent of diameter of eye. Snout in lateral profile bluntly rounded, in dorsal profile truncate (Fig. 8); canthus pro- nounced, rounded, not angular; loreal region slightly concave; lips thick, rounded, not flaring; nostrils slightly protuberant; internarial distance, 3.7 mm.; interorbital distance, 4.1 mm., somewhat broader than width of eyelid, 3.3 mm. A heavy dermal fold from posterior corner of eye above tympanum and then to insertion of forearm; tympanum round, its diameter three-fourths its distance from eye. Forearm rather slender; a short axillary membrane; no fold on wrist; no nuptial excrescence or spines on enlarged prepollex; subarticular tubercles moderately small, round; none bifid; few supernumerary tubercles on proximal segments of digits; large, flat palmar tubercle present; fingers long and slender; length of fingers from shortest to longest, 1-2-4-3; discs moderately large, that on third finger about equal to diameter of tympanum; rudimentary web between first and second fingers; web between third and fourth fingers extending to about one-fifth length of fourth finger (Fig. 1). Heels overlap by about one-third length of tibia when hind limbs adpressed; tibiotarsal artic- ulation extends to middle of eye; tarsal fold weak, extending from moderately large, flat, elliptical inner metatarsal tubercle to middle of tarsus; outer meta- tarsal tubercle minute, round, indistinct; subarticular tubercles round; single row of small, round supernumerary tubercles on proximal segments of each digit; toes moderately short and slender; length of toes from shortest to longest, 1-2-3-5-4, third and fifth about equal in length; toes about three-fourths webbed; web extending to base of terminal phalanx of fourth toe; discs small, about two-thirds size of those on fingers. Anal opening directed posteroventrally at middle of thighs; two transverse dermal folds above anus; short, thin anal sheath present; many small tubercles lateral and ventral to anal opening. Skin of dorsum, chin, and ventral surfaces of limbs smooth; belly moderately gran- ular; posterior surfaces of thighs heavily granular; no thoracic fold. Tongue nearly round, shallowly notched behind, free posteriorly for about one-fourth its length; vomerine teeth, 3-4, long, situated on posteroventral edges of narrow transverse vomerine ridges between moderately large, round inner nares; no vocal slits. Color (in alcohol) purplish brown on dorsal surfaces of head, body, and limbs; fine darker reticulations on dorsum; flanks pale tan with dark brown spots; posterior surfaces of thighs tan; chin creamy white with brown spots; belly dusty white; undersides of thighs and shanks yellow; webbing grayish brown; undersides of first two fingers dusty white, of third and fourth fingers and of feet brown; anal stripe absent; small white flecks above and below anal opening. Color (in life) dark green on dorsal surfaces of head, body, and limbs; darker green reticulations on back; flanks dusty white with dark olive-gray 480 UnIvErRsITY OF KANSAS Pusts., Mus. Nat. Hist. Fic. 3. Holotype of Hyla charadricola (KU 58414). x 1.5. mottling; dark olive-gray stripe from nostril through eye and then to insertion of forelimb; upper lips pale green; inguinal region, anterior and posterior sur- faces of thighs dark yellowish orange; ventral surfaces of tarsi and shanks, and webbing of feet dusty yellow; belly white; iris silvery gold. Variation—Adult males have snout-vent lengths from 35.3 to 44.4 mm., and adult females from 43.4 to 50.9mm. No notable variation in structure is dis- played in the type series. In some individuals the dark reticulation on the dorsum is faint. Juveniles in life had dorsal colorations varying from rich brown with darker reticulations to pale green or gray with dark green reticula- FROGS OF THE HYLA BISTINCTA GROUP 481 tions. Some adults when collected were pale green with faint or no dorsal reticulations; later these individuals darkened. In all specimens the anal stripe is absent, and the flanks are heavily spotted. Comparisons.—Hyla charadricola differs from all other members of the Hyla bistincta group in the following combination of characters: truncate snout, green dorsum, and absence of a thoracic fold. From other hylids that occur in the same area, Hyla charadricola differs from Hyla miotympanum and H. arborescandens, both of which are green dorsally, by having a truncate snout and longer fingers with less webbing. Hyla eximia, though green, has brown dorsal spots, shorter fingers, and a round snout. Hyla robertsorum differs in having a round snout and brown dorsum, and Hyla taeniopus is much larger, has transverse bands on the limbs, and has extensive webbing between fingers. Remarks.—At the type locality, a shallow rocky stream in pine forest, Hyla charadricola was found beneath rocks at the edge of fast moving sections of the stream and beneath rocks in shallow rifles in the stream. Most of the frogs were in water. At night they were found sitting on rocks in the stream. Hyla miotympanum, which is abundant at the type locality, lives in bushes and beneath rocks along the stream but usually is not found in the riffles in- habited by Hyla charadricola. At Lago de Tejocotal Hyla charadri- cola was found beneath rocks near the shore of the lake and by a stream in the pine forest. Individuals were found on low vegetation over-hanging a small stream in pine-oak forest four kilometers south- west of Tianguistengo. Five recently metamorphosed young (KU 58424-9) found at the type locality on June 8, 1960, have snout-vent lengths of 22.4 to 24.0 (average 23.2)mm. The young are colored like the adults, except that in life the dorsum is a brighter green and the flanks are more yellow than tan and have less dark spotting than in adults. Distribution—Hyla charadricola inhabits streams in pine and_pine-oak forests at elevations of 2000 to 2300 meters in northern Puebla and eastern Hidalgo (Fig. 2). Specimens examined.—Hmatco: Lago de Tejocotal, 11 km. E Acaxochitlan, KU 58488, UMMZ 104032, 118165; 4 km. SW Tianguistengo, KU 53811-2. Puesia: 11.7 km. W Huachinango, UMMZ 121567 (5); Rio Totolapa, 14.4 km. W Huachinango, KU 53813-5, 55624, 58414-37, 59813 (skeleton), 59886 es MCZ 34964-5, UIMNH 50966, UMMZ 118166 (5), S-2242 (skele- ton). Hyla rebertsorum Taylor Hyla robertsorum Taylor, Univ. Kansas Sci. Bull., 26:393-396, figs. 5-6, November 27, 1940 [Holotype—CNHM 100124 (formerly EHT-HMS 16264) from El Chico Parque Nacional, Hidalgo, México; Mr. and Mrs. Radclyffe Roberts and Edward H. Taylor collectors]; Univ. Kansas Sci. Bull., 28:310, November 15, 1942. Taylor and Smith, Proc. U. S. Natl. Mus., 95:589, June 30, 1945. Smith and Taylor, Bull. U. S. Natl. Mus., 194:87, June 17, 1948; Univ. Kansas Sci. Bull., 33:333, March 20, 1950. Rabb and Mosimann, Occas. Papers Mus. Zool. Univ. Michigan, 563:1-9, March 29, 1955. Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 15:48, December 20, 1961. 482, UNIVERSITY OF KAnsAS Pusts., Mus. Nat. Hist. Diagnosis—Maximum snout-vent length in males, 48 mm.; snout in dorsal profile rounded; tarsal fold short and weak; inner metatarsal tubercle moderate in size and elliptical; outer metatarsal tubercle small, round, and indistinct; webbing extending to base of penultimate phalanx of fourth toe; nuptial spines on thumb small; weak thoracic fold present; anal opening above level of mid- dle of thighs; dorsum dark brown with dark reticulations; venter brown with cream-colored flecks; flanks brown with creamy white spots; anal stripe ab- sent; small white spots in anal region; vocal slits absent. Description—The following description is based on KU 57651 from El Chico Parque Nacional, Hidalgo. Adult male having a snout-vent length of 45.lmm.; tibia length, 22.1 mm., 49.0 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe), 21.5mm.; greatest width of head, 13.7 mm., 30.4 per cent of snout-vent length; head length, 12.6 mm., 27.9 per cent of snout-vent length; diameter of eye, 4.0mm.; diameter of tympanum, 1.8mm., 45 per cent of diameter of eye. Snout short, in lateral profile blunt, in dorsal profile round; canthus rounded; loreal region slightly concave; lips thick, round, and not flaring; nostrils slightly protuberant; internarial distance, 8.6 mm.; interorbital distance, 4.0 mm., slightly broader than width of eyelid, 3.5mm. A moder- ately heavy dermal fold from posterior corner of eye above tympanum and curving downward towards insertion of forearm; tympanum nearly round, covered by dermal fold above, its diameter slightly less than its distance from eye. Forearm moderately robust; distinct fold on wrist; prepollex much en- larged with patch of small nuptial spines continuous on side of digit; similar line of nuptial spines on inner edge of second finger; subarticular tubercles round, moderate in size, none bifid; supernumerary tubercles small and present only proximally; fingers long and slender; length of fingers from shortest to longest, 1-2-4-3; discs moderately large, that on third finger about size of tympanum; no web between first and second fingers; rudimentary web between other fingers. Legs robust; heels over-lap by about one-fourth length of shank when hind limbs adpressed; tibiotarsal articulation extending to posterior corner of eye; tarsal fold weak, extending to about middle of tarsus; inner metatarsal tubercle moderately large, flat, and elliptical; outer metatarsal tubercle small, round, and indistinct; subarticular tubercles round; super- numerary tubercles small, in single row on proximal segments of each digit; toes moderately long and slender; length of toes from shortest to longest, 1-2-5-3-4, the fifth nearly as long as third; toes nearly fully webbed; web extending to base of penultimate phalanx of fourth toe and to discs on other toes; discs small, about two-thirds size of those on fingers. Anal opening above middle of thighs; anal sheath short, deeply creased medially; heavy transverse dermal fold above anus; no large anal tubercles. Skin of all dorsal surfaces, chin, and ventral surfaces of limbs, except proximal parts of thighs, smooth; belly and proximal parts of thighs areolate; thoracic fold present, weak. Tongue elliptical, slightly longer than wide, not notched behind, and free posteriorly for about one-fourth of its length; vomerine teeth 3-8, situated on small, widely separated, transverse ridges between rather small elliptical inner nares; no vocal slits. Color (in alcohol) dark brown with irregular darker reticulations on dorsal surfaces of head, body, and limbs; flanks brown with small creamy white spots; posterior surfaces of thighs dark brown; chin creamy tan; belly grayish brown FROGS OF THE HyLa BISTINCTA GROUP 483 with cream-colored flecks; ventral surfaces of limbs pale brown; webbing on feet gray; small white spots in anal region. Color (in life) chocolate brown with darker brown reticulations and ir- regular blotches above; flanks brown with yellow spots; belly gray to grayish brown with faint cream-colored spots; iris a deep bronze color. Variation.—In males the total number of vomerine teeth varies from 4 to 7. In many specimens the vomerine ridges are larger and more closely ap- proximated medially than in the specimen described above. Females attain snout-vent lengths of 51 mm., have as many as 9 vomerine teeth, and have a proportionately larger tympanum than males. Some of the largest specimens of both sexes have indistinct cream-colored pustules scattered on the ventral surface of the forearm. Some individuals have nearly uniform grayish brown ventral surfaces; in others the chin, as well as the abdomen, is brown with cream-colored spots. The dorsal surfaces of some specimens are nearly uni- form dark brown with no reticulations. In others the dorsum is paler brown with distinct darker mottling; in some of these there is little mottling laterally, so that there is the effect of an irregular, pale brown, dorsolateral stripe. Tadpoles—The tadpoles of this species were described by Rabb and Mosi- mann (1955). Tadpoles obtained from streams at 3.3 kilometers north and at 8.5 kilometers southeast of Zacualtipan, Hidalgo, are like those described by Rabb and Mosimann in having 2/3 tooth rows, peglike serrations on the beaks, and long rounded tails. The largest tadpole (KU 60078) has small hind legs, a body length of 22 mm., and a total length of 61 mm. Remarks.—Taylor (1940:393) found frogs of this species in plants along small spring-fed rivulets in an open meadow at El Chico Parque Nacional. Also, he noted that active frogs dove into the streams and took refuge in the mud on the bottom. Rabb and Mosimann (1955:1) found this species along banks of tiny streams in open meadows and noted that the frogs sought refuge in the water. At El Chico Parque Nacional on June 8, 1960, I found Hyla robertsorum under rocks along small rivulets by day; at night, when the temperature was 14° C., frogs were sitting on rocks and in junipers overhanging a small stream. At the same locality on June 23 and 24, 1962, frogs of this species were found on rainy nights, when the temperature varied from 10 to 12° C. At this time the frogs were sitting on the grassy banks of rivulets in the meadow. During the day Hyla robertsorum was found on the earthen banks of the rivulets in places where dense growths of grass overhung the streams. On December 23, 1959, one specimen of Hyla robert- sorum was found beneath a rock in a small stream in pine forest at 3.3 kilometers north of Zacualtipan. Rabb and Mosimann (1955:1) obtained tadpoles of Hyla robert- sorum from quiet pools of a stream at El Chico Parque Nacional. I found tadpoles in pools in rocky streams in pine forest at 3.3 kilo- meters north and at 8.5 kilometers southeast of Zacualtipan. Four 484 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist. completely metamorphosed juveniles obtained on June 8, 1960, at E] Chico Parque Nacional have snout-vent lengths of 380.6 to 32.0mm. Gravid females were found at the same locality on June 8, 1960, and June 23, 1962. A H. crassa w H. pachyderma @ H. robertsorum 20° Kilometers 98° 96° Fic. 4. Map showing locality records for Hyla crassa, Hyla pachyderma, and Hyla robertsorum. FRoGS OF THE HyLA BISTINCTA GROUP 485 Distribution —Hyla robertsorum inhabits streams in the pine and fir forests in the higher parts (2250 to 3050 meters) of the Sierra Madre Oriental in extreme northern Puebla and eastern Hidalgo (Fig. 4). Specimens examined.—Hmatco: 16km. W Agua Blanca, UMMZ 106432 (6); El Chico Parque Nacional, CNHM 75786, 100124, KU 57650-71, 59824-5 (skeletons), 59914-5 (skeletons), 71269-95, 71757 (skeleton), UIMNH 10349- 64, 27022-35, 39434-49, UMMZ 92462, 106401 (5), 106443 (tadpoles), USNM 114762-85, 1384268; 3.38km. N Zacualtipan, KU 53810, 60078 (tadpoles); 8.5km. SE ZacualtipAan, KU 60079 (tadpoles). Purpta: Honey, UMMZ 95245. Hyla pachyderma Taylor Hyla pachyderma Taylor, Univ. Kansas Sci. Bull., 28:308-310, pl. 27, figs. 1-4, November 12, 1942 [Holotype-—USNM 115029 from Pan de Olla, Veracruz, south of Tezuitlan, Puebla, México; Hobart M. Smith collector]. Taylor and Smith, Proc. U. S. Natl. Mus., 95:588, June 80, 1945. Smith and Taylor, Bull. U. S. Natl. Mus., 194:86, June 17, 1948; Univ. Kansas Sci. Bull., 83:350, March 20, 1950. Rabb and Mosimann, Occ. Papers Mus. Zool. Univ. Michigan, 563:7-8, March 29, 1955. Diagnosis.—Maximum snout-vent length in males 40 mm.; snout in dorsal profile round; tarsal fold strong; inner metatarsal tubercle round and moderate in size; outer metatarsal tubercle small and indistinct; webbing on foot ex- tending to middle of penultimate phalanx of fourth toe; nuptial spines on thumb large; thoracic fold present; anal opening at level of middle of thighs; dorsum dull grayish brown with scattered indistinct dark flecks; venter cream- color mottled with brown on throat and chest; flanks grayish brown with cream-colored reticulations; anal stripe distinct, creamy white, sometimes ex- tending outward on thighs; white spots or line below anus; vocal slits absent. Description—The following description is based on USNM 115028 from Pan de Olla, Veracruz. Adult male having a snout-vent length of 39.9 mm.; tibia length, 21.0 mm., 52.6 per cent of snout-vent Jength; foot length (meas- ured from proximal edge of inner metatarsal tubercle to tip of longest toe), 20.5 mm.; greatest width of head, 12.8 mm., 32.1 per cent of snout-vent length; head length, 12.8 mm., 30.8 per cent of snout-vent length. Snout short, in lateral profile bluntly rounded, in dorsal profile rounded; canthus rounded; loreal region slightly concave; lips thick, round, and not flaring; nostrils slightly protuberant; internarial distance, 2.7 mm.; interorbital distance, 3.7 mm., somewhat broader than eyelid, 2.9mm. A heavy dermal fold from posterior corner of eye above tympanic region and then to insertion of forearm; tympanum completely concealed. Forearm moderately robust; distinct fold on wrist; prepollex enlarged bearing cluster of moderate-sized, horny, nuptial spines continuous on edge of digit; row of spines present on second finger; subarticular tubercles round, small proximally and slightly larger distally; supernumerary tubercles small and indistinct; three palmar tubercles, median and outer partly fused; fingers long, moderately slender; discs moderately large; length of fingers from shortest to longest, 1-2-4-8; second and fourth fingers subequal in length; webbing between fingers rudimentary. Heels over- lap by about one-fourth length of shank when hind limbs adpressed; tibiotarsal articulation extends to anterior edge of eye; tarsal fold thick, low, extending nearly to heel; inner metatarsal tubercle moderately large and round; outer metatarsal tubercle small and indistinct; subarticular tubercles small and round; 486 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. supernumerary tubercles small, present on proximal segments of digits; toes moderately long and slender; length of toes from shortest to longest, 1-2-3-5-4; third and fifth toes subequal in length; toes about three-fourths webbed; web extending to middle of penultimate phalanx of fourth toe; discs rather small, about two-thirds size of those on fingers. Anal opening directed posteroventrally at level of middle of thighs; anal flap slightly elongate; thick, transverse dermal fold above anus. Skin of dorsum and ventral surfaces of limbs, except thighs, smooth; skin of chin, belly, and ventral surfaces of thighs granular; thoracic fold present. Tongue nearly round, slightly notched behind, and barely free posteriorly; vomerine teeth 3-3, situated on posteroventral edges of small, transverse vomerine ridges between rather large triangular inner nares; no vocal slits. Color (in alcohol) of dorsal surfaces of head, body, and limbs dull grayish brown with indistinct scattered darker flecks; flanks grayish brown with cream- colored reticulations; posterior surfaces of thighs tan; chin cream-color, mottled with brown; belly creamy yellow; anal stripe cream-color. Variation—In addition to the specimen described above three others are known. One is a juvenile having a snout-vent length of 29.5mm., and two are females having snout-vent lengths of 46.9 and 41.6mm. Variation in structure and coloration between the four specimens is slight. In the females the tympani are partly visible and are about one-third the diameter of the eye; the chest is mottled with brown; the anal stripe extends laterally in the form of a row of cream-colored dashes and spots onto the posterodorsal sur- faces of the thighs. Remarks.—On the basis of the four specimens available for study, Hyla pachyderma seems to be closely related to Hyla crassa and Hyla robertsorum. In the Hyla bistincta group, Hyla pachyderma is unique in having enlarged nuptial spines. Taylor and Smith (1945:588) stated that the frogs were found on bushes and weeds beside a small, bounding stream near Pan de Olla. I have searched unsuccessfully for this species in the area around Pan de Olla and Tezuitlan. Distribution.—This species is known only from a stream at an elevation of about 1600 meters on the Atlantic slopes of the Sierra Madre Oriental in cen- tral Veracruz (Fig. 4). Specimens examined.—VeEracruz: Pan de Olla, south of Tezuitlan, Puebla, USNM 115026-9. Hyla crassa (Brocchi) Cauphias crassus Brocchi, Bull. Soc. Philom. Paris, ser. 7, 1:180, 1877 [Holotype—MNHN 6331 from “Mexico;” Adolphe Boucard collector]. Cauphias crassum Brocchi, Etudes des batraciens de Amérique Centrale, p. 64, pl. 12, fig. 4, 1882. Diaz de Leén, Indice de los batracios que se encuentran en la Republica Mexicana, p. 21, June, 1904. Kellogg, Bull. U. S. Nat. Mus., 160:118-120, March 81, 1932. Taylor, Univ. Kan- sas Sci. Bull., 26:392, November 15, 1940. Rabb and Mosimann, Occas. Papers Mus. Zool. Univ. Michigan, 563:7, March 29, 1955. Hyla crassa, Boulenger, Catalogue Batrachia Salientia, 2nd. Ed., p. 896, February 1, 1882. Giinther, Biologia Centrali-Americana, Reptilia and Batrachia, p. 281, September, 1901. Nieden, Das Tierreich, Amphibia, FROGS OF THE HYLA BISTINCTA GROUP 487 Anura I, p. 248, June, 1923. Smith and Taylor, Bull. U. S. Natl. Mus., eee June 17, 1948. Taylor, Amer. Mus. Novitates, 1437:20, Decem- er 7, 1949. Hypsiboas crassus, Cope, Bull. U. S. Natl. Mus., 32:14, 1887. Hyla robustofemora Taylor, Univ. Kansas Sci. Bull., 26:389-893, figs. 3-4, November 27, 1940 [Holotype-—UIMNH 25050 (formerly EHT-HMS 16314) from Cerro San Felipe, 15 kilometers northeast of Oaxaca, Oaxaca, México; Edward H. Taylor collector]; Univ. Kansas Sci. Bull., 28:310, November 15, 1942. Smith and Taylor, Bull. U. S. Natl. Mus., 194:86, June 17, 1948. Taylor, Amer. Mus. Novitates, 1437:20, December 7, 1949. Smith and Taylor, Univ. Kansas Sci. Bull., 33:389, March 20, 1950. Rabb and Mosimann, Occas. Papers Mus. Zool. Univ. Michigan, 568:7, March 29, 1955. Plectrohyla crassa, Hartweg, Occas. Papers Mus. Zool. Univ. Michigan, 437:1, June 80, 1941. Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 455:6, January 5, 1942. Diagnosis —Maximum snout-vent length in males 54 mm.; snout in dorsal profile round; tarsal fold strong; inner metatarsal tubercle small and elliptical; outer metatarsal tubercle small, flat, and indistinct; foot fully webbed; nuptial spines on thumb small; thoracic fold absent; anal opening at level of middle of femur; dorsum dull olive-green; belly creamy yellow; chin gray with yellow flecks; flanks dull olive-green with scattered cream-colored spots; and stripe faint, cream-color; vocal slits absent. Description—tThe following description is based on UIMNH 25050 from Cerro San Felipe, Oaxaca. Adult male having a snout-vent length of 53.7 mm.; tibia length, 26.9 mm., 50.1 per cent of snout-vent length; foot length (meas- ured from proximal edge of inner metatarsal tubercle to tip of longest toe), 25.4 mm.; greatest width of head, 17.6 mm., 32.8 per cent of snout-vent length; head length, 16.0 mm., 29.8 per cent of snout-vent length; diameter of eye, 5.4mm.; diameter of tympanum, 1.5mm., 27.8 per cent of diameter of eye. Snout short, in lateral profile bluntly rounded, in dorsal profile broadly round; canthus absent; loreal region nearly flat; lips thick and not flaring; nostrils barely protuberant; internarial distance, 3.8 mm.; interorbital distance, 4.7 mm., somewhat broader than width of eyelid, 3.8mm. Heavy dermal fold from posterior corner of eye above tympanum and then to insertion of forearm; tympanum concealed above, its diameter about equal to its distance from eye. Forearm thick; distinct fold on wrist; prepollex enlarged bearing patch of small nuptial spines continuous on side of digit; similar patch on second finger; subarticular tubercles small and round, none bifid; few supernumerary tubercles on proximal segments of digits; large, flat palmar tubercle; fingers long and slender; length of fingers from shortest to longest, 1-2-4-3; discs moderately large; rudimentary web between second and third fingers and between third and fourth. Legs thick; heels overlap by about one-fourth length of shank when hindlimbs adpressed; tibiotarsal articulation extends to posterior corner of eye; tarsal fold thick, extending to heel; inner metatarsal tubercle small and elliptical; outer metatarsal tubercle small, flat, and indistinct; subarticular tubercles small and round; single row of supernumerary tubercles on proximal segments of each digit; toes moderately short and slender; length of toes from shortest to longest, 1-2-3-5-4; toes fully webbed; flap of skin on inner surface of first toe; discs about same size as those on fingers. Anal opening directed posteroventrally at middle of thighs; anal sheath moderately elongate; small tubercles below anal opening. Skin of dorsum rather smooth, somewhat 488 UNIVERSITY OF KANSAS PuB3s., Mus. Nat. Hist. granular on dorsal surfaces of limbs; skin of chin and belly moderately granular; that of posterior surfaces of thighs smooth; no thoracic fold. Tongue nearly round, shallowly notched posteriorly, and free for about one-fourth its length; vomerine teeth 5-5, situated on rounded ridges between small inner nares; no vocal slits. Color (in alcohol) dull olive-green on dorsal surfaces of head, body, and limbs; flanks dull olive-green with scattered cream-colored spots; posterior sur- faces of thighs grayish brown with faint creamy mottling; chin gray with cream- colored spots; belly creamy yellow, suffused with gray posteriorly; undersides of feet and webbing gray; anal stripe faint, pale cream-color. Variation —The only other known specimen (MNHN 6831) is a female having a snout-vent length of 53.7 mm. and resembling the specimen described above in most details of morphology. In MNHN 6331 the tympanum is com- pletely concealed, and the 8-7 vomerine teeth are arranged in two irregular rows. The female has more cream-colored mottling on the flanks and posterior surfaces of the thighs and more distinct mottling on the throat than the male described above. Remarks.—The systematic status of Cauphias crassus Brocchi has been in doubt since the time of the original description. Brocchi (1877:130) stated: “Les dernieres phalanges sont obtuses, tronqués a leur extrémité antérieure.” Brocchi placed the species in his genus Cauphias (type species, C. guatemalensis), which he considered to be related to Hylodes ( = Eleutherodactylus in the sense used by Brocchi); he thereby placed Cauphias in his Hylodidae ( = Lep- todactylidae, in part). This idea of relationships was perpetuated by Barbour (1927:96), who reported on the second known speci- men of Cauphias guatemalensis and stated: “When I dissected the sternum I was at once struck by its similarity to Noble’s figures of transitional types between arciferal and firmisternal forms. The Cauphias sternum recalls some of his figures for Sminthillus and Eleutherodactylus. This genus is probably most closely related to the latter and has probably become highly modified to meet some peculiar environmental condition or on account of some specialized habits as yet unknown.” Kellogg (1932:118) placed Cauphias in the Leptodactylidae and stated that the terminal phalanges are T-shaped. Hartweg (1941:1) considered Plectrohyla to be the cor- rect generic name for Cauphias guatemalensis; he thereby relegated Cauphias to the synonomy of Plectrohyla, Hartweg (1941:9) further showed that the terminal phalanges of Plectrohyla guatemalensis were not T-shaped and that intercalary cartilages were present. Thus, he correctly concluded that Plectrohyla guatemalensis (and P. crassa by implication) was a member of the family Hylidae. Stuart (1942:6) followed Hartweg’s allocations and further sug- gested that Plectrohyla crassa might be the same species as Hyla FRoGs OF THE HyLa BISTINCTA GROUP 489 robustofemora Taylor. In his description of H. robustofemora Taylor (1940:392), who had not examined the type of Cauphias crassus, stated that were it not for the statements of Brocchi and Kellogg that C. crassus has T-shaped terminal phalanges, “I might suspect I had before me a specimen of Cauphias closely related to crassum.” I have compared the type of Cauphias crassus with that of Hyla robustofemora. With the exception of the minor differences men- tioned in the preceding section on variation, the specimens are alike, leaving little doubt that they represent the same species. The statements of Brocchi and Kellogg to the contrary, the type of Cauphias crassus possesses intercalary cartilages between the pe- nultimate and terminal phalanges; the latter are not T-shaped, but as in the type of Hyla robustofemora, resemble those typical of Hyla. On the basis of the morphological characters, as pointed out for Hyla robustofemora by Taylor (1940:392), Hyla crassa is a member of the Hyla bistincta group. Distribution.—This species is definitely known only from a small stream at an elevation of 2300 meters in the mountains of central Oaxaca (Fig. 4). Specimens examined.—Oaxaca: Cerro San Felipe, UIMNH 25050. “Mex- ico,” MNHN 6331. RELATIONSHIPS The evolutionary trend in the members of the Hyla bistincta group is towards aquatic habits. Hyla bistincta, the least special- ized species in the group, has relatively short fingers, webbing be- tween the fingers, a truncate, high snout, and relatively large sub- articular and supernumerary tubercles. Hyla charadricola resembles bistincta in having relatively short fingers, a slight amount of web- bing, and a truncate snout. Apparently these two species are more closely related to one another than either is to the other species in the group. Hyla robertsorum, pachyderma, and crassa are the most aquatic members of the group. These species are closely related, possibly conspecific. All have round, sloping snouts, robust fore- arms, long, unwebbed fingers, and large webbed feet. Both H. pachyderma and H. crassa seem to be advanced beyond H. roberts- orum. If small nuptial spines, moderately webbed feet, and ab- sence of a well-defined thoracic fold are considered to be less ad- vanced than large nuptial spines and a strong thoracic fold, as in H. pachyderma, or fully webbed feet, as in H. crassa, then H. robertsorum must be considered to be less advanced than H. pachyderma or H. crassa. Members of the Hyla bistincta group inhabit mountain streams. 490 UNIvERSITY OF KAnsas Pusts., Mus. Nat. Hist. The frogs can be found along these streams throughout the year. Since in most stream-breeding hylids there is no migration to breed- ing sites, the breeding call does not function to attract females to the breeding site. Apparently voices are lacking in all members of the Hyla bistincta group, except in Hyla bistincta. The presence of vocal slits and the ability to call further indicate that Hyla bistincta is the primitive member of this group. Members of the Hyla bistincta group and the species of Plectro- hyla closely resemble each other in osteology and body form of the adults and in structure of the tadpoles. This resemblance suggests a close relationship between the two groups. Plectrohyla ap- parently evolved from an ancestral stock resembling the extant Hyla bistincta. Probably this stock gave rise independently to Plectrohyla and to the Hyla robertsorum-pachyderma-crassa com- plex. In the former the voice was retained, and a projecting prepollex spine developed, whereas in the latter the voice was lost, and the prepollex spine did not project. Plectrohyla lives in moun- tain streams in the Chiapan-Guatemalan highlands; the Hyla robert- sorum-pachyderma-crassa complex inhabits similar environments in the Sierra Madre Oriental in México. Hyla charadricola also lives in the Sierra Madre Oriental, whereas Hyla bistincta is widespread in the mountains of México southeastward to the Isthmus of Tehuantepec. FROGS OF THE HyLa BISTINCTA GROUP 491 LITERATURE CITED BARBOUR, T. 1927. Cauphias rediscovered. Copeia, no. 165:96-98, December 23. Broccut, P. 1877. Note sur quelques batraciens hylaeformes recueillis au Mexique et au Guatemala. Bull. Soc. Philom. Paris, ser. 7, 1 (3):122-182. DuELLMAN, W. E. 1961. The amphibians and reptiles of Michoacan, México. Univ. Kansas Publ. Mus. Nat. Hist., 15:1-148, pls. 1-6, December 20. Hartwec, N. 1941. Notes on the genus Plectrohyla, with descriptions of new species. Occas. Papers Mus. Zool. Univ. Michigan, 437:1-10, pl. 1, June 30. KEtLoce, R. 1932. Mexican tailless amphibians in the United States National Museum. Bull. U. S. Natl. Mus., 160:iv + 224 pp., pl. 1, March 31. Rasp, G. B. and Mosmanv, J. E. 1955. The tadpole of Hyla robertsorum, with comments on the affinities of the species. Occas. Papers Mus. Zool. Univ. Michigan, 563:1-9, March 29. SHANNON, F. A. 1951. Notes on a herpetological collection from Oaxaca and other locali- ties in Mexico. Proc. U. S. Natl. Mus., 101:465-484, May 17. Stuart, L. C. 1942. Descriptions of two new species of Plectrohyla Brocchi, with com- ments on several forms of tadpoles. Occas. Papers Mus. Zool. Univ. Michigan, 455:1-14, January 5. Tay.Lor, E. H. 1940. New species of Mexican Anura. Univ. Kansas Sci. Bull., 41:385- 405, November 27. 1948. A new hylid frog from eastern Mexico. Univ. Kansas Publ. Mus. Nat. Hist., 1:259-262, August 16. 1949. A new hylid frog from central Veracruz. Copeia, 1949 (4):272- 274, pl. 1, December 15. Tayior, E. H. and Smrrn, H. M. 1945. Summary of the collections of amphibians made in México under the Walter Rathbone Bacon Traveling Scholarship. Proc. U. S. Natl. Mus., 95:521-618, pls. 18-32, June 30. Transmitted June 24, 19683. 29-8590 So NPT BR fata ce] UNIVERSITY OF KANSAS PUBLICATIONS MusEuM OF NATURAL History MUS.\aGN Volume 15, No. 10, pp. 493-364, pls, 28-25, 14 fies any May 17,1965 ——p--—___— HA HVAT ry UNIVERSiTy An Ecological Study of the Garter Snake, Thamnophis sirtalis BY HENRY S. FITCH UNIVERSITY OF KANSAS LAWRENCE \ 1965 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Institutional libraries interested in publications exchange may obtain this series by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas. Copies for individuals, persons working in a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of. Kansas, Lawrence, Kansas. There is no provision for sale of this series by the University Library, which meets institutional requests, or by the Museum of Natural History, which meets the requests of individuals. Nevertheless, when individuals request copies from the Museum, 25. cents should be included, for each separate number that is 100 pages or more in length, for the purpose of defraying the costs of wrapping and mailing. * An asterisk designates those numbers of which the Museum’s supply (not the Library’s supply) is exhausted. Numbers published to date, in this series, are as follows: Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950. ®Vol. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text. Nos. 1-4 and index. Pp. 1-681. 1951. fi (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 figures in text. December 27, 1951. Vol. Nos. 1-87 and index. Pp. 1-676, 1951-1953. 2 Vol. 3 4 5. ®Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. or 8 9 1 » ®Vol. Durrant. Pp. 1-549; 91 figures in text, 30 tables. August 10, 1952. Nos. 1-15 and index., Pp. 1-651, 1952-1955. Nos. 1-10 and index. Pp. 1-675, 1954-1956. Vol. Vol. Vol. Vol. Nos. 1-23 and index. Pp. 1-690, 1955-1960. 0. 1. Studies of birds killed in’ nocturnal migration. By Harrison B. Tordoff and Hope M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 2. Comparative breeding behavior of Ammospiza caudacuta and A. maritima. ~ By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956. 8. The forest habitat of the University of Kansas Natural History Reservation. By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures in text, 4 tables. December $1, 1956. 4, Aspects of reproduction and development in the prairie vole (Microtus ochro- gaster).. By Henry S. Fitch. Pp. 129-161, 8 figures in text, 4 tables. De- cember 19, 1957. 5. Birds found on the Arctic slope of northern Alaska. By James W. Bee. Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958. ®6. The wood rats of Colorado: distribution and ecology. By Robert B. Finley, Jr. Pp. 213-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958. Home ranges and movements of the eastern cottontail in Kansas. By Donald W. Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, 1959. Natural history of the salamander, Aneides hardyi. By Richard F, Johnston and Gerhard A. Schad. Pp. 573-585. October 8, 1959. f . A new subspecies of lizard, Cnemidophorus*sacki, from Michoac4n, México. By William E. Duellman. Pp. 587-598, 2 figures in text. May 2, 1960. 10. A taxonomic study of the middle American snake, Pituophis deppei. By William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960. Index. Pp. 611-626. Vol.. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960. Vol. 12. 1. Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. July 8, 1959. *2. The ancestry of modern Amphibia: a review of the evidence. By Theodore H. Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, 1959, ; 8. The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49 figures in text... February 19, 1960. *4, A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By Theodore H. Eaton, Jr., and Peggy Lou-Stewart. Pp. 217-240, 12 figures in text. May 2, 1960. Natural history of the bell vireo. By Jon C. Barlow. Pp. 241-296, 6 figures in text. March 7, 1962. : Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C. Fox. Pp. 297-307, 6 figures in-text. May 21, 1962. h Vertebrates from the barrier island of Tamaulipas, México. By Robert K. Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 809- 345, plates 5-8. June 18; 1962. 8. Teeth of Edestid sharks. By Theodore H. Eaton, Jr. Pp. 847-362, 10 fig- ures in text. October 1, 1962. (Continued on inside of back cover) UNIVERSITY OF KANSAS PUBLICATIONS MuSEUM OF NATURAL HIsToRY Volume 15, No. 10, pp. 493-564, pls. 23-25, 14 figs. May 17, 1965 An Ecological Study of the Garter Snake, Thamnophis sirtalis BY, HENRY S. FITCH UNIVERSITY OF KANSAS LAWRENCE 1965 ¥ lf f FN I , yn f € i i UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HIsTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 15, No. 10, pp. 493-564, pls. 23-25, 14 figs. Published May 17, 1965 MUS. COM. -OOL LIBRARY DEC 3i 1y02 HARVARD UNIVERSITY UNVERSITY OF KANSAS Lawrence, Kansas PRINTED BY HARRY (BUD) TIMBERLAKE, STATE PRINTER TOPEKA, KANSAS An Ecological Study of the Garter Snake, Thamnophis sirtalis BY HENRY S. FITCH CONTENTS EN RRODEGELON ere ieee ee GO Ts Oe ee 495 DESCRIPTION OF Sarees fe. eos eMart e. OR 28, yin Aen eee 497 @rOCRAPHIG: V-AREA TION 4h tee fe se, on nce oe ae 501 ELARTEAT St) eR are ee oe ck OTRO ee ce eRe eee 505 iE MPERATURE) A\nEA TIONSHIPS’ 2.0... 0... aeons aloes eee ee 508 OODIELABTITS StU Piet henna A) ieee nn ys oe ee 513 IGREEDING 4 8 eee ee ee ee rn ee a er ae ce en ee 518 Number*of-young mm broodt...0 272 5.2. S2 a0. ee 521 dhiming-ofbreedingveycle ta... 0. wk kes. oe ee 526 GROWER ee ee hak cle ae pean eee 527 INIOVENENTS 8 Sar pee he ithe te Ce ee oe a ee ees 536 DDEFENSESAND? ESCAPE) (een AG i ee ee eee 540 INATURATES EGNEEIVIIEG) +e eS te 2 eg ees en oe Mee eee ee ee 544 PAR ASETES Me os, oo ee ee ORCA Sy oe ce eee eee 546 @OMPOSIEION OF THE POPULATION 2.5050. c5 ncn eee 548 INDREBERS A057 ia op ee, oe ORD lh” SE nn ee eet oe eee bol SUNENER ROT ia lige, So ll ae AO | aaa Rene eee 555 ETPERATORES CECED™ «221k et Oe ee ee eee 560 INTRODUCTION Over much of North America the most abundant and conspicuous species of snake is the garter snake, Thamnophis sirtalis. It attains remarkably high population densities and obviously it figures sig- nificantly in the economy of local ecosystems. Hence, it has im- portant economic bearing. Occasionally it preys upon pest species, (495 ) 496 UNIVERSITY OF KANsAS Puszs., Mus. Nat. Hist. but the bulk of its predation is upon earthworms and amphibians that certainly are not harmful to man’s interest and sometimes are highly beneficial to agriculture. The present account does not attempt to offer a final judgment as to the species’ economic role; obviously this varies according to the time and place. In my study no corroboratory evidence could be obtained of the oft-repeated statement that garter snakes feed on insects. Over the past 30 years I have collected and studied this garter snake in most of the states in the western half of the United States. From 1948 through 1963 I studied a local population on the Uni- versity of Kansas Natural History Reservation. In 1959 this study was intensified and extended to the adjacent Rockefeller Tract (hereinafter meant to be included whenever the Reservation is mentioned ), while concurrently similar field work was carried on at Harvey County Park, some 140 miles southwest of the Reservation. This field study of the garter snake was based primarily upon live-trapping. The traps used were wire cylinders of quarter-inch mesh, with entrance funnels having inward-projecting wire prongs. The first-year snakes were small enough to pass easily through the quarter-inch mesh; hence they were poorly represented in the catch. Traps of finer mesh were tested but were on the whole unsatis- factory because the more flexible wire permitted ingress of predators more readily, and because a variety of arthropods, some of kinds harmful to the snakes, were caught with them. By November 5, 1963, 1097 individual garter snakes had been captured a total of 1354 times on the Reservation, and 661 snakes of this species had been captured 812 times at Harvey County Park and neighboring areas. My conclusions are based primarily on the records of the recaptured snakes, for which growth, movement, and other information was available. However, it was not possible to achieve a marked population, and even in the later stages of the study the majority of records pertained to new individuals caught for the first time. For the last six years of the study at the Reserva- tion, the percentages of recapture records were as follows: 1958— 19 per cent; 1959—27 per cent; 1960—27 per cent; 1961—27 per cent; 1962—21 per cent; 1963—26 per cent. The rapid population turnover from year to year prevented obtaining a higher percentage of marked individuals. At the Reservation T. sirtalis was the only kind of garter snake present; although the locality is within the ranges of both T. radix and T. proximus, both are rare locally. On the Harvey County EcoLocIcaL Stupy oF GARTER SNAKE 497 study area T. radix also was present, but in relatively small numbers, and the findings concerning it have not been incorporated in the present report. The study of Thamnophis sirtalis herein reported upon was one phase of an ecological and life history study of Kansas snakes, which was supported by three successive grants from the National Science Foundation (G-3413, G-6158, G-16104) from 1957 through 1962. Many persons helped in various ways with the field work. In almost all instances these were the same individuals whose assistance has already been acknowledged in earlier papers con- cerning snakes of the Reservation (Fitch, 1960:92; 1963b:357). Mr. Dwight R. Platt is due special credit for his supervision of the field work at Harvey County Park, and active participation in it in 1959 through 1963. Mr. Richard S. Funk helped with the maintenance and study of a colony of captive garter snakes in the winter of 1963 and 1964. Dr, William R. Murchie kindly identified several earth- worms. Dr. William H. Coil and Mr. Willis A. Reid kindly identi- fied flukes found in the mouths of garter snakes in 1963 and pro- vided information concerning them. My wife, Virginia R. Fitch, read the manuscript critically and helped me in various other ways. My son, John H. Fitch, assisted me with the photography of living garter snakes. DESCRIPTION The scalation is that of a typical colubrid. The body scales are in 19 rows, of which those in the lowest row on each side are broadly oval and smooth or only weakly keeled, whereas those of the remaining rows are narrower and strongly keeled. At a point a little more than 61 per cent of the distance from the snout to the vent the scales of the fourth row on each side abruptly become smaller and then disappear altogether. Immediately beyond the point where the fourth row disappears scales of the adjacent third and fifth rows are somewhat enlarged to occupy the space of the missing fourth row. The remaining 17 rows extend to the posterior end of the body, but drop out in rapid succession on the tail. The cephalic scutellation is typical of colubrids in general, of natricines, and of the genus Thamnophis (Plate 23, fig. 1). The rostral plate is large and prominent. The scutes of the top of the head are: the internasals, the pre- frontals, the frontal (bordered on each side by a supraocular), and the paired parietals. The naris is situated between two nasals, near their upper edge. The posterior nasal is somewhat larger than the anterior. The loreal is rela- tively large. The eye is bordered anteriorly by a preocular and behind usually by three (sometimes only two) small postoculars, which decrease in size from the uppermost to the lowest. There is a single anterior temporal between the 498 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. parietal (above) and the fifth and sixth supralabials (below). Behind the anterior temporal there are usually two posterior temporals, one above the other, but the number may be increased to three or reduced to one. There are seven supralabials, increasing progressively in size from the anteriormost to the fifth then decreasing again. The third and fourth are in contact with the lower edge of the eye. The second supralabial is relatively elongate and is the homologue of both the second and the third in the many kinds of garter snakes that have eight pairs of supralabials. There are most often 10 pairs of infralabials. The first pair of infralabials is small but the second pair is even smaller. From the second to the fifth there is progressive increase in size and from the fifth to the tenth there is progressive decrease. The genials are in two pairs, anterior and posterior. Those of the posterior pair are markedly longer (usually from 20 to 40 per cent). The anal plate is undivided. The subcaudals are normally in two series—left and right—alternately placed. Each subcaudal is broadly hexagonal. The body scales are notched at the tip. In the adult male, scales of the chin region, the anterior genials and several anterior pairs of infralabials, have sensory pits which function during courtship, as the male presses his chin against the female’s back. Also, in the adult male, several scales near the anal plate on each side have their keels especially heavy, with knobs developed on them. These knobbed keels likewise function in courtship. The color pattern is, of course, subject to much geographic variation over the range as a whole. In some localities it is remarkably stable but in others including those of my study areas, it is subject to a wide range of individual variation. The dorsal stripe is the most prominent feature of the pattern. It begins on the nape and extends for the full length of the body and tail, oc- cupying the mid-dorsal scale row, and half (or a little more) of each adjacent row. It is buffy yellow, contrasting with the dark dorsolateral area on each side. These dorsolateral areas are almost black in some individuals but in others dark brown or olive, sufficiently pale so that a pattern of two alternating rows of black spots is faintly discernible. Also, there are much paler areas, which are confined chiefly to the skin between the scales, hence scarcely no- ticeable except when the skin is stretched. These marks are much more prominent on the lower part of each dorsolateral area, and they become smaller farther dorsally. The marks tend to be bluntly crescent-shaped, but the bulge of one crescent may contact that of the crescent above or below it, the two forming an H-shaped mark. In about ten per cent of the snakes examined, the pale markings were all yellowish or faintly suffused with bluish or greenish, but in most of the snakes at least the lower markings—those between rows 3 and 4, 4 and 5, and 5 and 6—were red, salmon, pink, or rusty. The less prominent marks between rows 6 and 7, 7 and 8, and 8 and 9 usually do not have well-defined reddish suffusion, and if they do, it is paler and more diffuse than in the lower markings. The lateral stripe occupies approximately the upper half of the second scale row and the lower half of the third. It is dull yellow, suffused with dusky pigment, and lacks well-defined borders, blending into the duskier areas above and below it. Below the lateral stripe, a darker area, partly suffused with dusky pigment, and also having scattered black spots, includes the lower half of the second scale row, all of the first row, and the adjacent tips of the ventral scutes. The ventral surface is white in the chin ECOLOGICAL STUDY OF GARTER SNAKE 499 region, but becomes increasingly grayish farther posteriorly, with a faint suffu- sion of greenish blue. In occasional individuals with erythristic tendencies, the red markings on the lower part of the dorsolateral area blend into the lateral stripe suffusing it with reddish, although it has no well defined markings. Reddish suffusion may even extend to the edge of the ventral surface. In broods of garter snakes that were born in captivity, the red lateral markings were much duller than in the mothers and were reddish tan or dull rusty, remarkably uniform among the members of a brood. Seemingly, the brighter, adult coloration is acquired early in development, well before sexual maturity. There is no noticeable difference between the sexes, nor between adults and partly grown individuals, either in pattern or in intensity of color. There is little evidence of geographic variation in size in Thamnophis sirtalis, but as yet no investigation has been made to determine whether such variation exists. Table 1 shows snout-vent lengths and weights of individuals of various categories. Because of allometric growth and slight sexual dimorphism, individual garter snakes, even from the same locality, differ considerably in proportions. Some of the most striking differences involve the proportions of the head, and the relative size of the head as compared with the body. The changes are gradual, but for the purpose of discussion four arbitrary ontogenetic stages may be recognized. First, in the late embryo, as figured by Zehr (1962:326 and 327) the head is relatively enormous and is bent at right angles to the body, the forehead bulges, and the eyes are remarkably large and protruding; second, in the newborn young the head is aligned with the body but is still relatively large causing the eyes to be conspicuous in profile in dorsal view, the rostrum is fully as wide as the posterior part of the head, and the head is conspicuously wider than any part of the body; third, in partly grown individuals or small adults, the head is conspicuously wider than the neck but not so wide as the TABLE 1.—SrzEs OF THAMNOPHIS SIRTALIS PARIETALIS FROM RESERVATION AND Harvey County PARK Adult females Adult males Newborn young Snout- Snout- Snout- vent | Weight | vent | Weight | vent | Weight (milli- | (grams) | (milli- | (grams) | (milli- | (grams) meters) meters) meters) Maxamumls: seas erte 950 410 678 110 191 25) Average (or typical)* 550 57 455 31.5 168 1.9 Minimum, a s.e2oeo- 504 21 387 14 134 sft ® For adult males and adult females the “typical” size shown is for two-year-olds, since these make up the most numerous size groups. 500 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. TABLE 2.—TAIL-LENGTH AS A PERCENTAGE OF SNOUT-VENT LENGTH IN MALE AND FEMALE THAMNOPHIS SIRTALIS PARIETALIS OF DIFFERENT SIZE GROUPS (FROM RESERVATION ) Tail-length: Size group: snout-vent length ra ro) snout-vent Up to 201 to 401 to 601 to | More than length 200 mm. | 400 mm. | 600 mm. | 800 mm. | 800 mm. of oN Ke Qo oS ot al aot Q 7 Ne ee ea a Mee ee Nd aa tT aes i: PA Aran a pee Wace oY Avil A 4 i Gite itoears ogee a LO sees 9 4 P(E Pay) Nee 7 os a | Pastel |e 40 3 28a obicss Gul 2 4) 5 44] 1 34 2 DOG + 7 Z 5 8 45]... 28 2 30) soe sees 4 a 2 8) 46 2 14 1 Slicisecaccce i 18 2 7 | 36 21 3 10 1 Oo see or 20 10} 3 5 | 38 16.) 3 3 1 Solna eter. 19 4] 11 6 | 67 3 2 1 Be SA eveds cht eo 11 1 6 3 | 52 4 1 Es BON saciek a 6 1 is fade 1 Oras steve 5 1 3 . | 25 < Dice sade 1 9 widest part of the body, the eyes, though large as compared with those of most other kinds of snakes, are only slightly noticeable in profile from dorsal view, and the proportions of the head are “normal”; fourth, in old adult fe- males, which attain by far the largest size, and in which allometric growth progresses farthest, the eye is relatively small, and does not appear in profile from dorsal view at all, or is not prominent, the temporal segment of the head is relatively wide as compared with the rostral portion causing the head to ap- pear to be triangular, the neck is thick, and the body is stout. In newborn males the tail averages approximately one-third of the snout- vent length, whereas it is approximately 31 per cent of the snout-vent length in newborn females. In adult males the relative length of the tail has changed slightly if at all, still averaging about one-third of the snout-vent length, whereas in females of corresponding size (400 to 600 millimeters) the tail averages only about 29 per cent of snout-vent length, and in larger females (more than 600 millimeters) it averages only a little more than 27 per cent. Large old females thus deviate from the bodily proportions of juveniles much more markedly than do other individuals (see Table 2). In Table 1 the snout-vent lengths and weights given as maximum (and those given as minimum) do not necessarily represent the same individual. For each length there is a wide range of variation in weights, and as length increases, the variability increases. Females are more variable than males, also, they attain greater lengths, and their weights are subject to more fluc- tuation being unusually high when the snakes are gravid and unusually low after parturition. The graph in Figure 1 shows correlation of weight with length in male and female garter snakes all from the Reservation and Rocke- feller Tract. ECOLOGICAL STuDY OF GARTER SNAKE 501 WEIGHT IN GRAMS 200 300 400 500 600 700 800 900 SNOUT-VENT LENGTH IN MILLIMETERS Fic. 1. Diagram showing correlation of weight with snout-vent length in live garter snakes on the Reservation. For each sample of 25-millimeter range in snout-vent length, the mean, minimum and maximum weights are shown. Standard error of the mean is shown for each series having a sufficient number of individuals. Males and females are combined in the samples shown here. The trends of correlation in length and weight are alike for the sexes, except that adult females tend to vary more in weight and, in early summer, to be slightly heavier than adult males of the same length, because of their develop- ing embryos. GEOGRAPHIC VARIATION Thamnophis sirtalis is a polytypic species with an enormous geo- graphic range (Fig. 2) which exceeds in area that of any other North American snake, extending from coast to coast and from the northernmost limits of areas habitable by snakes in the western hemisphere south to subtropical regions. The species as a whole has not been the subject of taxonomic revision since Ruthven’s 502 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. (1908) classic but much outdated monograph of the genus Tham- nophis. Eleven subspecies are currently recognized. Some of these are rather weakly differentiated, yet probably still others worthy of recognition would be discerned if all geographic populations were subjected to careful scrutiny. Among North American species of snakes only the Thamnophis elegans, Diadophis punctatus, and Lampropeltis doliata complexes exceed Thamnophis sirtalis in amount of racial differentiation, but in the first two of these some of the forms have become differentiated beyond the subspecific level and genetically isolated. Thamnophis sirtalis sirtalis is the best known subspecies and ranges over an extensive area, from New York and Florida west to Minnesota and Louisiana, with a considerable amount of variation, both individual and local. The dorsal stripe typically is yellow, well developed but with edges not sharply defined, and with dark spots encroaching. There are lateral stripes, less well defined than the dorsal, and the dorsolateral areas on each side are dark but with even darker areas in two alternating rows of spots. The numbers of ventrals and subcaudals are relatively low as compared with those in most of the other subspecies. Some of the northern popu- lations have a tendency to melanism. T. s. pallidula is a north- eastern race occurring entirely within recently glaciated regions in Maine, New Hampshire, Vermont, northern New York, and neigh- boring regions of southeastern Canada. In this subspecies the dorsal stripe is either completely lacking or is dull colored, grayish, and indistinct. Dorsolateral dark spots are conspicuous. Ventrals and subcaudals both average approximately seven or eight fewer than in neighboring populations of T. s. sirtalis. TL. s. semifasciata is a localized subspecies occurring in typical form only on or near the shores of Lake Michigan in the Chicago region. It differs from T. s. sirtalis which presumably is its parent subspecies in having the lateral stripes broken by a series of dark transverse markings. T. s. parietalis is the subspecies of the Great Plains. It differs from typical sirtalis in having from eight to ten more ventrals and subcaudals (when individuals of the same sex are compared ), but more especially in having the pale flecks of the dorsolateral area red, salmon, or orange, rather than yellow, bluish or greenish as in typical sirtalis. The transition from the pale flecks of sirtalis to the red flecks of parietalis is extremely gradual and intermediates or mixed populations occur from Michigan and Ohio to Minnesota ECOLOGICAL STuDY OF GARTER SNAKE 503 and Kansas. T. s. parietalis differs from the West Coast subspecies in having semicircular black spots on the edges of the ventrals. T. s. annectens replaces parietalis to the south of the latter’s range, in Oklahoma and Texas. In annectens the dorsal stripe is bright orange instead of yellow, and the dorsolateral flecks are pale as in T. s. sirtalis. T.s, ornatus, an isolated subspecies, inhabits the Rio Grande drainage of south-central Colorado, New Mexico and north- central Mexico and resembles parietalis in most respects, but has a relatively pale, grayish brown dorsolateral area. West of the Rocky Mountains in the Great Basin region and the eastern parts of Pacific states and provinces another subspecies, T. s. fitchi, occurs from Alaska and northern British Columbia to southern California. It resembles parietalis most closely but differs in lacking black lateral spots on the ventrals and in having brighter, scarlet marks which are limited to the lower half of the dorsolateral area on each side. In this latter respect it resembles all the West Coast subspecies. The subspecies T. s. pickeringi in northwestern Washington has melanistic tendencies; its dorsal stripe is narrowed and limited to the middorsal scale row, the lateral stripes are partly or entirely obscured, and the ventral surface is heavily clouded with black pigment. T. s. concinnus of the Williamette Valley in north- western Oregon is another melanistic subspecies having the lateral stripes partly or entirely obscured, but it differs from pickeringi in having the dorsal stripe of normal width (involving not only the middorsal scale row but half of the row adjacent on each side), and in having the head bright red rather than dark olive as in most other subspecies. T. s. concinnus and T. s. pickeringi are alike in having relatively low numbers of ventrals and subcaudals. T. s. infernalis occupies a narrow coastal strip for most of the length of California. It resembles fitchi in most respects but has on the aver- age a much paler dorsolateral area (gray rather than black) and on the average fewer ventrals and subcaudals. T. s. tetrataenia is another coastal subspecies, but is localized in a relatively small area—the San Francisco peninsula. It is perhaps the most distinc- tive of all the subspecies. Like concinnus it has a red head but lacks melanistic tendencies and instead has the red marks on the sides enlarged, and extended longitudinally forming bands running for the whole length of the body. Numbers of ventrals and subcaudals are subject to much geo- graphic variation over the range as a whole, but in the western three-fourths of the species’ range at least their changes do not 504 UNIvERSITY OF Kansas Pusts., Mus. Nat. Hist. correspond accurately with the subspecific boundaries outlined above. Instead, there are clines with lowest numbers of these scales in the population situated farthest north or along the coast, and highest numbers in populations living farther interiorly or farther south. In most taxonomic studies of garter snakes these meristic characters have been heavily relied upon, but experimental work by Fox (1948) has demonstrated that the differences between individ- uals and populations are not necessarily genetic. The temperature at critical stages of embryonic development (in Thamnophis ele- gans) was found by Fox to affect the numbers of scales. In view of these findings it seems best to de-emphasize numbers of ventral and subcaudal scales and concentrate attention on differences in color and pattern in defining geographic races. O 200 600 SCALE in MILES Fic. 2. Geographic range of Thamnophis sirtalis (shaded) with marginal or near-marginal records of occurrence (dots). More heavily shaded areas show known or presumed intergradation between subspecies: 1. T. s. pickeringi, 2. T. s. concinnus, 3. T. s. fitchi, 4. T. s. infernalis, 5. T. s. tetrataenia, 6. T. s. parietalis, 7. T. s. ornata, 8. T. s. annectens, 9. T. s. semifasciata, 10. T. s. sirtalis, 11. T. s. pallidula, Many disjunct populations occur in the south- western part of the range. The same statement probably applies also in the northern part, but as yet the distribution in Canada is poorly known. Prob- ably the ranges of T. s. parietalis and T. s. fitchi are to a large extent separated by uninhabitable areas along the Continental Divide. This map is based on information from Fitch (1941:576), Stebbins (1954:505), Logier and Toner (1961:69 and 71), Fitch and Maslin (1961:296), and Smith (1961:223). ECOLOGICAL STUDY OF GARTER SNAKE 505 HABITAT Thamnophis sirtalis parietalis prefers a habitat of mesic vegeta- tion. On the Reservation these garter snakes were caught in every habitat where traps were set, but the mixture of diverse habitats in juxtaposition probably often resulted in their wandering into situa- tions different from those they preferred. Judging from the relative numbers caught in different situations, some of the major habitats of my study areas ranked as follows in order of preference. 1. Margins of ponds (Plate 24, fig. 1), with rank low vegetation such as smartweed (Polygonum sp.), barnyard grass (Echinochloa crus-galli), and giant ragweed (Ambrosia trifida). In dry weather there was often a zone of dried mud several feet or several yards wide between the water level and the zone of rank vegetation. Garter snakes were often found in this barren zone hiding beneath boards or beneath a small rowboat beached there. 2. Silt flat about an acre in extent at upper end of Reservation Pond be- neath an open grove of willows and other trees, and having a thick ground vegetation of dayflower (Commelina communis), rice cutgrass (Leersia ory- zoides) and others. 8. Woodland edge, and isolated thickets of crab apple (Pyrus ioensis), plum (Prunus americanus), dogwood (Cornus drummondi), sumac (Rhus glabra), and elm (Ulmus americana) in prairie or pastureland. 4. Prairie with native tall-grasses, including bluestems (Andropogon ge- rardi and A. scoparius), switchgrass (Panicum virgatum) and Indian grass (Sorghastrum nutans) (Plate 24, fig. 2). 5. Meadow with introduced pasture grasses, especially awnless brome (Bromus inermis) and blue-grass (Poa pratensis). 6. Fallow field in bottomland, with rank growth of giant ragweed and sunflower (Helianthus annuus). 7. Woodland with elm, oak (Quercus muhlenbergii, QO. velutina, Q. borea- lis), hickory (Hicoria ovata), ash (Fraxinus americanus), honey locust (Gle- ditsia triacanthos), osage orange (Maclura pomifera), Kentucky coffee tree (Gymnocladus dioica), redbud (Cercis canadensis) and dogwood. 8. Upland fallow field with sparse weedy vegetation, three-awned grass (Aristida oligantha), common lespedeza (Lespedeza striata) and, at a later stage of succession, asters (Aster sp.) and goldenrods (Solidago sp.) 9. Disturbed and relatively barren situations such as gravel roads, rock quarry, yards and lawns. This listing of habitats applies primarily to garter snakes in their summer ranges. In autumn they were most often caught along hilltop limestone outcrops in woodland, situations rarely frequented in summer. Obviously they had repaired to the outcrops to hiber- nate. Probably some made the trip after forest trees had shed their leaves and the aspect of the forest was much altered from that pre- vailing in summer. The more moist the situation, the more favorable it seems to be 506 UNIVERSITY OF KANSAS Pusis., Mus. Nat. Hist. for the garter snake. In dry weather, especially, the two ponds on my main study area seemed to be especially favored locations, but there was little tendency for the snakes to concentrate about them. Instead, the snakes remained scattered over the entire area, in a variety of habitats. This lack of concentration is somewhat sur- prising because several of the main prey species on the area, espe- cially the leopard frog, do tend to remain concentrated near the ponds and streams at most times and disperse to other habitats only in wet weather. Smith (1956:292-293) wrote of Thamnophis sirtalis in Kansas“. . . one of the most ubiquitous snakes . . . It does not appear to be confined in particular to aquatic habitats, although it occurs there in perhaps greater abundance than elsewhere. It is the common garter snake in towns, where it may be found in vacant lots and overgrown yards.” However, farther west in the central part of Kansas this garter snake becomes increasingly rare and localized and limited to aquatic situations as woodlands give way to grasslands, and mesic vegetation becomes scarce. The subspecies of the Pacific Coast states and Great Basin are less likely to be found wandering far from water than is parietalis. In this far western part of its range sirtalis “ is primarily a swamp snake, and is most abundant on low wet ground where there is dense vegetation. Along the margins of sloughs, along slow-moving watercourses, and in lush meadows along small coastal streams, it occurs in abundance .. .”; “. pre-eminently a marsh- or meadow-inhabiting species. Everywhere that it occurs it seems to favor a habitat of thick vegetation in damp places . . .” (Fitch, 1941a:8; 1941b:590). Wright and Wright (1957:837) have reviewed an extensive literature con- cerning the habitat of T. sirtalis in eastern North America (chiefly pertaining to the subspecies T. s. sirtalis but including also populations of the subspecies pallidula, semifasciata, and annectens). In summarizing, these authors wrote, “Of the 154 habitat categories . . . the 20 found most frequently are: everywhere (all sorts of situations) 18; woods 16; near water 18; lakes 12; fields 10; meadows 8; swamps 8; borders of rivers 8; marshes 7; along streams 6; gardens 6; roadsides 6; parks 6; clearings 4; margins of ditches 4; margins of creeks 4; moist situations 4; in water swimming 4; in water (otherwise) 4; low ground 4.” Thamnophis sirtalis shares most of its geographic range except the extreme northern part with other garter snakes; sometimes two or more kinds occur in one locality. The many forms involved must offer varying degrees of competition to T. sirtalis and must exert different selective pressures on its several geographic populations. In almost every instance where T. sirtalis occurs together with an- other kind of garter snake, sirtalis occupies the more general eco- logical niche, that which is most typical of the garter snakes as a group, whereas its competitor is more of a specialist. Throughout most of the eastern United States the ribbon snakes (T. sauritus ECOLOGICAL STuDY OF GARTER SNAKE 507 and T. proximus) are sympatric with sirtalis but careful comparison of habitat and food (Carpenter, 1952:238) has shown that the rib- bon snakes are more aquatic, much more inclined to climb, con- fined to situations where there are thickets of shrubs, and more limited to amphibians as a food source. South of the Great Lakes sirtalis shares extensive areas with Butler’s garter snake (Tham- nophis butleri) and the short-headed garter snake (T. brachy- stoma), both stubby-bodied, terrestrial snakes adapted for feeding on earthworms. Throughout most of the Great Plains region T. radix is sympatric with sirtalis, but seemingly is less dependent on water. In the Southwest T. cyrtopsis and T. marcianus both over- lap the range of sirtalis but occur far beyond its limits, in desert regions. Both are more aquatic than sirtalis and their occupancy of the limited aquatic habitats may prevent sirtalis from extending its range in that direction. In the western states the many mem- bers of the elegans complex include specialists in both aquatic and terrestrial habits. In localities favorable for garter snakes as many Fic. 8. Map showing range of Thamnophis sirtalis; the part in the United States and the southern edge of Canada (heavy lines) is shared with many other kinds of snakes, but a substantial part of the range (fine lines) lies to the north of ranges of all other snakes. Known range limits of T. sirtalis on the north correspond approximately with the northern limits of the Canadian Life-zone as mapped by Muesebeck and Krombein (1952:25). Distributional data are chiefly from Wright and Wright (1957) and Logier and Toner (1961). 508 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. as three members of this complex may occur along with sirtalis and their combined competition must be severe. At any rate, sirtalis is relatively stable and conservative in its habits and habitats as compared with the members of these groups. Compared with its many partly sympatric competitors of other groups T. sirtalis has several traits that seem to confer distinct ad- vantage and may, in part, account for its more extensive geographic range and (often) greater abundance locally. These are: 1. seem- ingly greater tolerance of low temperature sometimes resulting in a longer growing season, 2. larger average and maximum size, 3. larger litters of young. TEMPERATURE RELATIONSHIPS The range of Thamnophis sirtalis extends north far beyond the limits of any other reptile in the Western Hemisphere (Fig. 8). hence it perhaps has some special attributes permitting it to resist low temperatures. In the northern and central United States, where many other reptiles occur along with Thamnophis sirtalis, the latter is notable for early emergence from hibernation in spring, and persistent activity into late autumn. Bailey (1949:238) studied minimum temperature tolerances in garter snakes (Thamnophis radix radix and T. sirtalis parietalis) in artificial hibernacula at various depths, equipped with thermocouples. At depths down to 16 inches and at temperatures lower than —8° C. no snakes sur- vived, but some did survive temperature as low as —2.2° and sur- vived temperature of —1.5° or below for as long as two weeks consecutively. A temperature of —2.2° seemed to be near the critical level for T. radix. Whether the same applied to T. sirtalis was not definitely determined because only two individuals were used and both survived a temperature of —2.2°. Bailey concluded that garter snakes could regularly hibernate above the frost line surviving a temperature of approximately —2° C. because of gradual physiological preconditioning, the low temperature of crystallization of protoplasm, and possibly a limited capacity for “undercooling” of the tissues comparable to that found in invertebrates. In February and March, 1964, I experimented with cold-tolerance in a group of T. s. parietalis from the Rockefeller Experimental Tract. The snakes were kept in a poorly heated room and usually their bodily temperatures were below 10° C. From time to time they were moved outdoors when the temperature of the air was near freezing or lower. Usually a Schultheis quick-reading ther- ECOLOGICAL Stupy OF GARTER SNAKE 509 mometer was taped in place in the snake’s cloaca and other ther- mometers were kept nearby to check environmental temperatures. On many occasions snakes survived bodily temperatures between —2° and —3° C. and seemingly the critical minimum was near the latter level. On several occasions snakes that survived tempera- tures below —1° C. had their bodies much stiffened by partial con- gealing of their body fluids, but those that were frozen and com- pletely rigid never survived. While cooling, the snakes moved about restlessly, trying to find shelter, and were capable of motion until bodily temperatures were at least one degree below freezing. However, they were soon immobilized and then gave the appear- ance of being anesthetized until they were warmed to several de- grees above freezing. Although a bodily temperature of 3° C. or a little lower was lethal, the snakes were found to be able temporarily to withstand air temperatures that were considerably lower. On one occasion a snake was exposed at an air temperature of —9° for more than half an hour with no apparent harm, but the substrate was somewhat warmer and the snake’s body was not completely frozen. Snakes that were too cold to move freely, and hence could not readily escape, were especially inclined to show fight, and would flatten their heads and bodies, coil, and strike with grotesquely slow motions. The lethal minimum for Thamnophis sirtalis is seemingly little, if at all, below those of other cold hardy snakes and does not suffice to explain the northward extent of the geographic range far beyond the limits of other species. Carpenter (1953:78) found garter snakes (T. s. sirtalis) hiber- nating in crayfish burrows and vole burrows, both solitarily and in groups, sometimes partly or wholly submerged in water. On the Reservation I found T, sirtalis to have a longer season of activity than any other kind of snake. Latest dates of record each year from 1949 through 1963 were in November, the 11th, 14th, 13th, 12th, 13th, 13th, 11th, 14th, 26th, 19th, 11th, 27th, 10th, 12th and 5th. All these late season records were of snakes caught in traps. An occasional individual may have been active on an even later date in some years, but certainly the bulk of the population retired somewhat earlier than indicated by the dates listed (see Fig. 4). In most years weather in late October and early November was often too cold for the snakes to be active, and their activity was therefore intermittent toward the end of the growing season, with an increasing proportion remaining in dormancy after the advent of each cold front. Earliest captures in spring were of snakes found 2—3965 510 UNIVERSITY OF KAnsAS PuBLs., Mus. Nat. Hist. SG, XQ_G XGI]W | Wh NUMBER OF RECORDS Y j j j j j j j j Z QQ GW 6 MI SS SS DEGREES C SEPTEMBER OCTOBER NOVEMBER Fic. 4. Diagram showing numbers of garter snakes recorded at Reservation in autumn, in fields (left-hand, darker column of each pair) and along wooded hilltop with limestone outcrops (right- hand, lighter columns). Records are grouped in ten-day intervals, beginning with September 1 to 10 and ending with November 20 to 29. Left-hand columns represent combined data for years 1958 through 1963. Upper (unshaded) portions of right-hand columns represent these same years, and lower (shaded) parts of right-hand columns represent the years 1949 through 1957—when the trap lines were maintained only along the hilltop outcrops. Lower part of figure shows averages for the maxima, means and minima of daily temperatures (1949 through 1962). in the open or beneath flat rocks, and trap lines were not set until well after the emergence of the snakes. Therefore in most years there was probably a considerable lag between the time garter snakes first became active and the time they were first recorded. On the other hand those snakes found in late February were cer- tainly forerunners whose emergence preceded that of the main population because their hibernacula were in sites where premature warming occurred. It is my impression that in most years there are almost always one or more days in late March when some garter DR SSOIO&i AAi ° aug S$QYO93Y JO YsEWNN Wea NO RQQQ_’_*’UVAW RM AAAANoN RP VX QQAAAQ RXXXtss SS ASS SASS SSS Rs _ 4nonnnnogw BSS |™|| QO©BOQOAQOaesssssssssy) RS AAAI hw1|qv_eq>ixr}449 QQ Ard A SN XX AAAAA SAWN NAY QQ ww RAB ag oe 40 ES CENTIGRADE DEGRE 10 | eo) I = ratt pal v t of figure shows ymewhat lower th: s Da S&S Yee SOY & qn “— U 4 et! OS aya) tn rats ~ 4 o wn Dera) - © nae) Sg n= = py CS OF +O =) 5°) lol) oD SH Orr SEA} += UN = 2 OG 0 4 ere —"_ 5°) n o py as) = Pe & ad “= = ae) 2O ral Ll LT a wf =r 4) UV Onn bd) ol =| a) Lila m! = Yu On oa 2 _o s°] wD 1 ao) ool Ov a > reais | 1eir te ate tl U & nity to regul = dortr Loon! Y ha some op to prefer a le Uy ch average temperatu =I Y el between 29° > eal = eratures, wh P< emy ~ cent air a U ys i ~Y the snake w] PLATE 2 u Fic. 1. Head of large adult female garter snake, Thamnophis sirtalis pa- rietalis from lateral view, 2% times natural size, The University of Kansas Natural History Reservation, May 18, 1964. Fic. 2. Ventral aspects of adult garter snakes at posterior ends of bodies with tails to the left, showing typical differences in size and shape between the sexes; in the male (above) the tail, lodging the hemipenes, is slightly wider than the body, but in the female the tail-base tapers abruptly from the diameter of the body; May 19, 1964. PLATE 24 Fic. 1. Woodland pond near headquarters at The University of Kansas Natural History Reservation, Douglas County, Kansas, May 16, 1964, a breeding site for several species of frogs that provided the chief food source of garter snakes on the area. The snakes were often found about the mar- gins of the pond, but also were well scattered over other habitats. Fic. 2. Small pond in tall-grass prairie on the Rockefeller Experimental Tract, Jefferson County, Kansas, May 17, 1964; like the pond on the Reser- vation, this one was productive of amphibian prey, and garter snakes were abundant throughout the surrounding prairie. PLATE 25 Fic. 1. Adult female garter snake in threatening defensive pose ready to strike, resting on patches of snow, illustrating the cold-hardiness of this species. Temperature of air was approximately 3° C. and that of the snake approximately 6° C. when this photograph was taken in March, 1964. Fic. 2. Large adult female garter snake 773 millimeters snout-vent, 150 grams, and first-year young of typical size, 220 millimeters snout-vent, 4.0 grams on April 18, 1964. ECOLOGICAL StupY OF GARTER SNAKE 511 snakes are active on the Reservation, and that there is intermittent but varying activity through early April. However, actual dates of first capture, 1950 through 1963, were as follows: April 10, Feb- ruary 28, May 7, March 25, February 24, April 3, March 26, April 10, April 15, April 4, April 12, April 18, May 12, May 8. For specimens from northeastern Kansas in the University of Kansas Museum of Natural History, the following dates represent the earliest records in several different years: March 28, 1910; March (exact date not given), 1912; March 2, 1927; April 1, 1930; April 9, 1933; April 12, 1934; April 9, 1939; April 11, 1941; March 31, 1947; April 6, 1949; April 12, 1950. In several winters garter snakes were kept near the Reservation headquarters in outdoor cages having artificial hibernacula at depths of 15 to 36 inches. Several other kinds of snakes kept overwinter in the same enclosure suffered heavy mortality, but the garter snakes were remarkably hardy and nearly always survived. On February 11, 1961, when air tem- perature was 15.3°C., on the warmest day of the winter, two hibernacula were exposed to de- termine the condition of the snakes in them. The frost line was 13 inches below the soil sur- face. In a hibernaculum box at NUMBER OF RECORDS D Uj IY QGCGG. MG DEGREES CENTIGRADE Fic. 6. Bodily temperatures of garter snakes confined in a cool cage, but with an electric lamp providing op- portunity to bask. Definite preference for temperatures between 29° and 30° is indicated. 30-inch depth temperature was approximately 4°, and snakes that were uncovered writhed feebly when touched. In a 15- inch hibernaculum temperature was 1.7°. A group of snakes were lying intertwined in a mass, but with their heads raised, seemingly aware of their surrounding and not fully torpid. All were in good condition. On May 20, 1960, tests of the relative cold-tolerance of several species of snakes were made. Two racers (young and adult), two adult ring-necked snakes, a small adult copperhead and a small adult garter snake were placed together in a plastic bag in a refrig- erator. After 45 minutes they were removed. All were stiff and virtually immobilized (body temperature of one, 4° C.) but the SlZ UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. garter snake was the least affected. After a few seconds it began to crawl forward slowly. The ring-necked snakes soon became more active than the garter snake and one tested was found to have raised its body temperature to 12° C. Relatively rapid recovery in the ring-necked snakes resulted from their small size and high sur- face-to-mass ratio permitting relatively rapid absorption of heat. The whole group was returned to the refrigerator in a slightly different spot. When they were removed after an hour body tem- peratures were 2° C. Again the garter snake seemed least affected in the first few minutes, but the ring-necked snakes, warming more rapidly at room temperature soon became more active and were able to craw] first. The group was returned to an even colder spot in the refrigerator and kept there for three and one-half hours. When removed all the snakes were partly frozen and appeared lifeless. Each of the two tested had a body temperature of —1° C. After 20 minutes the snakes were still partly frozen and were un- responsive to stimulation. After 50 minutes the garter snake and racers made slight response to stimulation, with slow, writhing movements. On the following day the garter snake and the smaller racer seemed to have recovered completely, but the larger racer was partly paralyzed, and the ring-necked snakes and copperhead were dead. In June 1960 experiments were performed to test the heat-toler- ance of several kinds of snakes. A garter snake was placed in a transparent plastic tube stoppered with cotton at each end, and with a Schultheis quick-reading thermometer inserted in the snake’s rectum and taped in place. The snake’s temperature was raised by exposing it to sunshine. As bodily temperature approached 40° C., the snake showed signs of discomfort and struggled violently to escape. At approximately 41° C. it collapsed and appeared to be dying, but was quickly revived by holding it in cold running water. A copperhead and ring-necked snakes subjected to the same treat- ment collapsed at approximately the same temperature, but a racer proved to be more resistant. Lueth (1941:130) experimented with the survival of snakes at temperatures near their limits of tolerance. Two Thamnophis sirtalis survived for 48 hours at 41° C., but another died after 24 hours at 38.5°.. On the Harvey County study area one garter snake when removed from a trap seemingly had died from overheating only a few minutes before and its rectal temperature was 40.5° C. Opportunities to record temperatures of active garter snakes under natural conditions were rare. At Harvey County Park bodily EcoLocicaL StuDyY OF GARTER SNAKE 518 temperatures of many of the snakes found in traps were recorded, but these temperatures in confined snakes are of doubtful signifi- cance. Certainly these snakes had some opportunity to regulate their body temperatures by utilization or avoidance of occasional patches of sunlight, and contact or avoidance of soil, metal, and plant debris. But such opportunity for thermoregulation was lim- ited and temperatures of the snakes tended to approximate ambient temperatures. Fig. 5 shows the distribution of these snakes’ tem- peratures. They extended over a wide range, from 13° C. to 39° C. but with the greatest concentration between 29° and 30°. On December 24, 1963, attempt was made to determine experi- mentally the preferred temperature level in a group of 21 adult Thamnophis sirtalis parietalis. The snakes were in a rectangular cage 36 inches long, 22 inches wide, and 28 inches deep, open on top and with a three-inch mat of straw on the bottom. The cage was in a poorly heated laboratory. A desk lamp was set in one corner of the cage, and turned on, with the bulb close to the bottom of the cage. Soon snakes attracted by the warmth emerged from the straw and approached the lamp to bask. A thermometer in the corner of the cage opposite from the lamp indicated a tempera- ture there of 16.0° C. at the start of the experiment, and probably it deviated only slightly from this level during the several hours that observations were made. From time to time snakes basking in the warmth of the lamp were momentarily removed from the cage and their bodily temperatures were recorded. The histogram of Fig. 6 shows the trend of the 60 readings taken. As in the rec- ords of snakes removed from traps in summer, readings are most concentrated in the one-degree range, 29° to 30°, suggesting that these temperatures are most preferred. In this series the records are much more concentrated than they were in the trapped garter snakes; more than 81 per cent are in the six-degree range, 27° to 33°. Admittedly temperature preference in this species and in other ectotherms may vary according to physiological conditions, season, sex or age, and, perhaps geographic origin, and the limited data presented here provide only a crude approximation of the temperature preferences in Thamnophis sirtalis. FOOD HABITS In the course of my study exactly 200 records of prey items were obtained, of which 188 were palped from stomachs, and 12 were identified from scats. Seventy of these records originated from the Reservation; the remaining 130 were from Harvey County Park. 38—3965 514 UNIVERSITY OF KAnsAS Pusts., Mus. Nat. Hist. These records were as follows: 120 leopard frogs (Rana pipiens), 7 bullfrogs (Rana catesbeiana, all juveniles), 25 unidentified frogs (probably all either Rana pipiens or R. catesbeiana, but too far digested to be readily distinguishable), 12 American toads (Bufo americanus—all juveniles), 8 gray tree frogs (Hyla versicolor), 6 cricket frogs (Acris crepitans), 5 earthworms (Allolobophora caliginosa), 5 wood mice (Peromyscus leucopus), 5 western nar- row-mouthed toads (Gastrophryne olivacea), 2 prairie voles (Microtus ochrogaster), and 1 each of harvest mouse (Reithro- dontomys megalotis), garden toad (Bufo woodhousei), striped chorus frog (Pseudacris triseriata), bird (unidentified) and juvenal copperhead (Agkistrodon contortrix). The trends of feeding differed somewhat on the two study areas, as might have been expected from the difference in habitat. At Harvey County 118 of the 180 items were ranid frogs, and pre- dominantly leopard frogs (only two definitely identified as bull- frogs); the remaining items were 4 cricket frogs, 2 toads and a bird. Thus at Harvey County Park 91 percent of the food items were ranid frogs (chiefly R. pipiens); at the Reservation these same frogs made up the most important component of the food, but amounted to only 41.4 percent of the total, with a greater variety of miscellane- ous items than were found at Harvey County. Besides the salientian amphibians, prey items consisted of voles, mice, a snake and a bird, TABLE 3.—DIsSTRIBUTION OF 55 Foop ITeMs oF GARTER SNAKES, ACCORDING TO SIZE OF SNAKE Snout-to-vent length of snakes in millimeters a TMOUBES,< esas Sere 4 .07 06 25 17 frog (Hana) sasee ae 28 .10 2 val .59 .50 . 66 tree-frog (Hyla)........ 8 .30 12 ae .18 325 Miscellaneous small salientian® ye acne 14 .20 .62 ) Aucust 25)... : 743 100 °) iWhylSor. cece = 623 72 November 11. . 670 98 °) JulysZaee a ae 617 83 November 2... 680 100 2 uleynseaihile og ce G22, 152) | August 26... <>. 750 120 2 Aprils sie os 748 148 | October 21.... 837 203 August 5—190 millimeters; September 24—256 millimeters; October 10—257 and 300 millimeters. Larger garter snakes also made substantial growth; records of all those recaptured after considerable intervals are shown in Table 6. Even more rapid gains, sometimes exceeding three millimeters per day, were made by some of the garter snakes at Harvey County Park. Records of seven individuals, all females, selected on the basis of their rapid growth rates, are shown in Table 7. When the data on snakes recaptured at Harvey County Park are arranged according to sex and size group the snakes are seen to have grown at approxi- mately twice the rate of those at the Reservation. Obviously food was more abundantly available at Harvey County Park, an area of low-lying prairie in sand dunes along the Little Arkansas River. Poorly drained areas among the dunes, which were often flooded with shallow water, provided breeding places for many species of frogs and toads. These amphibians and their tadpoles provided an abundant and varied food supply. Many times in the summer dispersing swarms of newly metamorphosed frogs or toads spread into all habitats. Usually amphibians of various sizes were abundantly available as food. However, at the Reservation the water supply was limited to two small ponds, and two intermittent streams. Growth in large adult garter snakes was found to be especially erratic and unpredictable. The oldest individuals of a population are probably in most 534 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist. TABLE 7.—GROWTH IN FEMALE GARTER SNAKES AT HARVEY COUNTY PARK Original Record Recapture Record Snout- Snout- vent | Weight vent | Weight Date length in Date length in in milli-| grams in milli- | grams meters meters Julyo> 1960525. 398 21 ‘| August 27, 1960:.... 581 65 June 25, 1960.... 428 28 | October 21, 1960.... 687 150 June 15, 1959.... 441 35 | August 20, 1959..... 676 96 April 19, 1960. %. .-? 703 172 May-os 19602 s42 4. 22 725 170* August 2, 1960...... 795 150 April 26, 1960... . 464 30 | October 17, 1960.... 771 185 May 28, 1959.... 480 5O © || uly 2) 1959). 0b. 574 73 September 27, 1959.. 672 110 June 12, 1959.... 537 AG Sulys23; V959.. = fae. 670 118 June-12, 1959... =. 578 do » | @uly LG; 1959.90 3. 650 130 > gravid TABLE 8.—GROWTH IN FEMALE GARTER SNAKES THAT ATTAINED UNUSUALLY LARGE SIZE Original Record Recapture Record ge |g pls n io] Qn Ss egs| & | & ees} & | & Date oe ie o 2 Date ae 2/ - | of Sway ie ~ SY P38 ~ ar Seal, = | fe 32S & | iss 6.2/8)! So: 7| ere S28| 3 | 38 D = Ay ND = a4 Nov. 14, 1956. ..| 450 29 15 Sept. 28, 1959...) 883 215 50 Nov. 11, 1955. ..| 378 16 15 Oct. 8, 1960... .| 900 410 75 Oct. 16, 1958....-;} 723 170 51 July 30, 1963. ..| 930 290 108 Sept. 2, 1958... .| 856 255 73 Oct. 10, 1959. . .| 902 227 86 April 138, 1959...) 748 148 56 Oct. 21, 1959. ..| 837 203 63 June 20, 1960...| 950 | 200 71 July 25, 1958. . .| 890 213 ? June 3, 1959... .| 935 285 2 May 3, 1960....| 940 [..... ? ECOLOGICAL STuDY OF GARTER SNAKE 535 instances near maximum size, and the growth recorded in large adults bears on the problem of longevity. The eleven largest garter snakes captured at the Reservation were all females in the size range 880 to 950 millimeters, and six of them had been captured and marked before they had reached their maxi- mum recorded size. Records of these individuals are presented in Table 8. The trend of these records indicates that the largest garter snakes at the Reservation, those more than 900 millimeters in snout-vent length, may be as much as nine years old or as little as five years old, but usually are of an inter- mediate age. In summary, it may be said that growth in these garter snakes is rapid as compared with growth in most other snakes, but that it is erratic and unpredictable, some individuals gaining more than three millimeters per day in snout-vent length while others remain static. A gain of 1.2 millimeters per day is fairly typical before the rate slows with approaching maturity. In one-year-old males the snout- vent length is usually between 350 and 400 millimeters and subse- quently the growth rate slows abruptly. In second-year females the rapid growth rate of the early months is maintained or even increased, although many are already gravid at an age of 21 months. Table 9 based on records of recapture of marked individuals com- pares the rate of growth (millimeters per month) in snakes of sev- eral series, bringing out differences that are caused by sex, size and locality. The largest snakes, females more than 900 millimeters in snout-vent length, grow much more slowly than small or medium- sized individuals, and their rate of growth is even more variable. TABLE 9.—GrowTH RATES, IN MILLIMETERS PER MONTH, OF GARTER SNAKES IN VARIOUS SIZE GROUPS, AT THE RESERVATION AND AT HARvEY COUNTY PARK Snout-vent Length of Snakes in Sample Sex and Locality 400 to | 450 to | 500 to | 600 to | 700 to | 800 to 449 499 599 699 799 899 milli- milli- milli- milli- milli- milli- meters | meters | meters | meters | meters | meters Males (Reservation). .| 16.6(10)| 7.2(8) | 6.6(19)| 5.1(7) |......2.[0..2.5.. Majles\(Harvey Co:):. :| 50:0(31))30:0(16)} 13:3(15)| 75(6) see. ee Females (Reservation)|........|........ 23.4(25)} 10.8(34)| 9.1(13)} 6.3(4) Females) (Harvey Cos). 2.2-2..|-....-.- AS THEO) 24-604)! I (Ost (As | emcees 536 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. MOVEMENTS Carpenter (1952:247) obtained records of 127 movements of the common garter snake at his Cherry Hill study area in southeastern Michigan over a three-year period. The maximum movement recorded was 984 feet and most were less than 600 feet. In general, the snakes tended to stay each in its own small area, which Carpenter termed an “activity range,” contrasting this with a “home range” which, according to the same author’s definition, must have a home base—a regularly used nest or shelter. No such feature is present in the range of a garter snake. Carpenter noted in those snakes recaptured re- peatedly that the pattern of movements tended to have one axis much more elongate than the other. He concluded that the average range is approxi- mately 2.07 acres—600 feet long and 150 feet wide. My own records lead to rather different conclusions which no doubt result from differences in the local populations of snakes, differences arising from field techniques, and differences in interpretation. I suspect the latter two are of greatest importance. Some of the garter snakes marked and recaptured at the Reservation were at hilltop rock outcrops where it was thought they had come to hibernate, perhaps having travelled beyond their usual spheres of activity. Eliminating these records of the snakes caught at hilltop outcrops, there are available 342 separate records of movements for the snakes on this study area and at Harvey County (Fig. 12). For a snake recaptured many times at random points, the area encom- passed would be sufficiently representative of the home range but would always be somewhat smaller. In actual practice most individuals were caught only once or a few times, and the movements recorded gave little indication of the shape, extent or size of the individual’s range. However, in the aggregate the movements measured do reflect the size of the home range. The longest movements probably represent shifts in range, but these are not readily dis- tinguished from movements within a range. On the average random move- NUMBER OF RECORDS 1000 2000 3000 DISTANCES BETWEEN CAPTURES: FEET Fic. 12. Movements within or between home ranges in garter snakes at the Reservation and Harvey County Park. Trends are similar in the sexes. Rela- tively few movements exceed 1250 feet (the approximate diameter of a home range); the longer movements represent shifts in home range. ECOLOGICAL StuDY OF GARTER SNAKE Bor ments within a home range should be equal to half the range’s diameter; some would connect points on opposite edges while others would connect points near together in the central portion. Table 10 shows certain trends revealed by grouping the records of movements according to sex of the snakes involved, according to length of time elapsed between captures and according to locality. Length of movement increases with elapsed time, showing that home ranges are not occupied indefinitely, but either they are gradually altered, or the snakes from time to time shift to new areas abandoning those formerly occupied. The change in time is much more marked in the female garter snakes than in the males, indicating that in the former the home range is altered more rapidly. The females have the smaller ranges. In general, the trends shown by the movements from the Reservation are borne out by the somewhat smaller sample from Harvey County Park. In Table 10 approximately 20 per cent of each sample, including both the longest and shortest movements were discarded in order to eliminate those that were unrepresentative of home ranges because they resulted from shift of range, and likewise to eliminate movements that were too short to be repre- sentative. Some of the snakes were recaptured before they had oppor- tunity to make extensive movements, and certain sites were so favorable that they might yield repeated captures of the same individual, with little or no movement demonstrated, even though the individual had travelled extensively between captures. Support for the procedure of eliminating the highest and lowest ten per cents of movements recorded is found in the graph of Fig. 13 showing the average movement for 102 females, and the shifting of the average as the extremes at each end of the series are progressively eliminated until only the median remains. The average movement of 554 feet indicates a home range diameter of 1108 feet. Obviously ranges of different individuals are not all ee TABLE 10.—AVERAGE DISTANCES IN FEET BETWEEN CAPTURE POINTS OF MALE AND FEMALE GARTER SNAKES FROM RESERVATION AND FROM HARVEY County Park, GROUPED ACCORDING TO ELAPSED TIME, SHOWING TENDENCY FOR MALES TO MovE FARTHER THAN FEMALES AND FOR DISTANCES TO IN- CREASE OVER LONGER PERIODS Time Sex Reservation Harvey County | Combined sample Up to one fot 675(for 31 of 38)|} 719(for 28 of 36)| 696(for 59 of 75) ment Q 542(for 60 of 76)| 605(for 26 of 32)| 562(for 86 of 108) Two to five aie fot 965(for 13 of 15)| 930(for 13 of 15)| 920(for 24 of 30) meat 9 814(for 19 of 23)| 1047(for 8 of 10)| 884(for 27 of 33) Sixto12 | o& | 753(for 9 of 11) | 775(for 11 of 13)| 765(for 20 of 24) eae Q 1091(for 25 of 31)| 1064(for 9 of 11)| 1075(for 32 of 40) More than io fa daria 781(for 8 of 10) | 1480(one) aie 859(for 9 of 11) months Ow MAT OGor 15: of U9) lee: se es ae ee 1110(for 15 of 19) 538 UNIVERSITY OF KAnsAS Pusts., Mus. Nar. Hist. 100 80 60 40 20 0 PERCENTAGE OF SAMPLE USED AVERAGE DISTANCE IN FEET Fic. 13. Diagram showing average movement in 102 female gar- ter snakes from the Reservation (lower line) and 39 males from Harvey County (upper line) and changes in the average as each series is reduced by eliminating the longest and shortest move- ments in successive pairs until only the medians remain. The series of movements of females included several that were excep- tionally long, representing shifts of range. The series of move- ments of males was more uniform and was less affected by removal of the extremes. precisely the same size, and though perhaps tending toward circular shape, all deviate in some degree with elliptical or irregular shapes. Therefore, it is to be expected that some movements within home ranges would exceed 1108 feet. The ten longest movements recorded in the series were: 3800, 3560, 3340, 3080, 2720, 2400, 2360, 1920, 1920, 1800. The remaining 92 move- ments were all less than 1500 feet, hence within the probable limits of a home range. Assuming that the average figures obtained of home range radius are correct —562 feet for females and 696 for males—and assuming that home ranges tend to be circular, approximate size of the average range may be obtained from the formula 7r2, for the area of a circle. A home range of 35.0 acres for males is indicated, and one of 22.7 acres for females. In other kinds of snakes also the males have been found to travel more extensively than the females and to have larger home ranges (Fitch, 1958:119; 1960:149; 1963a:391; 1963b:651). Breeding activity probably is not involved because in Thamnophis sirtalis mating takes place mainly in a period of a few days after emergence from hibernation, or in autumn shortly before retirement into dormancy, whereas the records that form the basis of my calculations of home ranges are based primarily on summer movements. In assessing the fact that the home range calculated in Carpenter’s study was less than one-tenth of that found in my own study, the sizes of the areas involved must be considered. Carpenter’s study area was a rectangle of 48 acres, approximately 1670 x 1250 feet. Nearly one-third of its central por- tion was rectangular marsh that was little used by the snakes, except along its border where the activity of the snakes and probably that of the collector was relatively concentrated. The maximum possible distance between points ECOLOGICAL STuDY OF GARTER SNAKE 539 in this area was approximately 2100 feet between opposite corners. Possibly many of the snakes had ranges extending beyond the limits of the study area, although Carpenter did check some of the peripheral areas, and concluded that there was little interchange. In the part of my own field study carried on at the Reservation a line of 200 live-traps a mile and a half in length and a mile in width at the maximum, was maintained on a study area totalling 750 acres. Ideally traps should have been uniformly spaced in a grid but practical considerations overruled such a procedure, because it was imperative to install the traps where they would be shaded, and where habitat conditions were favorable for the snakes. Actually some degree of randomness was achieved, as the traps were well scattered, and in fact, were set for snakes other than Thamnophis, primarily for copper- heads from 1949 through 1958 and in the same general area for racers from 1959 through 1962. With the shift in emphasis, from copperheads to racers, no great change in the trap lines or the procedure was involved, but many of the sets were shifted from woodland edge or dense thickets to more open situ- ations in the same general areas. The Harvey County study area was smaller —approximately 4,000 x 2,000 feet. At the Reservation those snakes caught at rock ledge locations in late Sep- tember, October or November were thought to be preparing to hibernate. A few of them were caught on either earlier or later occasions in what were pre- sumed to be their summer ranges. Throughout October some garter snakes were caught in the same situations frequented during the summer, and it is uncertain whether all repair to hilltop outcrops to hibernate. Certainly some do so, but others may utilize deep burrows in the meadow habitat where they spend the summer. The 24 females that had made trips between hilltop out- crops and meadows had moved, on the average, 1,745 feet, whereas the cor- responding figure for 13 males was 1,188 feet. Eight males and seven females were each caught twice along hilltop rock outcrops where they were preparing to hibernate in autumn or had emerged from dormancy in spring. One adult female was captured at the same crevice on September 24, 1959, and November 3, 1960. Also, she was captured three times in the summer of 1960 and once in the summer of 1961 in the small valley of the Reservation’s headquarters area, and seemingly she had moved some 800 feet up a wooded slope from the nearest part of her home range to reach the hibernaculum. Another female caught at an outcrop on September 28, 1952, was recaptured at part of the outcrop 570 feet from the first location on September 20, 1954. A. third female was captured at points 1,320 feet apart, along the ledge on October 27, 1959, and November 3, 1960. A male was caught at rock ledge locations 2,200 feet apart on November 2, 1959, and November 12, 1960. The remaining movements along the rock outcrops were seven for males and five for females. All were for short periods, and averaged 247 feet, indicating some wandering in search of suitable hibernacula even after the snakes had reached the sheltering outcrops. The records are too few to show the main trend of population movement in connection with dormancy, but do indicate that many, if not all individuals, migrate from their summer ranges in meadow habitat to hilltop outcrops— sometimes only after freezing weather has begun—and that some return to the same hibernacula in successive years whereas others find hibernacula remote from those previously used by them. 540 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. DEFENSE AND ESCAPE In its color pattern Thamnophis sirtalis is typical of a large group of colubrid snakes that are active and live in situations having dense ground vegetation. The sharply contrasting colors on the dorsal surface exemplify the “ruptive” type of pattern, breaking up the outlines of the body and hence rendering the animal more difficult to see under some conditions. The longitudinally continuous areas on the dorsal surface serve especially to conceal motion. Ordi- narily when a garter snake is seen, part of it is hidden from view by screening vegetation; motion can be readily detected only if the head is in view. Otherwise, as the snake moves forward the section of the body that is exposed seems to shrink and vanish as the ob- server watches, unaware that it is in motion. In tall grass, or ri- parian thickets with bramble and nettle these garter snakes are exceedingly elusive. In day-to-day checking of lines of live-traps on the Reservation I walked scores of miles without seeing a garter snake other than those that were trapped. Those that were seen amid dense vege- tation on warm days when they were fully active usually escaped. Often such a snake was barely glimpsed as its initial movement or the slight rustling sound produced drew attention to it. Darting through the dense vegetation with rapid and erratic movements and sudden pauses, the snake usually was beyond danger of capture within a few seconds. In all subspecies of Thamnophis sirtalis occurring to the west of the range of T. s. parietalis, the dorsal stripe is especially bright and its edges are so sharply defined that it creates the impression of being painted on the snake. In parietalis the stripe is not so bright; it may have a faint suffusion of dusky pigment and the edges are a little less sharply defined. In T. s. sirtalis the dorsal stripe is duller, with its edges blending into the adjacent darker areas. T. s. pallidula in the extreme northeastern part of the species’ range lacks most of the adaptive pattern characteristic of the species throughout the remainder of its range. The dorsal stripe is dull and faint, or absent, and the dorsolateral area is pale, marked with dark spots. Since the range of pallidula is entirely within recently glaciated areas, it seems logical to regard its racial characters as recently evolved, and adapted to the special conditions. Lacking field experience with pallidula, the writer is nevertheless impelled to speculate on the adaptive significance of its color pat- tern on the basis of experience with somewhat similarly patterned ECOLOGICAL STuDY OF GARTER SNAKE 541 garter snakes of the western United States. In the northern glaci- ated region inhabited by pallidula, summers are relatively cool and snakes must spend a greater part of their time basking in the open in order to maintain body temperatures near their preferred levels; the soil is mostly rocky and ground vegetation is relatively scanty. Against a background of granite boulders, or gravel and sand, the striped pattern would be less advantageous than in dense vegeta- tion, and a pale variegated pattern with spots such as has been developed by pallidula, would be more concealing. Although the garter snake is fast moving and usually is quick to escape at any alarm, it does not always attempt to flee. Instead it may show aggressive behavior with threat displays. The display consists of an extreme lateral flattening of the body and head as the snake coils in readiness to strike, and it may strike if the annoy- ing object comes too near. In the process of flattening, the skin is stretched and the red areas that are mainly confined to loose skin between the scales and are partly concealed at other times, are prominently displayed. The strike is mostly a bluff which in most instances, probably serves to intimidate the enemy rather than to cause any actual injury. I have often teased garter snakes that were engaged in this type of aggressive behavior, and usually the stroke fell just short of a hand presented to the snake at a distance of a few inches, or at the end of a strike the snake’s head might touch the hand, but without actually grasping it. The mouth may be opened so widely that if the stroke finds its mark contact with the teeth is unavoidable. Nevertheless there is usually no actual biting and the recoil is almost instantaneous. Factors that influence the response of the garter snake to threat of danger include type of cover available, type of enemy involved, size and sex of the snake, and temperature. Large garter snakes, especially females, are much more inclined to behave aggressively than are juveniles, and of course this attempt at bluff against a typical garter snake predator, for example a rail or a small heron, would have a much better chance of success. At relatively low temperatures, when the snake is least able to make a speedy escape, it is most inclined to use threatening behavior as an alternative. Captive garter snakes which had become more or less accustomed to handling, and normally evinced no defensive or aggressive be- havior, could be caused to flatten, coil, and attempt to strike by cooling them to within a few degrees of freezing. The defensive display would be least striking in T. s. pallidula, in which the flattening of the body and spreading of the skin would 542 UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist. result in little change in the appearance except that the surface area in sight would be increased. In T. s. sirtalis the partially concealed markings between the scales are pale, yellowish, greenish or bluish, and provide a far less striking display than the red markings of parietalis, at least for those natural enemies that have color percep- tion. Most reptilian and mammalian enemies probably have lim- ited powers to discriminate between colors and do not see red. The displays may be most effective against birds, which have color vision better developed. In T. s. tetrataenia of the San Francisco peninsula, having the red markings greatly enlarged and fused into continuous areas, the display is most impressive, and in fact the appearance is striking even when the snake is not engaged in a threat display. Whether the peculiarly striking pattern of tetra- taenia has any special adaptive value or is merely the chance product of a formerly insular habitat has not been demonstrated. However, potential avian enemies are certainly more concentrated there than over the garter snake’s range as a whole. The relatively small range of T. s. tetrataenia on the San Francisco peninsula has ocean on three sides, and sea birds, especially gulls, may be seen frequently in almost any part of it. At times these sea birds may resort to freshwater ponds or streams, and could then become a potential threat to garter snakes. Garter snakes are noted for the nauseous secretion of their anal musk glands. These paired glands are elongate and sausage-shaped and are situated in the base of the tail. The secretion is a tan-col- ored thick creamy liquid. It is exuded chiefly after the snake has been caught and is struggling to escape. As exudation occurs, the snake presses its vent against the annoying object (or against its own body) and then, with writhing movements, spreads the secre- tion as widely as possible in a thin film over its own body and that of its captor. Observations on the effectiveness of the musk glands in deterring attack or ingestion by a natural enemy have rarely been made. On the one hand there is a large body of evidence indicating that many kinds of predators do indeed eat garter snakes despite the musk, but on the other hand most human observers agree that the odor of the musk is highly offensive. Effectiveness of the musk probably depends on the kind of predator and the degree of hunger moti- vating it, but there can hardly be serious question that the musk does have some deterrent value. On one occasion when a short-tailed shrew (Blarina brevicauda ) and a garter snake were caught together in a trap, the shrew ~ EcoLoGIcAL Stupy OF GARTER SNAKE 543 attacked the garter snake at the tail end and ate its way toward the body up to the vent. Along with the remainder of the tail this shrew had eaten the musk glands. However, these glands had doubtless been emptied previously by the snake as it exuded musk in its struggles. On several occasions when predators including raccoons, opossums, and probably skunks, had broken open traps and killed the trapped garter snakes, parts of the snakes, including tail bases with musk glands, had not been eaten but it was uncer- tain whether these had been discarded because of the musk. Some of the garter snakes that were handled bit viciously. These were mostly large females. In biting, the snake would deeply embed the long recurved teeth at the rear of the maxillary bone, piercing the skin of a restraining hand in a way that caused pro- fuse bleeding. There is probably some geographic variation in disposition and aggressiveness. My field experience with the more western subspecies, especially fitchi, have indicated that these snakes are much less inclined to bite than are members of the sub- species parietalis. A garter snake that has been grasped roughly usually responds by almost instantly voiding the contents of the cloaca, writhing to smear itself and its captor with fecal material and musk. At the same time it twists and thrashes in an attempt to escape. If it is grasped by the tail, it twirls the body rapidly, and within a few seconds may twist the tail off. In garter snakes the tail is remark- ably fragile, just as it is in most lizards, and fragility serves the same purpose. When a break occurs, the broken end wriggles about in lively fashion, which might momentarily distract the atten- tion of a predator from the snake itself. Of 940 snakes examined at the Reservation, 17.9 per cent had incomplete tails, but the percentage was notably different in the sexes: 20.4 per cent in 524 females and only 14.6 per cent in 416 males. In the average male having an incomplete tail, approxi- mately 43.8 per cent of the tail was missing, whereas in the average female the corresponding figure was approximately 50 per cent. The larger snakes, on the average, had more injuries to the tail, and also tended to have more of the tail missing. In males the incidence of incomplete tails was: eight per cent in those of 300 to 400 millimeters (snout-vent), 11 per cent in those of 401 to 500 millimeters, 22 per cent in those of 501 to 600 millimeters, and 24 per cent in those of 601 to 700 millimeters. In females the corre- sponding figures were: 7, 12, 16, and 25 per cent; 30 per cent in those 701 to 800 millimeters and 41 per cent in those more than 800 544 UNIVERSITY OF Kansas PusLis., Mus. Nat. Hist. millimeters. Hence, the greater incidence of tail injuries in females seems to be correlated with their greater bulk rather than with any difference between the sexes in habits. The larger and heavier snakes when grasped by the tail, can snap or twist it off more easily than can smaller and lighter individuals. NATURAL ENEMIES In 1952, 1955, and 1958 through 1963, pellets of the red-tailed hawk (Buteo jamaicensis) were collected in early summer from beneath nests whenever the opportunity arose. Several of the nests were on the Reservation and Rockefeller Tract; a much larger number were at localities well scattered throughout the eastern part of Kansas. In the total of 1320 pellets examined there were exactly 100 occurrences of the garter snake, which was hence a fairly im- portant component of the food, although it ranked behind the cot- tontail (Sylvilagus floridanus), the prairie vole (Microtus ochro- gaster), the cotton rat (Sigmodon hispidus) and the black rat snake (Elaphe obsoleta). In a collection of 116 red-tailed hawk stomachs donated by the U. S. Fish and Wildlife Service from localities well scattered throughout the United States and southern Canada, there were 152 vertebrate prey items of which three were Thamnophis sirtalis: one from Shoal Lake, Manitoba, May 30, 1917; one from Oak Lake, Manitoba, October 2, 1921; and one from Lac de Nonne, Alberta, May 26, 1926. Another occurrence listed merely as Thamnophis, from Norwich, Connecticut, also may have been this species. In a smaller series of broad-winged hawk (Buteo platypterus) stomachs received from the U. S. Fish and Wildlife Service, there were three occurrences of natricines, some or all of which may have been the common garter snake. Twenty-five pellets collected be- neath the nest of a Swainson hawk (Buteo swainsoni) in July, 1962, in Miami County, Kansas, approximately 30 miles southeast of the Reservation, contained 57 prey items of which 6 were Thamnophis, presumably the present species. Guthrie (1932:102-103) mentioned instances of predation on Thamnophis sirtalis by Swainson’s hawk, the sparrow hawk (Falco sparverius) and the Virginia rail (Rallus virginianus ). Randall (1940:166) recorded garter snakes in the food of Penn- sylvania marsh hawk. Conant (1951:253) mentioned the finding of a garter snake in the stomach of a red-shouldered hawk in Mont- gomery County, Ohio, and Ernst (1945:453) in New York found EcoLoGICAL STUDY OF GARTER SNAKE 545 garter snakes and water snakes to constitute approximately three per cent of the food at the nests of 16 red-shouldered hawks. In Minnesota, Errington and Breckenridge (1938:113-121) recorded garter snakes in the food of the red-tailed hawk, broad-winged hawk, and Swainson’s hawk. Even the ruffed grouse has been re- corded preying upon garter snakes (Zorichak, 1953:2). Among mammals, raccoons, skunks, minks, foxes and badgers probably include the most important predators on the garter snake over its range as a whole. Unfortunately, workers who have studied the food habits of these animals have usually tended to lump garter snakes in such broad categories as “reptile” or “snake,” or at least have failed to distinguish species, so that the extent of predation on Thamnophis sirtalis is difficult to judge. Because of the small size of the newborn snakes, relatively small birds and mammals, which are not normally predators on other vertebrates, may occasionally eat them. McKeever (1958:170) recorded an instance, observed near Peterborough, Ontario, in which a chipmunk (Tamias striatus ) attacked a 14-inch garter snake. The snake was said to be slow and sluggish, presumably because of cold weather, as the date was April 20, 1957. The chipmunk dragged the snake into a hollow log as the observer watched. The snake was still writhing feebly, but when it was examined was found to be helpless, its head chewed to a pulp. As the observer left, the chipmunk returned to the at- tack. Even a “mouse” (Microtus sp.?) has been observed to make an unprovoked attack on a small garter snake. In the course of my study at the Reservation and Harvey County Park 1339 food records of the blue racer (Coluber constrictor) were accumulated, and 18 of these were garter snakes. Except for the relatively minute insect items, chiefly crickets, grasshoppers, and katydids, the garter snake was actually one of the more fre- quently occurring items, exceeded among vertebrates only by the six-lined racerunner (Cnemidophorus sexlineatus), prairie vole (Microtus ochrogaster), and wood mouse (Peromyscus leucopus). Other records of the racer preying upon this garter snake include those of Taylor (1892:330), Surface (1906:170), Ortenburger (1928:181, 228), Breckenridge (1944:118), Finneran (1948:124), and Duellman (1951:338). There is circumstantial evidence that other predators, for which no records were obtained, may have taken even heavier toll of garter snakes on the area. On numerous occasions the wire funnel traps in which most garter snakes were caught were disturbed by 5—8965 546 UNIVERSITY OF Kansas Pus.s., Mus. Nat. Hist. mammalian predators, but the type of disturbance varied. Some- times the trap had been merely rolled or pushed away from its position, but remained unopened, either with or without a trapped animal. In other instances the trap had one end, either the funnel entrance or the plug at the rear, wrenched away, or mashed and collapsed on one side permitting the predator to gain access to the animals caught. In still other instances, when the trap could not be opened a paw had been inserted through the funnel entrance and the trapped animal had been removed, sometimes only after a violent struggle, when the inserted paw had become snagged on the inward-projecting wire prongs of the funnel. Mammals known to have disturbed the traps from time to time include domestic dogs, raccoons, opossums, skunks (both striped and spotted) and long-tailed weasels. In many instances the evidence was insuf- ficient to identify definitely the predator involved, but raccoons were known to be the most habitual and skilled raiders of the trap lines. Most often insects, small mammals, or frogs were the prey sought from the traps, but on several occasions remains of garter snakes that had been removed from the traps and partly eaten were found nearby. Garter snakes in traps were seemingly more subject to attack than were some other larger and more formidable kinds of snakes, such as the copperhead and black rat snake, and probably they are occasionally caught and eaten under natural conditions by each one of the mammalian predators mentioned. At the pond on the Reservation green herons ( Butorides virescens) were often present in spring, and great blue herons (Ardea hero- dias) often visited both this pond and the smaller one on the Rockefeller Tract. Both may have caught garter snakes. The numerous large bullfrogs present at both ponds also were potential predators on small garter snakes that ventured into the vicinity. Although many records are available of mammalian, avian and reptilian predators feeding upon Thamnophis sirtalis, the role of predation in general or of any one predator species in regulating its populations are still unknown. Even less is known about mortality in hibernation, or from drought, or disease. PARASITES The only ectoparasites observed on the garter snakes of my study areas were chiggers. These larval mites varied greatly in abundance according to the time and place. Abundance was greatest in hot, humid weather of early summer in moist situations with mesic vegetation. The garter snakes handled usually were not thoroughly checked for chiggers, but in early sum- mer most individuals carried them. Sometimes many hundreds were attached to one snake, their close-packed bodies forming a reddish encrustation ECOLOGICAL STuDY OF GARTER SNAKE 547 over the loose skin between the scales. Even with such heavy infestations the snakes suffered no visible ill effects. Loomis (1956) made an intensive field study of chiggers from 1948 to 1952, which was most concentrated on the Reservation, but included many areas in other parts of Kansas and in neighboring states. Of the many species present on the Reservation, he found only three attached to garter snakes: Trombicula alfreddugési (1596 individuals), T. lipovskyana (40 individuals), and T. kansensis (4 individuals). T. alfreddugési is the common pest chigger that attacks humans; it is remarkably catholic in its choice of hosts, utilizing many species of mammals, birds and reptiles, and occasionally certain amphibians. T. lipovskyana is a less common species that is similar in its ecology. T. kansensis is a member of a different subgenus. It is known from only a few localities, chiefly from relatively dry and rocky habitat, and deer mice (Peromyscus maniculatus) are the most frequent hosts, but the chigger has been found also on four species of snakes, including Tham- nophis sirtalis. Styphlodorine trematodes, so-called lung flukes, are found in many kinds of snakes including garter snakes. Insofar as known all members of this group have in addition to the vertebrate host of the adult, intermediate hosts for the developmental stages—first a water snail and then a frog. The eggs escape through the digestive tract of the snake and in aquatic situations hatch producing free-swimming larvae which to develop further must find the right kind of snail and embed themselves in its tissues. After completing this phase of its development the parasite produces asexually numerous free- swimming larvae of a second type. These parasitize tadpoles, remain in them after the tadpoles metamorphose, and complete their development only when the frog is eaten by a snake. In keeping with its liking for wet places and its frog-eating habits, Tham- nophis sirtalis is a frequent host of various styphlodorine flukes. Kinds recorded in the literature as having been found in T. sirtalis include: Dasymetra longi- cirrus, Lechriorchis insignis; L. plesientera, L. primus, L. tygarti, Ochetosoma megametricum, Zeugorchis aequatus, Z. curinus,, Z. megacystis, Z. synto- mentera, and Z. syntomenteroides. In the summer of 1958, Miss Peggy Lou Stewart checked many of the garter snakes caught on the Reser- vation and Rockefeller Tract for flukes. She found that most of the snakes caught in June and July had Zeugorchis megacystis in their mouths and gullets. In other months flukes were rarely found, but they may have been present farther down the digestive or respiratory tracts where they could not be seen without sacrificing the snake. Miss Stewart examined 326 garter snakes (T. sirtalis, T. radix and T. proximus) in the University of Kansas Natural History Museum, from localities in various parts of Kansas. Six of these preserved snakes of all three species had Zeugorchis megacystis in their lungs. This fluke is about 2.5 millimeters long with body .7 millimeters wide, and attenuate anteriorly (Stewart, 1960). None of the live garter snakes that were found to be infested with Z. megacystis showed any pathologic effects. In November, 1963, a large female T. sirtalis that had been in confinement several weeks, but had re- fused food and was of emaciated and sickly appearance, was found to have many flukes in its mouth. These were identified by Dr, William H. Coil as Zeugorchis syntomentera. 548 UNIVERSITY OF Kansas Pusxs., Mus. Nar. Hist. COMPOSITION OF THE POPULATION In nine broods born in captivity to females from the Reservation and Harvey County Park, there were 68 males and 53 females, but the sample is too small to permit much confidence in the 1:.78 male-female ratio, or even to indicate that males are more numerous than females in broods. In south- eastern Michigan Carpenter (1952:253), with a larger sample of 360 young from 20 broods, found a male-female ratio of 1:.91. The trend of the figures indicates that males are slightly more numerous than females at birth. In adult garter snakes the sex ratio in samples varies according to the sea- son. In 519 snakes of my autumn sample from the Reservation, representing the combined data of 15 years of trapping, there were 301 males and 218 females. The 1:.72 sex ratio is probably distorted. In autumn there is some sexual activity; hence females caught in traps may serve as bait to attract males. Often a pair was found in a trap, and occasionally one female was accompanied by several males. In late spring and summer, however, the sex ratio of trapped samples can- not be influenced by sexual attraction, since sexual activity wanes rapidly after a short and intensive breeding period immediately after emergence from hibernation in early spring (ended before trapping began). The totals for the several years of trapping from May through August were: males—154, females—254. Many of these were second-year individuals, which in spring and early summer were still adolescent, and often were still small enough to pass through the quarter-inch wire of the traps. In this respect the traps were probably somewhat selective, because adolescent males, lagging behind the females of the same age in their growth, would be able to escape from the traps after the females were too large to do so. To avoid the bias arising from this selectiveness, all those snakes of both sexes known or supposed to be second year individuals were eliminated. There remained 255 adults, third year individuals or older, of which 99 were males and 156 were fe- males. The 1:1.58 sex ratio indicated by these figures may be somewhere near the correct one, but probably there are various distorting factors. The home ranges of males, more than 50 per cent larger than those of females ac- cording to my calculations, would cause males to be caught oftener and in more traps than their female counterparts. Hence, males may be even less numerous than the ratio of actual captures indicates. Age structure of the population was calculated from a combination of the records of marked individuals recaptured and the size distribution of all the snakes captured. Each individual recorded was assigned to an age group, either on the basis of its known growth, if it had been recaptured, or arbitrarily on the basis of its size if actual growth records for it were not available. In summer, individuals late in their first year or early in their second year were in many instances too small to be held in the traps and therefore were not adequately represented. The sample used is therefore that drawn from fall and spring, when the second-year snakes are nearly all large enough to be caught. The distribution of 650 adults according to age and sex is shown in Table 11. The trend of figures in Table 11 suggests that in adult garter snakes of the population studied there is an approximately 50 per cent loss from one year to the next in any age group. The rate of loss is not detectably ECOLOGICAL StuDY OF GARTER SNAKE 549 Taste 11.—NuMBERS AND PERCENTAGES OF ADULT GARTER SNAKES IN DiF- FERENT AGE GROUPS IN A SPRING AND FALL SAMPLE Both Sexes Comtined Females Males Age Group Num- | Percent-| Num- | Percent-| Num- | Percent- bers ages bers ages bers ages second year... «..:.«- 323 49.7 153 50.1 170 49.9 third-year. t.0..5. 23. 162 25.0 69 22.6 93 27.0 fourthsyeartus .) eco: 97 14.9 40 13:1 57 16.5 fifthyears. raci- oc, 34 5.2 17 5.6 itr 4.9 sixthtyear:: 62h. 22. 17 2.6 14 4.6 3 9 seventh year........ 11 ests 9 2.9 2 6 eighthiyear.....-... 6 9 3 1.0 3 9 Wotalss Mak. as. ce 650 | 100 305 | 100 345 | 100 different in males and females. In a cohort of 100 second-year individuals the life expectancy is such that typically only one or two would survive into their ninth year. The rate of attrition seems to be neither slowed nor in- creased with advancing age. Actually, snakes more than five years old were caught in such small numbers that chance played a major part in their distribution and it is un- certain whether the rate of loss characteristic of the younger adults was still maintained. Senility theoretically might be a factor in the dying off of the oldest snakes, but no evidence of it was noted. All of the largest and oldest garter snakes examined appeared healthy and vigorous. Their large size, especially in the females, would seem to confer a distinct advantage in de- fense against certain natural enemies. In calculating the productivity of a population it must be taken into account that the adults comprise at least seven discrete annual age classes, each class with a different number of individuals and differing somewhat from all the other classes in fecundity. Some of the younger adult females do not ovulate, and those that do produce fewer ova than older adults. Table 12 has been compiled with attempt to allow for these differences be- tween age groups, though admittedly the data on which it is based are im- perfect and almost surely the trends would be altered somewhat with larger samples. Actually the number of young produced by the cohort considered in Table 12 would be fewer than 776 because some of the females would be eliminated by natural causes before the time of parturition, and of those that survived some would lose fetuses by abortion or resorption, or death at the time of birth. In the absence of evidence to the contrary, it must 550 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. TABLE 12.—THEORETICAL PRODUCTIVITY IN A Conort oF 100 ApuLT FEMALES IN EARLY SUMMER Percent- | Percent- age of age of Number | Number rN adult gravid Number of eggs of a a female individ- gravid per young popula- uals in female | produced tion age group second year........ 50 42 21.0 11.4 240 third: years: +. 26. 25 58 14.5 12.3 178 fourth year. ....... 12.5 92 11.5 16.0 184 frit heyeate ori: fcc 6.25 92 5.75 15.7 90 sixth years 26.. 22 3.12 100 3.12 15.5 48 seventh year....... 1.56 100 L350 15.5 24 eighth year........ .78 100 78 15.5 12 MOGALE fc .eye.cio. 49 TOOK? Realtors Aiea 58.21 13.33 776 be assumed that the mortality rate is uniform throughout the year, and if this is true the cohort would sustain a loss of near ten per cent during ap- proximately ten weeks of gestation, reducing the estimate of 776 young to only 698. In those females surviving to parturition a further ten per cent reduction may be conservatively estimated from infertile eggs and from embryonic and fetal mortality during development and at the time of birth. Consequently the number of live young produced might be in the neighbor- hood of 628. The rate of loss by natural causes in juveniles is almost certainly higher than the rate of loss in adults, and the smaller the young the more rapid would be the expected death rate. If the male to female ratio at birth is 1 to .9, the 628 young would include 297 females, and in a 21%-month period these would be reduced to 50. If there is a 64 per cent reduction in the first-year young, leaving 107 surviving one-year-olds, and if these are sub- ject to a further 53 per cent reduction in the following nine and a half months, the requisite number of breeding second-year females (50, replacing the second-year females in the original cohort of 100) would be obtained. In summary, a cohort of 100 adult females in May might be reduced to 90 by early August, but by then they would have given birth to about 297 female young, and there would then be also about 107 one-year-old female survivors from the previous year’s brood. Calculation of the number of male counterparts represented by these females is probably subject to a wider range of error. There would be about 331 newborn males; about 57 adult males (accepting the 1:1.58 ratio indicated by trapping as correct), and perhaps 107 one-year-old males. According to my calculations the population is increased by about 173 per cent at the time young are born. ECOLOGICAL STuDY OF GARTER SNAKE 5b NUMBERS From 1949 through 1957 the garter snakes recorded in my study were essentially those trapped along hilltop limestone outcrops in autumn, with a relatively small number caught in summer. However, for the last six years of the study, 1958 through 1963, extensive summer trapping was carried on, and the catches of adult snakes at locations away from the rock outcrops, were as follows: 49 in 1958, 173 in 1959, 153 in 1960, 152 in 1961, 81 in 1962 and 72 in 1963. These figures do not satisfactorily reflect the actual numbers present. Both trapping effort and the area covered by the trap lines differed somewhat from year to year. The majority of individuals recorded were caught only once, and new individuals continued to be caught, throughout the course of the study; therefore it was obvious that most of the population had not been included in the samples. Under these conditions “census” could be carried out only by obtaining a ratio within a given period of individuals previously caught and those captured for the first time. Snakes are not especially well adapted to census by the “Lincoln Index” method. Secretiveness, seasonal changes in habits, and dif- ferences in behavior between the sexes are all complicating factors. The relatively long time required to capture snakes in sufficient numbers for a sample is a serious drawback of the method; in the period of weeks or months usually required the population is changed by immigration, emigra- tion, reproduction and mortality. Each year’s data were used as a separate unit in my census computation. Obviously in the period May through November some shifting of individuals occurred. This would constitute a source of error; some marked individuals would have moved out of the study area while others moved in. Under- representation of marked individuals in the later samples would result in an erroneously high figure being obtained for the total population present. While this factor undoubtedly influences the results and increases their margin of error, its effect must be relatively minor, because study of the records of movements of individuals has shown that most maintain home ranges and stay in the same small area over periods of months. The capture-recapture census involves two sampling periods—a preliminary period in which a number of the animals are caught and marked, and a fol- low-up period in which the ratio of those previously marked to others is determined. An entire summer’s records could be divided in various ways, and many different figures could be obtained for essentially the same population. In my census computations the season’s records were divided into monthly units. In 1959, for instance, 24 snakes were caught and marked in the month of May. In June 37 were caught of which 5 were recaptures from the original group of 24 marked in May. The total population (P) present on the study area in May is indicated by the formula — = = or, P= 178. In: July..40 snakes were captured of which two weresamone the 87 recorded in June. Hence, for June — == ee or P=740. Similarly the July catch, on the basis of August recaptures, Ste a population of 252, and for August and September, figures of 646 and 273, respectively, were obtained. Actually the 52. UNIVERSITY OF Kansas Pusts., Mus. Nar. Hist. population of June was the same as that present in May minus the small per- centage that had been eliminated through natural causes in the interval. Each month’s census figure should have indicated a slight decline from that of the previous month if the figures had been highly accurate, but no such trend was evident, and a wide range of fluctuations results from the small size of the samples. The average of all five samples was 422, and this figure is probably more nearly correct than any of the separate census computations. If periods longer than a month are used as sampling intervals, more sub- stantial numbers of recapture records are usable in the computations. For instance, of the 24 snakes caught in May, 1959, nine were recaptured along with 149 new snakes in the period June through October, and the formula may P 158 be stated: —— —=-—— or P= 421. Or the preliminary period used may be 24 9 May-June and the follow-up period July-August-September-October resulting P 131 in a slightly changed ratio as follows: ———=-—— or P=524. With pre- 56 14 liminary periods successively lengthened to May-June-July, May-June-July- August, and May-June-July-August-September and with the follow-up period in each instance consisting of the remainder of the growing season, through October, separate census figures of 531, 504, and 422 were obtained. For this whole series of five censuses there was a total of 66 recapture records, against only 16 from the same set of data divided into one-month census in- tervals as in the preceding paragraph. For the longer periods the average of five capture-recapture censuses was 480, with a markedly smaller devia- tion than was obtained with the shorter census periods and fewer re- captures. The smaller deviation in- dicates that for the longer periods more adequate samples were ob- 5859 60 Sl 62 63 tained. But the increased interval YEAR also allows more time for disappear- ance of the marked snakes, and the Fic. 14. Diagram showing population yelatively high census figure may re- density of garter snakes calculated for fect due tothis fact the Reservation study-area by Hayne oe ae ee oe pis aes method of capture-recapture census. In Lincoln Index censuses it is Lower line shows population of adults | commonly observed that those com- in early summer of each year calculated _ putations based on only one recap- from actual “census.” Middle line ture or a few, are subject to wide shows numbers estimated to have been ; : : present in early summer, making allow- fluctuation, while computations based ance for the juveniles that were not on more recaptures tend to fall within caught in the traps; upper line shows an expected range. Therefore, in numbers estimated to have been present averaging several such census com- in late summer at time of annual maxi- mum after birth of young. NUMBER PER ACRE putations it seems a reasonable pro- cedure to weight each one according to the number of recapture records on which it is based. The weighted average for the five 1959 censuses, based on a total of 66 recaptures, as explained in the preceding paragraph, is 492. ECOLOGICAL STUDY OF GARTER SNAKE 553 A refinement of the Lincoln Index method of census since it was proposed by Hayne (1949:399) has been widely used by vertebrate zoologists. In this method a series of population computations is made; in the case of my 1959 data the May population was computed from the capture-recapture ratio in June; the May-June population was computed from the ratio in July, and so on. With successive census samples the ratio of recaptures steadily rises, as the pool of marked individuals grows. By projecting the trend of these increasing ratios the total population can be estimated. The actual procedure is rather involved and will not be explained in detail here; the reader is referred to the original publication, for a fuller explanation of the procedure and mathematical theory. For my 1959 samples the ratios of recaptures to total captures in June, July, August, September and October, respectively, were 8.2, 11.6, 28.6, 20.0, and 39.3 per cent, and the total number of snakes indicated was 357. Still another method of using the Lincoln Index consists of combining the figures from several separate sampling periods before the census computation is made. Thus for the summer of 1959 in five preliminary sampling periods (May, May-June, May-June-July, May-June-July-August, May-June-July-Au- gust-September) there were caught 24, 56, 94, 124, and 156 snakes, totalling 454, and in the five corresponding follow-up periods there were caught 158, 131, 96, 65 and 7—totalling 477, with recaptures 9, 14, 17, 16 and 10, PS): 4T7 = — or P= 656. 454 66 All four of the census methods described above are included in Table 13 for the sake of comparison, The Hayne method is probably the most re- liable and in most instances, but not all, it yielded the lowest census figures. totalling 66. Hence, TABLE 13.—PoPULATION OF ADULT GARTER SNAKES ON RESERVATION STUDY AREAS CALCULATED FROM CAPTURE-RECAPTURE DaTA BY FouR DIFFERENT METHODS Number Estimated numbers of snakes of Number| Esti- |__ snakes of mated Year in recap- acres ; Average census ture in area | Hayne Ange 8°! Weighted of samples | records*| sampled | method means | 2Vverage combined 7 samples 1958 49 3 324 268 290 287 413 1959 173 | 44 374 357 480 492 656 1960 153 28 374 435 888 676 761 1961 152 10 324 993 1427 1035 1208 1962 81 9 176 345 284 285 324 1963 72 14 176 222 189 186 211 ® Recapture records within the same month are not included, nor are those of snakes from any previous years. 554 UNIVERSITY OF KANSAS PuBLS., Mus. Nat. Hist. In all four methods, the trends for the six-year period are similar, and indicate great changes in the number of snakes from year to year. Beginning with a relatively low population in 1958 the numbers increased steadily through 1961, with the greatest gain coming from 1960 to 1961 and more than doubling the population in a year. In the last two years the trend was re- versed, with an even more rapid decrease, and in 1963 the population had decreased to a level even lower than that of 1958 (Fig. 12). Drastic changes in population level from year to year are to be expected in species like this garter snake which have a high reproductive potential, with large broods and with a fairly short time (two years at the minimum) between generations. Klimstra (1958:232) presented figures intended to show “cyclic” changes in populations of snakes, but his evidence was not entirely convincing because data for several species and genera were combined and data for part of the period of his observations originated from Iowa, the remainder from Illinois. In any case Klimstra’s data did not afford evidence of a regular cycle, but merely of rather large changes in numbers from year to year, with progressive trends continuing over several years. In the garter snakes of the Reservation population changes are probably controlled by a complex combination of factors, and the available records of weather and of associated animals provide few clues as to causality, It might be expected that the food supply would be one of the most critical factors, but food supply is in turn largely controlled by the weather. Ample and well distributed rainfall would seem to render conditions favorable to garter snakes, partly by promoting the development of lush vegetation for shelter, but especially by promoting the successful reproduction and dispersal of the leopard frog and other important prey species. Although the figure for the annual total of rainfall might have some significance, the seasonal distribu- tion is more important. Precipitation during the time of year that the snakes are hibernating affects the snakes by replenishing the water supply of creeks and ponds, but precipitation during the warmer half of the year, when the snakes are active is much more critical. Especially after birth of the annual crop of young, in late July and early August, precipitation must be adequate in amount and well distributed for the small garter snakes to thrive. Available records indicate that earthworms are the most important food, but in dry weather such as that in August, September and October of 1953 earthworms are usually well beneath the soil surface, out of reach of the young snakes, which, as a result are undernourished and presumably die off faster than they TABLE 14.—PRECIPITATION (IN INCHES) ON THE UNIVERSITY OF Kansas NAT- URAL History RESERVATION IN WINTER, EARLY SUMMER, AND AUTUMN OF Six DIFFERENT YEARS 1958 | 1959 | 1960 | 1961 1962 | 1963 November through April....| 14.72 | 14.14 | 12.14 | 16.79 | 14.09 | 5.47 May-June-July............ 21.61 | 10.59 | 10.38 | 18.95 } 14.36 | 11.06 August-September-October..}| 9.69 | 15.39 | 8.76 | 17.33 | 14.85 Galt ECOLOGICAL STUDY OF GARTER SNAKE 555 would under more favorable conditions. The peak population of garter snakes present in 1961 consisted mostly of young adults, born in the summer of 1959. Rainfall in August, September and October, 1959, totalled 15.39 inches, more than for the corresponding periods of most other years, and those young snakes that were recorded had made unusually rapid growth. In the autumn of 1961 precipitation was even higher, but by 1963 the population had decreased as a result of other, unfavorable factors. Seasonal precipitation for six different years at the Reservation is shown in Table 14. SUMMARY Over a 15-year period, 1948 through 1963, a population of the garter snake, Thamnophis sirtalis parietalis, was studied on the Uni- versity of Kansas Natural History Reservation, the northeasternmost section of land in Douglas County, and from 1958 through 1963 on the adjacent Rockefeller Experimental Tract, a quarter-section in Jefferson County. An equally intensive field study was carried on 140 miles to the southwest at Harvey County Park, 1959 through 1963. The populations studied were near the center of the species’ geo- graphic range, and were typical of the subspecies parietalis. Adult females ranged from 504 millimeters (snout-vent) and 21 grams to 950 millimeters and 410 grams, and adult males ranged from 387 millimeters and 14 grams to 678 millimeters and 110 grams. New- born young typically are about 168 millimeters and 1.9 grams. The relatively small size of newborn young is correlated with large litters and high infant mortality. The range of the species is more extensive than that of any other American snake, and includes much of North America except for its arctic and tropical portions. Over this large area the garter snake is represented by 11 named subspecies. There is notable stability geographically in numbers of body scale rows, numbers of labial scales, and perhaps in body size, although these characters are highly variable in some other species of garter snakes. Geographic variation is striking in the color, width and distinctness of the dorsal stripe and lateral stripes. The dorsal stripe especially is usually bright yellow contrasting with the dark color of adjacent areas, oc- cupying the middorsal scale row and half of each adjacent row. But in sirtalis and pickeringi the stripe is reduced and in some indi- viduals of pallidula it is absent. In most subspecies the dorsolateral area of each side is dark, almost black, but is notably paler in sirtalis, pallidula, ornata and infernalis. Pale crescent-shaped flecks con- fined mainly to loose skin between the scales and arranged in several 556 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. longitudinal rows, are yellowish, greenish or bluish in the more eastern subspecies, but in parietalis at least the lower rows of flecks are usually some shade of red, and all the more western subspecies have red in the pattern. When the snake is cornered or captured the red marks are most prominently displayed. Lateral stripes are narrower and duller than the dorsal stripe and they may be broken by dark marks as in semifasciata or may be obscured or altogether lacking in the somewhat melanistic concinnus and pickeringi. These same two subspecies are heavily marked with black ventrally, but most others have whitish chins blending to pale blue or green (some- times tinged with red) farther posteriorly on the ventral surface. The pattern with longitudinal striping in contrasting colors has adaptive value; it effectively conceals motion in the habitat of dense vegetation where the snakes are usually found. In its habits and habitats T. sirtalis is fairly typical of its genus. Almost everywhere that it occurs (except in the far north) it is sympatric with more specialized competitors of the same genus which must sometimes place a heavy strain on its resources. In the eastern half of the United States T. sirtalis is characteristic of dense vegetation in mesic habitats. On my study areas it was most often caught in tall prairie grass or in brush. However, farther west it is increasingly confined to marshland or other riparian situations. Compared with other snakes, the common garter snake is exception- ally cold-hardy, and can survive temperature as low as —3° C. Some remain active until after the advent of freezing weather in autumn and emerge relatively early in spring. They are active over a wide range of air temperatures, but seem to prefer a bodily temperature between 29° C. and 30° C. The critical thermal maximum is near 41°C. Amphibians (chiefly frogs) and earthworms make up most of the food. In my field study 200 food records were obtained. The leopard frog was locally by far the most important food species, with 120 recorded occurrences. There was a variety of other salien- tians (ranid, hylid, bufonid, microhylid), 5 earthworms, 6 cricetid mice, 2 voles, 1 bird and 1 snake (juvenal copperhead). Earth- worms are especially important as food for the first-year young, whereas adult frogs and miscellaneous other vertebrates are eaten chiefly by large garter snakes. Composition of the food varies greatly from time to time and place to place, depending on the relative availability of the favorite types of prey. The supply of earthworms, and frogs (mostly im- mature), which are the chief food sources, are subject to consider- ECOLOGICAL STUDY OF GARTER SNAKE 5Da able fluctuation. Growth rate of the snakes is hence subject to much variation. Some young make little gain by the time of their first hibernation in late autumn and presumably most of these do not survive. A gain of approximately 1.1 mm. per day in snout-vent length is most typical for this autumn period. In the following growing season the rate is usually a little more rapid. By mid-June a typical first-year garter snake has more than doubled in length, measuring approximately 380 millimeters and weighing about 14 grams. At this age males and females have diverged but little in growth, but by then the males are approaching sexual maturity and slow their growth abruptly. By autumn, at an age of 15 months, they average about 455 millimeters in snout-vent length, whereas females of the same age having continued their early rapid growth, have reached approximately 550 millimeters, on the average. Although daily length increments of 1.1 to 1.3 millimeters per day are most typical for growing young, some thriving individuals grow at more than twice this rate for periods of weeks or months. Other garter snakes of all sizes fail to make any gains over periods of months. Growth rates are especially depressed in drought sum- mers when food is scarce. On the Reservation the largest garter snakes (all females) from 900 to 950 millimeters in snout-vent length were from five to nine years old, according to growth records avail- able for some of them. In Thamnophis sirtalis there is sporadic sexual activity after the onset of coo] weather in autumn, and there is a more concentrated breeding season in early spring. Copulation occurs chiefly on the first warm day of spring that the snakes are active, and sexual activity wanes rapidly. The males carry on active sexual search and females are found mainly by scent. The female’s role in mat- ing is almost entirely passive, and it is not known whether she has a definite oestrus period. Sexual relations are promiscuous, and several or many males may simultaneously court the same female. Typically she is larger than any of her suitors. The male lies on the female, or alongside her, with rhythmic writhing movements, his cloacal region pressed against hers. When intromission has been attained, the male’s movements cease and he is dragged be- hind the female for periods of minutes as she moves about slowly. Males have been known to breed with several females in succes- sion. Usually as copulation begins the unsuccessful suitors dis- perse. Congealed semen distends the female’s cloaca after mating, and may function as a copulatory plug. Ovulation occurs many weeks after insemination, in May. Palp- 558 Unrversity oF Kansas Pusts., Mus. Nat. Hist. ing of 132 gravid females on my study areas indicated an average of 14.5 eggs. Some of the eggs—an estimated 4.6 per cent—are lost during pregnancy. Number of young is closely correlated with size and age of the female, and varies from an average of 12.0 in two-year-olds to 16.8 in four-year-olds. Furthermore less than half the two-year-olds produce eggs, but 100 per cent of those examined that were six years old or older were fecund. An extensive litera- ture indicates that occasionally litters are much larger than those recorded by me, with a maximum of more than 80 young. Undue attention to unusually large litters has resulted in bias toward ab- normally high figures in the literature. Zehr (1962:324) found (by dissection) an average of 12.9 embryos in 104 gravid females from New Hampshire, by far the largest series yet recorded. The average distance between successive captures, 696 feet for males and 562 feet for females, were interpreted as representing home ranges of 35.0 and 22.7 acres respectively. All these move- ments were for relatively short intervals, one month or less. For longer intervals distances were greater showing that home ranges are not necessarily permanent but may be altered with time. On the Reservation many or all of the garter snakes made migrations from their summer ranges to hilltop limestone ledges where deep clefts and crevices provided abundant hibernacula. Several such movements were in the neighborhood of a quarter mile. Insofar as known, individuals did not return to the same hibernaculum or to the same area of ledge in successive years. The striped pattern of Thamnophis sirtalis is highly effective for concealing motion; the snake is capable of rapid movement and is elusive in its normal habitat of dense vegetation. When unable to make its usual rapid getaway, a snake slowed by low temperature or cornered, may resort to bluff, coiling, flattening the body, dis- playing conspicuously the red markings between the scales, and striking. If restrained, it may deliver a painful bite and further discourages a predator by smearing its own body and that of the captor with feces and musk. A garter snake that is inadvertently grasped too far posteriorly will adroitly twist off the tail and while the latter performs lively wriggling movements distracting the cap- tor, the snake itself may escape. As a result of such incidents incomplete tails were common in the populations studied, especially in large adults, Important natural enemies include the red-tailed hawk (100 garter snake occurrences in 1820 pellets), Swainson’s hawk (6 garter snake occurrences among 57 prey items), blue racer (18 garter ECOLOGICAL STUDY OF GARTER SNAKE 559 snakes among 1339 food records), raccoon and opossum. Hawks, water birds and carnivorous mammals certainly include some of the chief natural enemies of the garter snake throughout its range, but their effects on populations are not known and there is little evidence as to what factors actually control populations. At least three species of chiggers and two species of lung flukes parasitize the local populations of garter snakes studied. At least nine other kinds of lung flukes have been found in Thamnophis sirtalis from other localities. None of these parasites produces any obvious ill effects in the snake host. In the adult population studied, males were less numerous than females, in a ratio of .61 to 1. The adults represent at least seven annual age groups. Typically from one year to the next a cohort of adults is reduced by approximately 50 per cent by natural mor- tality factors. Seemingly the rate of mortality changes little with increased age and size. Because of rapid population turnover and lack of efficient means for capturing the snakes a majority re- mained unmarked even after many years of field work on the areas most intensively studied. “Census” therefore had to be based on capture-recapture ratios. It was based entirely on the adults, but was projected to include first-year and second-year young poorly represented in the actual catch. Various errors inherent in the method and the material prevented a highly accurate census, but trends of the figures indicated constant change in population on the Reservation with a minimum of less than three per acre at the time of the annual low in the summer of 1963 and a maximum of ap- proximately 18 per acre at the time of the annual population peak in early autumn of 1961. The population made steady increase through 1958, 1959 and 1960 reaching a high point in 1961, then declined in 1962 and reached its lowest point in 1963. 560 UNIvERSITY OF Kansas Pusts., Mus. Nat. Hist. LITERATURE CITED ANDERSON, P. 1942. Amphibians and reptiles of Jackson County, Missouri. Bull. Chicago Acad. Sci., 6(11):203-220. BalLey, R. M. 1949. Temperature toleration of gartersnakes in hibernation. Ecology, 30: 238-242. BarBour, R. W. 1950. The reptiles of Big Black Mountain, Harlan County, Kentucky. Copeia, 1950: 100-107. BarTon, A. J. 1948. Snake litters. Herpetologica, 4:198. Batts, B. S. 1961. 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Zool., 4 (4 and 5):113-208. TaAyLor, W. E. 1892. Ophidia of Nebraska. Ann. Rept. Nebraska State Board Agric.:310- 357. TRIPLEHORN, C. A. 1955. Notes on the young of some North American reptiles. Copeia, 1955(3):248-249. Unter, F. M., Cotram, C. and Cuarkg, T. E. 1939. Food of snakes of the George Washington National Forest, Virginia. Trans. 4th Amer. Wildlife Conf., Amer. Wildlife Inst., Washing- ton D. C., 1939:605-632. WALLACE, G. J. 1938. A garter snake with a litter of 73 young. Copeia, 1938(4):203. WARBURTON, F. 1950. Notes on the vertebrates of the Sault Ste. Marie region of Ontario. Canadian Field Nat., 64:192-200. WELTER, W. A. and Carr, K. 1939. Amphibians and reptiles of northeastern Kentucky. Copeia, 1939 (3):128-180. WERLER, J. E. 1951. Miscellaneous notes on the eggs and young of Texas and Mexican reptiles. Zoologica, 36(1):37-48. WERNER, W. E., JR. 1959. Amphibians and reptiles of the Thousand Islands region, New York. Copeia, 1959(2):170-172. Woon, J. T. 1945. Variation in lengths of newly born garter snakes. Copeia, 1945: Waricut, A. H. and Bisuop, S. C. 1915. A biological reconnaissance of the Okefinokee Swamp in Georgia. II. Snakes. Proc. Acad. Nat. Sci. Philadelphia, 67:139-192. Waricut, A. H. and Wright, A. A. 1957. Handbook of snakes of the United States and Canada. Comstock Publ. Associates, Cornell Univ. Press, Vol. 2, pp. i-ix and 565-1106. ZeuHR, D. R. j 1962. Stages in the normal development of the common garter snake, Thamnophis sirtalis sirtalis. Copeia, 1962(2):322-329. ZORICHAK, J. L. 1953. The ruffed grouse as a snake eater. Flicker, 25(1):2-3. Transmitted May 22, 1964. L] 30-3965 Vol. 138. Vol. 14. (Continued from inside of front cover) Variation in the muscles and nerves of the leg in two genera of grouse (Tympanuchus and Pedioecetes). By E. Bruce Holmes. Pp. 363-474, 20 figures. October 25, 1962. A new genus of Pennsylvanian Fish (Crossopterygii, Coelacanthiformes) from Kansas. By Joan Echols. Pp. 475-501, 7 figures. October 25, 1963. Observations on the Mississippi Kite in "southwestern Kansas. By Henry S. Fitch. Pp. 5038-519. October 25, 1968. . Jaw musculature of the Mourning and White-winged doves. By Robert L. Merz. Pp. 521-551, 22 figures. October 25, 1968. Thoracic and coracoid arteries in two families of birds, Columbidae and See: By Marion Anne Jenkinson. Pp. 553-5738, 7 figures. March The breeding birds of Kansas. By Richard F. Johnson. Pp. 575-655, 10 figures. May 18, 1964. The adductor muscles of the jaw in some primitive reptiles. By Richard C, Fox. Pp. 657-680, 11 figures in text. May 18,.1964. Index. Pp. 681-694. us 2. 3. Five natural hybrid combinations in minnows (Cyprinidae). By Frank B, Cross and W,. L. Minckley, Pp. 1-18. June 1, 1960. A. distributional study of the amphibians of the Isthmus of Tehauntepec, México. By William E. Duellman. Pp. 19-72, plates 1-8, 3 figures in text. August 16, 1960. A new subspecies of the slider turtle (Pseudemys scripta) from Coahuila, México. By John M. Legler. Pp. 78-84, plates 9-12, 3 figures in text. August 16, 1960. Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, plates 13-20, 26 figurs in text. November 30, 1960. Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and Rocky Mountains. By Henry ’S. Fitch and T. Paul Maslin. Pp. 289-308, 4 figures in text. February 10, 1961. Fishes of the Wakarusa river in Kansas. By James E. Deacon and Artie L. Metcalf. Pp. 309-322, 1 figure in text. February 10, 1961. Geographic variation in the North American cyprinid fish, Hybopsis gracilis. Y By Leonard J. Olund and Frank B. Cross. Pp. 328- 348, plates 21-24, 2 figures in text. February 10, 1961. Descriptions of two species of frogs, genus Ptychohyla; studies of Ameri- can hylid frogs, V. By William E, Duellman. Pp. 349-357, plate 25, 2 figures in text. April 27, 1961. Fish populations, following a drought, in the Neosho and Marais des Cygnes rivers of Kansas. By James Everett Deacon. Pp. 359-427, plates 26-30, 8 figures. August 11, 1961. Recent soft-shelled turtles of North America (family Trionychidae). By Rope Webb. Pp. 429-611, plates 31-54, 24 figures in text. February Index. Pp. 613-624. 1. 2, 8. Neotropical bats from western México. By Sydney Anderson. Pp. 1-8. October 24, 1960. Geographic variation in the harvest mouse. peabody megalotis, on the central Great Plains and in adjacent regions, y J. Knox Jones, Jr., and B. Mursaloglu. Pp. 9-27, 1 figure in text. July a 1961. Mammals of Mesa Verde National Park, Colorado. By. recited Anderson, Pp. 29-67, plates 1 and 2, 3 figures in text. July 24, 196 4, A new subspecies of the black myotis (bat) fom ee Mexico. By E. 6. ah 12. See Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December North American yellow bats, ‘““Dasypterus,” and a list of the named kinds of the genus Lasiurus Gray. By E, Haymes. Hall and J. Knox Jones, Jr. Pp. 78-98, 4 figures in text. December 29, 1961 Natural history of the brush mouse (Peromyscus boylii) in Kansas with description of-a new subspecies. By Charles A. Long. Pp. 99-111, 1 figure in text. December 29, 1961. Taxonomic status of some mice of the Peromyscus boylii group in eastern Mexico, with description of a new subspecies... By Ticul Alvarez.. Pp. 113- , 1 figure in text. December 29, 1961. r%e es subspecies of ground squirrel (Spermophilus spilosoma) from Ta- maulipas, Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962. Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J. ens Jones, Jr., and Ticul Alvarez. Pp. 125-133, 1 figure in text. March 7, A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene, of Texas. By E. Raymond Hall and Walter W. Dalquest. Pp. 135-138, 2 figures in text.. April 30, 1962. A new subspecies of wood rat (Neotoma) from northeastern Mexico. By Ticul Alvarez. Pp. 139-148, April 30, 1962. Noteworthy mammals from Sinaloa, "Mexico. By J. Knox Jones, Jr., Ticul een and M. Raymond Lee. Pp. 145-159, 1 figure in text. May 18, (Continued on outside of back cover ) 18. 14, 15. 16. Lhe Vol. 15.1. 2. 3. 4, 5. 6. 7. 8. 9. 10. Vol. 16. 1. (Continued from inside of back cover) A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, 1 figure in text. May 21, 1962. The mammals of Veracruz. By E. Raymond Hall and Walter W. Dalquest. Pp. 165-862, 2 figures. May 20,1963. The recent mammals of Tamaulipas, México. By Ticul Alvarez. Pp. 363- 473, 5 figures in text. May 20, 19638. f£ A new subspecies of the fruit-eating bat, Sturnira ludovici, from western Meno: By J. Knox Jones, Jr., and Gary L. Phillips. Pp. 475-481, March Records of the fossil mammal Sinclairella, Family Apatemyidae, from the ees and Orellan. By William C. Clemens. Pp. 483-491. March 2, More numbers will appear in volume 14, The amphibians and reptiles of Michoacan, México. By William E. Duell- man. Pp. 1-148, plates 1-6, 11 figures in text. December 20, 1961. Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962. ‘A new species of frag (Genus Tomodactylus) from western México. By Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962. Type specimens of amphibians and reptiles in the Museum of Natural His- tory, the University of Kansas. By William E. Duellman and Barbara Berg. Pp, 183-204. October 26, 1962. Amphibians and Reptiles of the Rainforests of Southern El Petén, Guatemala. By William E. Duellman. Pp. 205-249, plates 7-10, 6 figures in text. Oc- tober 4, 1963. A revision of snakes of the genus Conophis (Family Colubridae, from Middle seca): By John Wellman. Pp. 251-295, 9 figures in text. October 4, A review of the Middle American tree frogs of the genus Ptychohyla. By brace pas Duellman. Pp. 297-349, plates 11-18, 7 figures in text. October Natural history of the racer Coluber constrictor. By Henry S. Fitch. Pp. 351-468, plates 19-22, 20 figures in text. December 30, 1963. A review of the frogs of the Hyla bistincta group. By William E. Duellman. Pp. 469-491, 4 figures in text. March 2, 1964. An ecological study of the garter snake, Thamnophis sirtalis. By Henry S. Fitch. Pp. 493-564, plates 23-25, 14 figures in text. May 17, 1965. More numbers will appear in volume 15. Distribution and taxonomy of Mammals of Nebraska. By J. Knox Jones, Jr. Pp. 1-356, pls. 1-4, 82 figures in text. October 1, 1964. More numbers will appear in volume 16. MUS. COMF. - yoo kOe. ARTs Sn) pie wa) OS ae eerie LL Pee ae UNIVERSITY OF KANSAS PUBLICATIONS j)p 24 4 MvusEUM OF NATURAL HISTORY . HARVARD Volume 15, No. 11, pp. 565-575, 3 figs. UNIVERSITY May 17, 1965 Breeding Cycle in the Ground Skink, Lygosoma laterale BY HENRY S. FITCH AND HARRY W. GREENE UNIVERSITY OF KANSAS LAWRENCE 1965 UNIVERSITY OF KANSAS PUBLICATIONS, MusEUM OF NATURAL HIsTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 15, No. 11, pp. 565-575, 3 figures in text Published May 17, 1965 UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY HARRY (BUD) TIMBERLAKE, STATE PRINTER TOPEKA, KANSAS TY, MUS. COMPF. =OC e . . | IR DAR » Breeding Cycle in the Ground Skink,“ Lygosoma laterale DEC 31 1900 BY HARVARD HENRY S. FITCH and HARRY W. GREENE UNIVERSITY, The biology of the ground skink has been made fairly well known through the efforts of many workers, and especially the studies of Lewis (1951) and Johnson (1953). These latter studies were made at Houston, Texas, and New Orleans, Louisiana, respectively, both localities near the Gulf Coast in the southern part of the species’ range. Certain important questions were raised but not definitely answered by these studies. Our study was therefore undertaken with the hope of attaining a better knowledge of the breeding cycle of this small skink. We were especially interested in the repro- ductive potential, specifically, in the time required to attain breed- ing maturity, the effect of size of female on the number of eggs in her clutch, geographic variation in size of clutch, and the seasonal timing of breeding. Because of the great abundance and extensive geographic range of the ground skink, it was considered an almost ideal subject for such investigation. In our joint project a total of 523 specimens in The University of Kansas Natural History Museum, the Fort Worth Children’s Museum, the Stephen F. Austin State Teachers College collection, and the private collection of Harry W. Greene were examined, from Kansas, Oklahoma, Texas, Missouri, Arkansas, Louisiana, Mississippi, and Georgia. These included several substantial series, notably 22 from near Lewisville, Lafayette County, Arkansas, August 16 and 20, 1926, 31 from Texas, June 22, 27 and 28, 1930, 44 from Huntsville, Walker County, Texas, April 2 and 3, 1963, and 91 from Wister Dam, Le Flore County, Oklahoma, August 11 to 15, 1963. Reproductive cycles differ in details in different genera and species of reptiles. In the continental United States the prevailing pattern is that of a spring (and/or fall) breeding season, and ovulation in early summer, with the hatchlings or newborn young appearing in Jate summer or early autumn. One clutch or litter per year has been found to be the most general rule. However, in recent years many species of common lizards, especially iguanids occurring in the southern half of the country, have been found to deviate from this pattern in producing two or more clutches in a single season, for example Holbrookia texana (Cagle, 1950:230), Anolis carolinensis (Hamlett, 1952:184), Sceloporus undulatus (Crenshaw, 1955:273), Crotaphytus collaris (Fitch, 1956: 237), Cnemidophorus sp. (Milstead, 1957:439; Fitch, 1958:36), Sceloporus olivaceus (Blair, 1960:89), Uta stansburiana (Tinkle, 1961:230), and Ger- rhonotus liocephalus ( Burkett, 1962:211). In his study of Lygosoma, Johnson (op. cit.) found that some adult females (567 ) 568 UNIVERSITY OF KANSAS PuBLs., Mus. Nar. Hist. contained simultaneously both oviducal eggs and enlarging follicles (two milli- meters or more in diameter) and he suggested the possibility that such indi- viduals might produce two clutches per season. This idea was further supported by Johnson’s finding of females containing oviducal eggs in every month from March through August. There is no true hibernation in this species in the southern part of its range. Lewis (op. cit.:284) wrote that at Houston even in mid-winter on warm sunny afternoons the skinks could be found active in the open. Although intermittent periods of inactivity are enforced by the arrival of winter storms and cold fronts, the populations along the Gulf Coast have a relatively long growing season. In winter both Lewis and Johnson found their samples to consist almost entirely of adult skinks, with a few well grown young. Since Johnson found two sizes of young in summer, he was uncertain whether young attained adult size in the first year or in the second year. In a series of 72 skinks that we examined from Texas, collected in late March and April (mostly from Huntsville, April 2 and 3) only three were below minimum adult size and these were well grown (34, 31 and 29 milli- meters snout-vent). These undersized individuals may be interpreted as young that hatched unusually late in the season or that failed to grow at the usual rate. The young present in summer have hatched at different times, and hence do not constitute a clear cut size group, especially since growth is re- markably rapid in the early weeks of life. Also, a series of 64 skinks collected in Texas in June are all adults. A series collected in Lafayette County, southwestern Arkansas, August 20, 1926, and another series collected by the senior author at Wister Dam, Le Flore County, southeastern Oklahoma, August 11 to 15, 1963, are combined because they represent approximately the same latitudes and times of year, The combined series numbers 113 skinks. Thirty-nine per cent are young of the year and make up a size group fairly distinct from the adults but covering a wide size range—from hatchlings to adolescents (Fig. 2). Since egg-laying was not quite completed at the time of collection, it is evident that many young of the year, perhaps the majority of the second brood, were still not hatched, whereas the oldest young, having hatched in late June, as forerunners of the first brood, had grown for six or more weeks and were already approaching the size of the smallest adults. The young of this combined series show some tendency to bimodality, but the tendency is slight, and perhaps most of these young or all of them represent the hatch of the first brood. The rate of growth and maturation in the developing ova cannot be definitely determined from the available records, but can be inferred from the information available for other kinds of lizards. In the collared lizard (Crotaphytus collaris) a female living under natural conditions was recorded to have laid two successive clutches with an interval of three weeks or a little more (Fitch, 1956:237). Blair (1960:89) found that an interval of about a month between clutches was usual in the rusty lizard (Sceloporus olivaceus). Johnson (1960:298) found that in north-central Texas the greater earless lizard (Holbrookia texana) produced on the average five clutches of eggs in the course of the breeding season, with an average interval of about 35 days be- tween clutches. When the Huntsville series of Lygosoma was collected, several adults in- cluding three females of typical appearance, were kept alive to determine when they would produce eggs. Doubtless their normal cycles were disturbed BREEDING CYCLE IN THE GROUND SKINK 569 by the conditions of captivity. One produced a clutch in late April and another laid on May 31. Other clutches may have been laid and eaten before they were discovered as all the lizards were kept in the same container. When the eggs deposited on May 31 were discovered on the following day, a skink was eating one of them. On August 5 all three females were killed and examined and their ovaries were found to be small. Available evidence suggests that the eggs enlarge rapidly after the breeding season has begun. Of Kansas females two collected on April 1, 1930, had oviducal eggs and a third had ovarian eggs four millimeters in diameter, although these lizards must have emerged from hibernation only a short time before. In normal years in northeastern Kansas these skinks first appear near the end of March. In 1930 mean temperature for March deviated only .1 degree Fahrenheit from the sixty-year average (Flora, 1948:181). Of the 71 adult females of Lygosoma examined and collected in May, June, and July, 42 had oviducal eggs, 16 had enlarged ovarian eggs, and 13 had small ovaries, suggesting that the period of rapid growth of the eggs in the ovaries is shorter than their period of retention in the oviducts. Johnson (1953:22) reported embryos 1.54 to 3.10 millimeters in length in oviducal eggs, indicating that the early stages of embryonic development are passed in the oviducts before the eggs are laid, and some other species of Lygosoma are viviparous. In L. laterale oviducal eggs have been reported as early as March 25 in Mis- sissippi (Cook, 1943:19) and April 7 and April 10 in Texas (Lewis, op. cit.: 235). Werler (1951:39) recorded clutches laid at San Antonio on April 19, April 27, and May 1. Recorded incubation periods are somewhat variable but most of those recorded approximate one month. Therefore early broods of young might be expected to appear in late May in the vicinity of the Gulf Coast. Johnson (op. cit.:18) found no hatchlings before June, but his May sample was relatively small (15). Lewis (op. cit.:237) wrote that at Houston in 1947 hatchlings were first noted on June 30. Johnson found that females having oviducal eggs often also had several ovarian eggs considerably enlarged, evidently destined to form a second clutch, with their development already well underway. All things considered, there must be an interval of somewhat more than a month, perhaps five weeks, between successive clutches in the breeding season. In the region of the Gulf Coast such an interval would allow ample time for four broods that might be spaced about as follows: late April, end of May, beginning of July, first half of August. Because most females have enlarged ova in either ovaries or oviducts or both in this entire period, it seems unlikely that fewer than three clutches would be produced by most individuals. Farther north the breeding season becomes progressively shorter. Of 19 females collected in Kansas in April, five had oviducal eggs, five had ovarian eggs two to four millimeters in diameter, and the remaining nine had small ovaries. Of six collected in May two had oviducal eggs, two had enlarging ovarian eggs and the remaining two had small ovaries. Two females collected on July 17 and July 19 and three collected on August 17 all had small ovaries. On July 9 at The University of Kansas Natural History Reservation a clutch of three eggs of Lygosoma was found associated with a communal nest of the five-lined skink (Fitch, 1954:70). The eggs of the ground skink were within a few days of hatching and therefore must have been laid not later than mid- June. On the Reservation a gravid female was recorded on May 29, 1958. Two other females collected in Douglas County, Kansas, on May 30, 1960, 570 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. were both gravid but in one of them the ova were only 3.5 millimeters in diameter, hence probably several weeks short of laying. On the Reservation a hatchling was seen on July 29, 1957, and a juvenile of 29 millimeters was caught on August 20. Of adult females collected in southern Missouri, six taken in the latter half of April all had ovarian follicles 2.5 to 4.5 millimeters in diameter, and others with oviducal eggs were collected on May 3, June 19, and June 26 (3). Of 16 ground skinks collected in southern Missouri in late March and April, six were immature, having the following snout-vent lengths: 27, 30, 32, 34, 34, 37. Smith (1961:169) stated that in southern Illinois eggs are laid in July and hatchlings are abundant in late August. These scanty records indicate that even in the northern part of the range egg-laying probably extends from some time in April into July—allowing ample time for at least LATE AUGUST two broods. The relatively long and (ARKANSAS AND severe winter in the northern part of OKLAHOMA) the range has the effect of compressing the breeding season by preventing breeding in early spring and prevent- ing it in late summer and early au- tumn, thus enforcing on the popula- tion a more uniform breeding schedule than prevails in the southern part of the range. Fic. 1. Histograms showing size and breeding status of adult female ground skinks in three collections. Open col- umns represent specimens in which ova are small (all less than two milli- meters in diameter )—non-breeders or those in an early stage of the breeding cycle; stippled columns represent specimens having enlarged ovarian follicles (two to six millimeters in diameter); black columns represent specimens having oviducal eggs. In early April (lowest figure) most fe- EARLY APRIL males are in an early stage of the (TEXAS) breeding cycle, with ovarian eggs still not mature. Ten weeks later most females are in a late stage of the breeding cycle, with eggs, presumably their second clutches for the season, nearly ready to be laid. By late Au- gust, in Arkansas and Oklahoma, the breeding season is over except for a few of the smallest and youngest fe- males, which lag behind the majority 40 50 60 SNOUT VENT LENGTH: mm. in their breeding schedules. In the series of skinks from Huntsville, Texas, representing early spring, the 28 adult females fall into three distinct groups as regards their reproductive status. Two of near average size (44 and 45 millimeters snout-vent) have NUMBER OF SPECIMENS oviducal eggs that seem almost ready to be laid. Twenty-one others have smaller, ovarian eggs. Those eggs obviously represent clutches that would have ~ BREEDING CYCLE IN THE GROUND SKINK 571 matured and would have been laid somewhat later in the season. In this group the ova tend to fall in a fairly narrow size range; in 19 skinks they are from three to five millimeters in diameter. In two others eggs are two and one-half and two millimeters in diameter, In the third group, with six females, ovaries are still small, and three of these females are the smallest of the entire series; hence their retardation can probably be attributed to immaturity. Of the 37 adult female skinks from Texas collected in late June 35 (94.5 per cent) were gravid. Of these 35, 28 had oviducal eggs and probably were near the time of egg-laying, and the remaining individuals had ova 5, 5, 5, 3, 2% 2% and 2 millimeters in diameter. First clutches probably had already been laid, and some individuals, notably the two with small ovaries, may have already laid their second clutches. Oviducal eggs in late June probably repre- sented clutches that would have been deposited in early July. Development of the later clutches may be more rapid than development of first clutches, because of the higher temperatures prevailing in summer. Of six adult females collected in Arkansas in July (no definite dates re- corded) four had oviducal eggs and the other two were not breeding. In a female from Texas obtained on July 16 and in three collected on August 17, there were neither enlarging ova nor oviducal eggs, but two others collected on July 20 and August 7, both had oviducal eggs. In the Arkansas-Oklahoma series, representing dates from August 11 to August 20, only the two smallest (40 and 42 millimeters snout-vent) of the 42 adult females had oviducal eggs representing late clutches, and none had enlarging ovarian eggs, indicating that at these localities the breeding season was ending by mid-August. In the ground skink three is the most frequent number of eggs produced at one laying, but recorded clutches vary from one to seven. Johnson (op. cit.:19) found an average of 3.3 + .05 oviducal eggs in 31 females from Louisiana. Ten additional females from Louisiana examined in our study contained an average of 2.9 eggs. We examined 57 females from Texas that were gravid (having either oviducal eggs or enlarged ovarian follicles) and these had an average clutch of 3.02 + .15. Lewis (op. cit.:235) found an average of 2.82 eggs in 11 females from the vicinity of Houston. But, in 17 females from Arkansas and Oklahoma the average was only 2.35 + .20. In 17 females from Kansas and 14 from Missouri the clutch averaged 3.77 + .24. Although smaller than might be desired, this northern series seemed to have a significantly larger number of eggs per clutch than any from the more southern populations. Johnson (op. cit.:19) concluded that a significant correlation between Fic. 2. Lengths of immature ground skinks in a collection from Arkansas and Oklahoma in late August. Those of sizes up to 36 millimeters are al- most certainly young of the year hatched early in the summer. At the time of collection some eggs were still to be laid, and doubtless many others laid in late July or early August had not yet hatched. Hence young of the second brood probably are poorly 20 30 40 represented or not represented at all in SNOUT VENT LENGTH: mm. sree NO. OF SPECIMENS 5t2 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. number of eggs per clutch and snout-vent length of female did not exist in the series that he examined, as the regression coefficient was .069. However, the trend of his data did suggest a correlation, with the two-egg clutches found mostly in the smaller females, and the four- or five-egg clutches found in the larger females. In our study we found a definite positive correlation between size of female and number of eggs in clutch, as follows: 11 females 54 to 59 millimeters averaged 3.63 + .41 eggs 45 females 48 to 53 millimeters averaged 3.76 + .16 eggs 61 females 42 to 47 millimeters averaged 2.65 + .11 eggs 13 females 36 to 41 millimeters averaged 1.84 + .18 eggs Or, arranging the same data differently, grouped according to the number of eggs in the clutch: 10 females with l1-egg clutches averaged 43.3 + 1.31 millimeters 86 females with 2-egg clutches averaged 44.6 + .57 millimeters 43 females with 3-egg clutches averaged 47.0 + .62 millimeters 26 females with 4-egg clutches averaged 49.4 + .59 millimeters 11 females with 5-egg clutches averaged 50.5 + 1.78 millimeters 8 females with 6-egg clutches averaged 52.6 millimeters 1 female with 7-egg clutch was 52 millimeters The correlation between number of eggs in clutch and snout-vent length of female is further clarified by Fig. 3. It would perhaps be correct to say that the smallest adult females always have minimal numbers of eggs per clutch, and that with increasing average size proportionately larger clutches are eo 0e 00 Bo NUMBER OF OVA 40 45 50 55 60 SNOUT-VENT LENGTH: mm. Fic. 3. Number of ova and size in fecund ground skinks. On the average, the larger females produce more eggs, but occasionally large females produce small clutches. produced, but large females occasionally produce clutches with a small number of eggs perhaps because of senility, lateness of season or for other reasons. The lack of opportunity for individuals to produce successive clutches in one season in the northern part of the range because of the relatively short BREEDING CYCLE IN THE GROUND SKINK 51S breeding season is in part compensated for by a greater number of eggs per clutch. Larger size in northern populations may be a factor also. Over the extensive geographic range of Lygosoma laterale no subspecies have been named and geographic variation has not been demonstrated. In our study we found that ground skinks from Kansas differed slightly in general appearance from those of the Gulf Coast, and averaged slightly larger but the difference is not necessarily indicative of significant geographic variation. Table 1 shows the average sizes of adult males and females (those 36 millimeters or more in snout-vent length) in the larger series examined by us. It is shown that fe- males are approximately four millimeters longer than their male counterparts. Differences in age structures of the population samples involved are probably more important than geographic variation in affecting the figures. This is illustrated by the two series from Texas. Almost all of the skinks collected in early April were adults, but many of them were recently matured indi- viduals of the previous years’ brood. The June sample represents essentially the same population after approximately ten weeks of additional growth; both sexes had made substantial gains, but the females had grown more than the males. TABLE ]1.—SNOUT-VENT LENGTHS IN MILLIMETERS OF MALE AND FEMALE GROUND SKINKS IN SEVERAL SERIES Locality Females Males RAN BAS SR re vteteiss toiaity repcionle’ ha aged a haeess 48.9= .8lin37 | 43.0+ .84in 21 OKiaghomay es. c seca nc Loree 46.3 .47in 37 | 42.3 .53:in 27 IATKANISA SR shat: te ocieeraks Atel ween 46.4 .74in 27 | 42.1+ .69 in 22 Louisiana (Johnson, 1953:23; females with ovidueal eggs only)......... A735 881m 32y | oree ieee MPOKAS| (ANCA PTIL) ec toctve oc Yano yee as ae 45.3+1.24in 19 | 42.6+ .46in 31 Alexais | (im"JUNE) ah rateroees atten once eh 47.2= .67in 37 | 48.2 .62 in 27 In keeping with their larger size, the June-taken females from Texas averaged slightly more eggs per clutch—3.0, as compared with 2.76 for the April-taken females. In this southern part of the range June is the peak of the breeding season and the clutches produced that month are often both preceded and followed by others, Late clutches, produced as the breeding season wanes, probably tend to be smaller than others produced earlier. In our Arkansas- Oklahoma sample, eight females collected in April averaged 2.88 enlarged follicles or oviducal eggs, but in six females collected in July and August there was an average of only 1.83. It might be expected that at latitudes where the growing season is shortened to the extent that broods are eliminated, the size of clutch would increase abruptly. In summary, the ground skinks of the Gulf Coast have a long growing season with hibernation short and intermittent. There is ample time for at least four broods and some individuals possibly produce more. The majority of females have developing ovarian follicles in early April and first clutches 574 UNIVERSITY OF KANSAS PuBis., Mus. Nat. Hist. of eggs are laid from late April through May. Some young must hatch before the end of May but none has been recorded until June. A few females are several weeks ahead of the majority in their breeding schedule; others are still not sexually mature in early spring and their breeding schedule is several weeks behind that of the majority. In the northern part of the range at the latitude of Kansas the breeding season is relatively short, but the average female probably produces at least two clutches in the seven-months growing season. At this latitude egg-laying occurs in late April, June and July. At the latitude of southern Oklahoma and Arkansas egg-laying extends into early August. Three is the modal number of eggs per clutch but in the north clutches with four, five, and even six eggs are common. There is positive correlation between size of female and number of eggs per clutch. Those females that are of minimum adult size or only slightly larger produce clutches with fewer than the average number of eggs. The largest females produce the largest clutches but occasionally some of them produce small clutches. The time of year also affects number of eggs per clutch; toward the end of the breeding season clutches have fewer eggs on the average, even though females average larger then. Financial assistance from the National Science Foundation (G-16104) is acknowledged. For the loan of specimens from the collections in their care, and for other courtesies extended to us in the course of this study we are indebted to Mr. John R. Preston of the Childrens’ Museum, Fort Worth, Texas, and to Dr. William E. Duellman and Mr. Charles J. Cole of The University of Kansas Museum of Natural History. LITERATURE CITED Bua, W. F. 1960. The rusty lizard. Univ. Texas Press, Austin, xvi + 185 pp. BurkKETT, R. D. 1962. Two clutches of eggs in the lizard, Gerrhonotus liocephalus infer- nalis. Herpetologica, 18(3):211. CAGLE, FR. 1950. Notes on Holbrookia texana in Texas. Copeia, 1950(38) :230. Cook, F. A. 1948. Alligators and lizards of Mississippi. Surv. Bull., Mississippi State Game and Fish Comm., pp. 1-20. CRENSHAW, J. W., JR. 1955. The life history of the southern spiny lizard, Sceloporus undulatus undulatus Latreille. American Midl. Nat., 54( 2) :257-298. imc, HS: 1954. Life history and ecology of the five-lined skink, Eumeces fasciatus. Univ. Kansas Publs. Mus. Nat. Hist. 8( 1) :1-156. 1956. An ecological study of the collared lizard (Crotaphytus collaris). Univ. Kansas Publs. Mus. Nat. Hist. 8(3):218-274. 1958. Natural history of the six-lined racerunner (Cnemidophorus sex- lineatus). Univ. Kansas Publs. Mus. Nat. Hist. 11(2):11-62. Fora, S. D. 1948. Climate of Kansas. Report Kansas State Board Agric., 67, xii + 320 pp. HAMLETT, G. W. D. 1952. Notes on breeding and reproduction in the lizard Anolis carolinen- sis. Copeia, 1952 (3):183-185. BREEDING CYCLE IN THE GROUND SKINK 575 JouHNson, C. 1960. Reproductive cycle in females of the greater earless lizard, Hol- brookia texana. Copeia, 1960(4):299-300. JoHNsON, R. M. 1953. A contribution on the life history of the lizard Scincella laterale (Say). Tulane Studies in Zoology, 1(2):11-27. Lewis, T. H. 1951. The biology of Leiolopisma laterale (Say). American Midl. Nat., 45:232-240. MitsteEap, W. W. 1957. Some aspects of competition in natural populations of whiptail lizards (genus Cnemidophorus). Texas Jour. Sci. 9(4) :410-447. SmiTH, P. W. 1961. The amphibians and reptiles of Illinois. Bull. Illinois Nat. Hist. Surv., 28(1):1-298. TINKLE, D. W. 1961. Population structure and reproduction in the lizard, Uta stans- buriana stejnegeri. American Midl. Nat., 66(1):206-234. WERLER, J. E. 1951. Miscellaneous notes on the eggs and young of Texas and Mexican reptiles. Zoologica, 36(1):37-48. 80-3963 — - ol ar: a Stacuy a a is - buat h a a oF JT: , ieee an of ww ae 2d : 4 —— oe ene bri ce, sian ARS Rae kG wild © Galé br Ja men td > & aE _ ve - im Oe ~ .' . gy ara ler lan ap) A =. wey : : es ih Lewy Cdelaent ares pie wen Shit“ ot, oe oe aa ¢ me oe 6 4a- - 2 ow batts eh ieee ap 4.0 tar Grea ; x Sane “eaeT (gas ive srl So psf alow “et i] - AS y e ¢ _ = _ - ms > - : : “e UNIVERSITY OF KANSAS PUBLICATIONS MusEuM OF NATURAL HISTORY Volume 15, No. 12, pp. 577-614, 1 fig. June 22, 1965 ————ne—romr. 6c LIBRARY DEES LAses HARVARD Amphibians and Reptiles univessizy From the Yucatan Peninsula, Mexico BY WILLIAM E. DUELLMAN UNIVERSITY OF KANSAS LAWRENCE 1965 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Institutional libraries interested in publications exchange may obtain this series by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas. Copies for individuals, persons working in a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of Kansas, Lawrence, Kansas. There is no provision for sale of this series by the University Library, which meets institutional requests, or by the Museum of Natural History, which meets the requests of individuals. Nevertheless, when individuals request copies from the Museum, 25 cents (for each 100 pages or part thereof) should be included for the purpose of defraying the costs of wrapping and mailing. For certain longer papers an additional amount indicated below, is to be included toward the cost of production. * An asterisk designates those numbers of which the Museum’s supply (not necessarily Hee Library’s supply) is exhausted. Materials published to date, in this series, are as ollows: Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950. ®Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text. April 9, 1948. *Vol. 3. Nos. 1-4 and index. Pp 1-681. 1951. ®Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 figures in text. December 27, 1951. Vol. 5. Nos. 1-87 and index. Pp. 1-676, 1951-1953. ®Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952. Vol. 7. Nos. 1-15 and index. Pp. 1-651, 1952-1955. Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956. Vol. 9. Nos. 1-23 and index. Pp. 1-690, 1955-1960. Vol. 10. 1. Studies of birds killed in nocturnal migration. By Harrison B. Tordoff and Hopet M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. September 12, 2. Comparative breeding behavior of Ammospiza caudacuta and A. maritima. By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956. *3. The forest habitat of the University of Kansas Natural History Reservation. By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures in text, 4 tables. December 31, 1956. 4, Aspects of reproduction and development in the prairie vole (Microtus ochro- gaster). By Henry S. Fitch. Pp. 129-161, 8 figures in text, 4 tables. De- cember 19, 1957. 5. Birds found on the Arctic slope of northern Alaska. By James W. Bee. Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958. ®6. The wood rats of Colorado: distribution and ecology. By Robert B. Finley, Jr. Pp. 218-552, 34 plates, 8 figures in text, 35 tables. November 7, 1958. 7. Home ranges and movements of the eastern cottontail in Kansas. By Donald W. Janes. Pp. 553-572, 4 plates, 8 figures in text. May 4, 1959. 8. Natural history of the salamander, Aneides hardyi. By Richard F. Johnston and Gerhard A. Schad. Pp. 578-585. October 8, 1959. 9. A new subspecies of lizard, Cnemidophorus sacki, from Michoacan, México. By William E. Duellman. Pp. 587-598, 2 figures in text. May 2, 1960. 10. A taxonomic study of the middle American snake, Pituophis deppei. By William E. Duellman. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960. Index. Pp. 611-626. Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960. Vol. 12. #1. Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry A. Vaughan. Pp. 1-158, 4 plates, 24 figures in text. July 8, 1959. ©2. The ancestry of modern Amphibia: a review of the evidence. By Theodore H. Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, 1959. 8. The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49 figures in text. February 19, 1960. °4, A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in text. May 2, 1960. Natural history of the bell vireo. By Jon C. Barlow. Pp. 241-296, 6 figures in text. March 7, 1962. : 6. Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C. Fox. Pp. 297-307, 6 figures in text. May 21, 1962. . 7. Vertebrates from the barrier island of Tamaulipas, México. By Robert K. Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 809- 845, plates 5-8. June 18, 1962. 8. Teeth of Edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 fig- ures in text. October 1, 1962. (Continued on inside of back cover) UNIVERSITY OF KANSAS PUBLICATIONS MuSEUM OF NATURAL HISTORY Volume 15, No. 12, pp. 577-614, 1 fig. June 22, 1965 Amphibians and Reptiles From the Yucatan Peninsula, Mexico BY WILLIAM E. DUELLMAN UNIVERSITY OF KANSAS LAWRENCE 1965 er 6 ae i ~ UNIVERSITY OF KANSAS PUBLICATIONS, MuSEUM OF NATURAL HIsTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 15, No. 12, 577-614, 1 fig. Published June 22, 1965 MUS. COM... -OOL LIBRARY DEC 31 1Ss9 HARVARD UNIVERSITY. UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY HARRY (BUD) TIMBERLAKE, STATE PRINTER TOPEKA, KANSAS 1965 Amphibians and Reptiles From the Yucatan Peninsula, México BY WILLIAM E. DUELLMAN CONTENTS UINGERODUGCTION. Far ce oye oe ne oe A ee eee 579 Acknowledements: oo. fic eecc . cee oe Oe eee ee 582 SUMMARYJOF PEELED IVWVORK) o...f:1. 00a. ae cae ee 582 Gazetteer css hse eae 52g a ee ee 583 ANNOTATED IST OF. SPECIES). 45. -., + a aoe 585 Amphibia: o.0e cs 6 go gue ois «5 Facets an tne .. 586 G@aadatar (os, oe oe ede ae 586 Salientia’ a. pen eet cae aa er ee 586 Reptiliae pene. ses ee. cee ee 590 MEStUGINES: 2.5.5 cect. ox et ee ee 590 CKOCOGIT etc cms ate cand ocho ehh Lee eee 593 GATTI et en ck sts OR od os gee ne Ss ge 593 Serpentes 212. > ee ie : te + Aa Te ee eee Lah A ats : _ aT a ats Tee a a vy : ’ oo Bas oe - } al pn. ae & ya Sa Py Dae! a ve = i ? _ 7 oe Cyr, ‘ Ty A oe 7 ; y oe Fae : 7 | i ? mh 9) . ford af hy ee ; v\ i | : ine a Reis SS te iz eh ‘| is 1 a ® te a. Th Pia > oe te seas. tans Ia’ ites = Vol. 14, 15. (Continued from inside of front cover) Variation in the muscles and nerves of the leg in two genera of grouse (Tympanuchus and Pedioecetes). By E. Bruce Holmes.. Pp. 363-474, 20 figures. October 25, 1962. . A new genus of Pennsylvanian Fish (Crossopterygii, Coelacanthiformes) from Kansas. By Joan Echols. Pp. 475-501, 7 figures. October 25, 1963. Observations on the Mississippi Kite in southwestern Kansas. By Henry S. Fitch. Pp. 503-519. October 25, 1963, Jaw musculature of the Mourning and White-winged doves. By Robert L. Merz. Pp. 521-551, 22 figures. October 25, 1963. Thoracic and coracoid arteries in two families of birds, Columbidae and pura aginigre: By Marion Anne Jenkinson. Pp. 553-573, 7 figures. March The breeding birds of Kansas. By Richard F. Johnson. Pp. 575-655, 10 figures. May 18, 1964. The adductor muscles of the jaw in some primitive reptiles. By Richard C. Fox. Pp. 657-680, 11 figures in text. May 18, 1964. Index. Pp. 681-694. Vol. 18. 1, Five natural hybrid combinations in minnows (Cyprinidae). By Frank B. 2. 8. 4, 5. 6. Ts Cross and W. L. Minckley. Pp. 1-18. June 1, 1960. A distributional study of the amphibians of the Isthmus of Tehuantepec, México. By William E. Duellman. Pp. 19-72, plates 1-8, 3 figures in text, August 16, 1960. 50 cents. A new subspecies of the slider turtle (Pseudemys scripta) from Coahuila, México. By John M. Legler. Pp. 73-84, plates 9-12, 3 figures in text. August 16, 1960. Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, plates 13-20, 26 figures in text. November 30, 1960. Occurrence of the garter snake, Thamnophis sirtalis, in the Great Plains and Rocky Mountains. By Henry S. Fitch and T. Paul Maslin. Pp. 289-308, 4 figures in text. February 10, 1961. Fishes of the Wakarusa river in Kansas. By James E. Deacon and Artie L. Metcalf. Pp. 309-322, 1 figure in text. February 10, 1961. Geographic variation in the North American cyprinid fish, Hybopsis gracilis. By Leonard J. Olund and Frank B. Cross. Pp. 823-3848, plates 21-24, 2 figures in text. February 10, 1961. Descriptions of two species of frogs, genus Ptychohyla; studies of Ameri- can hylid frogs, V. By William E. Duellman. Pp. 849-357, plate 25, 2 figures in text. April 27, 1961. Fish populations, following a drought, in the Neosho and Marais des Cygnes rivers of Kansas. By James Everett Deacon. Pp. 859-427, plates 26-30, 8 figures. August 11, 1961. 75 cents. . Recent soft-shelled. turtles of North America (family Trionychidae). By Robert G. Webb. Pp. 429-611, plates 31-54, 24 figures in text. February 16, 1962. $2.00. Index. Pp. 613-624. & 1. 2. Neotropical bats from western México. By Sydney Anderson. Pp. 1-8. October 24, 1960. Geographic variation in the harvest mouse. Reithrodontomys megalotis, on the central Great Plains and in adjacent regions. _By J. Knox Jones, Jr., and B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961. Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson. " Pp, 29-67, plates 1 and 2, 3 figures in text. July 24, 1961. 4. A new subspecies of the black myotis (bat) from eastern Mexico. By. E. 6. il. 12. Pena Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December North American yellow bats, ‘‘Dasypterus,” and a list of the named kinds of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr. Pp. 73-98, 4 figures in text. December 29, 1961. Natural history of the brush mouse (Peromyscus boylii) in Kansas with description of a new subspecies. By Charles A. Long. Pp. 99-111, 1 figure in text. December 29, 1961. Taxonomic status of some mice of the Peromyscus boylii group in eastern Mexico, with description of a new subspecies. By Ticul Alvarez. Pp, 118- 120, 1 figure in text. December 29, 1961. A new subspecies of ground squirrel (Spermophilus spilosoma) from Ta- maulipas, Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962. Taxonomic status of the free-tailed: bat, Tadarida yucatanica Miller. By J. pots Jones, Jr., and Ticul Alvarez. Pp. 125-133, 1 figure in text. March 7, > A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene, of Texas. By E. Raymond Hall and Walter W. Dalquest. Pp. 135-138, 2 figures in text. April 30, 1962. } A new subspecies of wood rat (Neotoma) from northeastern Mexico. By Ticul Alvarez. Pp. 139-143, April 30, 1962. ’ Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul Alvarez, and M. Raymond Lee. Pp. 145-159, 1 figure in text. May 18, 1962. (Continued on outside of back cover) Vol. 15. I. (Continued from inside of back cover) A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, 1 figure in text. May 21, 1962. The mammals of Veracruz. By E. Raymond Hall and Walter W. Dalquest. Pp. 165-362, 2 figures. May 20, 1963. $2.00. The recent mammals of Tamaulipas, México. By Ticul Alvarez. Pp. 868- 473, 5 figures in text. May 20, 1968. $1.00. A new subspecies of the fruit-eating bat, Sturnira ludovici, from western Mexico. By J. Knox Jones, Jr., and Gary L. Phillips. Pp. 475-481, 1 figure in text. March 2, 1964. Records of the fossil mammal Sinclairella, Family Apatemyidae, from the Chadronian and Orellan. By William C, Clemens. Pp. 483-491. 2 figures in text. March 2, 1964. More numbers will appear in volume 14. The amphibians and reptiles of Michoacan, México. By William E. Duell- man. Pp. 1-148, plates 1-6, 11 figures in text.. December 20, 1961. $1.50. Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox Jones, Jr., and George W. Byers. Pp. 149-178. January 31, 1962. A new species of frog (Genus Tomodactylus) from western México. By Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962. Type specimens of amphibians and reptiles in the Museum of Natural His- tory, the University of Kansas. By William E. Duellman and Barbara Berg. Pp. 183-204. - October 26, 1962. Amphibians and Reptiles of the Rainforests of Southern El Petén, Guatemala. By William E. Duellman. Pp. 205-249, plates 7-10, 6 figures in text. Oc- tober 4, 1963. A revision of snakes of the genus Conophis (Family Colubridae, from Middle ane he By John Wellman. Pp. 251-295, 9 figures in text. October 4, A review of the Middle American tree frogs of the genus Ptychohyla. By William E. Duellman. Pp. 297-349, plates 11-18, 7 figures in text. October 18, 1968. 50 cenis. Natural history of the racer Coluber constrictor. By Henry S. Fitch. Pp. 351-468, plates 19-22, 20 figures in text. December 30, 1963. $1.00. 9. A review of the frogs of the Hyla bistincta group. By William E. Duellman. Vol._16. 1. 2. 3. Vole P71. Pp. 469-491, 4 figures in text. March 2, 1964 An ecological study of the garter snake, Thamnophis sirtalis. By Henry S. Fitch. Pp. 493-564, plates 23-25, 14 figures in text. May 17, 1965. Breeding cycle in the ground skink, Lygosoma laterale. By Henry S. Fitch and Harry W. Greene. Pp. 565-575, 3 figures in text. May 17, 1965, Amphibians and reptiles from the Yucatan Peninsula, México. By William E, Duellman. Pp. 577-614, 1 figure in text. June 22, 1965. More numbers will appear in volume 15. Distribution and taxonomy of Mammals of Nebraska. By J. ox Jones, Jr. Pp. 1-356, pls. 1-4, 82 figures in text.. October 1, 1964. $3.50. Synopsis of the lagomorphs and rodents of Korea. By J. Knox Jones, Jr. and David H. Johnson. Pp 354-407. February 12, 1965. J Mammals from Isla Cozumel, Mexico, with description of a new species of harvest mouse.- By J. Knox Jones, Jr. and Timothy E. Lawlor. Pp. 409-419, 1 fig. April 18, 1965. More numbers will appear in volume 16. Localities of fossil vertebrates obtained from the Niobrara Formation (Creta- ceous) of Kansas. By David Bardack. Pp. 1-14. January 22, 1965. More numbers will appear in volume 17. UNIVERSITY OF KANSAS PUBLICATIONS MusEuM OF NATURAL HISTORY Volume 15, No. 18, pp. 615-625, pls. 26-28, 2 figs!S. CON, 00 July 20, 1965 LIBRARY DEC 31 1965 HAR VARD A New Species of Turtle, Genus Kinostemon,” From Central America IBY, JOHN M. LEGLER UNIVERSITY OF KANSAS LAWRENCE 1965 UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HisTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Frank B. Cross Volume 15, No. 13, pp. 615-625 Published July 20, 1965 UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY ROBERT R. (BOB) SANDERS, STATE PRINTER TOPEKA, KANSAS ; 9 (V } i ye [ax OK ii MUS, COMP. 2¢ LIBRARY A New Species of Turtle, Genus Kinosternom 3 5 From Central America : HARVARD BY UNIVERSITY JOHN M. LEGLER In 1957 Dr. Edward H. Taylor asked me to study and report on a specimen of Kinosternon, obtained by him in Costa Rica, that differed from other known species. Description was delayed for want of comparative material. In the three years 1961-63 I col- lected, prepared, and studied some 700 specimens of the genus Kinosternon from Central America. Among these are 13 additional individuals of the species represented by Dr. Taylor’s specimen. The species is named and described as follows: Kinosternon angustipons new species Holotype.—University of Kansas 43631, adult female, alcoholic; Los Dia- mantes, Limén Province, Costa Rica; obtained by Edward H. Taylor and John Baker, August 13, 1952; original number 8507. Paratypes (total of 13).—University of Utah 37562, 13 mi. S. San Juan del Norte, Nicaragua in Limon Prov., Costa Rica; UU 8757-60,62-64, KU 84882 6 6, UU 8765-662 2, .6 mi. NNW Puerto Viejo, Heredia Prov., Costa Rica; UU 37672, 2 mi. E and 1% mi. S Guabito, Bocas del Toro Province, Pan- ama and, UU 418946, 2 mi. NW Almirante, Bocas del Toro Province, Panama. Diagnosis—A small species of Kinosternon, most closely resembling K. dunni Schmidt (1947), and having: 1) a flattened, noncarinate carapace; 2) a narrow plastron with anal notch and interlaminal seams that tend to fill with soft tissue in older individuals; 3) a narrow bridge (20% or less of length of carapace); 4) an unstriped head; 5) a maxillary beak that is neither hooked nor notched; 6) clasping organs on posterior limbs of males; and, 7) tip of tail unmodified (neither horn-covered nor clawlike) in both sexes. Description of species (based upon type series ).—Carapace relatively low (highest point on posterior part of fourth central), evenly arched or flat-topped in cross section, oval in dorsal aspect, greatest width usually at level of bridge. No pronounced sculpturing, even in smallest individuals; faint suggestion (creases) of mid-dorsal and dorsolateral keels in small specimens; no trace of dorsal keels in older, larger specimens. Anterior margin of carapace smooth, slightly indented; posterior margin shallowly notched (between postcentrals ). Carapacal scutes imbricated (juxtaposed only in old, worn specimens); first or third central longest and broadest, fifth central shortest and narrowest; centrals 1 to 3 approximately as long as broad, 4th and 5th significantly broader than long. Precentral wedge-shaped, narrowed anteriorly, usually broader than long, shorter than seam between first and second marginals. First central in narrow contact (or not in contact) with Me. Tenth marginal highest, sloping gradually up from point of contact with Mg, abruptly higher than postcentrals; other marginals flat-topped. Two parallel lateral ridges along margin of carapace, the lower ridge continuous with anterior and posterior free edges of carapace and passing approximately through centers of marginals 4 to 7, the upper ridge continuous with upper borders of mar- ginal scutes and terminating on Myo; a distinct marginal bulge between mentioned lateral ridges. (617) 618 UNIVERSITY OF KANSAS Pusxs., Mus. Nat. Hist. Plastron narrow, its freely movable lobes incompletely closing orifices of shell (extent of closure approximately 55 per cent for anterior lobe, 50 per cent for posterior); bridge narrow (17 to 20 per cent of carapace length), imparting subcruciform appearance to plastra of largest males. Plastral lobes constitute, respectively, approximately 33, 29, and 38 per cent of plastral length; posterior lobe onic at hinge and having wide, shallow anal notch (more pro- nounced in males than in females). Interlaminal seams (especially those over hinges and the interfemorals) of older individuals containing varying amounts of soft pale tissue. Axillary and inguinal scutes in contact on bridge (narrowly separated in one old male); axillary scute narrowly in contact with M4 and Mg; inguinal scute in broad contact with Mg and M7 (forming narrow contact with Mg in two specimens). Plastral scutes, in order of length— abdominal, anal, humeral, femoral, gular, pectoral. Head slightly broadened; snout wide and blunt in dorsal aspect as well as profile; maxillary sheath flat or slightly concave in premaxillary region, neither hooked nor notched. Tip of mandibular sheath blunt. Inner crushing surfaces of jaws weakly developed and chiefly smooth. Tomial edges of both sheaths nearly straight in Safle, Snout, in general, evenly tapered, lacking pinched appearance characteristic of most other species of Kinosternon. Top of head evenly rounded in anterior view, lacking raised brows. Mature males having bosslike enlargement of snout in prefrontal region. Dorsal head shield present but indistinct, blending gradually with softer skin in temporal region. Internal choanae broadly oval (approximately twice as long as broad); choanal flaps bearing a single papilla on anterior half of flap; free edge of flap (when closed) longitudinally bisecting choanal opening and papilla extending up to one half of remaining distance from edge of flap to medial border of opening. External narial openings directed anteriorly, situated just below tip of snout, round in cross section; floor of each narial passage having distinct, blunt, longitudinal ridge or papilla (clearly visible as a bulge just inside opening and slightly lateral to midventral floor of passage). A linear series (usually three to six) of small barbels on each side of throat from mandibular symphysis to end of hyoid bar (and following course of that element), concentrated chiefly in triangular area bounded by mandibular rami; usually two or three smaller barbels on or near midline just posterior to symphysis; gular barbels variable in size but never so large as in K. leucosto- mum, and never consisting of enlarged mental pair with smaller ones behind. Skin of neck studded with distinct papillae having rounded white tips; papillae arranged, more or less regularly, in 14 to 16 longitudinal rows. Papillae numerous and distinct also on posterior surfaces of limbs and on tail (especially in perianal region). Hands and feet fully webbed, free edges of webs strongly fimbriated; antebrachium having three falciform scales on anterior surface. Discrete, apposed patches of specialized scales (clasping organs) well de- veloped on posterior thighs and legs of males; individual scales in each patch spadelike (not “tuberculate,” not pointed); spadelike scales angled dorsally at approximately 45 degrees. Tail of mature males elongate (equal to or longer than posterior plastral lobe), heavy at base, prehensile, and having blunt unmodified tip (not horn- covered, not clawlike). Tail of females not or barely extending to posterior margin of carapace. Coloration —The following descriptions are based upon live adults, viewed in clear water with a beam of incandescent light. Colors of live specimens are not significantly different from those which have been fixed and preserved in ethyl alcohol (no formalin used). In general the colors of both sexes are drab and neutral. Adjacent pale and dark colors are never sharply delimited but grade gradually into one another. The pale snout, pale upper eyelids, and dark eye stand out clearly in lateral or dorsolateral views of the head and an observer's attention is drawn to them at first glance. This is not so in K. leucostomum in which the upper eyelids are either not pale or are joined in a continuously pale or mottled field with the frontal region and snout. A New Species OF KINOSTERNON 619 Female (UU 3767) Iris brown, flecked with golden yellow, the combination appearing to be uniform dark brown except in bright light. Soft skin around nostrils cream; upper eyelids grayish cream; horny sheaths of jaws yellowish cream, grading into pale brown, the darker color being near junction of skin with horn. Ground color of limbs, tail, and dorsal part of neck pale gray to brownish gray, slate in darkest areas. Inguinal packet grayish cream. Ventral surface of neck (except for a small gray area in front of plastron) yellowish cream, suffused with pink on gular region; a short, dull brown, indistinct stripe ex- tending from mandibular symphysis (not on horny sheath) onto gular region. Top ot head dark neutral brown with slight purplish cast, head shield slightly darker than softer skin behind it; side of head tannish above, grading gradually into paler color of throat; palest area on side of head over tympanum, same color as horny sheaths; a few indistinct, irregular, pale brown marks on side of head, chiefly near corner of mouth, below tympanum, and between tym- panum and orbit. Plastron dark golden yellow; interlaminal seams narrowly edged with dark brown; soft skin of interlaminal seams (where present) gray, same color as limbs. Undersurfaces of marginals slightly Hae than plastron, pale color extending up to lower of two lateral marginal ridges. Carapace dark brown, having indistinct paler brownish areas near centers of scutes. Male (UU 3757) Iris somewhat paler than in female, having a pale reddish cast but con- sisting of brown and golden flecks (iris not distinct from pupil except in bright light). Jaw sheaths having fine, vertical, pale brown stripes. Ground color of plastron pale neutral yellow, much paler than in female. Osteology—Of Central American kinosternids, the skeleton of Kinosternon leucostomum bears the closest resemblance to K. angustipons. The following description is based upon two complete adult skeletons (UU 37604, UU 376692) and an adult shell (UU 41892) of angustipons; where skeletal characteristics are regarded as differing significantly from those of eight adult specimens of leucostomum (4 males and 4 females), the characteristics of leucostomum are given in brackets. In general, no significant differences were observed in the appendicular skeletons of the two species. Skull solidly built, greatest width 65 to 66 per cent of total length, height 71 to 73 per cent of width; supraoccipital process relatively short, eight per cent of total length of skull [13 to 17 per cent in leucostomum]. Orbit rela- tively small and nearly circular, having substantial overhang of bone above; vertical diameter as a percentage of least interorbital breadth, 56 (4) to 62 (9) [orbit larger, relatively higher due to deficiency in overhang of bone above, vertical diameter 58 to 83 per cent ( ¢ ¢ ) and 68 to 74 per cent ( 2 2 ) of least interorbital breadth]. Snout tapering anteriorly from base of post- orbital bar [from point above mid-orbit]; top of skull (anterior aspect) more or less evenly rounded [more nearly flat-topped], dorsal orbital rim not at all jutting or browlike in appearance [distinctly jutting and browlike, albeit smaller]. Stapediotemporal foramen indistinct [distinct]. Temporal arch weakly emarginate below; quadratojugal comprising nearly one-half of temporal arch, jugal excluded from upper free edge of arch [jugal entering upper free edge]; crushing surfaces of maxillae poorly defined, not concave, sloping dorsomedially, especially weak in premaxillary region [concave; inner border sharp, distinct and continuous across premaxillary region]; anteroventralmost part of vomer greatly expanded, this expansion, the vomeropremaxillary articu- lation, and incisive foramina visible in ventral exposure [not so expanded, and structures mentioned excluded from ventral view by posterior premaxillary ridge]; maxillae widely separated anteriorly [in contact or closely approxi- mated]; maxillary beak weakly or not at all developed; ventral part of snout lacking indented or “pinched” appearance in ventral view; profile of maxillary cutting edge forming nearly straight line [beak well developed; snout bilaterally 620 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. indented, having “pinched” appearance; profile of maxillary cutting edge recurvate]._ Angular and coronoid bones of mandible not or but scarcely visible in direct lateral view; angular separating surangular and prearticular ventrally. Cervical vertebrae typically kinosternid, second opisthocoelous, third bi- convex, and remaining five procoelous; of these, sixth and seventh doubled posteriorly, seventh and eighth doubled anteriorly. Phalangeal formula 2-3-3-3-3 on hand and foot (this assumes that meta- tarsal V is combined with a tarsal to form a single “hooked” element). Primary sacral ribs expanded nearly to width of ilial blade at sacroiliac joint, narrow at distal articulation; secondary sacral ribs not at all expanded. D Fic. 1. A and B, Ventral and lateral views of skull, Kinosternon angustipons, paratype (UU 37669 ) x 2; C and D, skull of K. leucostomum (UU 42839, Gamboa, Canal Zone) 2. PLATE 26 Comparison of holotype, Kinosternon angustipons (left column) and paratype K. dunni (right column); heads x .9; plastral views 5 and .6, respectively. (CNHM 42803 @ ) PLATE 27 Old male of K. angustipons (UU 3756), freshly killed; dorsal and ventral views - head slightly larger than actual size; all photographs from color trans- parencies. V 72 PLATE 28 Top. Habitat of K. angustipons, .6 Mi. NNW Puerto Viejo, Heredia, Costa Rica, 22 July 1961. Nine specimens were obtained from small pool in fore- ground. Bottom. Anterior view of live female (UU 3767) showing pale snout and eyelids and unhooked beak, x 13%. All photographs from color trans- parencies. Fic. 2. A, Ventral view of shell, Kinosternon angustipons, paratype, (UU 37662) x .8; B, same view, K. leucostomum (UU 42322, Puerto Viejo, Heredia, Costa Rica) x .6; C, dorsal view of scuteless shell, K. angustipons, paratype (UU 37604) x .8; D and E, posterior margin of carapace showing fifth central lamina and the relative heights of postcentral and 10th marginal laminae in K. angustipons (UU 37662) and K. leucostomum (UU 42324 ), respectively (approximately x .85). A New SPECIES OF KINOSTERNON 621 Caudal vertebrae 18 in ¢, 21 in @, distal two or three fused or nearly fused into blunt tip [terminal vertebrae fused into clawlike tip]. Five juxtaposed neurals, first neural separated from nuchal (by intercostal suture) by distance half its own length; shape of neurals basically hexagonal, long sides being anterior to an intercostal suture and short sides posterior in all specimens; pygal more or less rectangular, slightly notched; suprapygal six- sided, concave above, convex below, width approximately three times height [five-sided, width two times height]; small, subtriangular anterior suprapygal in two of three specimens [no anterior suprapygal]; of eight pairs of costals, last three pairs in mid-dorsal contact between last neural and anterior supra- pygal, eighth pair only narrowly so [three, sometimes four pairs in broad contact]; first peripheral about as high as second, separated from its fellow (by nuchal) by width equal to or greater than its own; first peripheral in contact with first costal, separating nuchal and second peripheral [first periph- eral much lower than second; nuchal and second in contact, excluding first peripheral from contact with first costal]. Comparison with other Central American species—In my opinion there are four full species of Kinosternon in Central America south of the Republic of Mexico. These are Kinosternon angustipons, K. acutum Gray, K. leucosto- mum (including K. postinguinal Cope and K. spurrelli Boulenger), and K. scorpioides (Linnaeus) (including K. panamensis Schmidt and K. cruentatum Duméril, Bibron and Duméril). In K. angustipons the axillary scute is in narrow contact with the fourth marginal, never with the third; in the other three species the axillary is in contact with Ms or narrowly separated from it, never separated by nearly the entire length of My. K. angustipons is further distinguished from these species in lacking a hooked, recurved maxillary beak and in lacking a distinct claw on the tip of the tail in both sexes. All these characteristics (except that of the maxillary beak) serve also to distinguish K. dunni from the other three species mentioned. Comparison with Kinosternon dunni.—Differences in the heads and necks of the two species are the most striking, yet the most subjective (Pls. I-III). In profile K. dunni has a moderately hooked beak and recurved tomial edge on the maxillary sheath whereas angustipons has a flattened beak and a tomial edge that is nearly straight. In dorsal and ventral views the snout of dunni is much more pronounced, having a pinched appearance and probably being somewhat larger because of greater bulging of soft, circumnarial tissues. The snout of angustipons tapers evenly (unpinched) and is generally blunt in appearance. The dorsal head shield of dunni is sharply delimited whereas that of angustipons grades subtly into the softer skin of the neck. The dermal papillae on the neck of angustipons are numerous, distinct, white-tipped, and arranged in regular longitudinal rows. There are also white-tipped papillae on the posterior surfaces of the limbs, the tail, and perianal region of angustipons. In dunni these dermal papillae are smaller, less numerous, less distinct, and irregularly arranged. Few papillae are dis- cernible on the holotype (CNHM 42804) of dunni and those on the neck of the paratype (CNHM 42803) are restricted chiefly to a lateral row on each side. The tails and perianal regions of the two species are sharply differen- tiated, that of angustipons being distinctly papillose, and that of dunni being nearly bare. Females of angustipons have relatively broad, short tails that are rounded (never pointed) and lack a terminal spine. Females of dunni have distinctly narrower tails, which are pointed. The holotype of dunni has a small terminal spine; the paratype does not. 622 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. On the holotype of dunni there is a deep distinct crease on each side of the middle plastral lobe (abdominal), beginning just behind the abdominal areola and extending to a point opposite the anterolateral corner of the posterior lobe. The middle lobe slopes angularly upward into the bridge lateral to the creases. The sides of the creases (especially the medial sides) form blunt, raised ridges. The creases are evident but weakly developed in the juvenal paratype of dunni and Medem’s photographs (1961:465, fig. 8) indicate that at least the ridges are present on his specimens. No specimen of K. angustipons has such creases or ridges; the middle plastral lobe lies in approximately the same plane as the bridge or, if slightly lower, slopes into the bridge more gradually than in dunni. This feature of the plastron accounts, in part, for the relatively lower shell of angustipons (Table 1). Consistent differences in shape of carapace are as follows: dunni tending to be widest at level of Msg, noticeably constricted opposite bridge (angusti- pons widest at point opposite bridge, not constricted, not flared posteriorly) ; large individuals of dunni having distinct, narrow, flattened area confined to central laminae, suggesting the weakly tricarinate condition found in some TABLE 1. PROPORTIONAL DIFFERENCES BETWEEN Kinosternon angustipons sp. Nov. AND K. dunni Scumipt. Data ARE BasED Upon 14 SPECIMENS OF angustipons (106 6 4292) ANd Upon THE Types (EXAMINED) AND Two OTHER SPECIMENS (Not SEEN, MEpEM, 1961, 1962) oF dunni (1¢ 8929). WueERE MEASUREMENTS OF dunni WrRE Nor MaAvE FROM SPECIMENS, PRO- PORTIONS WERE COMPUTED FROM PUBLISHED DATA, DRAWINGS, OR PHOTO- GRAPHS. STATEMENTS IN THE LEFT COLUMN REFER TO angustipons. ARITH- METIC MEAN, RANGE, AND STANDARD DEVIATION ARE GIVEN FOR EACH CHARACTER (SEPARATELY FOR EACH SEX IF CHARACTER JUDGED TO BE Dt- MORPHIC ). == Condition in K. angustipons, 2 : and Character Measured K. angustipons K. dunni Bridge relatively narrower; minimal width (anteroposterior) of bridge expressed as percentage of | carapace length 18 + 1.1 (17-20) 24 == () Shell relatively lower; height of shell expressed as percentage of |o'0155 = =+1.4 (51-56) oa 65 2 maximal width of carapace 9958 +8.6 (54-62) 9962 +3.16 (59-65) Abdominal and anal scutes relatively shorter, pectora! and femoral scutes relatively longer; interlaminal length of each scute expressed as percentage of sum of median interlaminal lengths: Pectoral | 8.3 + 1.28 (6.8-10.8) 6.3 4 :97:(5.2=7.1) Abdominal 29.8 + 1.44 (26.7-31.7) 32.3 + 2.27 (29 .9-34.8) Femoral 15.7 + 1.88 (13.4-19.4) 11.7 + 1.24 (10.3-12.5) Anal 07 18.8 + 2.21 (15.6-22.1) o' 22.1 9 9 22.8 + 2.12 (19.6-23.9)| 9 9 25.7 = .64 (25.2-26.1) Head relatively narrower, es- pecially in females; width of head expressed as percentage of cara- | oo" 31 +1.5 (28-33) o' 32 Den wee pace width 9929 +1.0 (28-30) 99235 + 2.35 (31-35) Plastron constricted at pos- | terior hinge; width at hinge ex- pressed as percentage of width . (maximal) across femorals 94 +3.0 (90-102) 100 +1.6 (98-102) A New SPECIES OF KINOSTERNON 623 populations of K. scorpioides (shell of angustipons more evenly rounded, not distinctly flat-topped in region of centrals). Arithmetic expressions of these and other proportional differences between the two species are set forth in Table 1. The characters mentioned for the bridge, head, and posterior plastral hinge, however, are detectable without the aid of measurement and computation. Size.—Use of adult size as a taxonomic character to compare small samples of turtles is hazardous; but, in some chelonians growth is determinate to the extent that: 1) it tapers off rapidly at a certain age or size and proceeds slowly or not apparently thereafter; 2) old individuals at or near maximum size are recognizable by external characteristics of the shell. It seems clear that the size attained by adults is at least partly under genetic control and that, if differences in size between two species can be demonstrated, they should be exploited as taxonomic characters. The largest male of angustipons has a carapace length of 112 mm (nine other males range from 82 to 101) and the largest female a length of 119 (three others 90 to 109). Corresponding figures for known specimens of dunni are as follows: @ 171; 2 2 150, 149; a shell of unknown sex 175, The largest specimens of angustipons are clearly adults and, on the basis of appear- ance of shell, are past the age of normal regular growth; hence, they are near the maximum size that they would have attained. The second largest male (UU 3756, 101 mm) is likewise near maximum size. The paratype of dunni (CNHM 42803,2 93 mm) is probably immature, whereas all females of angustipons are mature. I therefore regard the difference in adult size of angustipons and dunni as too great to be attributed to fortuitous sampling; angustipons is much the smaller of the two species and probably never attains the relatively ponderous bulk of dunni. Localities and Habitat. Localities mentioned for the paratypes are reck- oned, for clarity, in terms of airline miles from geographic points which appear on most maps. The locality for UU 3756 lies on the southern edge of a large island formed by the Bravo and Colorado channels of the Rio San Juan at its delta, just west of the confluence of the Rio Chirripéd and the Colorado Channel. Actual place of collection was 4% mile E of a farm referred to locally as “Valiente.” Specimens from Bocas del Toro Province, Panama, were taken along the railroad of the Chiriqui Land Company. The localities, as marked along the right of way, are: Mile 22% (UU 3767) and Mile 2 (UU 4189), the “miles” being reckoned from the railroad shops in Almirante. Los Dia- mantes is a farm on the railroad, approximately two miles east of Guapiles. All specimens were collected by means of baited hoopnets in shallow per- manent swamps or slow, scummy backwaters of streams. Traps set nearby in clearer, flowing water yielded specimens of K. leucostomum but no angusti- pons. Geographic range.—K. angustipons is now known only from the specimens herein reported; hence, the known range of the species extends, in the At- lantic lowlands, approximately from the delta of the Rio San Juan (on the boundary between Nicaragua and Costa Rica) to Almirante, Bocas del Toro, Panama. Most of the localities from which specimens are known are near sea level (less than 100 M); Los Diamantes is the highest, having an elevation of approximately 260 meters. Satisfactory microhabitats for the species prob- ably exist, almost continuously, from northeastern Honduras to Colon, Panama. Discussion.—It is evident from the data presented that K. dunni and K. angustipons are closely related but taxonomically distinguish- 624 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. able. There is insufficient evidence at present to evaluate judi- ciously the standing of these taxa, in terms of species vs. subspecies. The apparent rarity of both taxa and general lack of field work in critical areas combine to make the problem an irresolvable one now. K, angustipons is here regarded as a full species because it is mor- phologically distinctive and is not known to interbreed with K. dunni nor other kinosternids. The gap separating the known ranges of dunni and angustipons is at least 400 miles (from Almirante to the mouth of the Rio Atrato) and includes the whole of the Isthmian region of Panama. Possibly the gap is more apparent than real. Until better evidence is avail- able, I predict (by inference from studies of other aquatic che- lonians in Central America) that dunni and angustipons constitute an example of a pair of closely related species of recent origin whose ranges are separated narrowly in the Isthmian region of Panama. The species Geoemyda funerea (Cope) and G. punctularia (Daudin) constitute another such pair in the same region whereas Stauro- typus triporcatus (Wiegmann) and S. salvini Gray probably con- stitute a like example in northwestern Central America. Relationships.—Except for its evident close relationship to K. dunni, the relationship of K. angustipons to other members of the genus is not clear. The combination of narrow plastron and narrow bridge is a striking feature and is seen also, to variable degrees, in Kinosternon bauri Garman, K. herrerai Stejneger, K. hirtipes Wag- ler, and K. subrubrum (Lacépéde). My own studies of these species indicate that none of them is especially closely related to angustipons or dunni, in spite of the plastral similarity. Beyond this, I am unable to assess the relationships of K. angustipons at the moment. Remarks.—Kinosternon angustipons is seemingly nowhere well known by natives. The species is not utilized for food (kinosternids rarely are) and, to my knowledge, it has not been given a common name even in local areas where many persons are familiar with turtles. In western Panama and northern Costa Rica, K. angusti- pons and K. leucostomum are called “Galapago.” Most persons who confused the two species in this manner were quick to see the differences I pointed out to them. The specific name angustipons is from the Latin angustus (nar- row) and pons (bridge) meaning narrow bridged, Acknowledgments.—I am grateful to Dr. William E. Duellman (University of Kansas) and Dr. Robert F. Inger (Chicago Natural History Museum) for the loan of specimens. The authorities of the Gorgas Memorial Laboratory in A New Species oF KINOSTERNON 625 Panama (especially Drs. C. F. Johnson, M. Hertig, G. B. Fairchild, and P. Galindo) did much to facilitate field work in that country as did Dr. Alvaro Wille (University of Costa Rica) in Costa Rica. Messrs. C. M. Keenan and E. Mendez have provided additional specimens from Panama, Messrs. Lee Bell and Nowlan Dean rendered capable assistance in the field. Elizabeth Lane made the drawings of the skull. Research on which this account is based was supported by the Society of Sigma Xi (RESA Grant) and the University of Utah Research Committee in 1960, and thereafter by the National Science Foundation (Grant # B-10178). In August of 1964, after the above account was in press, I discovered that Mr. Wilfred T. Neill had prepared a description of the species newly named in this paper. My thanks go to Mr. Neill for having withdrawn his manuscript when he learned of mine. LITERATURE CITED MeEpEM, F. 1961. Contribuciones al conocimiento sobre la morfologia, ecologia y distribucién geografica de la tortuga Kinosternon dunni K. P. Schmidt. Novedades Colombianas, 1(6):446-476, 13 figs., 2 tables, 1 map, 1 September. 1962. La distribucién geografica y ecologia de Los Crocodylia y Testu- dinata en el departamento del Chocd. Rev. Acad. Colombiana Ciéncias Exactas, Fisicas y Nat., 11(44):279-303; 56 figs. (com- prising 20 pls.), folding map, December. ScumwT, K, P. 1947. A new kinosternid turtle from Colombia. Fieldiana Zoology, 31(18):109-119, fig. 14, 11 April. Museum of Zoology, University of Utah, Salt Lake City, Utah. Transmitted to editor January 21, 1964. O 80-5832 ;Y - ae, : aes eh awe ry ‘A eee D ay ale = bas ney “ma et PU sant +e ten el ifs nv 4 aoe en 2 Pee fi ; »y < wow Gt fale At erin Ritedeeerereed q . ce ee aid ies fo cena ae eT oed si rps pte sau fe mpes oe r Ve e @ > »f AY 15 4 a> ind &f co ti ade i pa 6) 6 Ne BALI oy ito OT arwtraiy > joe ay j et fe ro. oy? ey b jer we = or Uy CPR Sov aeir™: + tie : > yy ’ a ~~ oe 7 mh MW Grgh| age? 4 ridee “Fi — Y Wire ay a hi vay as vin : ~ ! . =a) ni? ; 6) bk ma Ms. 7 : e biel wpe a o8) (UL errs) wh Kase al i 7 ‘ s ; 7¢ cement z% A ' 6 1] tz ar c * siz ine a a 1 ty ’ sat ah Ql ieee eh Be ae y, Ag rf "] é = | Se OLLI se & mi 6 PS | 7 Vek amp lsh ae ae - » * OU ey wl nie =I" a . . "y as P \ ‘ we _—— sts paris} aig nue! a j < me rg ir - Le a Ti ke , j marr 4 y = Ay a i een ah Ta qm = 5 ihe IR = KL PRN an NA Be. me Ny Ne ViLS: COMP. ZOOL ee UNIVERSITY OF KANSAS PUBLICATIONS[B | {OF§ _ Museum oF Natura. History HARVARD UNIVERSITY, Volume 15, No. 14, pp. 627-709, pls. 29-36, 5 figs. December 30, 1965 A Biogeographic Account Of the Herpetofauna of Michoacan, Mexico Bx WILLIAM E. DUELLMAN UNIVERSITY OF KANSAS LAWRENCE 1965 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Institutional libraries interested in publications exchange may obtain this series by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas, Copies for individuals, persons working in a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of Kansas, Lawrence, Kansas. When individuals request copies from the Museum, 25 cents should be included, for each 100 pages or part thereof, for the purpose of defraying the costs of wrapping and mailing. For certain longer papers an additional amount, indicated below, toward some of the costs of production, is to be included. ® An asterisk designates those numbers of which the Museum’s supply is exhausted. Vol. 1. Nos. 1-26 and index.. Pp. 1-638, 1946-1950. *Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text. April 9, 1948. Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, and distribution. By Rol- lin H. Baker. Pp. 1-359, 16 figures-in text. June 12, 1951. °2. A quantitative study of the nocturnal migration of birds. By George H. Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951. 8. Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473- 530, 49 figures in text, 13 tables. October 10, 1951. °4, Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and peer W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, Index. Pp. 651-681. ®Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 figures in text. December 27, 1951. Vol. 5. Nos. 1-87 and index. Pp. 1-676, 1951-1953. ®Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952. Vol. 7. Nos. 1-15 and index. Pp. 1-651, 1952-1955. Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956. Vol. 9. Nos. 1-23 and index. Pp. 1-690, 1955-1960. Vol. 10. Nos. 1-10 and index. Pp. 1-626, 1956-1960. Vol. 11. Nos. 1-10 and index. Pp. 1-708, 1958-1960. Vol. 12. *1. Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. July 8, 1959. ®2. The ancestry of modem Amphibia: a review of the evidence. By Theodore H. Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, 1959. 8. The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49 figures in text. February 19, 1960. *4, A new order of fishlike Amphibia from the Pennsylvanian of Kansas. By Theodore H. Eaton, Jr., and Peggy Lou Stewart. Pp. 217-240, 12 figures in text. May 2, 1960. 5. Natural history of the Bell Vireo, Vireo bellii Audubon. By Jon C. Barlow. Pp. 241-296, 6 figures in text. March 7, 1962. 6. Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C. Fox. Pp. 297-307, -6 figures in text. May 21, 1962. 7. Vertebrates from the barrier island of Tamaulipas, México. By Robert K. Selander, Richard F. Johnston, B. J. Wilks, and Gerald G..Raun. Pp. 309- 345, pls. 5-8. June 18, 1962. 8. Teeth of edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 figures in text. October 1, 1962. . Variation in the muscles and nerves of the leg in two genera of grouse (Tym- panuchus and Pedioecetes). By E. Bruce Holmes. Pp. 363-474, 20 figures in text. October 25, 1963. $1.00. 10. A new genus of Pennsylvanian fish (Crossopterygii, Coelacanthiformes) from Kansas. -By Joan Echols. Pp. 475-501, 7 figures in text. October 25, 1963. 11. Observations on the Mississippi kite in southwestern Kansas. By Henry S. Fitch. Pp. 503-519. October 25, 1963. 12. Jaw musculature of the Mourning and White-winged doves. By Robert L. Merz. Pp. 521-551, 22 figures in text. October 25, 1963. 18. Thoracic and coracoid arteries in two families of birds, Columbidae and Hirundinidae. By Marion Anne Jenkinson. Pp. 553-573, 7 figures in text. March 2, 1964. , 14.. The breeding birds of Kansas. By Richard F. Johnston. Pp. 575-655, 10 figures in text. May 18, 1964. 75 cents. 15. The adductor muscles of the jaw in some primitive reptiles. By Richard C. Fox. Pp. 657-680, 11 figures in text. May 18, 1964. Index. Pp. 681-694. (Continued on inside of back cover) UNIVERSITY OF KANSAS PUBLICATIONS Museum oF NATURAL HiIsTOoRY Volume 15, No. 14, pp. 627-709, pls. 29-36, 5 figs. December 30, 1965 A Biogeographic Account Of, the Herpetofauna of Michoacan, Mexico BY WILLIAM E. DUELLMAN UNIVERSITY OF KANSAS LAWRENCE 1965 UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL History Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Frank B. Cross Volume 15, No. 14, pp. 627-709, pls. 29-36, 5 figs. Published December 30, 1965 muUS. COMP. ZOOL LIBRARY FEB 4 Iyco HARVARD UNIVERSITY UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY ROBERT R. (BOB) SANDERS, STATE PRINTER TOPEKA, KANSAS A Biogeographic Account Of the Herpetofauna of Michoacan, México BY WILLIAM E. DUELLMAN CONTENTS PAGE PN FRODUGCIIONS 44 4G. Soca th fore cre aks Sk ig es He eee 629 Aicknow led GImeMtS tats gerbe 62 uy dat eRe ee ee 631 [DESCRIP TIONJOFSIAE PARE AG. pscrd du Aa eee ee, ee ee 631 Physioerapay ands Geoloty: (..148-2ta15 estes ace = ae 632 Fy COO TaD Vyagta eis By ai yo OP Gone i Poe, a 635 Geological SHIStOry ki gcc Ase est ca a ee 636 Clinnate} ect en Sa As ee ee ee 637 WMECAtONi a 32.046 cole oti Alt oe, eke as cas GF rae ere 639) GOMPOSITION ‘OF THE HIERPETOPAUNA, =... .. 0-4 See ee 650: E-COLOGY.OF ‘THE HIERPETOFAUNA. 4. «.sa56.5 gs.) + Stein s cs oe 650) Distribution Within: Habitats: «6 -. c.s0 0.50 29: ol ea ee 651 Altitudinal: Wistributiony -49. 965 eee eee 659) BactorseAttectingy DishabutiOne 22 rie ees te ee 662) nmtluencesob+ Matic... Bityed ore oro 2s ee gig ae .... 669 Analysis of Ecological) Distributiom 2.4.0 >. 22650: ci ae 670 CEOGRAPHY OF THE HIERPETOFAUNA ..0 ). 0 44606085 uo sh se ae 673 Methods of Zoogeographic Analysis ................. ede: Distnibution-or-the. Herpetotauna™ «n,n. sen ee 677 Patterns Oke Distt Dut OMe aoe yt ee ee ee 6seG ORIGIN) OF "THE HIERPEROFAUNA, =..,.... 65 586.0 oe Jee Oe 691 ORIGIN OF PRESENT PATTERNS OF DISTRIBUTION ......... ee OUD! Pleistocene Biogeosraphy 2.0.0. ace ee ee Pasa OS Dispersal Routes in Southwestern México .......... eos Speciation in Southwestern México ................. a enrktt) SURENEARY ©1520. ante eee ane nee Me a Ae re ere Eee ae 702’ EITERA TURES GITED 5) teeth es eaten ee ee eae 704 INTRODUCTION In a previous paper (Duellman, 1961) I presented a systematic account of the amphibians and reptiles of Michoaca; at that time I noted that a report on the biogeography of the herpetofauna was in (629) 630 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. preparation. The following is the faunal analysis anticipated at that time. The biogeographical synthesis presented here is, for the most part, based on data obtained in about twelve months of field work in Michoacan from 1951 to 1960 and on data obtained from the exami- nation of 9676 specimens from Michoacan and countless individuals from other regions. I have utilized all kinds of biogeographic evi- dence from various sources and have drawn freely on literature pertaining to Michoacan and on any other sources that offered sub- stantial evidence and ideas. All species of amphibians and reptiles that occur in Michoacan have been included in this synthesis, not only as they occur in Michoacan, but from beyond the limits of the state. Also, species that inhabit areas adjacent to Michoacan but that do not occur in that state have been included. Consequently, most of the species of amphibians and reptiles that inhabit south- western México have been taken into consideration. Even so, the present work is definitive only for the herpetofauna of Michoacan. I hope that this work can be utilized as a basis for comparative faunal syntheses of adjacent regions in southwestern México. In the preparation of this report I have given considerable thought to various zoogeographic principles and to methods of zoogeographic analysis. I have taken the opportunity to discuss various methods and their applicability to the herpetofauna of Michoacan. The result is a modified system of faunal analysis, pro- posed in the section on the geography of the herpetofauna. Hope- fully, this system will be applicable to various groups of animals in other regions. Since 1931 herpetologists have been following Dunn’s historical classification of the American herpetofaunas. On the basis of new historical evidence and modern ideas of phylogeny within the amphibians and reptiles I propose some modifications of Dunn’s classification. A comment is needed regarding terminology. Throughout this report I use the term “herpetofauna” to mean the total aggregate of species of amphibians and reptiles in a given region, without regard to ecological communities or historical origins. This term has been in use for many years; I think it is more meaningful than the terms “herpetogeny, herpetozoa, and herptiles.”. Many writers have used the terms “faunal assemblage” and “faunal element” synonymously. For the sake of convenience, I have used “faunal assemblage” to mean a geographic or ecological division of the herpetofauna, and I have used the term “faunal element” to mean a group of species BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 631 having the same geographic history, without regard to present as- semblages. The ecological terminology used here is that of Ken- deigh (1961). In order to be consistent and avoid confusing non-taxonomists, I have used here the same names for taxa that were used in the systematic account (Duellman, 1961), even though some taxonomic changes have been proposed since that date. In the list below the names given in the left hand column are those used by Duellman (1961) and the names given in the right hand column are those names currently in use. Bufo compactilis compactilis=Bufo compactilis Hyla baudini=Smilisca baudini Phyllodactylus homolepidurus=Phyllodactylus davisi Phyllodactylus lanei=Phyllodactylus lanei rupinus Cnemidophorus scalaris=Cnemidophorus scalaris scalaris Conophis vittatus vittatus=Conophis vittatus Acknowledgments Since the synthesis presented here is based on my previous work on Michoacan, all persons listed in the acknowledgments in that paper were in some way helpful in the accumulation of the data used here. I am especially grateful to Donald D. Brand of the University of Texas for his advice and helpful criticism of parts of the manuscript. Rogers McVaugh of the Univer- sity of Michigan Herbarium patiently spent many hours helping me with bo- tanical data and literature. Also, I am indebted to him for critically reading the section on vegetation. Much of the work on this report was done while I was associated with the Museum of Zoology at the University of Michigan. There and at the University of Kansas I profited from many discussions on Mexican biogeography with numerous colleagues. Which of the ideas pre- sented here are theirs and which are mine I can no longer say, but I claim full responsibility for the statements in the following pages. Lastly, I thank Ann S. Duellman who aided me in gathering data, discussed with me many of the ideas presented here, and critically read the manuscript. I respectfully dedicate this work to the memory of Professor Norman E. Hartweg, who fostered my early ambitions in Neotropical herpetology, offered guidance to this neophyte, and set an example of scholarly endeavor; were it not for him, this work might never have been initiated, let alone brought to completion. I only hope that he would have been pleased with the results. DESCRIPTION OF THE AREA The state of Michoacan comprises an area of 60,093 square kilometers (Vivo, 1958), lies between 17° 54’ and 20° 13’ North Latitude and 100° 02’ and 103° 47’ West Longitude, and borders on the Pacific Ocean for a distance of about 200 airline kilometers. The rugged terrain has a total relief of nearly 4000 meters. 632 UNIVERSITY OF KANSAS Pusts., Mus. Nat. Hist. Physiography and Geology If a person were to view Michoacan from an altitude of several thousand meters at a point off the coast of southwestern México, he would see in some places a narrow coastal plain with mountains rising to an elevation of about 3000 meters, then an abrupt drop into a low broad valley followed by a chain of volcanoes, some of which rise to elevations of nearly 4000 meters. Beyond these volcanoes at a somewhat lower elevation he would see a plateau, the rolling terrain of which is frequently broken by small mountain ranges and lakes. My purpose here is to correlate the physiographic features of Michoacan with those of all southwestern México, especially those thought to be important in interpreting the distribution of the herpetofauna in the region. Thayer (1916) outlined the physiographic regions of México. His work subsequently was modified by other geographers, such as Ordonez (1941), Tamayo (1941), and Robles (1942). Although the terminology of the prov- inces differs in these works, the general conclusions of each are the same. Tamayo (1949) summarized the earlier work and presented a synthesis of Mexican physiography. The following classification of the physiographic prov- inces of Michoacan and surrounding area is only slightly modified from the synthesis given by Tamayo; the Spanish names for the provinces (if different ) are given in parentheses (Fig. 1). LOWwLANDs.— Pacific Coastal Plain (Planicie Costera del Pacifico ) Balsas-Tepalcatepec Basin ( Depresi6n del Balsas ) HiGHLANDS.— Mexican Plateau (Mesa Central or Altiplanicie Mexicana ) Cordillera Volcanica Sierra de Coalcoman ELEVATION IN METERS 4- 3000 2250 500 600 Sea Level 5 0 25 50 75 100 on ——— 25 50 5° Fic. 1. Physiographic map of southwestern México. BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 633 The coastal region of Michoacan has been studied in detail by Brand (1957, 1958); most of the information presented here is a summary of his extensive reports. Although some areas seem to have had a recent history of submer- gence, most tof the coast seems to be rising. Only in the northwestern and southeastern coastal areas in Michoacan does an actual coastal plain exist. The plain is broad at the Rio Coahuayana (the boundary between Colima and Michoacan); from this point the plain becomes narrower until Punta San Juan de Lima is reached about 10 kilometers airline from the mouth of the Rio Coahuayana (Boca de Apiza). From Punta San Juan de Lima to Las Penas (about 150 kilometers airline) the coast consists of rocky promontories and sea cliffs separated by sandy or shingle beaches (PI. 1, fig. 1). Some of the prom- ontories, such as Punta San Juan de Lima, rise abruptly from the sea to eleva- tions of more than 200 meters. Many of the beaches are short and crescent- shaped; others are five to eight kilometers long. From Las Penas to the delta of the Rio Balsas, a distance of about 30 kilometers, there is a coastal plain; this is broadest near the Rio Balsas, where the hills rise some 12 kilometers inland from the sea. Small lagoons are found in areas where a coastal plain exists. Some of the promontories are composed of limestone. Others are composed of granites, andesites, and diorites; these are a part of an intrusive mass underlying the limestone of the Sierra de Coalcoman and coming to the surface in places as far as 20 kilometers inland. Some of the beaches are composed entirely of sand; others are composed of a mixture of sand and rock, whereas still others are rocky (shingle beaches). According to Swadley (1958), the beach sands are predominately of igneous and metamorphic sources, with minor traces of volcanic sands. His analysis of beach sands indi- cates a southeast long-shore current. Lying north of the Sierra de Coalcoman and the Sierra del Sur and south of the Cordillera Volcanica, and extending WNW-ESE is a broad structural depression, the Balsas-Tepalcatepec Basin. The western part, beginning in southern Jalisco and about 130 kilometers in length, is occupied by the Rio Tepalcatepec, a major tributary of the Rio Balsas, which occupies the eastern part of the depression. From the point of junction of the Rio Tepalcatepec with the Rio Balsas, the basin extends eastward for about 400 kilometers, ter- minating in southwestern Puebla and northwestern Oaxaca. In Michoacan the floor of the valley varies from 200 to about 700 meters above sea level. The terrain is relatively flat or rolling, frequently interrupted by barrancas formed by the many tributaries of the major rivers. The Rio Tepalcatepec closely follows the front of the Sierra de Coalcoman, which rises abruptly from the floor of the valley (Pl. 1, fig. 2). The scarp on the north side of the basin is more gentle; its original nature has been obscured by extensive volcanism and erosion. Apparently the Tepalcatepec Valley and the lower east-west versant of the Balsas Valley represent a graben formed by faulting along what is now the northem scarp of the Sierra de Coalcoman and Sierra Madre del Sur, and a similar scarp on the north side of the basin. The presence of numerous hanging valleys in the north-central part of the Sierra de Coalcomaén and in the Guerreran section of the Sierra Madre del Sur help substantiate this hy- pothesis. Seemingly the base rock of the depression is shale and sandstone; both are found as extensive exposures in the barrancas of the Rio Marquéz and Rio Cancita. Superimposed on the base rock is a variety of alluvium eroded from the surrounding mountains. Volcanic rocks are abundant, espe- cially in the Tepalcatepec Valley. The headwaters of the Rio Tepalcatepec 634 UNIVERSITY OF KANSAS PuBLS., Mus. Nat. Hist. in southeastern Jalisco, especially the tributary, Rio San Gerdénimo, are only narrowly separated from the headwaters of the Rio Ahuijullo, a tributary of the Rio Coahuayana. Ridges less than 600 meters in elevation intervene be- tween these river systems. The Ahuijullo depression presents a continuous lowland connection between the Tepalcatepec Valley and the lowlands of Colima; possibly in the past this depression was the drainage outlet of the present Balsas-Tepalcatepec Basin (see following section of the geological his- tory). Shortly below the confluence with the Rio Tepalcatepec, the Rio Balsas flows in a southerly course through a narrow gorge separating the Sierra de Coalcoman from the Sierra Madre del Sur and emerges onto the coastal plain and flows into the Pacific Ocean (PI. 2, fig. 1). The Mexican Plateau, which comprises the central part of México, reaches its highest elevations in the south; in Michoacan the plateau varies from about 1500 to 1900 meters above sea level. It is bordered on the south by a chain of volcanoes making up the Cordillera Volcdnica. Between the western ex- tension of the main part of the Cordillera Volcanica (the Volcan and Nevado de Colima are outliers to the south) and the southern extension of the Sierra Madre Occidental into northern Jalisco the central tableland is not separated from the coastal lowlands and the upper Tepalcatepec Valley by high moun- tains; instead the plateau gradually breaks down into rolling hills descending into the lowlands. Apparently the chief base rock of the plateau is limestone; conglomerates, rhyolitic and andesitic volcanic rocks, and sandstones are super- imposed on the base rock. Lacustrine deposits surround the large lakes indi- cating that these lakes once were more extensive than at present. The nearly unbroken chain of volcanoes comprising the Cordillera Vol- canica extends along the southern scarp of the Mexican Plateau from Jalisco to central Veracruz. These volcanoes, the highest in México, are on the southern edge of the plateau and lie roughly along the nineteenth parallel. They are Nevado de Colima (4300 m.), Nevado de Toluca (4560 m.), Popocatépetl (5400 m.), Ixtaccihuatl (5150 m.), La Malinche (4460 m.), and Citlaltépetl (5600 m.), as well as the highest mountains in Michoacan: Cerro San Andrés (3930 m.) and Cerro de Tancitaro (3870 m.). The most recent cones are Volcan Jorullo, lying to the south of the Cordillera Volcdnica and formed in 1759, and Volcan Paricutin, formed in 1943. Different parts of the Cordillera Volcanica are known by different names. The area between Cerro de Tanci- taro and Patzcuaro is called the Sierra de los Tarascos; the range immediately to the east of Morelia is called the Sierra de Ozumatlan; the area to the north and east of the Sierra de Ozumatlan and including the Cerro San Andrés is frequently referred to as the Serrania de Ucareo; the high range just east of Zitacuaro and extending along the western border of the state of México is called the Sierra de Temazcaltepec; the high volcanoes in the central part of the state of México, in the Distrito Federal, and in northern Morelos are re- ferred to as the Sierra Ajusco. The rugged terrain of the Cordillera Volcanica is formed by hundreds of volcanic cones, especially in the Sierra de los Tarascos (Pl. 2, fig. 2). These mountains are invaded from the lowlands by numerous barrancas resulting from the headward erosion of many of the larger tributaries of the Rio Balsas and Rio Tepalcatepec. According to studies of the volcanoes of the Paricutin region by Williams (1950), some intrusive granitic rocks are present on the surface, but most of the rocks are extrusive andesites and ba- saltics. The soils are principally volcanic sands and ash. BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 635 Lying between the Rio Tepalcatepec and the Pacific Ocean and extending from the Rio Coahuayana and Rio Ahuijullo on the west to the Rio Balsas on the east is the Sierra de Coalcoman, a mountain mass some 200 kilometers in length and 80 kilometers in width. Apparently the highest mountain in this range is Cerro de Barolosa slightly more than 3000 meters above sea level. Structurally this range is part of the Sierra Madre del Sur Physiographic Prov- ince of Guerrero and Oaxaca (Tamayo, 1941, and Robles, 1942), but physio- graphically the Sierra de Coalcoman is isolated from the Sierra Madre del Sur by the low valley of the Rio Balsas. The seaward base of the Sierra de Coal- coman is made up of intrusive granites. The remainder of the sierra is mostly limestone. The central part of the range is a karst terrain of sink holes, caves, underground rivers, and dogtooth limestone. For the most part, the seaward slopes of the Sierra de Coalcoman are well eroded by the numerous streams and rivers flowing into the Pacific Ocean. Contrariwise, the northern face of the sierra, at least in the central and highest part, is an escarpment dropping into the Tepalcatepec Valley. A. few small streams flow through hanging val- leys and cascade into the valley. The principal soils in the Sierra de Coal- coman are clays. Hydrography With the exception of the numerous coastal rivers, the drainage of Michoacan falls into two river systems, the Lerma-Santiago and the Balsas-Tepalcatepec. The Rio Lerma forms most of the northern boundary of the state and flows into Lago de Chapala. The Rio Duero in the extreme northwestern part of the state is the only important tributary of the Rio Lerma in Michoacan; most of the other tributaries are intermittent streams. Lago de Chapala is drained by the Rio Grande de Santiago which flows northwestward into the Pacific Ocean. Essentially all of the Cordillera Volcanica and most of the Mexican Plateau in Michoacan are drained by the Rio Balsas and its principal tributary the Rio Tepalcatepec. The major streams flowing southward from the Cordi- llera Volcanic into the Balsas-Tepalcatepec Basin are from west to east: Rio Cancita, Rio Cupatitzio (= Marquéz), Rio Tacambaro (= Turicato), Rio Chinapa, Rio Tuxpan, and Rio Zitacuaro; the last three rivers flow into the Rio Cutzamala, which in tum flows into the Rio Balsas. No large streams flow northward from the Sierra de Coalcoman into the basin. As mentioned previously, after receiving the Rio Tepalcatepec, the Rio Balsas flows south- ward through a narrow gorge into the Pacific Ocean. With the exception of the Rio Coahuayana and the Rio Balsas, which drain relatively little of the Coalcoman block, the major streams draining the Sierra de Coalcoman and flowing into the Pacific Ocean are from west to east: Rio del Ticuiz, Rio Ostula, Rio Cachan, Rio Tupitina, Rio Nexpa, Rio Mexcalhuacan, Rio Chuta, and Rio del Carrizal. All of these originate in the highlands and flow in a southerly course to the sea. On the southern part of the Mexican Plateau are several lakes, the three largest of which in Michoacan are: Lago de Chapala in the northwestern part of the state and partly in Jalisco, Lago de Cuitzeo in the north-central part of the state and partly in Guanajuato, and Lago de Patzcuaro at the edge of the Cordillera Volcdnica (pl. 3, fig. 1). Other lakes in the state include Lago de Zirahuan at the edge of the Cordillera Volcanica and Lago de Camécuaro near Tangancicuaro. Although Lago de Cuitzeo has an extensive lakebed, in dry years only a small part of it contains water. 636 UNIVERSITY OF KANSAS PusBts., Mus. Nat. Hist. Geological History The following is a brief summary of the historical events that have formed the present terrain of Michoacan and that are important in understanding the distribution of plants and animals in the region. During the Cretaceous the area that is now Michoacan may have been part of a seaway known as the Balsas Portal that connected the Pacific Ocean and the Atlantic Ocean in the region of the southern part of the Mexican Geosyncline (Schuchert, 1935, and Imlay, 1944). Coinciding with the Laramide Revolution at the end of the Cretaceous there was an uplift of the Mexican Plateau. Following this uplift was a period of peneplanation; a subsequent orogeny with uplift took place again in the Miocene and Pliocene and raised the Mexican Plateau to its present elevation. The development of the Cordillera Volcanica is correlated with the last period of orogeny. According to Flores (1946) and Blasquez and Garcia (1946), the oldest volcanic rocks in the Cordillera Volcanica in eastern Michoacan are of Miocene age. Alfonso de la O. Carreno (1943) as- signed the great cones of the cordillera, including Tancitaro, Nevado de Colima, Nevado de Toluca, Ixtaccthuatl, and Citlaltépetl, to the Pliocene. Ap- parently the volcanism initiated in the Miocene has continued without notable interruption to the present time. The Sierra Madre del Sur was elevated at the end of the Cretaceous; dur- ing the Miocene another period of orogeny took place accompanied by ex- trusive and intrusive activity; this was followed by a period of volcanism and elevation again in the late Pliocene and Pleistocene (Garfias and Chapin, 1949). The historical geology of southern Michoacan has been summarized by Brand (1958:150-2): “The major tectonic developments in the Coalcoman region seem to have been as follows. We start by assuming an old landmass (sometimes called by the uncouth name of Del Sur Borderland) on the seaward side of the present shore or overlapping slightly on the present land. In Paleozoic and Mesozoic times this borderland varied from a narrow band that completely separated the Pacific from the Atlantic, through a series of long islands, to a peninsula—so far as our southwest Mexico is concerned. This borderland may have pro- vided a basement of pre-Cretaceous crystallines if we attribute to it some of the metamorphics which are found along the coast from Jalisco to Chiapas. In Guerrero, Oaxaca and Chiapas the Jurassic-Cretaceous formations lie dis- cordantly upon a basal complex of metamorphics. At any rate, this border- land contributed to the Cretaceous deposits which (especially in the Middle Cretaceous) have a marine fauna with a marked inshore or littoral nature. In the late Cretaceous-Paleocene there was an extensive granitic intrusion, up- lifting, folding, overthrusting, and faulting. Great sinkings took place parallel to the folds, both on the seaward and landward sides. By the end of the Eocene the Mexcala-Balsas-Tepalcatepec graben had been down faulted as well as the Manzanillo and Acapulco troughs. So far the nature and chronology of the tectonic activity seem to be fairly well indicated by evidence from various portions of the socalled Sierra Madre del Sur Province, along the coast from Cape Corrientes to the Isthmus of Tehuantepec. “During the Tertiary there were two outstanding times of volcanic erup- tions and extrusions along fault lines—during the Paleocene-Eocene and dur- ing the Miocene-Pliocene. Apparently only the earlier of these extrusive periods was important in the Coalcomaén region. There followed widespread denudation and erosion which stripped off most of the Tertiary volcanic mantle as well as the Turonian sedimentaries, and developed a peneplain surface. The remnants of this peneplain are seen best in the accordant summits between the Ostula and Tupitina rivers and inland past Tehuantepec BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 637 and San José de la Montafa. Again there was uplift, accompanied by local intrusions that resulted in dikes that cut the granitics as well as the Cretaceous sedimentaries. As a result of this uplift much of the seaward slope of the Coalcoman region was dissected to a late youth stage. “Towards the end of the Tertiary, during the Miocene and Pliocene when some other parts of Mexico were being mantled with extrusives, there was a sinking along the Pacific coast. Maria Madre Island in the Tres Marias, for example, shows Miocene and Pliocene deposits, and probably deposits of this time underlie the Pleistocene and Recent deposits along the costs of Jalisco and Colima and in the Balsas delta. It is possible that at this time a head- ward-eroding stream tapped a great lake occupying the Mexcala-Balsas- Tepalcatepec graben and formed the Balsas river. This great lake, which did not last long enough to leave much evidence, may have been draining earlier to the west via the Ahuijullo depression. “More recently there was a down warping of the entire coastal section of Michoacan along a hinge line that can be traced some 15 or 20 miles inland from and parallel to the shore. This hinge line is marked by such items as rapids in the Cachan and Balsas rivers, seismic epicenters, some affluents that enter their streams at nearly right angles, and possibly one of the two hot springs in southwestern Michoacan. This down warping probably produced the drowned effect at Calestas de Campos and other spots on the Michoacan coast and probably also on the Costa Grande of Guerrero. Also this down yang increased the erosive action and the carrying capacity of the streams along the coast, which probably explains the pebble and boulder conglomerates in the Balsas delta and elsewhere. “Today the southwest coast of Michoacan is characterized by little vul- canism, great earthquake activity, little or no coastal plain, and a positive or upward movement. The Coalcoman block especially typifies these charac- teristics. There are no recent volcanic cones or basaltic flows, and there are but two hot springs (at Cuchumala northeast of Coire, and the Aguas Calientes de Cuilala) and the warm mineralized spring at La Majahuita.” The historical events that seem to have had the greatest effect on the present distribution of the herpetofauna in southwestern Michoacan are, of course, the more recent ones. Among these are the more recent independent histories of the Cordillera Volcanica (characterized by extensive volcanism) and the Sierra de Coalcoman (without volcanism), the relatively late con- nection of the Balsas-Tepalcatepec Basin to the westem lowlands via the Ahuijullo depression and only recent connection of this interior basin with the coastal lowlands of much of Michoacin. Climatic fluctuation during the Pleistocene no doubt was important in establishing many of the present pat- terns of distribution. The Pleistocene biogeography will be discussed in a subsequent section of this paper. Climate The great amount of physical relief in Michoacan not only produces con- siderable climatic variations, but delimits these variations along sharp bound- aries. Temperature within the region depends primarily on altitude (the cool- ing effect of the onshore breezes along the coast being an exception), and precipitation depends on the prevailing winds and the terrain (Fig. 2). An analysis of vertical temperature gradients in Michoacan indicates that on an average the mean annual temperature decreases slightly less than 1° C. per 200 meter increase in elevation. Adequate meteorological records are not available for coastal Michoacan, but records for Manzanillo, Colima, and La Unién, Guerrero, show that at Manzanillo the mean annual temperature is 26.1° C. with a range from the coolest month (March) to the warmest 638 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. 35 — T SS Sra aa tt ats lami A Bi OR L { be | | a cn, Sear cA Sates ee EL UGONGING te ec ae oe a or ee Halarbistindia., Merc. eae ee TL GSCLUMN AE so He aa et oe athe MOOR FR lareniey ee ccs ees ec a Hyla microcephala sartori................ Hala/smaragding. 5 ose een ie rae Hla Bria ee RRR ee ee eee Gastrophrynemsta Usta's. \ 304 0) oe sin as oe Hypopachus caprimimus.........00.005- Hypopachus oryrrhinus ovis... .........- ONE QUINT: ss Pcie ae oo ones ee TAY, MeEGADOOM. BRA Oe Ace sien ese Rana Montezumaes «owns oe ee TEONGDUDTENS oR 8h Os OR ee ee Arid tropical scrub forest deciduous forest Tropical semi- Mesquite- r Ie uaigr™ Setpoint al AIT alg hea wi OIG eel) Wezel el aMelb Teele fe sliele| | seercl)seaie de ee [ee) aed > | J ie lle sees | ee a eit ah abt tea sss le ha cage grassland PSssro| |x| Pine-oak forest fell Sees ares ccoce lass! fs ea al cable eal) Bsa gelto elle Nhe tie Fir forest a < heerel Se leah ovale Se Mee ie PSEC | lp le) alse eas a onl Altitudinal range In meters 2300-2800 2165 2400-3000 2000-2200 1500-2900 2900-3000 0-10 1500-2500 335-640 1525-2050 0-1000 0—1000 900-2400 180-1630 180-1050 0—1050 1700 1450-2750 20-1500 130-1630 1500-2500 2100-2200 1200-1800 20-300 335-1630 1600-2100 1600-2400 1500-2300 Vey 150-1500 0-1000 ? 200-1800 1500-2200 2000-2165 1525 1500-2000 0—-2800 BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 653 TABLE 2.—ECOLOGICAL AND ALTITUDINAL DISTRIBUTION OF AMPHIBIANS AND REPTILES IN MicuoacAn.—Continued SPECIES LRA J OISMORO sol to0.5.006-010.6.0. 000, 6080 OG Kinosternon hirtipes hirtipes............- IKGNOStERILONUANLEG HUTT itis acreeiele © eid = ae Geoemyda pulcherrima pulcherrima........ Geoemyda rubida perixantha...........-- PRSCuUdeMYS:SChUDLGONTUALG:s ae Neias ts 4 1-0 1 Crocodylis acutus AcCutuse oso. es ne ee BSUDESICO LGU CULES seo eraleteyt shal re Seerere sce Coleonyx elegans nemoralis..............- Bhyllodactylusduellmant.... .2=..5.5-.:-- Phyllodactylus homolepidurus............ phy lodactyLUsilanetceh. mace elrne tick cate Phyllodactylus paucituberculatus.......... PA NOLISEAUTUN Up teg dowehare. cetera gets Eee etter Iguana iguana rhinolopha............... OUP UTRHUIRE WAG, SOS aoe oaoebos 6 odes EIU COSCAUT US ECLON IG ile terrence ateteie etek ZRTUMOSOTIO OSLO aE Wetereeliete <1 seecie Cte Sceloporus aeneus aeneus............-+-- IS COLO DONUS AS DEN 7 cor uit, Seater a - oee eta 3.55858 IS CCLODOWUSIOULLEN I = her: eregeke REA eee G Sceloporus dugesi intermedius............ ISCCLOPOTUSIGACOWAE an reenter eee oe Sceloporus grammicus microlepidotus... ... ISGELOPOMUSINCLETOLEDIS Ys Ayaan -itarie Siecle ve oe Sceloporus horridus oligoporus............ Sceloporus melanorhinus calligaster........ ISCELODORUSEDUNOCEDILALUS asus ransg servos, tele Sceloporus scalarts scalaris............... Sceloporus siniferus siniferus............. Sceloporus spinosus spinosus...........-. Sceloporus torquatus torquatus............ ISCCLONORUSMULU ONIN ISHN: ira teetacs iets eer Urosaurus bicarinatus tuberculatus........ WROSCUTUSOAdOWU ee toe ae ects ee ee Mabuya brachypoda. oa a uss. ots oe: 21> SCUNCELUG ASSOLGILAYLONUe mea a te oe ae eet OVATE HOT OUR 005.6 &00.4.0.0 oo date Oaoe EUIMECES COLLIMLENSTSen ee sues ee IHAUTCCES COPE HMer ares xae ete SOME EUTLECES OULD CSUR EERE a Paseheisee i oernic ie: EU CCES ANG ULUULUSeE ae cael eer eerettieke he oe IB QUNECESEPOLVULLS emp ienne = Cae rere ae Amewa Uunaidata sintstras . 2 Bh os see eo Cnemidophorus calidipes............++-- Cnemidophorus communis communis...... Cnemidophorus costatus occidentalis....... Arid tropical scrub forest ml. Tropical semi- deciduous forest Mesquite- tite AC Th letccll Pach aril) a8 Rca | be sls ed scans cates oI ae] grassland PSS Vico OO Tie TST Mecha SPe eS PO Ae ies | ea seers a Al essen teal Gl sede at ea tlre sisal mal aa) alee - = —— Selita Weleda ae eal sslealesl tts) Pine-oak forest Fir forest Altitudinal range In meters Pet AL eTRSID I sco TS re Se Se Te eres cl a Rete 8 850-2150 1525-2160 Ce 0-635 ? 0-10 ? ? 180-500 1850-3100 335-1630 2000-2350 1500-2200 250-1050 1550-3100 1800-2700 0-1600 30-1500 0-1000 1550-2300 0-150 1500-2300 1450-3000 0-1550 0-1700 180-1200 0-1050 120-850 350-1050 130-950 1800-2700 1550-1850 2750 30-500 10-950 200-650 10-950 1570 654 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist, TABLE 2.—ECOLOGICAL AND ALTITUDINAL DISTRIBUTION OF AMPHIBIANS AND REPTILES IN MicHoacAn.—Continued SPECIES Arid tropical scrub forest deciduous forest Tropical semi- Mesquite- grassland Pine-oak forest Fir forest Altitudinal range in meters Cnemidophorus costatus zweifeli.......... Cnemidophorus deppet deppet............ Cnemidophorus deppei infernalis.......... Cnemidophorus lineatissimus exoristus..... Cnemidophorus lineatissimus lineatissimus. . Cnemidophorus lineatissimus lividus....... Cuemidophorus scalarts |". t. uc 2 veoh ess os: Heloderma horridum horridum............ Gerrhonotus «mbricatus imbricatus......... LM PNLODSHONQMANUS a=. ec. crcteke oats ae sisie: Leptotyphiops bressont.... 2... 5.58 5.505585 Bentotiyphilops GAdowt:. 0s. eles seams we ws Leptotyphlops phenops bakewelli.......... EGxOCeniis O1COlOT SF). 28 3 ia es oe ok eee (BOONCOStTLCLOT, VN. DET GLOT Ax «ls Bas cre 8 te oe GUAT NS COT IE RENAE ere CaP ig Sterne aera ae Coniophanes fissidens dispersus........... Coniophanes lateritius lateritius........... Conophis vittatus vittatus................ Conopsis, Ot8ertaliss 3. sca ine cee hs Fyne CONOPEIS ASUS. 4.2 Si. sed eas TS oe ite DUGdODNTS QUGES1). sc noha secs eee . eets <4 Dryadophis melanolomus stuart... ....... Drymarchon corais rubidus.............- Drymobius margaritiferus fistulosus....... Elaphe triaspis intermedius.............. MAUS ECOLOT 2 hte Ciera sete escent Geagnasinedvmitus tues Sie Oe A Geatractus tecpanensis.............2.045- GeODNISHGUGOSUR cache Senos RAE aes, oe Geenhisinconiplus:. 280% eee... ee aes Geophisemaculaferus: <2. eou5 ee ames =) Geophis NignOcinClustues. 42s aes cies «te Geophis Merer syn dn. Mgt ee ee Pee GeEODRISHOTGSCHE ja Fass ee sees aoe Hypsiglena torquata ochrorhyncha......... Hypsiglena torquata torquata............. Imantodes gemmistratus gracillimus....... Imantodes gemmistratus latistratus........ LGiMpnropellts GOUGIGs cic). oan nee eee QM DLODelUUSEVUL VEN Ae Bian o eee eh aus entodema:latufasctata: oo. ...9¢ «eee 32 vases eptodema maculatat.w. etek ee ones Leptodeira septentrionalis polysticta....... Leptodeira splendida bressoni............. Leptophis duplotropiss. a. a- feck. 2. eae Moanolenis putnamir. 3 st acter hoes © tise Masticophis striolatus striolatus........... Masticophis taeniatus australis........... Oxybelts aeneus auratus inn cee eee Peco er MMe ewee APSE Drea (etna Meee ess fe aT | tal Satie Aes eae se Ie 200-1100 130-500 200-1050 170-900 0-10 0-500 1600-1900 0-1000 1500-3500 335-850 1050 800 0-850 0-640 0-1000 850 900 800-1100 1550-2800 1900-2450 1900-2200 0-50 0-1350 0-1150 350-1350 900-1800 2 1750-2050 2100 1900-2200 335-1050 Veet 950-1650 850-1700 20-120 0-1650 1570-2000 0-1700 BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 655 TABLE 2.—ECOLOGICAL AND ALTITUDINAL DISTRIBUTION OF AMPHIBIANS AND REPTILES IN MicHoacAn.—Concluded = 84 g a 2/83] 3| 5 a% SPECIES Se Sl Seah hae le Ee ee ee yh tes =< E2tuophis dep perdeppery oun . staan oe ee — M | — | — | 1900-2200 Pituophis lineaticollis lineaticollis......... —= |— |.B | M | — | 1500-2300 PPSEUCOPLCITIVIG J RONUGIIS os 8) sue os ey nF M 330-1050 Pseudoficimia pulcherrima.............-. R 330 Rhadinaea hesperia hesperia............. R|M — }— | 850-1500 eRQOINQCOIAURCALERTS ears can era —}|— A | — | 1800-2300 LUROAINGCO (AGRI ALO meray ie Sea —/}/—/;—;R}i— 1850 ISETUTE LA GMAT UIT 8 ety OC es OOO IS — |—]|A | M |} — } 1550-2500 SOAWACOTEMNCTICON a0. scr. cent wo si no eae oes A }—{]—|]—/]— 0—1000 IS PUOT LCDI LUST CeCe — | R | =| =) => 150 Sonora michoacanensis michoacanensis.... . M | — | — | — | — | 380-1000 GNIAUG.DOCONTDA. Aiwosins casdascrdecaetes ene — |— | — | M |} R__| 2000-2200 ATOLL CON OINAT TIVO ojo e122 che oss aueie * Sr Rea | | | 20-330 TL OLUCT AN EOUE LN EQUA. aval, cmos Si se) 2 ik — |— | — | A | M | 1550-2800 Trimorphodon biscutatus biscutatus........ A |— |— 20-1050 UiVMOnpROdOnlaltfasctas «5.4.4. we ee ae M | — | — | — | — | 300-1480 LGLIMOTPROCOTOTAU. sie... 2 TERY aes, oo eet — |—]/R | — | — | 1600-2000 Tropidodipsas occidentalis................ —|Ri—|]—/]— 950 NGtrte UGA ISAVEEGE) on nae ee es nee es M | —/—/]—/]— 0-10 ISLONENID SLONETVOLNES oe eA sae sore oe — |—/M | R |} 1550-2800 Thamnophis dorsalis cyclides............. — |—]M|A /R | 1550-2800 Thamnophis dorsalis postremus........... A |R |—|— |—| 200-1550 DRAMNGPRISKEQUCS EQUES 1 Ais oe Pid ois ys — A | M | — | 1550-2300 Thamnophis melanogaster canescens....... — | — | M | — |} — | 1550-2200 Thamnophis scalaris scaliger............. — |—]/]—|R | R | 1800-3400 Micrurus disians michoacanensis.........| M | — | — | — | — 350 VEG CHINES HOLL COLLCT IS «lan dte ene a ial etere: Reka acl R |—/—/]—!]— | 400-1050 Agkistrodon bilineatus bilineatus.......... M. | — | — | — | — } <300-1500 Crotalus basiliscus basiliscus............. M/}—/]—/|—!/— 20-1070 Crotalus durissus culminatus............. Me th | 1050 Crotalus intermedius intermedius.......... —}—|]R | — 2200 Crotalus molossus nigrescens...........-- — |— | — | M | — | 1550-2300 CEOLOVUS DOLUSUCLUS sor 2 ity cialis sae suse —|—]R | — | — | 1450-2400 Crolalustpusiliisnn ee an iea oe bk noe — |— | — | A _ |} — | 1550-2300 Crotalus triseriatus aquilus............... —}—]|M | M | — | 1600-2000 Crotalus triseriatus triseriatus............ —|—]|—|A | — | 1600-3270 In the same habitat on the Pacific coastal plain the following species are characteristic: Bufo marinus Bufo marmoreus Leptodactylus melanonotus Hyla baudini Hyla smithi Anolis nebulosus Basiliscus vittatus Ctenosaura pectinata Sceloporus horridus Sceloporus pyrocephalus Sceloporus siniferus Leptodeira maculata Characteristically the species inhabiting the arid tropical scrub forest are adapted for life in open, subhumid conditions. Diurnal species usually are fast and agile, such as Drymobius margaritiferus, Masticophis striolatus, Salva- dora mexicana, Basiliscus vittatus, Sceloporus siniferus, and the various species 656 UNIveRSITY OF KANSAS Pusis., Mus. Nat. Hist, of Cnemidophorus. Saxicolous diurnal species inhabiting natural rock out- croppings or stone walls include, among others, Ctenosaura pectinata, Sceloporus gadowae, Sceloporus horridus, and Sceloporus pyrocephalus. Few diumal species are arboreal, but Enyaliosaurus clarki lives in scrubby trees, and Urosaurus gadowi inhabits the trunks of small trees in the scrub forest. Many species inhabiting the arid tropical scrub forest are nocturnal. In this category are all of the anurans, of which Bufo marinus, Leptodactylus mela- nonotus, Hyla baudini, and Hyla smithi are the most widespread. Common nocturnal snakes are Leptodeira latifasciata, Leptodeira maculata, Pseudoficimia frontalis, Thamnophis dorsalis, Trimorphodon biscutatus, Trimorphodon lati- fascia, Micrurus distans, Micrurus laticollaris, and Crotalus basiliscus. Of the lizards, only Phyllodactylus duellmani and Phyllodactylus paucituberculatus are noctural. A higher percentage of species in the arid tropical scrub forest is nocturnal than in the other habitats in Michoacdn, probably because the nocturnal temperatures in that forest are optimal for many species. Some, for example Drymarchon corais and Drymobius margaritiferus, are diurnal in many parts of their geographic ranges but are at least partly noctural in the arid tropical scrub forest in the Balsas-Tepalcatepec Basin. In the pine-oak forest Thamnophis dorsalis cyclides is strictly diurnal, whereas the subspecies, T. dorsalis postremus, is nocturnal in the arid tropical scrub forest in the low- lands. The lack of a well-developed humus layer on the ground in the arid tropical scrub forest apparently excludes some of the small semi-fossorial species, such as Eumeces colimensis, Eumeces parvulus, Scincella assata, and Enulius uni- color. Likewise, the absence of an extensive arboreal habitat precludes the presence of such species as Sceloporus melanorhinus, Leptophis diplotropis, and Oxybelis aeneus, or greatly restricts their distributions. Tropical Semi-deciduous Forest The tropical semi-deciduous forest occurs in small isolated patches on the windward slopes at low elevations. Sixty-two species occur in this habitat; of these, only 11 can be considered abundant there. The great amount of vegeta- tional cover in this kind of forest provides ready concealment for most small animals, consequently many species that seem to be uncommon possibly are abundant. Characteristic inhabitants of the tropical semi-deciduous forest include: Microbatrachylus pygmaeus Eumeces parvulus Eleutherodactylus rugulosus Amevia undulata Phyllomedusa dacnicolor Cnemidophorus lineatissimus Phyllodactylus lanei Dryadophis melanolomus Anolis nebulosus Leptophis diplotropis Basiliscus vittatus Manolepis putnami Sceloporus melanorhinus Oxybelis aeneus Sceloporus utiformis Rhadinaea hesperia Most species inhabiting the tropical semi-deciduous forest are tolerant of hot, humid conditions. Many species, such as Microbatrachylus pygmaeus, Eumeces parvulus, Scincella assata, Enulius unicolor, and Rhadinaea hesperia, in- habit the mulch on the forest floor. Others, including Ameiva undulata, Cnemidophorus lineatissimus, and Dryadophis melanolomus, actively move about on the forest floor, whereas Phyllomedusa dacnicolor, Anolis nebulosus, BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 657 Leptophis diplotropis, and Oxybelis aeneus are primarily arboreal. The well- developed arboreal microhabitat in the tropical semi-deciduous forest is in- habited by approximately 25 per cent of the species there; this is a higher percentage than in other habitats in the state. Apparently, because of the rather equable terrestrial climate in the tropical semi-deciduous forest, a relatively high percentage of species there are diurnal. The shade provided by the dense foliage results in a darkened forest floor having minimal evapora- tion; consequently, small amphibians, such as Eleutherodactylus rugulosus and Microbatrachylus pygmaeus often are abroad by day. Mesquite-Grassland The scrubby vegetation occurring on the Mexican Plateau provides a def- initely subhumid habitat characterized by warm days and cool nights. Of the 39 species of amphibians and reptiles comprising the impoverished fauna there, only nine can be considered abundant. Among the characteristic species in this ecological assemblage are: Scaphiopus hammondi Cnemidophorus scalaris Bufo compactilis Pituophis deppei Hyla eximia Salvadora bairdi Hypopachus oxyrrhinus Thamnophis eques Sceloporus dugesi Thamnophis melanogaster Sceloporus spinosus Crotalus triseriatus Most of the species inhabiting the mesquite-grassland are terrestrial; the uncommon Hyla smaragdina, which takes refuge in small bromeliads in low trees, is arboreal. The primarily terrestrial Anolis nebulosus, an invader from the lowlands, often sleeps in bushes at night. The anurans are active chiefly at night; some snakes, such as Elaphe triaspis, Hypsiglena torquata, and Trimorphodon tau, characteristically are nocturnal. In the mesquite-grassland the margins of the several lakes on the Mexican Plateau form an important microhabitat. Rana dunni, Rana megapoda, Rana montezumae, Thamnophis eques, and Thamnophis melanogaster are typical inhabitants of those aquatic margins. Pine-Oak Forest The pine-oak forest is the most varied and extensive habitat in Michoacan and occurs in two areas—the Cordillera Volcanica and the Sierra de Coalcoman. Seventy species are known to inhabit the pine-oak forest; of these, only 17 are considered abundant there. In the pine-oak forest in the Cordillera Volcanica the following species are characteristic: Pseudoeurycea belli Gerrhonotus imbricatus Bufo occidentalis Conopsis biserialis Microbatrachylus hobartsmithi Geophis petersi Tomodactylus angustidigitorum Pituophis lineaticollis Hyla bistincta Rhadinaea laureata Hyla eximia Toluca lineata _ Sceloporus aeneus Storeria storerioides Sceloporus grammicus Thamnophis dorsalis Sceloporus torquatus Crotalus molossus Eumeces dugesi Crotalus pusillus 658 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. In the Sierra de Coalcoman the following species inhabit the pine-oak forest: Bufo occidentalis Gerrhonotus imbricatus Tomodactylus rufescens Geophis incomptus Hyla bistincta Geophis nigrocinctus Anolis nebulosus Pituophis lineaticollis Sceloporus bulleri Storeria storerioides Sceloporus heterolepis Thamnophis dorsalis Crotalus pusillus In the pine-oak forest the well-developed broad-leaf and pine needle mulch on the forest floor forms an important microhabitat for such semi-fossorial species as Pseudoeurycea belli, Microbatrachylus hobartsmithi, Eumeces copei, Eumeces dugesi, Eumeces indubitus, Gerrhonotus imbricatus, Conopsis_bi- serialis, Geophis dugesi, Geophis incomptus, Geophis nigrocinctus, Geophis petersi, Rhadinaea laureata, Tantilla bocourti, Toluca lineata, and Storeria storerioides. Lizards of the genus Sceloporus are the most conspicuous herpetological in- habitants of the pine-oak forest. In the Cordillera Volcanica Sceloporus aeneus is the common terrestrial species; S. grammicus is the ever-present arboreal species, whereas the larger S. torquatus lives on tree trunks and the ground alike. In the Sierra de Coalcoman Sceloporus heterolepis is the strictly arboreal species, and S. bulleri occurs as frequently on the ground as in the trees. Of the snakes and lizards occurring in the pine-oak forest, only about one- third of the species tend to be nocturnal. Many of these, such as Rhadinaea laureata and Toluca lineata, are mulch-inhabitants that apparently move about mostly at night. The four species of Crotalus inhabiting the pine-oak forest are partly nocturnal. Phyllodactylus lanei, the only nocturnal lizard in this habitat, occurs only at low elevations in the pine-oak forest. As in the mesquite- grassland, the relatively low nocturnal temperatures preclude much activity by snakes at night. Fir Forest This cool, moist habitat characterized by a deep, moist layer of mulch on the ground is discontinuous in Michoacan. Patches of fir forest of varying areal dimensions occur on the highest mountains in the Cordillera Volcanica and in the Sierra de Coalcoman. Of the 23 species of amphibians and reptiles known to occur in the fir forests in Michoacan, only four can be considered abundant there. Characteristic inhabitants of the fir forests in the Cordillera Volcanica are: Ambystoma ordinarium Sceloporus grammicus Pseudoeurycea belli Sceloporus torquatus Pseudoeurycea robertsi Eumeces dugesi Hyla lafrentzi Gerrhonotus imbricatus Sceloporus aeneus Conopsis biserialis Toluca lineata In the fir forests of the Sierra de Coalcoman the list is much smaller; pos- sibly this is due to less extensive collecting there. Known inhabitants of the fir forest in the Sierra de Coalcoman are Bufo occidentalis, Sceloporus hetero- lepis, Gerrhonotus imbricatus, and Storeria storerioides. The only arboreal inhabitants of the fir forests are Sceloporus grammicus, S. heterolepis, and S. torquatus, all of which are more widespread and abundant in pine forests at lower elevations. The other inhabitants of the fir forests are BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 659 equally divided between mulch inhabitants, such as the various salamanders, Eumeces dugesi, Conopsis biserialis, and Toluca lineata, and terrestrial species, such as Sceloporus aeneus and the various species of Thamnophis. All of the lizards are strictly diurnal, as are all of the snakes with the possible exception of some of the mulch inhabitants. Altitudinal Distribution In an area having such varied topography as Michoacan, different altitudinal distributions among the species are expected. The known altitudinal ranges of the amphibians and reptiles in the state are given in Table 2. Examination of this table reveals that of the 183 species and subspecies listed therein, only 43 have altitudinal ranges of 1000 meters or more; of those, only five species (Rana pipiens, Kinosternon integrum, Anolis nebulosus, Gerrhonotus imbricatus, and Thamnophis dorsalis) have altitudinal ranges of 2000 meters or more. An analysis of the limits of altitudinal distribution of the various species (Fig. 4) shows that the upper and lower limits of distribution of many species are reached at certain elevations, whereas fewer distributional limits occur at intermediate elevations. Of course, a large number (53) of species have the lower limits of their distributions at or near sea level. The lower limits of the altitudinal distributions of several other species approach sea level, but terminate between 100 and 200 meters above it on the slopes of the Sierra de Coalcoman. In this group are some frogs, such as Eleutherodactylus rugulosus and Hyla smaragdina, which live along streams having moderately steep gradients, and certain lizards and snakes, such as Scincella assata, Eumeces colimensis, and Sibon nebulatus, which do not inhabit the arid tropical scrub forest on the coastal plain. The group of species having their lower limits of altitudinal distribution at about 300 meters is primarily made up of those species that reach the lower limits of their distributions in the Tepalcatepec Valley and do not live on the coastal lowlands. Common species in this group are: Bufo perplexus Cnemidophorus costatus Hypopachus caprimimus Hypsiglena torquata Phyllodactylus duellmani Leptodeira latifasciata Enyaliosaurus clarki Pseudoficimia frontalis Sceloporus gadowae Sonora michoacanensis Eumeces altamirani Trimorphodon latifascia Cnemidophorus calidipes Thamnophis dorsalis Miscrurus distans Fewer species reach the lower limits of their distributions primarily between 800 and 1000 meters. These are species that mainly inhabit the pine-oak forests and also occur in the higher parts of the tropical semi-deciduous forest; among species in this group are Rana pustulosa, Leptodeira splendida, Lep- tophis diplotropis, and Rhadinaea hesperia. Next to the group of 53 species reaching their lower limits of distribution at or near sea level, the largest group contains 33 species reaching their lower altitudinal limits between 1400 and 1600 meters. Included in this group are species that are typical inhabitants of pine forest, such as: 4— 8425 660 UNIVERSITY OF KANSAS Pusis., Mus. Nat. Hist. Pseudoeurycea belli Eumeces dugesi Scaphiopus hammondi Gerrhonotus imbricatus Microbatrachylus hobartsmithi Conopsis biserialis Tomodactylus angustidigitorum Toluca lineata Tomodactylus rufescens Storeria storerioides Hyla bistincta Crotalus molossus Sceloporus grammicus Crotalus pusillus Also included in this group are species inhabiting the Mexican Plateau, which in Michoacan descends to an elevation of about 1500 meters. Plateau inhabitants reaching the lower limits of their distributions at this elevation include: Bufo compactilis Sceloporus spinosus Hypopachus oxyrrhinus Cnemidophorus scalaris Rana megapoda Masticophis taeniatus Rana montezumae Trimorphodon tau Kinosternon. hirtipes Thamnophis melanogaster Crotalus polystictus A small group of species reach the lower limits of their distributions at about 1900 meters; most of these are species that inhabit the cool montane forests, such as Ambystoma tigrinum, Tomodactylus fuscus, Sceloporus bulleri, Conopsis nasus, Diadophis dugesi, and Tantilla bocourti. Pseudoeurycea robertsi, which insofar as is known is restricted to the fir forest, reaches its lower limits at 2900 meters. All other species in the state occur at lower elevations. The upper limits of distribution of nine species are at about 100 meters; these species are restricted to the coastal plain and include: Rhinophrynus dorsalis Dryadophis melanolomus Phrynohyas inflata Geagras redimitus Crocodylus acutus Manolepis putnami Anolis schmidti Natrix valida The small group of species having their upper altitudinal limits between 300 and 500 meters are mostly those inhabiting only the lower parts of the Balsas- Tepalcatepec Basin and include: Diaglena reticulata Pseudoficimia pulcherrima Phyllodactylus duellmani Tantilla calamaria Enyaliosaurus clarki Micrurus distans The upper limits of altitudinal distribution for the greatest number of species (35) is between 900 and 1100 meters, where the arid tropical scrub forest of lower elevations is replaced at higher elevations by pine-oak forest. Typical of the species that characteristically inhabit the lowlands and reach the upper limits of distribution at this altitude are: Bufo marinus Ameiva undulata Leptodactylus melanonotus Cnemidophorus communis Phyllomedusa dacnicolor Cnemidophorus deppei Hyla smithi Heloderma horridum Phyllodactylus lanei Boa constrictor Ctenosaura pectinata Drymobius margaritiferus Sceloporus pyrocephalus Leptodeira maculata Mabuya brachypoda Salvadora mexicana Trimorphodon biscutatus Ten species reach the upper limits of their distributions at about 1600 meters; most of these species inhabit the lowlands and invade the lower parts BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 661 of the Mexican Plateau. Included in this group are Tomodactylus nitidus, Hyla smaragdina, Hypopachus caprimimus, Urosaurus bicarinatus, and Masti- cophis striolatus. Another group of species reaches the upper limits of their distributions at about 2200 meters. Most of these are inhabitants of the Mexican Plateau; included in this group are: Ambystoma tigrinum Sceloporus scalaris Hypopachus oxyrrhinus Lampropeltis ruthveni Rana montezumae Pituophis deppei Kinosternon hirtipes Trimorphodon tau Sceloporus dugesi Thamnophis eques Thamnophis melanogaster 20 ai EE ale ay rE fe) 7 a fal a ae a em female eal Pals tu Bo. Se eee oO. He secon ma a [re a a J Seo aaa ro} 1000 1500 2000 2500 ao Ww = =)5 Pa lO I5 ake LOWER LIMITS Fic. 4. Numbers of species of amphibians and reptiles reaching their upper and lower altitudinal limits at various elevations in Michoacan. 662 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. The group of species apparently reaching the upper limits of their dis- tributions at about 2800 meters possibly is the result of inadequate collecting, rather than actual upper limits of distribution, for few areas above this eleva- tion have been studied thoroughly. Eight species are known to occur at ele- vations of 3000 meters or more. Two of these are amphibians—Ambystoma ordinarium and Pseudoeurycea robertsi; both inhabit the fir forest. The six reptiles inhabiting the highest elevations are Sceloporus torquatus (3000 m.), Scleoporus aeneus (3100 m.), Sceloporus grammicus (3100 m.), Crotalus tri- seriatus (8270 m.), Thamnophis scalaris (3400 m.), and Gerrhonotus imbricatus (3500 m.). Factors Affecting Distribution Within the various vegetation types discussed previously and within the altitudinal limits set forth above, each species possesses certain characters that enable it to survive under a given set of environmental conditions and prob- ably preclude its occurrence in other habitats. Little is known of ecological tolerances of the species of amphibians and reptiles inhabiting Michoacan, so interpretations of ecological factors that seem to restrict the distributions of species must be made on the basis of observations in the field. Some species obviously are closely associated with certain ecological conditions that can be observed readily; for example, Cnemidophorus calidipes is always found in the Acacia-Cercidium-Prosopis association in the Tepalcatepec Valley. Although Acacia and Prosopis are widespread in the lowlands, Cnemidophorus calidipes occurs only in that part of the scrub forest where Cercidium grows. What the ecological relationships are remains to be discovered, but the distributional limits of Cnemidophorus calidipes are obvious. Analysis of distributional data and ecological data shows that several ecological factors seem to limit the distributions of some species either within a major habitat or within the entire region. The factors that I think important are discussed below. Microhabitats Subdivisions can be recognized within each of the five major habitats in Michoacan. For most of the species under consideration the recognition of five microhabitats is sufficient—arboreal, terrestrial, fossorial, riparian, and aquatic. Some of these could be divided further; for example, in the arboreal habitat it is possible to discern species that inhabit tree trunks and others that inhabit branches. However, such refinement is unnecessary to demonstrate the effects of the presence or absence of certain microhabitats in a given habitat on the distribution of amphibians and reptiles. The distribution of the numbers of species by microhabitats in the major habitats is shown in Table 3. Although the arboreal microhabitat is well developed in the tropical semi- deciduous forest, the pine-oak forest, and the fir forest, a much higher per- centage of species is arboreal in the tropical semi-deciduous forest. In that habitat Phyllomedusa dacnicolor, Sceloporus melanorhinus, Leptophis diplotro- pis, and Oxybelis aeneus are abundant arboreal species. Five of the ten arboreal species in the pine-oak forest and all three of the arboreal species in the fir forest are species of Sceloporus. Other arboreal species in the pine-oak forest are lowlanders that extend into the lower altitudinal range of the pine- oak forest; these include Anolis nebulosus, Leptophis diplotropis, and Oxybelis BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 663 TABLE 3.—COMPARISON OF ACTUAL AND RELATIVE NUMBERS OF SPECIES OF AMPHIBIANS AND REPTILES FOUND IN VARIOUS MICROHABITATS IN THE Major HaBiraTs IN MICHOACAN. Arid Semi- : Mircee tinct) peel | condi | esate neo aoa forest forest Arborealisc ... e54'% 7 (9.2%)|15 (24.6%)| 2 (5.1%)|10 (14.8%)}| 3 (13.0%) Rerrestrial $2.40 53 (69.7%)|382 (52.4%)|28 (71.8%)|35 (50.0%)| 9 (39.0%) Hassorialcy acl. css 7 (9.2%)| 6 (9.8%)} 1 (2.5%)|18 (25.7%)| 9 (39.0%) FaipAari ales ty. te. 6 (7.9%)| 7 (11.6%)| 6 (15.6%)| 5 (7.0%)| 2 (9.0%) AQUATIC oar alice. 3 (4.0%) 1 (1.6%)) 2 (5.0%)| 2 (3.0%)| 0 (0.0%) aeneus. The lack of cover due to deciduousness apparently precludes or greatly restricts the habitation of the scrubby trees in the arid tropical scrub forest by most of the species living in the tropical semi-deciduous forest. Although nine species are partly arboreal in the arid tropical scrub forest, only Urosaurus gadowi, which lives on the trunks of small trees, Enyaliosaurus clarki, which lives on the branches of trees and takes refuge in hollow branches, and Phyllodactylus duellmani, which lives in the columnar cacti and standing dead trees, are primarily arboreal. In the mesquite-grassland on the Mexican Plateau, the arboreal habitat is poorly represented. Only two species there are partly arboreal; Hyla smaragdina inhabits bromeliads on small trees, and Anolis nebulosus sleeps at night on low bushes. Terrestrial species, as classified here, include those that live on the ground but do not characteristically burrow or inhabit the mulch layer. Excluded from this category are species that normally inhabit the edges of streams, ponds, or other bodies of water thoughout the year. In the fir forest 39 per cent of the amphibians and reptiles are terrestrial, but in the other habitats at least 50 per cent of the species live on the ground. The highest percentage of terrestrial species is in the open habitats—arid tropical scrub forest and mesquite-grassland. The small amount of vegetational cover and near absence of mulch in these habitats is correlated with the presence of many fast and agile species there: Masticophis striolatus, M. taeniatus, Salvadora bairdi, S. mexicana, and several species of Cnemidophorus are characteristic. Many terrestrial species inhabit rocky areas, often natural outcrops or rock fences. In the arid tropical scrub forest Ctenosaura pectinata, Sceloporus gadowae, S. horridus, and S. pyrocephalus are common saxicolous species. In the mesquite- grassland S. dugesi and S. spinosus are among the conspicuous inhabitants of rock fences. Included as fossorial species are those that burrow or inhabit the mulch on the forest floor. Few fossorial species are known from the arid tropical scrub forest and mesquite-grassland, where a layer of mulch is almost non-existent. Although mulch is present in the tropical semi-deciduous forest, only a few species live in it, such as Scincella assata, Leptotyphlops phenops, Enulius uni- color, Geagras redimitus, and Rhadinaea hesperia. In the cooler pine-oak and 664 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. fir forests the mulch is well developed, and in these habitats many species live in the mulch. Characteristic inhabitants of the mulch are salamanders of the genera Ambystoma and Pseudoeurycea, lizards of the genus Eumeces, and snakes of the genera Conopsis, Diadophis, Geophis, Rhadinaea, Tantilla, and Toluca. Mulch apparently is a positive factor responsible for the distributions of many of these animals. Water, either in permanent ponds or flowing streams, is an important factor in the distributions of many species. Species characteristically found near water throughout the year are grouped into two categories—riparian and aquatic. The riparian inhabitants include various species of Rana and Thamnophis that live around the lakes and marshes on the Mexican Plateau. They make up 15 per cent of the herpetofauna of the mesquite-grassland. Also in this group are frogs, such as Eleutherodactylus rugulosus, Hyla areni- color, Hyla bistincta, and Rana pustulosa, which always are found along streams, and the lizards Basiliscus vittatus and Iguana iguana, which live at the edge of permanent bodies of water. Aquatic members of the herpetofauna include the salamanders of the genus Ambystoma. (Ambystoma dumerili is known only from neotenic individuals, and some populations of the other species of Ambystoma are neotenic.) Turtles and the crocodile make up the rest of the aquatic herpetofauna. The majority of the aquatic species live in lakes or lagoons, but Kinosternon integrum lives also in quiet streams, and neotenic individuals of Ambystoma ordinarium also inhabit streams. Streams having steep gradients are important microhabitats for Hyla arenicolor and Rana pustulosa, but these same streams are barriers to the distribution of in- habitants of quiet water, such as Kinosternon integrum. Reproductive Habits Diversity of reproductive habits among amphibians and reptiles indicates a variety of factors necessary for the species to maintain populations in a given environment. Although detailed information on the life histories is available for only a few species, data concerning sites of egg deposition and oviparity versus ovoviviparity are available. Analysis of these data provides evidence that the availability of suitable breeding sites and the methods of producing young importantly affects the ecological distributions of many species. Since the modes of reproduction in amphibians differ from those of reptiles, the two groups will be considered separately. Some amphibians utilize a variety of breeding sites; usually such species are widespread ecologically and geographically. For example, Rana pipiens, the most widespread amphibian in Michoacan and the only species that lives in all five major habitats, breeds and deposits its eggs in almost any aquatic situation except fast-moving streams. Other species are more restricted as to their breeding sites; Hyla bistincta deposits its eggs only in fast-moving mountain streams, and Phyl- lomedusa dacnicolor suspends its eggs from bushes and trees over shallow ponds. The eggs of two plethodontid salamanders of the genus Pseudoeurycea and of nine species of leptodactylid frogs of the genera Eleutherodactylus, Microbatrachylus, and Tomodactylus develop on land. Habitats in which the layer of mulch is well developed provide favorable sites for deposition of eggs. In the pine-oak forests, eight species (40 per cent of the amphibians living in that habitat) lay their eggs in the well-developed layer of mulch, whereas in the arid tropical scrub forest, where the mulch layer is poorly developed, only BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 665 two species (8.7%) lay terrestrial eggs. The abundance of streams varies in different habitats; in the mesquite-grassland on the Mexican Plateau, only one species of amphibian breeds in the streams, most of which are intermittent, but in the pine-oak forest and tropical semi-deciduous forest, where small streams are numerous, six species of amphibians deposit their eggs in streams. The correlation between egg deposition sites and major habitats is shown in Table 4. TABLE 4.—COMPARISON OF ACTUAL AND RELATIVE NUMBERS OF SPECIES OF AMPHIBIANS THAT Use Various Ecc Deposition SITES IN THE Major HaBITATs IN MICHOACAN. Arid Semi- DEPOSITION tropical | deciduous | Mesquite- | Pine-oak Fir SITE serub tropical | grassland forest forest forest forest Temporary Pond.. .|/14 (60.8%)| 8 (36.4%)| 7 (50.0%)| 4 (20.0%)| 2 (28.5% Permanent Pond...| 2 (8.7%)| 2 (9.1%)| 4 (28.6%) 2 (10.0%)| 1 (14.38% Stream aero eke 4 (17.4%)} 6 (27.2%)| 1 (7.1%) 6 (80.0%)| 3 (42.9%) Merrestrialese ce. - 2 (8.7%)| 5 (22.7%)| 2 (14.8%) 8 (40.0%)) 1 (14.8%) Arboreal/Pond..... 1 (44%)| 1 (4.6%)| 0 (0.0%)} 0 (0.0%) 0 (0.0%) Most reptiles lay eggs, but some lizards and snakes give birth to young. Although information concerning the reproductive habits is lacking for some species, data are available for enough species to draw some conclusions about the reproductive habits as they relate to distribution. Most species of snakes and lizards in the lowlands below an elevation of 1000 meters are oviparous and deposit their eggs in rotting logs and stumps, in mulch, in loose soil, or beneath stones. Solar radiation and the continuously warm substrate provide sufficient heat for incubation. At successively higher elevations the percentage of oviparous species decreases. Six species of reptiles (Sceloporus grammicus, S. aeneus, S. torquatus, Gerrhonotus imbricatus, Thamnophis scalaris, and Crotalis triseriatus) occur at elevations in excess of 3000 meters; all are ovoviviparous. The activities of these animals are restricted to the warm parts of days in a relatively short summer. Apparently the habit of basking in the sunshine provides sufficient warming of the body to allow development of the eggs in the females, whereas eggs laid in the continuously cool mulch or soil would not receive sufficient heat to develop. The correlation between ovoviviparity and altitude parallels that between ovoviviparity and northward extent of range of some species, such as Gerrhonotus multicarinatus and Thamnophis sirtalis in North America and Lacerta vivipera and Vipera berus in Eurasia. Diel Activity Conditions in the various major habitats and included microhabitats control the diel activities of the amphibians and reptiles living there. Throughout Michoacan most amphibians are nocturnal. The low intensity of light on the 666 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. floor of the tropical semi-deciduous forest, combined with the relatively high humidity and moist layer of mulch, provides conditions favorable for the diurnal activities of some small amphibans, such as Eleutherodactylus occi- dentalis and Microbatrachylus pygmaeus. Other amphibians, such as Eleuthero- dactylus rugulosus, Hyla arenicolor, and Rana pustulosa, inhabit stream beds in ravines, where the amount of light is reduced in the daytime; consequently, these species often are diurnal. The crocodile and the turtles of the genus Kinosternon are sometimes active by day but are chiefly nocturnal. Lizards and snakes are characterized by a wide range of diel activity. Most lizards are diurnal; in Michoacan geckos of the genus Phyllodactylus are an exception. In the arid tropical scrub forest few reptiles are active during the hottest part of the day when temperatures at mid-day sometimes exceed 43° C. Cnemidophorus calidipes is a notable-exception. Most active terrestrial species, such as Drymobius margaritiferus, Masticophis striolatus, Saluadora mexicana, Sceloporus pyrocephalus, S. siniferus, and the various species of Cnemidophorus, have two daily peaks of activity, the early daylight hours until about 10 AM and the late afternoon from about 5 PM until dusk. In the mesquite-grassland on the Mexican Plateau, where temperatures at mid-day are moderate, dis- tinct morning and late afternoon periods of activity are lacking; instead, most diurnal species are active throughout the day. In the tropical semi- deciduous forest, where dense foliage provides shade, agile species, such as Ameiva undulata, Cnemidophorus lineatissimus, Dryadophis melanolomus, Leptophis diplotropis, and Oxybelis aeneus, are active throughout the day. Increase in altitude is correlated with decrease in duration of diurnal activity, at least in the lizards of the genus Sceloporus. At Uruapan at an elevation of 1630 meters, Sceloporus grammicus and S. torquatus often were active before 8 AM, but at Mil Cumbres at an elevation of 2800 meters, these lizards were not seen before 10 AM, and then only on clear days. In the arid tropical scrub forest, where nocturnal temperatures are moderate relative to the diurnal temperatures, more than 50 per cent of the lizards and snakes are nocturnal (Table 5). In the hot Balsas-Tepalcatepec Basin some species, such as Drymarchon corais, are nocturnal, whereas in other parts of their ranges they normally are diurnal. In the mesquite-grassland and pine- oak forest Thamnophis dorsalis is diurnal, but in the Tepalcatepec Valley is nocturnal. Many of the small mulch-inhabiting snakes, such as Conopsis and Geophis, in the pine-oak forest apparently are nocturnal. Lower nocturnal temperatures on the Mexican Plateau apparently restrict nocturnal activity there; only Elaphe triaspis, Thamnophis melanogaster, and Trimorphodon tau commonly are found at night. Possibly the low nocturnal temperature limits the altitudinal distribution of such strictly nocturnal animals as lizards of the genus Phyllodactylus and snakes of the genus Leptodeira. Likewise, the excessive heat and light in the arid tropical scrub forest possibly precludes the occupation of that habitat by lizards such as Ameiva undulata and Cnemidophorus lineatissimus, both of which live in the adjacent tropical semi-deciduous forest. An adequate under- standing of the significance of diel activity as related to distribution must await data from detailed studies of temperature responses of the species con- cerned. BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 667 TABLE 5.—COMPARISON OF ACTUAL AND RELATIVE NUMBERS OF SPECIES OF Lrzarps AND SNAKES Havinc NocTruRNAL AND DiuRNAL HABITS IN THE Various Major HABITATS IN MICHOACAN. Arid Semi- PERIOD OF tropical | deciduous | Mesquite- | Pine-oak Fir ACTIVITY scrub tropical | grassland forest forest forest forest Diumnal ee eyedacke 25 (47.2%)|29 (65.9%)|17 (70.8%)|31 (67.4%)|10 (71.4%) Nocturnali’ =<: 2. 28 (52.8%)|15 (84.1%)| 7 (29.2%)|15 (32.6%)| 4 (28.6%) Ecological Segregation Similar adaptations and requirements in closely related species affect the distributions of some species within and between communities. Many ob- servations in nature, as well as experimental evidence, have demonstrated that two or more species having closely similar niche-requirements cannot exist as stabilized populations in the same area; this concept, which has been known as Gause’s Rule, was thoroughly discussed and defined as the “Competitive Exclusion Principle” by Hardin (1960). Observations in the field indicate that in Michoacan several species-pairs seem to have closely similar niche- requirements. Members of each of these pairs are segregated geographically. For example, Sceloporus bulleri and S. torquatus closely resemble each other morphologically, and each inhabits pine and fir forests, where both live on the ground and on the trunks of the large trees, but the former species occurs in the Sierra de Coalcoman and the latter in the Cordillera Volcanica. Although these two species are segregated geographically, each occurs sympatrically with other species of Sceloporus, which, because of differences in morphology or habits, have different niche-requirements. Data to support concepts of ecological segregation of animals in nature are difficult to obtain, especially on amphibians and reptiles, many of which are secretive in their habits. The abundant and conspicuous iguanids, Scelop- orus and Urosaurus, are notable exceptions. In a transect from the Mexican Plateau, over the Cordillera Volcdnica, down to the Tepalcatepec Valley, 12 species of Sceloporus and one of Urosaurus can be found. Three or more species occur in each of the five major habitats included in the transect. Sym- patric species differ in size or in their microhabitats. On the Mexican Pla- teau two species are broadly sympatric, and each is terrestrial and saxicolous; of these, S. spinosus is large, and S. dugesi is small. Another small species, S. scalaris, is restricted to bunch-grass associations on the plateau and in the Cordillera Volcénica. In the coniferous forests on the Cordillera Volcanica there are two small species—the terrestrial S. aeneus and the arboreal S. grammicus. Also living there is the large S. torquatus, which lives on the ground and on trees. At an elevation of about 1000 meters on the slopes of the Cordillera Volcdnica the highland species of Sceloporus are replaced in the broad-leafed oak or tropical semi-deciduous forest by an assemblage of lowland species, of which S. asper is small and arboreal, S. melanorhinus is large and arboreal, and S. utiformis is terrestrial. At lower elevations in the 668 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist, arid tropical scrub forest S. melanorhinus is still the large arboreal species, but Urosaurus gadowi has replaced S. asper as the small arboreal species. The terrestrial species include the large S. horridus and the small S. pyroceph- alus. On conglomerate cliffs the latter species is replaced by S. gadowae. In another transect, one from the high ridges of the Sierra de Coalcoman to the Pacific coast, many of the species found in the lowlands are the same as those listed in the previously described transect, but some additional species are present on the Pacific coastal lowlands. Entirely different species of lizards occur in the two highland areas. In the coniferous forest in the Sierra de Coalcoman the small arboreal Sceloporus heterolepis is the ecolog- ical equivalent of S. grammicus in the Cordillera Volcanica. Sceloporus bul- leri, which lives on the ground and on the trees in the Sierra de Coalcoman, is the ecological equivalent of S$. torquatus in the Cordillera Volcanica. The niche filled by S. aeneus in the Cordillera Volcdnica apparently is vacant in the Sierra de Coalcoman. In the tropical semi-deciduous forest on the sea- ward slopes of the Sierra de Coalcoman only two species of Sceloporus occur —the large arboreal S. melanorhinus and the terrestrial S. utiformis. In the arid tropical scrub forest on the Pacific lowlands $. melanorhinus is the large arboreal species, Urosaurus bicarinatus is the small arboreal species, and S. horridus is the large terrestrial species. In rocky areas in the Pacific low- lands S. pyrocephalus is the small terrestrial species, but in open areas there it is replaced by S. siniferus. Similar selection of microhabitats that apparently allow several related species to live in a single community has been demonstrated in other groups of animals, for example, lizards of the genus Anolis (Duellman, 1963). Direct competition between arboreal and terrestrial species obviously is slight, but competition may exist between two arboreal, or between two terrestrial, species in a given community. The distinct difference in size between species such as Sceloporus grammicus and S. torquatus, which is about twice the length and about five times the bulk of the former, suggests that, at least for adults, food items probably differ. However, juveniles of S. torquatus may consume the same kinds of food as does S. grammicus, at which time the two species may be in direct competition. Studies on the autecology of these lizards are needed to clarify their ecological interrelationships. Differences in size in sympatric pairs of species have been noted in Chiropterotriton by Rabb (1958) and in Cnemidophorus by Zweifel (1960) and Duellman (1960a). Additional evidence on sizes of ecological pairs of teiid lizards of the genera Ameiva and Cnemidophorus supports observations on ecological segregation in these lizards. In certain areas in the Tepalcatepec Valley five species of Cnemidophorus and one species of Ameiva are sympatric. Ameiva undulata and the smaller Cnemidophorus lineatissimus are widespread in the tropical semi-deciduous forest on the seaward slopes of the Sierra de Coalcoman, but in the Tepcalcatepec Valley these species occur only on the shady floor of the gallery forest along major streams. In the ecotone between the gallery forest and the arid tropical scrub forest A. undulata and C. lineatissimus are sometimes found with other species of Cnemidophorus—two large species (C. communis and C. costatus) and two small species (C. cali- dipes and C. deppei). Although both of the small species live in open areas in the arid tropical scrub forest in the Tepalcatepec Valley, each has notice- able ecological preferences. Cnemidophorus deppei is most abundant in open grassy areas, especially in arroyos where the vegetation is more lush than BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 669 on the adjacent flats, where C. calidipes lives on nearly barren. ground support- ing only sparse stands of grass below the scattered scrubby trees. The ecological segregation between C. communis and C. costatus is not so well marked. Cnemidophorus costatus is abundant throughout the Tepalcatepec Valley and is closely associated with both C. calidipes and C. deppei, but does not occur on the coastal lowlands. Cnemidophorus communis is uncommon and only locally present in the Tepalcatepec Valley, is more abundant on the coastal lowlands, and is the only species of Cnemidophorus in some of the valleys in the Sierra de Coalcoman. Probably C. communis is a recent in- vader in the Tepalcatepec Valley, where it may be in direct competition with C. costatus. Other ecological species pairs or equivalents are present in Michoacan. In the Tepalcatepec Valley Bufo perplexus replaces Bufo marmoreus, which is restricted to the coastal lowlands. In the Sierra de Coalcoman Geophis nigrocinctus and G. incomptus replace G. tarascae and G. maculiferus of the Cordillera Volednica. On the Mexican Plateau Hypopachus oxyrrhinus, Masti- cophis taeniatus, and Salvadora bairdi are the respective ecological equivalents of Hypopachus caprimimus, Masticophis striolatus, and Salvadora mexicana in the Tepalcatepec Valley. The meager data on ecological segregation and replacement indicate that in well-known groups the species are segregated ecologically within major habitats, and that in ecotones between major habitats the species comprising a community in one habitat are replaced by another group of species in the second habitat. Interspecific competition may be significant in maintaining distributional limits in a climax environment and in effecting dispersal in areas of ecological succession. Influence of Man Much of Michoacan has been inhabited by man since pre-historic times. Ancient Aztecs and Tarascans cultivated extensive areas on the Mexican Plateau. Pine forests in the Cordillera Volcdnica were cut to provide fuel for the perpetual fires that were a part of Tarascan religion. Probably the amount of cultivation and cutting by means of crude indian implements was slight in comparison with that accomplished by use of the iron plows and axes intro- duced by the Spaniards in the Sixteenth Century. Now, these tools seem crude as compared with modern mechanized equipment. Obviously man has altered habitats in many areas in Michoacan. Marshes on the Mexican Plateau have been partly drained; lakes and irrigation ditches have been created in the lowlands. Many forested areas have been cleared, and over-grazing probably has altered the mesquite-grassland on the Mexican Plateau. Agricultural malpractices have resulted in widespread erosion, and irrigation probably has modified the level of the water table in some areas. There is no evidence that any major habitat has been eliminated by man in any physiographic area, but some habitats have been partly altered or reduced in area. Constriction of one habitat results in the expansion of an adjacent habitat or the creation of a new habitat, and the distributions of some species have been altered accordingly. In other instances the habitats of some species probably have been modified, but with minor adjustments the species are able to survive in them. Among the probably more important results of 670 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist, human alteration of natural habitats in Michoacan are the following: change in vegetation on the Mexican Plateau from grasses and mesquite to a legume- cacti-agave association due to over-grazing, formation of pine savannas having little or no understory due to heavy grazing by goats, reduction of forested areas because of lumbering, reduction of marshes by draining, creation of aquatic and aquatic margin microhabitats in arid areas through irrigation, and creation of an extensive network of microhabitat for saxicolous species by the construction of rock fences. The patterns of ecological and geographical dis- tribution probably have not been altered significantly by man. Analysis of Ecological Distribution The ecological distribution of amphibians and reptiles in Michoacan is closely correlated with the limits of the major types of vegetation or animal habitats as defined previously on page 641. Of the 183 species and subspecies analyzed, 115 (62.8%) are restricted to one habitat; 55 (30%) occur in only two habitats, whereas only seven (3.8%) occur in three habitats, five (2.8%) in four habitats, and one species (0.6%) in all five habitats. The number of species that occupy more than one habitat decreases as the habitats become more diverse. For example, of the 77 species inhabiting the arid tropical scrub forest, 26 also live in the tropical semi-deciduous forest, ten in the mesquite grassland, nine in the pine-oak forest, and only one in the fir forest. Of the TABLE 6.—ANALYSIS OF DISTRIBUTIONS OF AMPHIBIANS AND REPTILES BY MAJOR TAXONOMIC GROUPS IN THE Major HABITATS IN MICHOACAN. (Percentages are the proportion of the taxonomic group in a given habitat. Numbers in parentheses after the name of the group indicate the number of species and subspecies belonging to that group in Michoacan. ) Arid Tropical G tropical semi- Mesquite- | Pine-oak Fir eres scrub deciduous | grassland forest forest forest forest Salamanders (6).... 0 ) 0 5 4 0.0% 0.0% 0.0% 83.3% 66.8% Frogs and toads (42) 18 17 13 18 5 42.9% 40.5% 30.9% 42.9% 11.9% Murtles'(S) 2as-28 2: 4 1 2 1 0 80.0% 20.0% 40.0% 20.0% 0.0% Crocodilians (1).... 1 0 0 0 0 100.0% 0.0% 0.0% 0.0% 0.0% Lizards,(52)) <5. as << 22 21 7 17 7 42.3% 40.4% 13.5% 32.6% 13.5% Snakesi(77) tee. 32 23 17 29 7 41.6% 29.9% 22.1% 37.7% 9.1% Total species per habitatesse. . 2o8 77 62 39 70 23 42.1% 39.4% 21.3% 38.3% 13.7% BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 671 62 species inhabiting the tropical semi-deciduous forest, 17 also live in the pine-oak forest, eight in the mesquite-grassland, and only one in the fir forest. Of the 89 species inhabiting the mesquite-grassland, 19 also live in the pine- oak forest, and three in the fir forest. The largest amount of ecological inter- change is between the fir forest and the pine-oak forest; of the 23 inhabitants of the fir forest, 22 are among the 70 species that occur in the pine-oak forest. TABLE 7.—ACTUAL AND RELATIVE NUMBERS OF SPECIES AND SUBSPECIES OF Mayor Groups OF AMPHIBIANS AND REPTILES LIvING IN VARIOUS HABITATS IN MicuoacANn. (Percentages are the proportion of the total herpetofauna in a habitat comprised by the taxonomic group. Numbers in parentheses after the name indicate the number of species and subspecies belonging to that group in Michoacan. ) Arid Tropical | tropica semi- Mesquite- | Pine-oak Fir Group scrub deciduous | grassland forest forest forest forest Salamanders (6).... 0 0 0 5 4 0.0% 0.0% 0.0% Cols 17.4% Frogs and toads (42) 18 17 13 18 23.4% 27.4% 33.3% 257 e 21.8% Purtles(S ie. sete «2s 4 1 2 1 0 IIE, 1.6% 5.1% 1.4% 0.0% Crocodilians (1).... 1 0 0 0 0 1.3% 0.0% 0.0% 0.0% 0.0% hizardsi(52). 22. 22 21 7 17 7 28.6% 33.9% 18.0% 24.4% 380.4% Snaikesi(77) esc 32 23 17 29 a 41.5% BYE IG 43.6% 41.4% 30.4% MotaliGlss) nas ae. Cah 62 39 70 23 100.0% 100.0% 100.0% 100.0% 100.0% As expected, the ecological distributions of the major taxonomic groups of amphibians and reptiles differ. In Table 6 the ecological distributions are analyzed on the basis of the groups. For example, in Michoacan only one crocodilian occurs. It lives in the lagoons in the arid tropical scrub forest; thus, one hundred per cent of the crocodilian fauna occurs in that habitat. In Table 7 the distributions are analyzed on the basis of habitats. In this analysis the one crocodilian inhabiting the arid tropical scrub forest accounts for only 1.3 per cent of the species of amphibians and reptiles in that habitat. Salamanders are absent in the tropical habitats, but frogs are about equally distributed in the arid tropical scrub forest, tropical semi-deciduous forest, and pine-oak forest. Lizards are about equally diverse in the two tropical habitats, but there are fewer species in the pine-oak forest and far fewer species in the mesquite-grassland and fir forest. The species of snakes are nearly as numerous in the pine-oak forest as in the arid tropical scrub forest; fewer species occur UNIvERSITY OF Kansas Pusxs., Mus. Nat. Hist. 672 *SIND90 UOTZLYBZAA JO add} JSOIOF PNOTD B YOIYAA Ul SvOIV O}BOTpU sjoquds pros ayL ‘syeyqey Jofeur jo uOYNqIAsIp SurMmoys UvBO Igoe oY} OF OAVJOURT, OL) WOIF UPOVOTY UJOYINOS JO IYOIg °“G “OY S19J9WO|\ GZ = XOE = UO!jDsaHHoxy [Dd 10/A TID UDIIO. DiflI04~». HOD, rapes 00S So) =) fais (oyele) ; } oY ( 4 oos| 0002 t 00Sz 0) Or “Th ise104 4 | | JOOOE ispi03 1p0=eUla d oos¢ jsa10.4 SNONpo@p-Iwas |DI!dOd, ik p 000b SIOJO//\ 4 I8dH{DI/DdA Oley "ANY a aye DSO/ODG 04422 OIDJIIUDL 0149. Sas04 qnuos |podod, Puy -*&é BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 673 in the tropical semi-deciduous forest and in the mesquite-grassland, and far fewer live in the fir forest. The good correlation between distributions of the components of the herpetofauna and the habitats suggests that the definition of habitats proposed here is real in that it rather accurately reflects the ecological factors that are significant in limiting the distributions of most of the species. The analysis of altitudinal distribution shows that most species reach the upper or lower limits of their distributions at elevations approximating the altitudinal limits of the major habitats. It should be noted that the altitudinal limits of the habitats vary from windward to leeward slopes and differ in the Cordillera Volcanica from those in the Sierra de Coalcoman (Fig. 5). These variations account, in part, for some of the discrepancies in altitudinal distributions of the amphibians and reptiles. In some aquatic species and others inhabiting microhabitats that are not directly dependent on a given kind of vegetation, the overall ecological dis- tributions are not well correlated with the limits of the major habitats. Never- theless, the physiognomy of the vegetation and resulting microhabitats seem to be the most important factors in determining the ecological distributions of amphibians and reptiles in Michoacan. GEOGRAPHY OF THE HERPETOFAUNA The geographical analysis presented here is centered on the herpetofauna of Michoacan but applies to the herpetofauna of a large part of southwestern México. An attempt is made to answer four questions—1.) How can the geographical relationships of the various faunal assemblages in southwestern México best be described and analyzed? 2.) What is the distribution of each of the species of amphibians and reptiles in Michoacan and adjacent areas? 3.) What are the patterns of distribution of amphibians and reptiles in south- western México? 4.) How are these patterns related to one another and to the patterns in outlying regions? Methods of Zoogeographic Analysis Zoogeographers work with facts and inferences drawn from a variety of sources; in most cases no one set of data is complete. Consequently, many generalizations are arrived at by deductive extrapolation. Zoologists since Humboldt in the early Nineteenth Century have tried with various degrees of success to analyze regional (as opposed to world-wide) faunas. Three methods of regional analysis have been used, and Kendeigh (1954) has reviewed their history and evaluated each. The oldest and first method utilizes the Life-zone Concept of Merriam (1890, 1894); although tempera- tures (isotherms) are the theoretical basis for this classification, in practice floristic indices are utilized. Life-zones are discontinuous areas. They have been used more satisfactorily in the mountainous regions of western North America, where floristic and temperature breaks are sharp, than in the eastern part of North American where gradual changes in elevation result in less sharp floristic, faunal, or climatic boundaries. Goldman (1951) applied the Life- zone Concept in his discussion of the distribution of vegetation, birds, and mammals in México, and he recognized six zones in Michoacan. According 674 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hist. to him, the Lower Tropical Life-zone is divided into the Humid Lower Tropical Subzone and the Arid Lower Tropical Subzone; the latter occurs on the Pacific Coastal Plain and slopes of adjacent mountains to elevations of about 2,500 feet and extends from Nayarit to Chiapas. The Upper Tropical Life-zone also is divided into two subzones, of which the Humid Upper Tropical Subzone corresponds to what I have defined as cloud forest in the earlier section on vegetation. This kind of forest does not occur in Michoacan, but cloud forest does occur in the Sierra Madre del Sur and possibly in the Sierra de Autlan in Jalisco. Goldman’s Lower Austral Life-zone includes the slopes of the western Mexican highlands, whereas the Transition Life-zone in- cludes most of the mountainous regions above 7,000 feet, and the Canadian Life-zone includes only the highest peaks, where fir forest occurs. Essentially this classification of Life-zones is nothing more than the application of names to habitats or vegetational formations. This aspect of the distribution of the herpetofauna in Michoacan already has been presented in the section on ecology of the herpetofauna. It is significant to note that the ecological dis- tribution of a species does not necessarily reflect the geographical distribution of either the species or its Life-zone. The plotting of Life-zones and associated species of animals on the Pacific slopes of the Sierra de Coalcoman reveals the altitudinal distribution of environments and associated animals, but a similar transect on the slopes of the Sierra Madre del Sur in Guerrero or on the Sierra Madre Oriental in Veracruz shows the same Life-zones, but most species of animals inhabiting these Life-zones are different from those in Michoacan. Therefore, for understanding patterns of animal distribution on a spatial (horizontal) scale the Life-zone Concept is unsatisfactory. The second method of faunal analysis is the Biotic Province Concept, which was proposed by Vestal (1914) and refined for North America by Dice (1943), who defined a biotic province as a considerable and continuous geographic area that is characterized by the occurrence of one or more important ecological associations that differ, at least in proportional area covered, from the associa- tions of adjacent provinces. Theoretically, this concept has much in its favor, since it is based on principles of community ecology and finds its application in the common limits of distribution of biological taxa. Objectons to the concept of biotic provinces have been many. Johnson, Bryant, and Miller (1948), Miller (1951), and Kendeigh (1954) have pointed out difficulties in applying nebulous principles of community ecology to patterns of distribution and have emphasized the importance given to geographical boundaries in- stead of actual animal distributions. Smith (1939, 1940, and 1949) mapped the Biotic Provinces of México; his earlier works were based solely on the distribution of the species of Sceloporus, whereas the last paper was based on the distributions of all amphibians and reptiles. Moore (1945) similarly mapped the Biotic Provinces of México on the basis of avian distribution, and Goldman and Moore (1956) repeated the process for mammalian distribution. Although there is considerable agreement in the various maps, there are im- portant differences. The availability of neatly delimited regions drawn on maps seems to have contributed to the popularity of the Biotic Province Concept. This popularity has resulted in widespread use and misuse of the concept, espe- cially by workers who were not familiar with the premises on which the con- cept is based. The establishment of a biotic province on distributions of a BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 675 few species (Firschein and Smith, 1956) and the establishment of a system of biotic provinces on the distribution of a limited taxonomic group of animals (Smith, 1939, 1940, and Stuart, 1943) are unwarranted. Equally misleading are descriptions of geographic distributions of species on the basis of biotic provinces, when the actual known range of the species in- cludes only a small part of the biotic province. Peters (1955) discussed various uses and misuses of the Biotic Province Concept, especially as it has been applied to southwestern México. Biotic provinces are best applied on a horizontal scale; in mountainous regions the biotic provinces have little use. Smith, an early proponent of the Biotic Province Concept, recently (1960) registered his disillusionment and suggested disregarding the concept. Axtell (1962) argued for the use of the concept and suggested the inclusion of an “endemism clause,” which would seem further to complicate the use- fulness of the concept. According to Goldman and Moore (1946), the Sierra de Coalcoman and all of the lowlands in Michoacan are part of the Nayarit- Guerrero Province extending along the Pacific coast from Nayarit to Guerrero, and all of the rest of the state is contained in the Transverse Volcanic Prov- ince, which extends from Nayarit and Jalisco to Veracruz. According to Smith (1949), the coastal lowlands and the Sierra de Coalcoman are part of the Acapulcan Province, which consists of the southern coastal part of Goldman and Moore’s Nayarit-Guerrero Province. Smith placed the Balsas Basin in a separate Balsan Province, the Cordillera Volcdnica as part of the Austro- Occidental Province, and the Mexican Plateau in the Austro-Central Province. The lack of agreement on boundaries of provinces can be attributed in part to the fact that these authors were treating different groups of animals, but more important, the disagreement of boundaries reflects differences in the interpretation and application of the concept of biotic provinces. The third method of faunal analysis is the ecological concept of the biome (Clements and Shelford, 1939). This concept has been thoroughly reviewed by Kendeigh (1961), according to whom a biome is “a biotic com- munity characterized by distinctiveness in life-forms of the important climax species.” Terrestrial biomes include such categories as coniferous forest, grassland, desert, and tropical forest, among others. These are, of course, discontinuous in their distribution on the surface of the earth. The “key” to the definition of the biome is the “life-form” of the vegetation, Thus, the biome is an ecological classification of major vegetational formations. Ken- deigh (1961) refined the concept of biomes and distinguished various sub- divisions, one of which, the biociation is defined as “a subdivision of a biome distinguished by uniformity and distinctiveness in the species composition of the climax community, particularly the animal predominants.” This defini- tion resembles that of the biotic province and is equally difficult to apply. In the over-all scheme of geographic distribution, major assemblages of animals representing large faunal units show some agreement with the distribution of biociations; for example, the distributional limits of North American Herpetofaunas (Savage 1960) coincide at least in part, with the distribution of North American biociations as mapped by Kendeigh (1961:278). When describing the distribution of a fauna in a given region, three salient features need to be taken into consideration—1.) The distribution of the animal. 2.) The distribution of the animal’s habitat. 3.) The distribution of the animal within its habitat. If the distributions of all animals in a given 5—8425 676 UNIVERSITY OF Kansas Pusts., Mus. Nat. Hist. habitat were the same as the limits of the habitat, then the concept of biotic provinces (biociations) would be perfectly applicable. But, such instances are rare. The distribution of a species of animal often is determined by factors other than the distribution of the animal’s habitat. Means and rates of dispersal, presence of allopatric competitors, and presence of physical barriers, as well as various historical events may account for discrepancies in distributions. Objections to the use of each of the major methods of faunal analysis, each of which is based primarily on ecological premises, have been noted above. Actually, geographical analysis of faunas should be based primarily on physiographic data and secondarily on ecological premises. Zoogeographers have divided the earth into faunal realms, each of which is divided into faunal regions, and each of which, in turn, is divided into faunal provinces. The faunal provinces are circumscribed by the limits of distributions of most of the members of a fauna that inhabits a certain usually continuous physiographic region. Thus, in western México there are three major physiographic provinces—the coastal plain, the highlands of the western sierras, and the tableland of the Mexican Plateau. Each of these three physiographic provinces has a distinctive herpetofauna, respectively, the Neotropical, the Sierra Madrean, and the Desert and Plains herpetofauna (Savage, 1960). In analyzing the patterns of distribution on a local basis, such as in Michoacaén and adjacent areas of southwestern México, a more refined division is desirable. Faunal area has been used as a subdivision of a faunal province. For the purposes of the following analysis a faunal area is defined as a part of a faunal province characterized by an assemblage of species that differs from assemblages found elsewhere in the same faunal province. The faunal areas in Michoacan approximate the physiographic areas outlined in the preceding description of the geography (see page 632). Each faunal area has characteristic species and/or assemblages of species different from species and/or assemblages in the other faunal areas. These faunal areas are discussed in detail below, where the fauna of each is described and analyzed. Each faunal area can be described, and the species known to occur in the area can be listed, but such descriptions and lists do not afford an analysis of the fauna. In analyzing the faunas of adjacent or separated areas zoogeog- raphers are interested to know the degree of faunal resemblance between the areas being considered. Several methods of determining the degree of faunal or taxonomic relationship have been proposed. Burt (1958) and Simpson (1960) have evaluated the various methods; Smith (1960) and Long (1963) have made further suggestions. Formulas that take into consideration both geographic and taxonomic levels are useful for measuring faunal resem- blances. A comparison of the faunas of Eurasia and North America at the species level would indicate only a negligible amount of resemblance, and a comparison of the faunas of two mountain ranges in southern México at the family level would show only negligible differences, if any. In choosing a formula for determining faunal resemblances it is desirable to take into consideration the size of the faunas of each area, as well as the number of taxa common to both. Simpson (1960) evaluated various formulas and presented some new ideas for showing faunal similarities between regions. His methods of analysis based both on the presence and absence of taxa and their ecological distribution and abundance will be of use in areas where such detailed information is available. Sufficient data on local abundance of taxa BrioGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 677 in Michoacan are lacking for that kind of analysis. The ecological distribu- tion of a species is important for understanding its local distribution; paleo- ecological data are useful to draw inferences as to how the species could have established its present pattern of distribution. Neither this information, nor that concerning the relative abundance of taxa are of primary importance in determining the present faunal resemblance of one area to another. Only the presence or absence of taxa are of great significance. For this purpose the rather simple formula of Pirlot (1956) seems perfectly applicable, for it takes into account the number of taxa in each fauna and the number of taxa common to both faunas. The results of a formula used to show faunal resemblances have been termed amount of similarity, degree of relationship, and faunistic relation factor, among others. Smith (1960) proposed the last term, which, although it has merit, includes the word relation that can be confused with relationships in a strict phylogenetic sense. I suggest the modification of Smith’s term to Faunal Resemblance Factor, which still retains his abbreviation FRF. By slightly modifying Pirlot’s formula to reduce the factors to per- centages, one step in the calculation can be omitted. The formula then reads: 2G where C = the number of taxa in common to the two areas, Ni = the num- ber of taxa present in one area, Ne = the number of taxa present in the second area, and FRF = the Faunal Resemblance Factor. In this formula an FRF of 0 means that no taxa are common to the two areas, and an FRF of 1 means that all taxa are common to the two areas. In an effort to present analyses of faunal resemblance in a graphic or tabular manner readily comprehensible to the reader, I have presented information on which the FRF is based, as well as the FRF itself (Tables 9-11). In these tables the geographic areas that are being analyzed are listed vertically on the left and horizontally in the same order across the top. In their common squares on the resulting grids the total number of species known from that area is set in bold-faced type. To the right and above the totals the numbers of species in common to each combination of faunal areas are set in roman type. To the left and below the totals the Faunal Resemblance Factors for each of the combinations are set in italic type. Throughout the following analysis of geographic distribution I have dealt only with species. Use of this method reveals degrees of resemblance or dissimilarity be- tween faunal areas, faunal provinces, or more diverse categories, and also shows the relative amount of resemblance within categories, as compared to that occurring between them. For example, examination of Table 9 shows that the FRF for the species of amphibians and reptiles in the three faunal provinces in Michoacan is low, ranging from .17 to .32, whereas the FRF for faunal areas within the Neotropical Province (Table 10) is much higher, ranging from .55 to .86. = FRE, Distribution of the Herpetofauna In the following account the distribution of marine turtles and the sea snake in Michoacan is not included, but species that have not been found in Michoacan, but whose occurrence there is probable have been included. This review of distributions is on a geographic, not a systematic, basis. Many 678 UNIVERSITY OF KANsAs PuBis., Mus. Nat. Hist. species of amphibians and reptiles have similar distributions in Michoacan and adjacent areas; consequently, species with like distributions are grouped. Distributions are grouped into three basic patterns that roughly correspond to parts of three of the eight North American herpetofaunas recognized by Savage (1960); these are the Neotropical Herpetofauna in the lowlands, the Sierra Madrean Herpetofauna in the mountains, and the Desert and Plains Herpeto- fauna on the Mexican Pleateau. Two of these herpetofaunas contain assem- blages of species having different patterns of distribution. For this reason the various assemblages are discussed separately, as outlined below: NEOTROPICAL HERPETOFAUNA Widespread Pacific Lowland Assemblage Pacific Coast Assemblage Balsas-Tepalcatepec Basin Assemblage SreERRA MADREAN HERPETOFAUNA Widespread Montane Assemblage Sierra Madre Occidental Assemblage Cordillera Volcanica Assemblage Sierra de Coalcoman Assemblage Sierra Madre del Sur Assemblage DESERT AND PLAINS HERPETOFAUNA The distributions of many species fit well into the general patterns outlined above; others do not. Our knowledge of several of the latter is incomplete, or the species seemingly range across faunal boundaries is an incomprehensible pattern. In southwestern México there is faunal interchange between the Mexican Plateau and the Pacific lowlands, including the Balsas-Tepalcatepec Basin. On the basis of their over-all distributions or interspecific relationships, most of the species the distributions of which include the Mexican Plateau and the Pacific lowlands in southwestern México can be assigned to either the Desert and Plains Herpetofauna or to the Neotropical Herpetofauna. This aspect of distribution is discussed in detail in the section dealing with the Desert and Plains Herpetofauna. Three species (Rana pipiens, Kinosternon integrum, and Lampropeltis doliata) are wide-ranging and occur in all of the physiographic areas in Michoacan. In the lists of species given below they are omitted, but they are included in the succeeding analysis of the fauna. Typhlops braminus apparently is an introduced species and has not been in- cluded in the following discussion. The bases for accounts of distribution are preserved specimens listed in the systematic study of the amphibians and reptiles of the state (Duellman, 1961). Distributional data for species in other parts of southwestern México were taken from unreported specimens and from a variety of published works, notably Duellman (1958) for Colima, Davis and Dixon (1959 and 1961) for Guerrero, and Davis and Smith (1953 a-c) for Morelos, as well as Smith and Taylor (1945, 1948, and 1950). Neotropical Herpetofauna In México the Neotropical Herpetofauna is composed of those species liv- ing in the tropical lowlands and the distributions of which, or the distributions of their relatives, extend southward into Central and South America. In Michoacan and adjacent parts of southwestern México species comprising this herpetofauna are found principally at elevations below 1000 meters. Three BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 679 faunal assemblages have different patterns of distribution in southwestern México—one assemblage in all of the lowland regions, another only on the coastal lowlands and lower mountain slopes, and a third in the Balsas- Tepalcatepec Basin and to a greater or lesser extent on the coastal lowlands. Each assemblage is discussed separately. Widespread Pacific Lowland Assemblage—Included in this assemblage are 85 species that are more or less widely distributed on the Pacific lowlands of México, including the Balsas-Tepalcatepec Basin. Many of the species occur in other lowland regions as well. For example, the distributions of five of the species extend to South America, 12 extend northward into the Mexican state of Sonora, and 18 include the lowlands in eastern México. On the other hand, some of the species range no farther northward than Colima and some no farther southward than Oaxaca, but all are found both to the north and to. the south of Michoacan. The species included in this assemblage are: Bufo marinus Sceloporus melanorhinus Eleutherodactylus rugulosus Leptodactylus melanonotus Diaglena reticulata Phyllomedusa dacnicolor Sceloporus pyrocephalus Mabuya brachypoda Ameiva undulata Heloderma horridum Smilisca baudini Leptotyphlops phenops Hyla smithi Boa constrictor Geoemyda pulcherrima Loxocemus bicolor Geoemyda rubida Conophis vittatus Coleonyx elegans Drymarchon corais Phyllodactylus lanei Drymobius margaritiferus Anolis nebulosus Imantodes gemmistratus Basiliscus vittatus Leptophis diplotropis Iguana iguana Masticophis striolatus Ctenosaura pectinata Salvadora mexicana Phrynosoma asio Trimorphodon biscutatus Sceloporus horridus Agkistrodon bilineatus Crotalus basilicus Pacific Coastal Assemblage——Seventeen species occur on the coastal low-- lands and adjacent foothills, but do not occur in the Balsas-Tepalcatepec Basin. Most of these species are widely distributed throughout the coastal lowlands; the distributions of seven of the species extend southward into South America, four extend northward into the Mexican state of Sonora, and nine include the lowlands in eastern México. The species included in this assemblage are: Bufo marmoreus Urosaurus bicarinatus Phrynohyas inflata Scincella assata Dryadophis melanolomus Clelia clelia Geagras redimitus Geatractus tecpanensis Hyla microcephala Leptodeira septentrionalis Gastrophryne usta Manolepis putnami Pseudemys scripta Oxybelis aeneus Crocodylus acutus Sibon nebulatus Natrix valida Two other species (Rhinophrynus dorsalis and Coniophanes fissidens) are widespread in the lowlands of northern Central America and México, but reach the northern limits of their distributions on the coast of Michoacan. Four species (Phyllodactylus homolepidurus, Anolis schmidti, Eumeces coli- mensis, and Eumeces parvulus) have restricted distributions on the Pacific low- lands from Nayarit to Michoacan, where they reach the known southern limits of their ranges. 680 UNIVERSITY OF KANSAS PuBts., Mus. Nat. Hist. Twenty-four species occur on the coastal lowlands of Guerrero and south- ward, but are unknown from Michoacan; these are: Dermophis multiplicatus Cnemidophorus sacki Bufo gemmifer Ficimia publia Syrrhophus pipilans Leptodeira annulata Hyla staufferi Leptodeira nigrofasciata Bipes tridactylum Leptophis mexicanus Hemidactylus frenatus Masticophis mentovarius Phyllodactylus delcampi Salvadora lemniscata Anolis gadowi Stenorrhina freminvilli Anolis subocularis Tantilla martindelcampoi Anolis taylori Xenodon. rhabdocephalus Sceloporus stejnegeri Micrurus nigrocinctus Cnemidophorus guttatus Micrurus nuchalis On the coastal lowlands of Colima and northward there are six species, the distributions of which approach Michoacan, but do not cross the Rio Coa- huayana; these are Bufo mazatlanensis, Syrrhophus modestus, Hypopachus oxyrrhinus, Leptotyphlops humilis, Dipsas gaigeae, and Micrurus diastema. Balsas-Tepalcatepec Basin Assemblage-—The large, arid, lowland basin lying inland to the Sierra de Coalcoman and the Sierra Madre del Sur has a distinctive herpetofaunal assemblage, including 13 endemic species. Eight of these endemics are found only in the Tepalcatepec Valley (indicated by an asterisk in the following list): Bufo perplexus Sceloporus gadowae Hypopachus caprimimus Urosaurus gadowi * Bipes canaliculatus Eumeces altamirani * Phyllodactylus duellmani * Cnemidophorus calidipes * Phyllodactylus paucituberculatus * Leptotyphlops bressoni * Enyaliosaurus clarki * Leptotyphlops gadowi * Pseudoficimia pulcherrima Eleven species are widely distributed in the Balsas-Tepalcatepec Basin and extend through the Ahuijullo depression to the lowlands of Colima and thence northward; five of these species range northward into Sonora; the species in this group are: Tomodactylus _nitidus Leptodeira splendida Pternohyla fodiens Pseudoficimia frontalis Coniophanes lateritius Sonora michoacanensis Hypsiglena torquata Trimorphodon latifascia Leptodeira latifasciata Micrurus distans Micrurus laticollaris Four species have distributions similar to those listed above, but also occur on the coast of Michoacan; these are Cnemidophorus communis, Cnemidophorus lineatissimus, Leptodeira maculata, and Tantilla calamaria. Five species are widely distributed on the coastal lowlands from Guerrero southward and occur also in the Balsas-Tepalcatepec Basin; these are Bufo coccifer, Leptodactylus labialis, Crotalus durissus, Sceloporus siniferus, and Cnemidophorus deppei. The last two species also occur on the southeastern part of coastal Michoacan. Four species are known from the upper Balsas Basin in Morelos and Guerrero, but are unknown in the lower basin in Michoacan; these are Hypopachus alboventer, Sceloporus ochoterenai, Cnemidophorus sacki, and Leptotyphlops maximus. BroGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 681 Sierra Madrean Herpetofauna The mountains of southwestern México can be divided into four separate highland masses—the Sierra Madre Occidental that parallels the Pacific coast from Sonora to Jalisco, the Sierra de Coalcoman in southern Michoacan, the Sierra Madre del Sur in southern Guerrero and Oaxaca, and the Cordillera Volcanica that extends from Michoacan to Veracruz. For purposes of faunal analysis the Cordillera Volcanica is divided into three regions—the Sierra de los Tarascos in central Michoacan, the Sierra Temazcaltepec in eastern Michoa- can and the western part of the state of México, and the Sierra Ajusco in southern México and northern Morelos. Inhabiting the mountains of south- western México is a rather distinctive assemblage of species that I consider to be a part of the Sierra Madrean Herpetofauna, Generally these species are distributed at elevations of more than 1000 meters; many do not descend to that elevation. Mountainous regions often are characterized by a high amount of ende- mism in their faunas. The mountains in southwestern México are no exception. Fifty-one of the 98 species of amphibians and reptiles found there are not known to occur in other highland regions in Mexico. A finer analysis reveals 19 endemic species in the Sierra Madre del Sur, 12 in the Cordillera Vol- canica, three in the Sierra de Coalcoman, and one in the southern part of the Sierra Madre Occidental. Widespread Montane Assemblage.—Because of the many discontinuities in distributions, the description of distributional patterns in the highlands is more difficult than for the lowland fauna. The distribution of each of the 98 species known from the mountains of southwestern México is shown in Table 8. Of these, only six species occur in all of the highland regions—Bufo occidentalis, Eleutherodactylus augusti, Hyla arenicolor, Hyla bistincta, Rana pipiens, and Rana pustulosa. Twenty-one species are widespread in the mountainous regions but are ab- sent from one or both of the isolated mountain ranges, Sierra de Coalcoman and Sierra Madre del Sur. In the following list a single asterisk denotes species occurring in the Sierra de Coalcoman and two asterisks denote species occur- ring in the Sierra Madre del Sur; species not so marked are absent from both of the aforementioned mountain ranges, but occur widely through the Sierra Madre Occidental and the Cordillera Volcanica. Pseudoeurycea belli ** Tantilla bocourti ** Microbatrachylus hobartsmithi Storeria storerioides * Phrynosoma orbiculaire Sceloporus scalaris Sceloporus aeneus Sceloporus torquatus Sceloporus grammicus ** Eumeces brevirostris Sceloporus heterolepis * Eumeces dugesi Gerrhonotus imbricatus * Eumeces indubitus Conopsis biserialis Thamnophis dorsalis * Rhadinaea laureata Crotalus intermedius ** Rhadinaea taeniata Crotalus molossus Crotalus triseriatus Sierra Madre Occidental Assemblage —The Sierra Madre Occidental extends roughly along a north-south axis for a distance of more than 1300 kilometers. The distributions of many species, such as Bufo occidentalis, Eleutherodac- tylus augusti, Hyla arenicolor, Gerrhonotus imbricatus, Storeria storerioides, Thamnophis dorsalis, and Crotalus molossus, include nearly the entire moun- tain mass. Other species, such as Hyla bistincta, Sceloporus heterolepis, and 682 UNIVERSITY OF KANSAS PuBLs., Mus. Nat. Hisr, TABLE 8.—DISTRIBUTION OF AMPHIBIANS AND REPTILES IN THE MOUNTAINS OF SOUTHWESTERN MExIco. oO Ss 4| 3 SMS g a Sw a| & 5 < Pi es DME eo i I SPECIES e = iS Sore. 5 = '}) as (5) OS oO q CO) | Benge, Jans Boall Fe il po | oe Adalat tht wD Cordillera Volednica| ® 2) Ambystoma amblycephalum........ —_ x — -- a — Ambystoma ordinarium........... x xX xX = — a Rhyacosiredon altamirani.......... = = = xX — — Rhyacosiredon rivularis............ — — xX — — Rhyacosiredon zempoalensis........ — — — xX _ — Chiropterotriton chiropterus........ — = X — — Pseudoeurycea altamontana........ —_ — — x — -— Pseudoeunyceqnbellt... 3)s0.eyceye oe sac x x xX xX — xX Pseudoeurycea cephalica........... — — — xX — — Pseudoeurycea leprosa............. — — — x — Pseudoeurycea robertsi............ — — xX —_— — — Bugo oceidentas. osc 2c cne en ese nes x xX xX xX x x Eleutherodactylus augustt.......... xX xX x x xX xX Eleutherodactylus calcitrans........ — —_ — — — xX Eleutherodactylus saltator.......... — — — — — xX Microbatrachylus hobartsmithi...... xX xX xX — — — Microbatrachylus pygmaeus........ — — — — x xX Tomodactylus albolabris........... — —_ — —_ x Tomodactylus angustidigitorum..... — D4 — == = = Tomodactylus dilatus.............. — — — — xX POMOGACHLUS FUSCUS <6 6050 6,00 © 0 60 — — xX = — — Tomodactylus grandis............. — —- -~ xX — — Tomodactylus rufescens............ — — x — Hyla arboricola... . cece ccc cence — — — — — xX Elalaranenicolor. =.\.s eek aoe oe. ee xX xX xX x xX xX EDU DUSTCLGR. mk aia ss dene 5 hoe 8k xX xX xX xX x xX Hyla erythromma............0.00. — a — — — x Fla TOfrentet. .. cacc sage ve vance — xX x Dx — == Hyla melanomma.............024. — -- a —- _ xX FLO STOROGOUNG, s 256 se i eee ee — xX — — x — Ptychohyla leonhardschultzet....... —_— —- — — — xX Cochranella viridissima............ — — — —_ — xX TEONG BIDIENS 5.2 5 isis aby oo oe ema x xX xX xX x x ECOG DUSVUNOS Cio lsu esis onnanee nos xX xX -— — xX xX Rana sierramadrensis............. — — _- —_ xX Kinosternon integrum..........0.. -- xX —- — xX xX AMOI QUANT ics oust eAacteres amok — — — —_ x x AMOS WOGAStens «oc nlc. ay, Gussie cron —_ = — or x Anolis microlepidotus............. — — = = = x Anolis omiltemanus............... — -- — -= _ xX Phrynosoma orbiculaire...........| X — = D.€ = Phrynosoma taurus. .............. — — — — — D.€ Sceloporus aeneus...........0.00. xX xX x x = = Seeloporus Ospere 25. dacacn wes wc — x = 773 x a Sceloporus bullert................. x — — — x — Sceloporus formosus.............. ae ~— -= —_ —_— x Sceloporus grammicus............4. x xX xX x = x Sceloporus heterolepis.............] X xX = — x Tan Sceloporus mucronatus. . — — = — — x BIOGEOGRAPHIC ACCOUNT OF HERPETOFAUNA 683 TABLE 8.—DISTRIBUTION OF AMPHIBIANS AND REPTILES IN THE MOUNTAINS OF SOUTHWESTERN Mrxico.—Concluded SPECIES Sceloports scalars aoe ole aes CELONOTUSILOTGUCLLS,.. Nevertire riers Sceloporus YOrTOWts 12.52.01.) ae Eumeces brevirostris yes «se eas EUMeCESICOPED. es send ciseiehal- 2 oer EI WIMECES LAUD ESt ceo ie craters eel Eumeces indubitus. ..<..:....+5--- Eumeces ochoterenai..........---- Albroniaidep pets serene seer Gerrhonotusigadowt..&. se. sae Gerrhonotus imbricatus...........-. Gerrhonotus liocephalus............ Chersodromus annulatus.........-. Gonopsisibisenvaliss rcictas a tere ae Cononsisinasustere cc ae BMUAUS UNTCOLOT Me aa Geophisibicolon. mies se een eae GEODNtSANCOMPUUS ae eee Geophis\macuiijerus..-0 3. 26-55. Geophisnigrocinetusey. we aase oe Geophis omiltemanus.............- Geophis peterste «oe ae Geonhisitarascaene- Seyret ie Bampropelitsidoatae Patuophisiineaticolliss... eee a Rhadinaeaaemiulaar we. <2 was eee TRGAINOCON OURO Gas eae ee Rhadinaea omiltemana............ Rhadinaecatiaenivata® Se ee ee Mantillaibocourte cts 4 oa ce Mantua conOnadovi.. verre ic) selene ARGH IING) CA) NG B tec en ane Oe Oe MM OVUCOACONUCG ree ccvorevs oe ciao e e.e Ones NOULCO ANCHO ea sce clc oac ene Tropidodipsas guerreroensis........ Tropidodipsas occidentalis......... SLONETTOSLOTENIOLAES: @ ie eee ae Thamnophis chrysocephalus........ Thamnopnrsidorsais ya 57.0001. Thamnophis scalars. st) -)eke.- a ee MierirusOrOwn erie ier iets Bothrops DOrCOUTt..< cc <. ee _ . 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